The Descent of Man

Chapter XIII: Secondary Sexual Characters of Birds

Sexual differences—Law of battle—Special weapons—Vocal organs—Instrumental music—Love-antics and dances—Decorations, permanent and seasonal—Double and single annual moults—Display of ornaments by the males.

Secondary sexual characters are more diversified and conspicuous in birds, though not perhaps entailing more important changes of structure, than in any other class of animals. I shall, therefore, treat the subject at considerable length. Male birds sometimes, though rarely, possess special weapons for fighting with each other. They charm the female by vocal or instrumental music of the most varied kinds. They are ornamented by all sorts of combs, wattles, protuberances, horns, air-distended sacks, top-knots, naked shafts, plumes and lengthened feathers gracefully springing from all parts of the body. The beak and naked skin about the head, and the feathers, are often gorgeously coloured. The males sometimes pay their court by dancing, or by fantastic antics performed either on the ground or in the air. In one instance, at least, the male emits a musky odour, which we may suppose serves to charm or excite the female; for that excellent observer, Mr. Ramsay (1. ‘Ibis,’ vol. iii. (new series), 1867, p. 414.), says of the Australian musk-duck (Biziura lobata) that “the smell which the male emits during the summer months is confined to that sex, and in some individuals is retained throughout the year; I have never, even in the breeding-season, shot a female which had any smell of musk.” So powerful is this odour during the pairing-season, that it can be detected long before the bird can be seen. (2. Gould, ‘Handbook of the Birds of Australia,’ 1865, vol. ii. p. 383.) On the whole, birds appear to be the most aesthetic of all animals, excepting of course man, and they have nearly the same taste for the beautiful as we have. This is shewn by our enjoyment of the singing of birds, and by our women, both civilised and savage, decking their heads with borrowed plumes, and using gems which are hardly more brilliantly coloured than the naked skin and wattles of certain birds. In man, however, when cultivated, the sense of beauty is manifestly a far more complex feeling, and is associated with various intellectual ideas.

Before treating of the sexual characters with which we are here more particularly concerned, I may just allude to certain differences between the sexes which apparently depend on differences in their habits of life; for such cases, though common in the lower, are rare in the higher classes. Two humming-birds belonging to the genus Eustephanus, which inhabit the island of Juan Fernandez, were long thought to be specifically distinct, but are now known, as Mr. Gould informs me, to be the male and female of the same species, and they differ slightly in the form of the beak. In another genus of humming-birds (Grypus), the beak of the male is serrated along the margin and hooked at the extremity, thus differing much from that of the female. In the Neomorpha of New Zealand, there is, as we have seen, a still wider difference in the form of the beak in relation to the manner of feeding of the two sexes. Something of the same kind has been observed with the goldfinch (Carduelis elegans), for I am assured by Mr. J. Jenner Weir that the bird-catchers can distinguish the males by their slightly longer beaks. The flocks of males are often found feeding on the seeds of the teazle (Dipsacus), which they can reach with their elongated beaks, whilst the females more commonly feed on the seeds of the betony or Scrophularia. With a slight difference of this kind as a foundation, we can see how the beaks of the two sexes might be made to differ greatly through natural selection. In some of the above cases, however, it is possible that the beaks of the males may have been first modified in relation to their contests with other males; and that this afterwards led to slightly changed habits of life.

LAW OF BATTLE.

Almost all male birds are extremely pugnacious, using their beaks, wings, and legs for fighting together. We see this every spring with our robins and sparrows. The smallest of all birds, namely the humming-bird, is one of the most quarrelsome. Mr. Gosse (3. Quoted by Mr. Gould, ‘Introduction to the Trochilidae,’ 1861, page 29.) describes a battle in which a pair seized hold of each other’s beaks, and whirled round and round, till they almost fell to the ground; and M. Montes de Oca, in speaking or another genus of humming-bird, says that two males rarely meet without a fierce aerial encounter: when kept in cages “their fighting has mostly ended in the splitting of the tongue of one of the two, which then surely dies from being unable to feed.” (4. Gould, ibid. p. 52.) With waders, the males of the common water-hen (Gallinula chloropus) “when pairing, fight violently for the females: they stand nearly upright in the water and strike with their feet.” Two were seen to be thus engaged for half an hour, until one got hold of the head of the other, which would have been killed had not the observer interfered; the female all the time looking on as a quiet spectator. (5. W. Thompson, ‘Natural History of Ireland: Birds,’ vol. ii. 1850, p. 327.) Mr. Blyth informs me that the males of an allied bird (Gallicrex cristatus) are a third larger than the females, and are so pugnacious during the breeding-season that they are kept by the natives of Eastern Bengal for the sake of fighting. Various other birds are kept in India for the same purpose, for instance, the bulbuls (Pycnonotus hoemorrhous) which “fight with great spirit.” (6. Jerdon, ‘Birds of India,’ 1863, vol. ii. p. 96.)

[Fig. 37. The Ruff or Machetes pugnax (from Brehm’s ‘Thierleben’).]

The polygamous ruff (Machetes pugnax, Fig. 37) is notorious for his extreme pugnacity; and in the spring, the males, which are considerably larger than the females, congregate day after day at a particular spot, where the females propose to lay their eggs. The fowlers discover these spots by the turf being trampled somewhat bare. Here they fight very much like game-cocks, seizing each other with their beaks and striking with their wings. The great ruff of feathers round the neck is then erected, and according to Col. Montagu “sweeps the ground as a shield to defend the more tender parts”; and this is the only instance known to me in the case of birds of any structure serving as a shield. The ruff of feathers, however, from its varied and rich colours probably serves in chief part as an ornament. Like most pugnacious birds, they seem always ready to fight, and when closely confined, often kill each other; but Montagu observed that their pugnacity becomes greater during the spring, when the long feathers on their necks are fully developed; and at this period the least movement by any one bird provokes a general battle. (7. Macgillivray, ‘History of British Birds,’ vol. iv. 1852, pp. 177-181.) Of the pugnacity of web-footed birds, two instances will suffice: in Guiana “bloody fights occur during the breeding-season between the males of the wild musk-duck (Cairina moschata); and where these fights have occurred the river is covered for some distance with feathers.” (8. Sir R. Schomburgk, in ‘Journal of Royal Geographic Society,’ vol. xiii. 1843, p. 31.) Birds which seem ill-adapted for fighting engage in fierce conflicts; thus the stronger males of the pelican drive away the weaker ones, snapping with their huge beaks and giving heavy blows with their wings. Male snipe fight together, “tugging and pushing each other with their bills in the most curious manner imaginable.” Some few birds are believed never to fight; this is the case, according to Audubon, with one of the woodpeckers of the United States (Picu sauratus), although “the hens are followed by even half a dozen of their gay suitors.” (9. ‘Ornithological Biography,’ vol. i. p. 191. For pelicans and snipes, see vol. iii. pp. 138, 477.)

The males of many birds are larger than the females, and this no doubt is the result of the advantage gained by the larger and stronger males over their rivals during many generations. The difference in size between the two sexes is carried to an extreme point in several Australian species; thus the male musk-duck (Biziura), and the male Cincloramphus cruralis (allied to our pipits) are by measurement actually twice as large as their respective females. (10. Gould, ‘Handbook of Birds of Australia,’ vol. i. p. 395; vol. ii. p. 383.) With many other birds the females are larger than the males; and, as formerly remarked, the explanation often given, namely, that the females have most of the work in feeding their young, will not suffice. In some few cases, as we shall hereafter see, the females apparently have acquired their greater size and strength for the sake of conquering other females and obtaining possession of the males.

The males of many gallinaceous birds, especially of the polygamous kinds, are furnished with special weapons for fighting with their rivals, namely spurs, which can be used with fearful effect. It has been recorded by a trustworthy writer (11. Mr. Hewitt, in the ‘Poultry Book’ by Tegetmeier, 1866, p. 137.) that in Derbyshire a kite struck at a game-hen accompanied by her chickens, when the cock rushed to the rescue, and drove his spur right through the eye and skull of the aggressor. The spur was with difficulty drawn from the skull, and as the kite, though dead, retained his grasp, the two birds were firmly locked together; but the cock when disentangled was very little injured. The invincible courage of the game-cock is notorious: a gentleman who long ago witnessed the brutal scene, told me that a bird had both its legs broken by some accident in the cockpit, and the owner laid a wager that if the legs could be spliced so that the bird could stand upright, he would continue fighting. This was effected on the spot, and the bird fought with undaunted courage until he received his death-stroke. In Ceylon a closely allied, wild species, the Gallus Stanleyi, is known to fight desperately “in defence of his seraglio,” so that one of the combatants is frequently found dead. (12. Layard, ‘Annals and Magazine of Natural History,’ vol. xiv. 1854, p. 63.) An Indian partridge (Ortygornis gularis), the male of which is furnished with strong and sharp spurs, is so quarrelsome “that the scars of former fights disfigure the breast of almost every bird you kill.” (13. Jerdon, ‘Birds of India,’ vol. iii. p. 574.)

The males of almost all gallinaceous birds, even those which are not furnished with spurs, engage during the breeding-season in fierce conflicts. The Capercailzie and Black-cock (Tetrao urogallus and T. tetrix), which are both polygamists, have regular appointed places, where during many weeks they congregate in numbers to fight together and to display their charms before the females. Dr. W. Kovalevsky informs me that in Russia he has seen the snow all bloody on the arenas where the capercailzie have fought; and the black-cocks “make the feathers fly in every direction,” when several “engage in a battle royal.” The elder Brehm gives a curious account of the Balz, as the love-dances and love-songs of the Black-cock are called in Germany. The bird utters almost continuously the strangest noises: “he holds his tail up and spreads it out like a fan, he lifts up his head and neck with all the feathers erect, and stretches his wings from the body. Then he takes a few jumps in different directions, sometimes in a circle, and presses the under part of his beak so hard against the ground that the chin feathers are rubbed off. During these movements he beats his wings and turns round and round. The more ardent he grows the more lively he becomes, until at last the bird appears like a frantic creature.” At such times the black-cocks are so absorbed that they become almost blind and deaf, but less so than the capercailzie: hence bird after bird may be shot on the same spot, or even caught by the hand. After performing these antics the males begin to fight: and the same black-cock, in order to prove his strength over several antagonists, will visit in the course of one morning several Balz-places, which remain the same during successive years. (14. Brehm, ‘Thierleben,’ 1867, B. iv. s. 351. Some of the foregoing statements are taken from L. Lloyd, ‘The Game Birds of Sweden,’ etc., 1867, p. 79.)

The peacock with his long train appears more like a dandy than a warrior, but he sometimes engages in fierce contests: the Rev. W. Darwin Fox informs me that at some little distance from Chester two peacocks became so excited whilst fighting, that they flew over the whole city, still engaged, until they alighted on the top of St. John’s tower.

The spur, in those gallinaceous birds which are thus provided, is generally single; but Polyplectron (Fig. 51) has two or more on each leg; and one of the Blood-pheasants (Ithaginis cruentus) has been seen with five spurs. The spurs are generally confined to the male, being represented by mere knobs or rudiments in the female; but the females of the Java peacock (Pavo muticus) and, as I am informed by Mr. Blyth, of the small fire-backed pheasant (Euplocamus erythrophthalmus) possess spurs. In Galloperdix it is usual for the males to have two spurs, and for the females to have only one on each leg. (15. Jerdon, ‘Birds of India’: on Ithaginis, vol. iii. p. 523; on Galloperdix, p. 541.) Hence spurs may be considered as a masculine structure, which has been occasionally more or less transferred to the females. Like most other secondary sexual characters, the spurs are highly variable, both in number and development, in the same species.

[Fig.38. Palamedea cornuta (from Brehm), shewing the double wing-spurs, and the filament on the head.]

Various birds have spurs on their wings. But the Egyptian goose (Chenalopex aegyptiacus) has only “bare obtuse knobs,” and these probably shew us the first steps by which true spurs have been developed in other species. In the spur-winged goose, Plectropterus gambensis, the males have much larger spurs than the females; and they use them, as I am informed by Mr. Bartlett, in fighting together, so that, in this case, the wing-spurs serve as sexual weapons; but according to Livingstone, they are chiefly used in the defence of the young. The Palamedea (Fig. 38) is armed with a pair of spurs on each wing; and these are such formidable weapons that a single blow has been known to drive a dog howling away. But it does not appear that the spurs in this case, or in that of some of the spur-winged rails, are larger in the male than in the female. (16. For the Egyptian goose, see Macgillivray, ‘British Birds,’ vol. iv. p. 639. For Plectropterus, Livingstone’s ‘Travels,’ p. 254. For Palamedea, Brehm’s ‘Thierleben,’ B. iv. s. 740. See also on this bird Azara, ‘Voyages dans l’Amerique merid.’ tom. iv. 1809, pp. 179, 253.) In certain plovers, however, the wing-spurs must be considered as a sexual character. Thus in the male of our common peewit (Vanellus cristatus) the tubercle on the shoulder of the wing becomes more prominent during the breeding-season, and the males fight together. In some species of Lobivanellus a similar tubercle becomes developed during the breeding-season “into a short horny spur.” In the Australian L. lobatus both sexes have spurs, but these are much larger in the males than in the females. In an allied bird, the Hoplopterus armatus, the spurs do not increase in size during the breeding-season; but these birds have been seen in Egypt to fight together, in the same manner as our peewits, by turning suddenly in the air and striking sideways at each other, sometimes with fatal results. Thus also they drive away other enemies. (17. See, on our peewit, Mr. R. Carr in ‘Land and Water,’ Aug. 8th, 1868, p. 46. In regard to Lobivanellus, see Jerdon’s ‘Birds of India,’ vol. iii. p. 647, and Gould’s ‘Handbook of Birds of Australia,’ vol. ii. p. 220. For the Hoplopterus, see Mr. Allen in the ‘Ibis,’ vol. v. 1863, p. 156.)

The season of love is that of battle; but the males of some birds, as of the game-fowl and ruff, and even the young males of the wild turkey and grouse (18. Audubon, ‘Ornithological Biography,’ vol. ii. p. 492; vol. i. pp. 4-13.), are ready to fight whenever they meet. The presence of the female is the teterrima belli causa. The Bengali baboos make the pretty little males of the amadavat (Estrelda amandava) fight together by placing three small cages in a row, with a female in the middle; after a little time the two males are turned loose, and immediately a desperate battle ensues. (19. Mr. Blyth, ‘Land and Water,’ 1867, p. 212.) When many males congregate at the same appointed spot and fight together, as in the case of grouse and various other birds, they are generally attended by the females (20. Richardson on Tetrao umbellus, ‘Fauna Bor. Amer.: Birds,’ 1831, p. 343. L. Lloyd, ‘Game Birds of Sweden,’ 1867, pp. 22, 79, on the capercailzie and black-cock. Brehm, however, asserts (‘Thierleben,’ B. iv. s. 352) that in Germany the grey-hens do not generally attend the Balzen of the black-cocks, but this is an exception to the common rule; possibly the hens may lie hidden in the surrounding bushes, as is known to be the case with the gray-hens in Scandinavia, and with other species in N. America.), which afterwards pair with the victorious combatants. But in some cases the pairing precedes instead of succeeding the combat: thus according to Audubon (21. ‘Ornithological Biography,’ vol. ii. p. 275.), several males of the Virginian goat-sucker (Caprimulgus virgianus) “court, in a highly entertaining manner the female, and no sooner has she made her choice, than her approved gives chase to all intruders, and drives them beyond his dominions.” Generally the males try to drive away or kill their rivals before they pair. It does not, however, appear that the females invariably prefer the victorious males. I have indeed been assured by Dr. W. Kovalevsky that the female capercailzie sometimes steals away with a young male who has not dared to enter the arena with the older cocks, in the same manner as occasionally happens with the does of the red-deer in Scotland. When two males contend in presence of a single female, the victor, no doubt, commonly gains his desire; but some of these battles are caused by wandering males trying to distract the peace of an already mated pair. (22. Brehm, ‘Thierleben,’ etc., B. iv. 1867, p. 990. Audubon, ‘Ornithological Biography,’ vol. ii. p. 492.)

Even with the most pugnacious species it is probable that the pairing does not depend exclusively on the mere strength and courage of the male; for such males are generally decorated with various ornaments, which often become more brilliant during the breeding-season, and which are sedulously displayed before the females. The males also endeavour to charm or excite their mates by love-notes, songs, and antics; and the courtship is, in many instances, a prolonged affair. Hence it is not probable that the females are indifferent to the charms of the opposite sex, or that they are invariably compelled to yield to the victorious males. It is more probable that the females are excited, either before or after the conflict, by certain males, and thus unconsciously prefer them. In the case of Tetrao umbellus, a good observer (23. ‘Land and Water,’ July 25, 1868, p. 14.) goes so far as to believe that the battles of the male “are all a sham, performed to show themselves to the greatest advantage before the admiring females who assemble around; for I have never been able to find a maimed hero, and seldom more than a broken feather.” I shall have to recur to this subject, but I may here add that with the Tetrao cupido of the United States, about a score of males assemble at a particular spot, and, strutting about, make the whole air resound with their extraordinary noises. At the first answer from a female the males begin to fight furiously, and the weaker give way; but then, according to Audubon, both the victors and vanquished search for the female, so that the females must either then exert a choice, or the battle must be renewed. So, again, with one of the field-starlings of the United States (Sturnella ludoviciana) the males engage in fierce conflicts, “but at the sight of a female they all fly after her as if mad.” (24. Audubon’s ‘Ornithological Biography;’ on Tetrao cupido, vol. ii. p. 492; on the Sturnus, vol. ii. p. 219.)

VOCAL AND INSTRUMENTAL MUSIC.

With birds the voice serves to express various emotions, such as distress, fear, anger, triumph, or mere happiness. It is apparently sometimes used to excite terror, as in the case of the hissing noise made by some nestling-birds. Audubon (25. ‘Ornithological Biography,’ vol. v. p. 601.), relates that a night-heron (Ardea nycticorax, Linn.), which he kept tame, used to hide itself when a cat approached, and then “suddenly start up uttering one of the most frightful cries, apparently enjoying the cat’s alarm and flight.” The common domestic cock clucks to the hen, and the hen to her chickens, when a dainty morsel is found. The hen, when she has laid an egg, “repeats the same note very often, and concludes with the sixth above, which she holds for a longer time” (26. The Hon. Daines Barrington, ‘Philosophical Transactions,’ 1773, p. 252.); and thus she expresses her joy. Some social birds apparently call to each other for aid; and as they flit from tree to tree, the flock is kept together by chirp answering chirp. During the nocturnal migrations of geese and other water-fowl, sonorous clangs from the van may be heard in the darkness overhead, answered by clangs in the rear. Certain cries serve as danger signals, which, as the sportsman knows to his cost, are understood by the same species and by others. The domestic cock crows, and the humming-bird chirps, in triumph over a defeated rival. The true song, however, of most birds and various strange cries are chiefly uttered during the breeding-season, and serve as a charm, or merely as a call-note, to the other sex.

Naturalists are much divided with respect to the object of the singing of birds. Few more careful observers ever lived than Montagu, and he maintained that the “males of song-birds and of many others do not in general search for the female, but, on the contrary, their business in the spring is to perch on some conspicuous spot, breathing out their full and amorous notes, which, by instinct, the female knows, and repairs to the spot to choose her mate.” (27. ‘Ornithological Dictionary,’ 1833, p. 475.) Mr. Jenner Weir informs me that this is certainly the case with the nightingale. Bechstein, who kept birds during his whole life, asserts, “that the female canary always chooses the best singer, and that in a state of nature the female finch selects that male out of a hundred whose notes please her most. (28. ‘Naturgeschichte der Stubenvögel,’ 1840, s. 4. Mr. Harrison Weir likewise writes to me:—“I am informed that the best singing males generally get a mate first, when they are bred in the same room.”) There can be no doubt that birds closely attend to each other’s song. Mr. Weir has told me of the case of a bullfinch which had been taught to pipe a German waltz, and who was so good a performer that he cost ten guineas; when this bird was first introduced into a room where other birds were kept and he began to sing, all the others, consisting of about twenty linnets and canaries, ranged themselves on the nearest side of their cages, and listened with the greatest interest to the new performer. Many naturalists believe that the singing of birds is almost exclusively “the effect of rivalry and emulation,” and not for the sake of charming their mates. This was the opinion of Daines Barrington and White of Selborne, who both especially attended to this subject. (29. ‘Philosophical Transactions,’ 1773, p. 263. White’s ‘Natural History of Selborne,’ 1825, vol. i. p. 246.) Barrington, however, admits that “superiority in song gives to birds an amazing ascendancy over others, as is well known to bird-catchers.”

It is certain that there is an intense degree of rivalry between the males in their singing. Bird-fanciers match their birds to see which will sing longest; and I was told by Mr. Yarrell that a first-rate bird will sometimes sing till he drops down almost dead, or according to Bechstein (30. ‘Naturgesch. der Stubenvögel,’ 1840, s. 252.), quite dead from rupturing a vessel in the lungs. Whatever the cause may be, male birds, as I hear from Mr. Weir, often die suddenly during the season of song. That the habit of singing is sometimes quite independent of love is clear, for a sterile, hybrid canary-bird has been described (31. Mr. Bold, ‘Zoologist,’ 1843-44, p. 659.) as singing whilst viewing itself in a mirror, and then dashing at its own image; it likewise attacked with fury a female canary, when put into the same cage. The jealousy excited by the act of singing is constantly taken advantage of by bird-catchers; a male, in good song, is hidden and protected, whilst a stuffed bird, surrounded by limed twigs, is exposed to view. In this manner, as Mr. Weir informs me, a man has in the course of a single day caught fifty, and in one instance, seventy, male chaffinches. The power and inclination to sing differ so greatly with birds that although the price of an ordinary male chaffinch is only sixpence, Mr. Weir saw one bird for which the bird-catcher asked three pounds; the test of a really good singer being that it will continue to sing whilst the cage is swung round the owner’s head.

That male birds should sing from emulation as well as for charming the female, is not at all incompatible; and it might have been expected that these two habits would have concurred, like those of display and pugnacity. Some authors, however, argue that the song of the male cannot serve to charm the female, because the females of some few species, such as of the canary, robin, lark, and bullfinch, especially when in a state of widowhood, as Bechstein remarks, pour forth fairly melodious strains. In some of these cases the habit of singing may be in part attributed to the females having been highly fed and confined (32. D. Barrington, ‘Philosophical Transactions,’ 1773, p. 262. Bechstein, ‘Stubenvögel,’ 1840, s. 4.), for this disturbs all the functions connected with the reproduction of the species. Many instances have already been given of the partial transference of secondary masculine characters to the female, so that it is not at all surprising that the females of some species should possess the power of song. It has also been argued, that the song of the male cannot serve as a charm, because the males of certain species, for instance of the robin, sing during the autumn. (33. This is likewise the case with the water-ouzel; see Mr. Hepburn in the ‘Zoologist,’ 1845-46, p. 1068.) But nothing is more common than for animals to take pleasure in practising whatever instinct they follow at other times for some real good. How often do we see birds which fly easily, gliding and sailing through the air obviously for pleasure? The cat plays with the captured mouse, and the cormorant with the captured fish. The weaver-bird (Ploceus), when confined in a cage, amuses itself by neatly weaving blades of grass between the wires of its cage. Birds which habitually fight during the breeding-season are generally ready to fight at all times; and the males of the capercailzie sometimes hold their Balzen or leks at the usual place of assemblage during the autumn. (34. L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 25.) Hence it is not at all surprising that male birds should continue singing for their own amusement after the season for courtship is over.

As shewn in a previous chapter, singing is to a certain extent an art, and is much improved by practice. Birds can be taught various tunes, and even the unmelodious sparrow has learnt to sing like a linnet. They acquire the song of their foster parents (35. Barrington, ibid. p. 264, Bechstein, ibid. s. 5.), and sometimes that of their neighbours. (36. Dureau de la Malle gives a curious instance (‘Annales des Sc. Nat.’ 3rd series, Zoolog., tom. x. p. 118) of some wild blackbirds in his garden in Paris, which naturally learnt a republican air from a caged bird.) All the common songsters belong to the Order of Insessores, and their vocal organs are much more complex than those of most other birds; yet it is a singular fact that some of the Insessores, such as ravens, crows, and magpies, possess the proper apparatus (37. Bishop, in ‘Todd’s Cyclopaedia of Anatomy and Physiology,’ vol. iv. p. 1496.), though they never sing, and do not naturally modulate their voices to any great extent. Hunter asserts (38. As stated by Barrington in ‘Philosophical Transactions,’ 1773, p. 262.) that with the true songsters the muscles of the larynx are stronger in the males than in the females; but with this slight exception there is no difference in the vocal organs of the two sexes, although the males of most species sing so much better and more continuously than the females.

It is remarkable that only small birds properly sing. The Australian genus Menura, however, must be excepted; for the Menura Alberti, which is about the size of a half-grown turkey, not only mocks other birds, but “its own whistle is exceedingly beautiful and varied.” The males congregate and form “corroborying places,” where they sing, raising and spreading their tails like peacocks, and drooping their wings. (39. Gould, ‘Handbook to the Birds of Australia,’ vol. i. 1865, pp. 308-310. See also Mr. T.W. Wood in the ‘Student,’ April 1870, p. 125.) It is also remarkable that birds which sing well are rarely decorated with brilliant colours or other ornaments. Of our British birds, excepting the bullfinch and goldfinch, the best songsters are plain-coloured. The kingfisher, bee-eater, roller, hoopoe, woodpeckers, etc., utter harsh cries; and the brilliant birds of the tropics are hardly ever songsters. (40. See remarks to this effect in Gould’s ‘Introduction to the Trochilidae,’ 1861, p. 22.) Hence bright colours and the power of song seem to replace each other. We can perceive that if the plumage did not vary in brightness, or if bright colours were dangerous to the species, other means would be employed to charm the females; and melody of voice offers one such means.

[Fig. 39. Tetrao cupido: male. (T.W. Wood.)]

In some birds the vocal organs differ greatly in the two sexes. In the Tetrao cupido (Fig. 39) the male has two bare, orange-coloured sacks, one on each side of the neck; and these are largely inflated when the male, during the breeding-season, makes his curious hollow sound, audible at a great distance. Audubon proved that the sound was intimately connected with this apparatus (which reminds us of the air-sacks on each side of the mouth of certain male frogs), for he found that the sound was much diminished when one of the sacks of a tame bird was pricked, and when both were pricked it was altogether stopped. The female has “a somewhat similar, though smaller naked space of skin on the neck; but this is not capable of inflation.” (41. ‘The Sportsman and Naturalist in Canada,’ by Major W. Ross King, 1866, pp. 144-146. Mr. T.W. Wood gives in the ‘Student’ (April 1870, p. 116) an excellent account of the attitude and habits of this bird during its courtship. He states that the ear-tufts or neck-plumes are erected, so that they meet over the crown of the head. See his drawing, Fig. 39.) The male of another kind of grouse (Tetrao urophasianus), whilst courting the female, has his “bare yellow oesophagus inflated to a prodigious size, fully half as large as the body”; and he then utters various grating, deep, hollow tones. With his neck-feathers erect, his wings lowered, and buzzing on the ground, and his long pointed tail spread out like a fan, he displays a variety of grotesque attitudes. The oesophagus of the female is not in any way remarkable. (42. Richardson, ‘Fauna Bor. Americana: Birds,’ 1831, p. 359. Audubon, ibid. vol. iv. p. 507.)

[Fig. 40. The Umbrella-bird or Cephalopterus ornatus, male (from Brehm).]

It seems now well made out that the great throat pouch of the European male bustard (Otis tarda), and of at least four other species, does not, as was formerly supposed, serve to hold water, but is connected with the utterance during the breeding-season of a peculiar sound resembling “oak.” (43. The following papers have been lately written on this subject: Prof. A. Newton, in the ‘Ibis,’ 1862, p. 107; Dr. Cullen, ibid. 1865, p. 145; Mr. Flower, in ‘Proc. Zool. Soc.’ 1865, p. 747; and Dr. Murie, in ‘Proc. Zool. Soc.’ 1868, p. 471. In this latter paper an excellent figure is given of the male Australian Bustard in full display with the sack distended. It is a singular fact that the sack is not developed in all the males of the same species.) A crow-like bird inhabiting South America (see Cephalopterus ornatus, Fig. 40) is called the umbrella-bird, from its immense top knot, formed of bare white quills surmounted by dark-blue plumes, which it can elevate into a great dome no less than five inches in diameter, covering the whole head. This bird has on its neck a long, thin, cylindrical fleshy appendage, which is thickly clothed with scale-like blue feathers. It probably serves in part as an ornament, but likewise as a resounding apparatus; for Mr. Bates found that it is connected “with an unusual development of the trachea and vocal organs.” It is dilated when the bird utters its singularly deep, loud and long sustained fluty note. The head-crest and neck-appendage are rudimentary in the female. (44. Bates, ‘The Naturalist on the Amazons,’ 1863, vol. ii. p. 284; Wallace, in ‘Proceedings, Zoological Society,’ 1850, p. 206. A new species, with a still larger neck-appendage (C. penduliger), has lately been discovered, see ‘Ibis,’ vol. i. p. 457.)

The vocal organs of various web-footed and wading birds are extraordinarily complex, and differ to a certain extent in the two sexes. In some cases the trachea is convoluted, like a French horn, and is deeply embedded in the sternum. In the wild swan (Cygnus ferus) it is more deeply embedded in the adult male than in the adult female or young male. In the male Merganser the enlarged portion of the trachea is furnished with an additional pair of muscles. (45. Bishop, in Todd’s ‘Cyclopaedia of Anatomy and Physiology,’ vol. iv. p. 1499.) In one of the ducks, however, namely Anas punctata, the bony enlargement is only a little more developed in the male than in the female. (46. Prof. Newton, ‘Proc. Zoolog. Soc.’ 1871, p. 651.) But the meaning of these differences in the trachea of the two sexes of the Anatidae is not understood; for the male is not always the more vociferous; thus with the common duck, the male hisses, whilst the female utters a loud quack. (47. The spoonbill (Platalea) has its trachea convoluted into a figure of eight, and yet this bird (Jerdon, ‘Birds of India,’ vol. iii. p. 763) is mute; but Mr. Blyth informs me that the convolutions are not constantly present, so that perhaps they are now tending towards abortion.) In both sexes of one of the cranes (Grus virgo) the trachea penetrates the sternum, but presents “certain sexual modifications.” In the male of the black stork there is also a well-marked sexual difference in the length and curvature of the bronchi. (48. ‘Elements of Comparative Anatomy,’ by R. Wagner, Eng. translat. 1845, p. 111. With respect to the swan, as given above, Yarrell’s ‘History of British Birds,’ 2nd edition, 1845, vol. iii. p. 193.) Highly important structures have, therefore, in these cases been modified according to sex.

It is often difficult to conjecture whether the many strange cries and notes uttered by male birds during the breeding-season serve as a charm or merely as a call to the female. The soft cooing of the turtle-dove and of many pigeons, it may be presumed, pleases the female. When the female of the wild turkey utters her call in the morning, the male answers by a note which differs from the gobbling noise made, when with erected feathers, rustling wings and distended wattles, he puffs and struts before her. (49. C.L. Bonaparte, quoted in the ‘Naturalist Library: Birds,’ vol. xiv. p. 126.) The spel of the black-cock certainly serves as a call to the female, for it has been known to bring four or five females from a distance to a male under confinement; but as the black-cock continues his spel for hours during successive days, and in the case of the capercailzie “with an agony of passion,” we are led to suppose that the females which are present are thus charmed. (50. L. Lloyd, ‘The Game Birds of Sweden,’ etc., 1867, pp. 22, 81.) The voice of the common rook is known to alter during the breeding-season, and is therefore in some way sexual. (51. Jenner, ‘Philosophical Transactions,’ 1824, p. 20.) But what shall we say about the harsh screams of, for instance, some kinds of macaws; have these birds as bad taste for musical sounds as they apparently have for colour, judging by the inharmonious contrast of their bright yellow and blue plumage? It is indeed possible that without any advantage being thus gained, the loud voices of many male birds may be the result of the inherited effects of the continued use of their vocal organs when excited by the strong passions of love, jealousy and rage; but to this point we shall recur when we treat of quadrupeds.

We have as yet spoken only of the voice, but the males of various birds practise, during their courtship, what may be called instrumental music. Peacocks and Birds of Paradise rattle their quills together. Turkey-cocks scrape their wings against the ground, and some kinds of grouse thus produce a buzzing sound. Another North American grouse, the Tetrao umbellus, when with his tail erect, his ruffs displayed, “he shows off his finery to the females, who lie hid in the neighbourhood,” drums by rapidly striking his wings together above his back, according to Mr. R. Haymond, and not, as Audubon thought, by striking them against his sides. The sound thus produced is compared by some to distant thunder, and by others to the quick roll of a drum. The female never drums, “but flies directly to the place where the male is thus engaged.” The male of the Kalij-pheasant, in the Himalayas, often makes a singular drumming noise with his wings, not unlike the sound produced by shaking a stiff piece of cloth.” On the west coast of Africa the little black-weavers (Ploceus?) congregate in a small party on the bushes round a small open space, and sing and glide through the air with quivering wings, “which make a rapid whirring sound like a child’s rattle.” One bird after another thus performs for hours together, but only during the courting-season. At this season, and at no other time, the males of certain night-jars (Caprimulgus) make a strange booming noise with their wings. The various species of woodpeckers strike a sonorous branch with their beaks, with so rapid a vibratory movement that “the head appears to be in two places at once.” The sound thus produced is audible at a considerable distance but cannot be described; and I feel sure that its source would never be conjectured by any one hearing it for the first time. As this jarring sound is made chiefly during the breeding-season, it has been considered as a love-song; but it is perhaps more strictly a love-call. The female, when driven from her nest, has been observed thus to call her mate, who answered in the same manner and soon appeared. Lastly, the male hoopoe (Upupa epops) combines vocal and instrumental music; for during the breeding-season this bird, as Mr. Swinhoe observed, first draws in air, and then taps the end of its beak perpendicularly down against a stone or the trunk of a tree, “when the breath being forced down the tubular bill produces the correct sound.” If the beak is not thus struck against some object, the sound is quite different. Air is at the same time swallowed, and the oesophagus thus becomes much swollen; and this probably acts as a resonator, not only with the hoopoe, but with pigeons and other birds. (52. For the foregoing facts see, on Birds of Paradise, Brehm, ‘Thierleben,’ Band iii. s. 325. On Grouse, Richardson, ‘Fauna Bor. Americ.: Birds,’ pp. 343 and 359; Major W. Ross King, ‘The Sportsman in Canada,’ 1866, p. 156; Mr. Haymond, in Prof. Cox’s ‘Geol. Survey of Indiana,’ p. 227; Audubon, ‘American Ornitholog. Biograph.’ vol. i. p. 216. On the Kalij-pheasant, Jerdon, ‘Birds of India,’ vol. iii. p. 533. On the Weavers, Livingstone’s ‘Expedition to the Zambesi,’ 1865, p. 425. On Woodpeckers, Macgillivray, ‘Hist. of British Birds,’ vol. iii. 1840, pp. 84, 88, 89, and 95. On the Hoopoe, Mr. Swinhoe, in ‘Proc. Zoolog. Soc.’ June 23, 1863 and 1871, p. 348. On the Night-jar, Audubon, ibid. vol. ii. p. 255, and ‘American Naturalist,’ 1873, p. 672. The English Night-jar likewise makes in the spring a curious noise during its rapid flight.)

[Fig. 41. Outer tail-feather of Scolopax gallinago (from ‘Proc. Zool. Soc.’ 1858).

Fig. 42. Outer tail-feather of Scolopax frenata.

Fig. 43. Outer tail-feather of Scolopax javensis.]

In the foregoing cases sounds are made by the aid of structures already present and otherwise necessary; but in the following cases certain feathers have been specially modified for the express purpose of producing sounds. The drumming, bleating, neighing, or thundering noise (as expressed by different observers) made by the common snipe (Scolopax gallinago) must have surprised every one who has ever heard it. This bird, during the pairing-season, flies to “perhaps a thousand feet in height,” and after zig-zagging about for a time descends to the earth in a curved line, with outspread tail and quivering pinions, and surprising velocity. The sound is emitted only during this rapid descent. No one was able to explain the cause until M. Meves observed that on each side of the tail the outer feathers are peculiarly formed (Fig. 41), having a stiff sabre-shaped shaft with the oblique barbs of unusual length, the outer webs being strongly bound together. He found that by blowing on these feathers, or by fastening them to a long thin stick and waving them rapidly through the air, he could reproduce the drumming noise made by the living bird. Both sexes are furnished with these feathers, but they are generally larger in the male than in the female, and emit a deeper note. In some species, as in S. frenata (Fig. 42), four feathers, and in S. javensis (Fig. 43), no less than eight on each side of the tail are greatly modified. Different tones are emitted by the feathers of the different species when waved through the air; and the Scolopax Wilsonii of the United States makes a switching noise whilst descending rapidly to the earth. (53. See M. Meves’ interesting paper in ‘Proc. Zool. Soc.’ 1858, p. 199. For the habits of the snipe, Macgillivray, ‘History of British Birds,’ vol. iv. p. 371. For the American snipe, Capt. Blakiston, ‘Ibis,’ vol. v. 1863, p. 131.)

[Fig. 44. Primary wing-feather of a Humming-bird, the Selasphorus platycercus (from a sketch by Mr. Salvin). Upper figure, that of male; lower figure, corresponding feather of female.]

In the male of the Chamaepetes unicolor (a large gallinaceous bird of America), the first primary wing-feather is arched towards the tip and is much more attenuated than in the female. In an allied bird, the Penelope nigra, Mr. Salvin observed a male, which, whilst it flew downwards “with outstretched wings, gave forth a kind of crashing rushing noise,” like the falling of a tree. (54. Mr. Salvin, in ‘Proceedings, Zoological Society,’ 1867, p. 160. I am much indebted to this distinguished ornithologist for sketches of the feathers of the Chamaepetes, and for other information.) The male alone of one of the Indian bustards (Sypheotides auritus) has its primary wing-feathers greatly acuminated; and the male of an allied species is known to make a humming noise whilst courting the female. (55. Jerdon, ‘Birds of India,’ vol. iii. pp. 618, 621.) In a widely different group of birds, namely Humming-birds, the males alone of certain kinds have either the shafts of their primary wing-feathers broadly dilated, or the webs abruptly excised towards the extremity. The male, for instance, of Selasphorus platycercus, when adult, has the first primary wing-feather (Fig. 44), thus excised. Whilst flying from flower to flower he makes “a shrill, almost whistling noise” (56. Gould, ‘Introduction to the Trochilidae,’ 1861, p. 49. Salvin, ‘Proceedings, Zoological Society,’ 1867, p. 160.); but it did not appear to Mr. Salvin that the noise was intentionally made.

[Fig. 45. Secondary wing-feathers of Pipra deliciosa (from Mr. Sclater, in ‘Proc. Zool. Soc.’ 1860). The three upper feathers, a, b, c, from the male; the three lower corresponding feathers, d, e, f, from the female. a and d, fifth secondary wing-feather of male and female, upper surface. b and e, sixth secondary, upper surface. c and f, seventh secondary, lower surface.]

Lastly, in several species of a sub-genus of Pipra or Manakin, the males, as described by Mr. Sclater, have their SECONDARY wing-feathers modified in a still more remarkable manner. In the brilliantly-coloured P. deliciosa the first three secondaries are thick-stemmed and curved towards the body; in the fourth and fifth (Fig. 45, a) the change is greater; and in the sixth and seventh (b, c) the shaft “is thickened to an extraordinary degree, forming a solid horny lump.” The barbs also are greatly changed in shape, in comparison with the corresponding feathers (d, e, f) in the female. Even the bones of the wing, which support these singular feathers in the male, are said by Mr. Fraser to be much thickened. These little birds make an extraordinary noise, the first “sharp note being not unlike the crack of a whip.” (57. Sclater, in ‘Proceedings, Zoological Society,’ 1860, p. 90, and in ‘Ibis,’ vol. iv. 1862, p. 175. Also Salvin, in ‘Ibis,’ 1860, p. 37.)

The diversity of the sounds, both vocal and instrumental, made by the males of many birds during the breeding-season, and the diversity of the means for producing such sounds, are highly remarkable. We thus gain a high idea of their importance for sexual purposes, and are reminded of the conclusion arrived at as to insects. It is not difficult to imagine the steps by which the notes of a bird, primarily used as a mere call or for some other purpose, might have been improved into a melodious love song. In the case of the modified feathers, by which the drumming, whistling, or roaring noises are produced, we know that some birds during their courtship flutter, shake, or rattle their unmodified feathers together; and if the females were led to select the best performers, the males which possessed the strongest or thickest, or most attenuated feathers, situated on any part of the body, would be the most successful; and thus by slow degrees the feathers might be modified to almost any extent. The females, of course, would not notice each slight successive alteration in shape, but only the sounds thus produced. It is a curious fact that in the same class of animals, sounds so different as the drumming of the snipe’s tail, the tapping of the woodpecker’s beak, the harsh trumpet-like cry of certain water-fowl, the cooing of the turtle-dove, and the song of the nightingale, should all be pleasing to the females of the several species. But we must not judge of the tastes of distinct species by a uniform standard; nor must we judge by the standard of man’s taste. Even with man, we should remember what discordant noises, the beating of tom-toms and the shrill notes of reeds, please the ears of savages. Sir S. Baker remarks (58. ‘The Nile Tributaries of Abyssinia,’ 1867, p. 203.), that “as the stomach of the Arab prefers the raw meat and reeking liver taken hot from the animal, so does his ear prefer his equally coarse and discordant music to all other.”

LOVE ANTICS AND DANCES.

The curious love gestures of some birds have already been incidentally noticed; so that little need here be added. In Northern America large numbers of a grouse, the Tetrao phasianellus, meet every morning during the breeding-season on a selected level spot, and here they run round and round in a circle of about fifteen or twenty feet in diameter, so that the ground is worn quite bare, like a fairy-ring. In these Partridge-dances, as they are called by the hunters, the birds assume the strangest attitudes, and run round, some to the left and some to the right. Audubon describes the males of a heron (Ardea herodias) as walking about on their long legs with great dignity before the females, bidding defiance to their rivals. With one of the disgusting carrion-vultures (Cathartes jota) the same naturalist states that “the gesticulations and parade of the males at the beginning of the love-season are extremely ludicrous.” Certain birds perform their love-antics on the wing, as we have seen with the black African weaver, instead of on the ground. During the spring our little white-throat (Sylvia cinerea) often rises a few feet or yards in the air above some bush, and “flutters with a fitful and fantastic motion, singing all the while, and then drops to its perch.” The great English bustard throws himself into indescribably odd attitudes whilst courting the female, as has been figured by Wolf. An allied Indian bustard (Otis bengalensis) at such times “rises perpendicularly into the air with a hurried flapping of his wings, raising his crest and puffing out the feathers of his neck and breast, and then drops to the ground;” he repeats this manoeuvre several times, at the same time humming in a peculiar tone. Such females as happen to be near “obey this saltatory summons,” and when they approach he trails his wings and spreads his tail like a turkey-cock. (59. For Tetrao phasianellus, see Richardson, ‘Fauna, Bor. America,’ p. 361, and for further particulars Capt. Blakiston, ‘Ibis,’ 1863, p. 125. For the Cathartes and Ardea, Audubon, ‘Ornithological Biography,’ vol. ii. p. 51, and vol. iii. p. 89. On the White-throat, Macgillivray, ‘History of British Birds,’ vol. ii. p. 354. On the Indian Bustard, Jerdon, ‘Birds of India,’ vol. iii. p. 618.)

[Fig. 46. Bower-bird, Chlamydera maculata, with bower (from Brehm).]

But the most curious case is afforded by three allied genera of Australian birds, the famous Bower-birds,—no doubt the co-descendants of some ancient species which first acquired the strange instinct of constructing bowers for performing their love-antics. The bowers (Fig. 46), which, as we shall hereafter see, are decorated with feathers, shells, bones, and leaves, are built on the ground for the sole purpose of courtship, for their nests are formed in trees. Both sexes assist in the erection of the bowers, but the male is the principal workman. So strong is this instinct that it is practised under confinement, and Mr. Strange has described (60. Gould, ‘Handbook to the Birds of Australia,’ vol. i. pp. 444, 449, 455. The bower of the Satin Bower-bird may be seen in the Zoological Society’s Gardens, Regent’s Park.) the habits of some Satin Bower-birds which he kept in an aviary in New South Wales. “At times the male will chase the female all over the aviary, then go to the bower, pick up a gay feather or a large leaf, utter a curious kind of note, set all his feathers erect, run round the bower and become so excited that his eyes appear ready to start from his head; he continues opening first one wing then the other, uttering a low, whistling note, and, like the domestic cock, seems to be picking up something from the ground, until at last the female goes gently towards him.” Captain Stokes has described the habits and “play-houses” of another species, the Great Bower-bird, which was seen “amusing itself by flying backwards and forwards, taking a shell alternately from each side, and carrying it through the archway in its mouth.” These curious structures, formed solely as halls of assemblage, where both sexes amuse themselves and pay their court, must cost the birds much labour. The bower, for instance, of the Fawn-breasted species, is nearly four feet in length, eighteen inches in height, and is raised on a thick platform of sticks.

DECORATION.

I will first discuss the cases in which the males are ornamented either exclusively or in a much higher degree than the females, and in a succeeding chapter those in which both sexes are equally ornamented, and finally the rare cases in which the female is somewhat more brightly-coloured than the male. As with the artificial ornaments used by savage and civilised men, so with the natural ornaments of birds, the head is the chief seat of decoration. (61. See remarks to this effect, on the ‘Feeling of Beauty among Animals,’ by Mr. J. Shaw, in the ‘Athenaeum,’ Nov. 24th, 1866, p. 681.) The ornaments, as mentioned at the commencement of this chapter, are wonderfully diversified. The plumes on the front or back of the head consist of variously-shaped feathers, sometimes capable of erection or expansion, by which their beautiful colours are fully displayed. Elegant ear-tufts (Fig. 39) are occasionally present. The head is sometimes covered with velvety down, as with the pheasant; or is naked and vividly coloured. The throat, also, is sometimes ornamented with a beard, wattles, or caruncles. Such appendages are generally brightly-coloured, and no doubt serve as ornaments, though not always ornamental in our eyes; for whilst the male is in the act of courting the female, they often swell and assume vivid tints, as in the male turkey. At such times the fleshy appendages about the head of the male Tragopan pheasant (Ceriornis Temminckii) swell into a large lappet on the throat and into two horns, one on each side of the splendid top-knot; and these are then coloured of the most intense blue which I have ever beheld. (62. See Dr. Murie’s account with coloured figures in ‘Proceedings, Zoological Society,’ 1872, p. 730.) The African hornbill (Bucorax abyssinicus) inflates the scarlet bladder-like wattle on its neck, and with its wings drooping and tail expanded “makes quite a grand appearance.” (63. Mr. Monteiro, ‘Ibis,’ vol. iv. 1862, p. 339.) Even the iris of the eye is sometimes more brightly-coloured in the male than in the female; and this is frequently the case with the beak, for instance, in our common blackbird. In Buceros corrugatus, the whole beak and immense casque are coloured more conspicuously in the male than in the female; and “the oblique grooves upon the sides of the lower mandible are peculiar to the male sex.” (64. ‘Land and Water,’ 1868, p. 217.)

The head, again, often supports fleshy appendages, filaments, and solid protuberances. These, if not common to both sexes, are always confined to the males. The solid protuberances have been described in detail by Dr. W. Marshall (65. ‘Ueber die Schädelhöcker,’ etc., ‘Niederland. Archiv. fur Zoologie,’ B. I. Heft 2, 1872.), who shews that they are formed either of cancellated bone coated with skin, or of dermal and other tissues. With mammals true horns are always supported on the frontal bones, but with birds various bones have been modified for this purpose; and in species of the same group the protuberances may have cores of bone, or be quite destitute of them, with intermediate gradations connecting these two extremes. Hence, as Dr. Marshall justly remarks, variations of the most different kinds have served for the development through sexual selection of these ornamental appendages. Elongated feathers or plumes spring from almost every part of the body. The feathers on the throat and breast are sometimes developed into beautiful ruffs and collars. The tail-feathers are frequently increased in length; as we see in the tail-coverts of the peacock, and in the tail itself of the Argus pheasant. With the peacock even the bones of the tail have been modified to support the heavy tail-coverts. (66. Dr. W. Marshall, ‘Über den Vogelschwanz,’ ibid. B. I. Heft 2, 1872.) The body of the Argus is not larger than that of a fowl; yet the length from the end of the beak to the extremity of the tail is no less than five feet three inches (67. Jardine’s ‘Naturalist Library: Birds,’ vol. xiv. p. 166.), and that of the beautifully ocellated secondary wing-feathers nearly three feet. In a small African night-jar (Cosmetornis vexillarius) one of the primary wing-feathers, during the breeding-season, attains a length of twenty-six inches, whilst the bird itself is only ten inches in length. In another closely-allied genus of night-jars, the shafts of the elongated wing-feathers are naked, except at the extremity, where there is a disc. (68. Sclater, in the ‘Ibis,’ vol. vi. 1864, p. 114; Livingstone, ‘Expedition to the Zambesi,’ 1865, p. 66.) Again, in another genus of night-jars, the tail-feathers are even still more prodigiously developed. In general the feathers of the tail are more often elongated than those of the wings, as any great elongation of the latter impedes flight. We thus see that in closely-allied birds ornaments of the same kind have been gained by the males through the development of widely different feathers.

It is a curious fact that the feathers of species belonging to very distinct groups have been modified in almost exactly the same peculiar manner. Thus the wing-feathers in one of the above-mentioned night-jars are bare along the shaft, and terminate in a disc; or are, as they are sometimes called, spoon or racket-shaped. Feathers of this kind occur in the tail of a motmot (Eumomota superciliaris), of a king-fisher, finch, humming-bird, parrot, several Indian drongos (Dicrurus and Edolius, in one of which the disc stands vertically), and in the tail of certain birds of paradise. In these latter birds, similar feathers, beautifully ocellated, ornament the head, as is likewise the case with some gallinaceous birds. In an Indian bustard (Sypheotides auritus) the feathers forming the ear-tufts, which are about four inches in length, also terminate in discs. (69. Jerdon, ‘Birds of India,’ vol. iii. p. 620.) It is a most singular fact that the motmots, as Mr. Salvin has clearly shewn (70. ‘Proceedings, Zoological Society,’ 1873, p. 429.), give to their tail feathers the racket-shape by biting off the barbs, and, further, that this continued mutilation has produced a certain amount of inherited effect.

[Fig. 47. Paradisea Papuana (T.W. Wood).]

Again, the barbs of the feathers in various widely-distinct birds are filamentous or plumose, as with some herons, ibises, birds of paradise, and Gallinaceae. In other cases the barbs disappear, leaving the shafts bare from end to end; and these in the tail of the Paradisea apoda attain a length of thirty-four inches (71. Wallace, in ‘Annals and Magazine of Natural History,’ vol. xx. 1857, p. 416, and in his ‘Malay Archipelago,’ vol. ii. 1869, p. 390.): in P. Papuana (Fig. 47) they are much shorter and thin. Smaller feathers when thus denuded appear like bristles, as on the breast of the turkey-cock. As any fleeting fashion in dress comes to be admired by man, so with birds a change of almost any kind in the structure or colouring of the feathers in the male appears to have been admired by the female. The fact of the feathers in widely distinct groups having been modified in an analogous manner no doubt depends primarily on all the feathers having nearly the same structure and manner of development, and consequently tending to vary in the same manner. We often see a tendency to analogous variability in the plumage of our domestic breeds belonging to distinct species. Thus top-knots have appeared in several species. In an extinct variety of the turkey, the top-knot consisted of bare quills surmounted with plumes of down, so that they somewhat resembled the racket-shaped feathers above described. In certain breeds of the pigeon and fowl the feathers are plumose, with some tendency in the shafts to be naked. In the Sebastopol goose the scapular feathers are greatly elongated, curled, or even spirally twisted, with the margins plumose. (72. See my work on ‘The Variation of Animals and Plants under Domestication,’ vol. i. pp. 289, 293.)

In regard to colour, hardly anything need here be said, for every one knows how splendid are the tints of many birds, and how harmoniously they are combined. The colours are often metallic and iridescent. Circular spots are sometimes surrounded by one or more differently shaded zones, and are thus converted into ocelli. Nor need much be said on the wonderful difference between the sexes of many birds. The common peacock offers a striking instance. Female birds of paradise are obscurely coloured and destitute of all ornaments, whilst the males are probably the most highly decorated of all birds, and in so many different ways that they must be seen to be appreciated. The elongated and golden-orange plumes which spring from beneath the wings of the Paradisea apoda, when vertically erected and made to vibrate, are described as forming a sort of halo, in the centre of which the head “looks like a little emerald sun with its rays formed by the two plumes.” (73. Quoted from M. de Lafresnaye in ‘Annals and Mag. of Natural History,’ vol. xiii. 1854, p. 157: see also Mr. Wallace’s much fuller account in vol. xx. 1857, p. 412, and in his ‘Malay Archipelago.’) In another most beautiful species the head is bald, “and of a rich cobalt blue, crossed by several lines of black velvety feathers.” (74. Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 405.)

[Fig. 48. Lophornis ornatus, male and female (from Brehm).

Fig. 49. Spathura underwoodi, male and female (from Brehm).]

Male humming-birds (Figs. 48 and 49) almost vie with birds of paradise in their beauty, as every one will admit who has seen Mr. Gould’s splendid volumes, or his rich collection. It is very remarkable in how many different ways these birds are ornamented. Almost every part of their plumage has been taken advantage of, and modified; and the modifications have been carried, as Mr. Gould shewed me, to a wonderful extreme in some species belonging to nearly every sub-group. Such cases are curiously like those which we see in our fancy breeds, reared by man for the sake of ornament; certain individuals originally varied in one character, and other individuals of the same species in other characters; and these have been seized on by man and much augmented—as shewn by the tail of the fantail-pigeon, the hood of the jacobin, the beak and wattle of the carrier, and so forth. The sole difference between these cases is that in the one, the result is due to man’s selection, whilst in the other, as with humming-birds, birds of paradise, etc., it is due to the selection by the females of the more beautiful males.

I will mention only one other bird, remarkable from the extreme contrast in colour between the sexes, namely the famous bell-bird (Chasmorhynchus niveus) of S. America, the note of which can be distinguished at the distance of nearly three miles, and astonishes every one when first hearing it. The male is pure white, whilst the female is dusky-green; and white is a very rare colour in terrestrial species of moderate size and inoffensive habits. The male, also, as described by Waterton, has a spiral tube, nearly three inches in length, which rises from the base of the beak. It is jet-black, dotted over with minute downy feathers. This tube can be inflated with air, through a communication with the palate; and when not inflated hangs down on one side. The genus consists of four species, the males of which are very distinct, whilst the females, as described by Mr. Sclater in a very interesting paper, closely resemble each other, thus offering an excellent instance of the common rule that within the same group the males differ much more from each other than do the females. In a second species (C. nudicollis) the male is likewise snow-white, with the exception of a large space of naked skin on the throat and round the eyes, which during the breeding-season is of a fine green colour. In a third species (C. tricarunculatus) the head and neck alone of the male are white, the rest of the body being chestnut-brown, and the male of this species is provided with three filamentous projections half as long as the body—one rising from the base of the beak, and the two others from the corners of the mouth. (75. Mr. Sclater, ‘Intellectual Observer,’ Jan. 1867. Waterton’s ‘Wanderings,’ p. 118. See also Mr. Salvin’s interesting paper, with a plate, in the ‘Ibis,’ 1865, p. 90.)

The coloured plumage and certain other ornaments of the adult males are either retained for life, or are periodically renewed during the summer and breeding-season. At this same season the beak and naked skin about the head frequently change colour, as with some herons, ibises, gulls, one of the bell-birds just noticed, etc. In the white ibis, the cheeks, the inflatable skin of the throat, and the basal portion of the beak then become crimson. (76. ‘Land and Water,’ 1867, p. 394.) In one of the rails, Gallicrex cristatus, a large red caruncle is developed during this period on the head of the male. So it is with a thin horny crest on the beak of one of the pelicans, P. erythrorhynchus; for, after the breeding-season, these horny crests are shed, like horns from the heads of stags, and the shore of an island in a lake in Nevada was found covered with these curious exuviae. (77. Mr. D.G. Elliot, in ‘Proc. Zool. Soc.’ 1869, p. 589.)

Changes of colour in the plumage according to the season depend, firstly on a double annual moult, secondly on an actual change of colour in the feathers themselves, and thirdly on their dull-coloured margins being periodically shed, or on these three processes more or less combined. The shedding of the deciduary margins may be compared with the shedding of their down by very young birds; for the down in most cases arises from the summits of the first true feathers. (78. Nitzsch’s ‘Pterylography,’ edited by P.L. Sclater, Ray Society, 1867, p. 14.)

With respect to the birds which annually undergo a double moult, there are, firstly, some kinds, for instance snipes, swallow-plovers (Glareolae), and curlews, in which the two sexes resemble each other, and do not change colour at any season. I do not know whether the winter plumage is thicker and warmer than the summer plumage, but warmth seems the most probable end attained of a double moult, where there is no change of colour. Secondly, there are birds, for instance, certain species of Totanus and other Grallatores, the sexes of which resemble each other, but in which the summer and winter plumage differ slightly in colour. The difference, however, in these cases is so small that it can hardly be an advantage to them; and it may, perhaps, be attributed to the direct action of the different conditions to which the birds are exposed during the two seasons. Thirdly, there are many other birds the sexes of which are alike, but which are widely different in their summer and winter plumage. Fourthly, there are birds the sexes of which differ from each other in colour; but the females, though moulting twice, retain the same colours throughout the year, whilst the males undergo a change of colour, sometimes a great one, as with certain bustards. Fifthly and lastly, there are birds the sexes of which differ from each other in both their summer and winter plumage; but the male undergoes a greater amount of change at each recurrent season than the female—of which the ruff (Machetes pugnax) offers a good instance.

With respect to the cause or purpose of the differences in colour between the summer and winter plumage, this may in some instances, as with the ptarmigan (79. The brown mottled summer plumage of the ptarmigan is of as much importance to it, as a protection, as the white winter plumage; for in Scandinavia during the spring, when the snow has disappeared, this bird is known to suffer greatly from birds of prey, before it has acquired its summer dress: see Wilhelm von Wright, in Lloyd, ‘Game Birds of Sweden,’ 1867, p. 125.), serve during both seasons as a protection. When the difference between the two plumages is slight it may perhaps be attributed, as already remarked, to the direct action of the conditions of life. But with many birds there can hardly be a doubt that the summer plumage is ornamental, even when both sexes are alike. We may conclude that this is the case with many herons, egrets, etc., for they acquire their beautiful plumes only during the breeding-season. Moreover, such plumes, top-knots, etc., though possessed by both sexes, are occasionally a little more developed in the male than in the female; and they resemble the plumes and ornaments possessed by the males alone of other birds. It is also known that confinement, by affecting the reproductive system of male birds, frequently checks the development of their secondary sexual characters, but has no immediate influence on any other characters; and I am informed by Mr. Bartlett that eight or nine specimens of the Knot (Tringa canutus) retained their unadorned winter plumage in the Zoological Gardens throughout the year, from which fact we may infer that the summer plumage, though common to both sexes, partakes of the nature of the exclusively masculine plumage of many other birds. (80. In regard to the previous statements on moulting, see, on snipes, etc., Macgillivray, ‘Hist. Brit. Birds,’ vol. iv. p. 371; on Glareolae, curlews, and bustards, Jerdon, ‘Birds of India,’ vol. iii. pp. 615, 630, 683; on Totanus, ibid. p. 700; on the plumes of herons, ibid. p. 738, and Macgillivray, vol. iv. pp. 435 and 444, and Mr. Stafford Allen, in the ‘Ibis,’ vol. v. 1863, p. 33.)

From the foregoing facts, more especially from neither sex of certain birds changing colour during either annual moult, or changing so slightly that the change can hardly be of any service to them, and from the females of other species moulting twice yet retaining the same colours throughout the year, we may conclude that the habit of annually moulting twice has not been acquired in order that the male should assume an ornamental character during the breeding-season; but that the double moult, having been originally acquired for some distinct purpose, has subsequently been taken advantage of in certain cases for gaining a nuptial plumage.

It appears at first sight a surprising circumstance that some closely-allied species should regularly undergo a double annual moult, and others only a single one. The ptarmigan, for instance, moults twice or even thrice in the year, and the blackcock only once: some of the splendidly coloured honey-suckers (Nectariniae) of India and some sub-genera of obscurely coloured pipits (Anthus) have a double, whilst others have only a single annual moult. (81. On the moulting of the ptarmigan, see Gould’s ‘Birds of Great Britain.’ On the honey-suckers, Jerdon, ‘Birds of India,’ vol. i. pp. 359, 365, 369. On the moulting of Anthus, see Blyth, in ‘Ibis,’ 1867, p. 32.) But the gradations in the manner of moulting, which are known to occur with various birds, shew us how species, or whole groups, might have originally acquired their double annual moult, or having once gained the habit, have again lost it. With certain bustards and plovers the vernal moult is far from complete, some feathers being renewed, and some changed in colour. There is also reason to believe that with certain bustards and rail-like birds, which properly undergo a double moult, some of the older males retain their nuptial plumage throughout the year. A few highly modified feathers may merely be added during the spring to the plumage, as occurs with the disc-formed tail-feathers of certain drongos (Bhringa) in India, and with the elongated feathers on the back, neck, and crest of certain herons. By such steps as these, the vernal moult might be rendered more and more complete, until a perfect double moult was acquired. Some of the birds of paradise retain their nuptial feathers throughout the year, and thus have only a single moult; others cast them directly after the breeding-season, and thus have a double moult; and others again cast them at this season during the first year, but not afterwards; so that these latter species are intermediate in their manner of moulting. There is also a great difference with many birds in the length of time during which the two annual plumages are retained; so that the one might come to be retained for the whole year, and the other completely lost. Thus in the spring Machetes pugnax retains his ruff for barely two months. In Natal the male widow-bird (Chera progne) acquires his fine plumage and long tail-feathers in December or January, and loses them in March; so that they are retained only for about three months. Most species, which undergo a double moult, keep their ornamental feathers for about six months. The male, however, of the wild Gallus bankiva retains his neck-hackles for nine or ten months; and when these are cast off, the underlying black feathers on the neck are fully exposed to view. But with the domesticated descendant of this species, the neck-hackles of the male are immediately replaced by new ones; so that we here see, as to part of the plumage, a double moult changed under domestication into a single moult. (82. For the foregoing statements in regard to partial moults, and on old males retaining their nuptial plumage, see Jerdon, on bustards and plovers, in ‘Birds of India,’ vol. iii. pp. 617, 637, 709, 711. Also Blyth in ‘Land and Water,’ 1867, p. 84. On the moulting of Paradisea, see an interesting article by Dr. W. Marshall, ‘Archives Neerlandaises,’ tom. vi. 1871. On the Vidua, ‘Ibis,’ vol. iii. 1861, p. 133. On the Drongo-shrikes, Jerdon, ibid. vol. i. p. 435. On the vernal moult of the Herodias bubulcus, Mr. S.S. Allen, in ‘Ibis,’ 1863, p. 33. On Gallus bankiva, Blyth, in ‘Annals and Mag. of Natural History,’ vol. i. 1848, p. 455; see, also, on this subject, my ‘Variation of Animals under Domestication,’ vol. i. p. 236.)

The common drake (Anas boschas), after the breeding-season, is well known to lose his male plumage for a period of three months, during which time he assumes that of the female. The male pin-tail duck (Anas acuta) loses his plumage for the shorter period of six weeks or two months; and Montagu remarks that “this double moult within so short a time is a most extraordinary circumstance, that seems to bid defiance to all human reasoning.” But the believer in the gradual modification of species will be far from feeling surprise at finding gradations of all kinds. If the male pin-tail were to acquire his new plumage within a still shorter period, the new male feathers would almost necessarily be mingled with the old, and both with some proper to the female; and this apparently is the case with the male of a not distantly-allied bird, namely the Merganser serrator, for the males are said to “undergo a change of plumage, which assimilates them in some measure to the female.” By a little further acceleration in the process, the double moult would be completely lost. (83. See Macgillivray, ‘Hist. British Birds’ (vol. v. pp. 34, 70, and 223), on the moulting of the Anatidae, with quotations from Waterton and Montagu. Also Yarrell, ‘History of British Birds,’ vol. iii. p. 243.)

Some male birds, as before stated, become more brightly coloured in the spring, not by a vernal moult, but either by an actual change of colour in the feathers, or by their obscurely-coloured deciduary margins being shed. Changes of colour thus caused may last for a longer or shorter time. In the Pelecanus onocrotalus a beautiful rosy tint, with lemon-coloured marks on the breast, overspreads the whole plumage in the spring; but these tints, as Mr. Sclater states, “do not last long, disappearing generally in about six weeks or two months after they have been attained.” Certain finches shed the margins of their feathers in the spring, and then become brighter coloured, while other finches undergo no such change. Thus the Fringilla tristis of the United States (as well as many other American species) exhibits its bright colours only when the winter is past, whilst our goldfinch, which exactly represents this bird in habits, and our siskin, which represents it still more closely in structure, undergo no such annual change. But a difference of this kind in the plumage of allied species is not surprising, for with the common linnet, which belongs to the same family, the crimson forehead and breast are displayed only during the summer in England, whilst in Madeira these colours are retained throughout the year. (84. On the pelican, see Sclater, in ‘Proc. Zool. Soc.’ 1868, p. 265. On the American finches, see Audubon, ‘Ornithological Biography,’ vol. i. pp. 174, 221, and Jerdon, ‘Birds of India,’ vol. ii. p. 383. On the Fringilla cannabina of Madeira, Mr. E. Vernon Harcourt, ‘Ibis,’ vol. v. 1863, p. 230.)

DISPLAY BY MALE BIRDS OF THEIR PLUMAGE.

Ornaments of all kinds, whether permanently or temporarily gained, are sedulously displayed by the males, and apparently serve to excite, attract, or fascinate the females. But the males will sometimes display their ornaments, when not in the presence of the females, as occasionally occurs with grouse at their balz-places, and as may be noticed with the peacock; this latter bird, however, evidently wishes for a spectator of some kind, and, as I have often seen, will shew off his finery before poultry, or even pigs. (85. See also ‘Ornamental Poultry,’ by Rev. E.S. Dixon, 1848, p. 8.) All naturalists who have closely attended to the habits of birds, whether in a state of nature or under confinement, are unanimously of opinion that the males take delight in displaying their beauty. Audubon frequently speaks of the male as endeavouring in various ways to charm the female. Mr. Gould, after describing some peculiarities in a male humming-bird, says he has no doubt that it has the power of displaying them to the greatest advantage before the female. Dr. Jerdon (86. ‘Birds of India,’ introduct., vol. i. p. xxiv.; on the peacock, vol. iii. p. 507. See Gould’s ‘Introduction to Trochilidae,’ 1861, pp. 15 and 111.) insists that the beautiful plumage of the male serves “to fascinate and attract the female.” Mr. Bartlett, at the Zoological Gardens, expressed himself to me in the strongest terms to the same effect.

[Fig. 50. Rupicola crocea, male (T.W. Wood).]

It must be a grand sight in the forests of India “to come suddenly on twenty or thirty pea-fowl, the males displaying their gorgeous trains, and strutting about in all the pomp of pride before the gratified females.” The wild turkey-cock erects his glittering plumage, expands his finely-zoned tail and barred wing-feathers, and altogether, with his crimson and blue wattles, makes a superb, though, to our eyes, grotesque appearance. Similar facts have already been given with respect to grouse of various kinds. Turning to another Order: The male Rupicola crocea (Fig. 50) is one of the most beautiful birds in the world, being of a splendid orange, with some of the feathers curiously truncated and plumose. The female is brownish-green, shaded with red, and has a much smaller crest. Sir R. Schomburgk has described their courtship; he found one of their meeting-places where ten males and two females were present. The space was from four to five feet in diameter, and appeared to have been cleared of every blade of grass and smoothed as if by human hands. A male “was capering, to the apparent delight of several others. Now spreading its wings, throwing up its head, or opening its tail like a fan; now strutting about with a hopping gait until tired, when it gabbled some kind of note, and was relieved by another. Thus three of them successively took the field, and then, with self-approbation, withdrew to rest.” The Indians, in order to obtain their skins, wait at one of the meeting-places till the birds are eagerly engaged in dancing, and then are able to kill with their poisoned arrows four or five males, one after the other. (87. ‘Journal of R. Geograph. Soc.’ vol. x. 1840, p. 236.) With birds of paradise a dozen or more full-plumaged males congregate in a tree to hold a dancing-party, as it is called by the natives: and here they fly about, raise their wings, elevate their exquisite plumes, and make them vibrate, and the whole tree seems, as Mr. Wallace remarks, to be filled with waving plumes. When thus engaged, they become so absorbed that a skilful archer may shoot nearly the whole party. These birds, when kept in confinement in the Malay Archipelago, are said to take much care in keeping their feathers clean; often spreading them out, examining them, and removing every speck of dirt. One observer, who kept several pairs alive, did not doubt that the display of the male was intended to please the female. (88. ‘Annals and Mag. of Nat. Hist.’ vol. xiii. 1854, p. 157; also Wallace, ibid. vol. xx. 1857, p. 412, and ‘The Malay Archipelago,’ vol. ii. 1869, p. 252. Also Dr. Bennett, as quoted by Brehm, ‘Thierleben,’ B. iii. s. 326.)

[Fig. 51. Polyplectron chinquis, male (T.W. Wood).]

The Gold and Amherst pheasants during their courtship not only expand and raise their splendid frills, but twist them, as I have myself seen, obliquely towards the female on whichever side she may be standing, obviously in order that a large surface may be displayed before her. (89. Mr. T.W. Wood has given (‘The Student,’ April 1870, p. 115) a full account of this manner of display, by the Gold pheasant and by the Japanese pheasant, Ph. versicolor; and he calls it the lateral or one-sided display.) They likewise turn their beautiful tails and tail-coverts a little towards the same side. Mr. Bartlett has observed a male Polyplectron (Fig. 51) in the act of courtship, and has shewn me a specimen stuffed in the attitude then assumed. The tail and wing-feathers of this bird are ornamented with beautiful ocelli, like those on the peacock’s train. Now when the peacock displays himself, he expands and erects his tail transversely to his body, for he stands in front of the female, and has to shew off, at the same time, his rich blue throat and breast. But the breast of the Polyplectron is obscurely coloured, and the ocelli are not confined to the tail-feathers. Consequently the Polyplectron does not stand in front of the female; but he erects and expands his tail-feathers a little obliquely, lowering the expanded wing on the same side, and raising that on the opposite side. In this attitude the ocelli over the whole body are exposed at the same time before the eyes of the admiring female in one grand bespangled expanse. To whichever side she may turn, the expanded wings and the obliquely-held tail are turned towards her. The male Tragopan pheasant acts in nearly the same manner, for he raises the feathers of the body, though not the wing itself, on the side which is opposite to the female, and which would otherwise be concealed, so that nearly all the beautifully spotted feathers are exhibited at the same time.

[Fig. 52. Side view of male Argus pheasant, whilst displaying before the female. Observed and sketched from nature by T.W. Wood.]

The Argus pheasant affords a much more remarkable case. The immensely developed secondary wing-feathers are confined to the male; and each is ornamented with a row of from twenty to twenty-three ocelli, above an inch in diameter. These feathers are also elegantly marked with oblique stripes and rows of spots of a dark colour, like those on the skin of a tiger and leopard combined. These beautiful ornaments are hidden until the male shows himself off before the female. He then erects his tail, and expands his wing-feathers into a great, almost upright, circular fan or shield, which is carried in front of the body. The neck and head are held on one side, so that they are concealed by the fan; but the bird in order to see the female, before whom he is displaying himself, sometimes pushes his head between two of the long wing-feathers (as Mr. Bartlett has seen), and then presents a grotesque appearance. This must be a frequent habit with the bird in a state of nature, for Mr. Bartlett and his son on examining some perfect skins sent from the East, found a place between two of the feathers which was much frayed, as if the head had here frequently been pushed through. Mr. Wood thinks that the male can also peep at the female on one side, beyond the margin of the fan.

The ocelli on the wing-feathers are wonderful objects; for they are so shaded that, as the Duke of Argyll remarks (90. ‘The Reign of Law,’ 1867, p. 203.), they stand out like balls lying loosely within sockets. When I looked at the specimen in the British Museum, which is mounted with the wings expanded and trailing downwards, I was however greatly disappointed, for the ocelli appeared flat, or even concave. But Mr. Gould soon made the case clear to me, for he held the feathers erect, in the position in which they would naturally be displayed, and now, from the light shining on them from above, each ocellus at once resembled the ornament called a ball and socket. These feathers have been shown to several artists, and all have expressed their admiration at the perfect shading. It may well be asked, could such artistically shaded ornaments have been formed by means of sexual selection? But it will be convenient to defer giving an answer to this question until we treat in the next chapter of the principle of gradation.

The foregoing remarks relate to the secondary wing-feathers, but the primary wing-feathers, which in most gallinaceous birds are uniformly coloured, are in the Argus pheasant equally wonderful. They are of a soft brown tint with numerous dark spots, each of which consists of two or three black dots with a surrounding dark zone. But the chief ornament is a space parallel to the dark-blue shaft, which in outline forms a perfect second feather lying within the true feather. This inner part is coloured of a lighter chestnut, and is thickly dotted with minute white points. I have shewn this feather to several persons, and many have admired it even more than the ball and socket feathers, and have declared that it was more like a work of art than of nature. Now these feathers are quite hidden on all ordinary occasions, but are fully displayed, together with the long secondary feathers, when they are all expanded together so as to form the great fan or shield.

The case of the male Argus pheasant is eminently interesting, because it affords good evidence that the most refined beauty may serve as a sexual charm, and for no other purpose. We must conclude that this is the case, as the secondary and primary wing-feathers are not at all displayed, and the ball and socket ornaments are not exhibited in full perfection until the male assumes the attitude of courtship. The Argus pheasant does not possess brilliant colours, so that his success in love appears to depend on the great size of his plumes, and on the elaboration of the most elegant patterns. Many will declare that it is utterly incredible that a female bird should be able to appreciate fine shading and exquisite patterns. It is undoubtedly a marvellous fact that she should possess this almost human degree of taste. He who thinks that he can safely gauge the discrimination and taste of the lower animals may deny that the female Argus pheasant can appreciate such refined beauty; but he will then be compelled to admit that the extraordinary attitudes assumed by the male during the act of courtship, by which the wonderful beauty of his plumage is fully displayed, are purposeless; and this is a conclusion which I for one will never admit.

Although so many pheasants and allied gallinaceous birds carefully display their plumage before the females, it is remarkable, as Mr. Bartlett informs me, that this is not the case with the dull-coloured Eared and Cheer pheasants (Crossoptilon auritum and Phasianus wallichii); so that these birds seem conscious that they have little beauty to display. Mr. Bartlett has never seen the males of either of these species fighting together, though he has not had such good opportunities for observing the Cheer as the Eared pheasant. Mr. Jenner Weir, also, finds that all male birds with rich or strongly-characterised plumage are more quarrelsome than the dull-coloured species belonging to the same groups. The goldfinch, for instance, is far more pugnacious than the linnet, and the blackbird than the thrush. Those birds which undergo a seasonal change of plumage likewise become much more pugnacious at the period when they are most gaily ornamented. No doubt the males of some obscurely-coloured birds fight desperately together, but it appears that when sexual selection has been highly influential, and has given bright colours to the males of any species, it has also very often given a strong tendency to pugnacity. We shall meet with nearly analogous cases when we treat of mammals. On the other hand, with birds the power of song and brilliant colours have rarely been both acquired by the males of the same species; but in this case the advantage gained would have been the same, namely success in charming the female. Nevertheless it must be owned that the males of several brilliantly coloured birds have had their feathers specially modified for the sake of producing instrumental music, though the beauty of this cannot be compared, at least according to our taste, with that of the vocal music of many songsters.

We will now turn to male birds which are not ornamented in any high degree, but which nevertheless display during their courtship whatever attractions they may possess. These cases are in some respects more curious than the foregoing, and have been but little noticed. I owe the following facts to Mr. Weir, who has long kept confined birds of many kinds, including all the British Fringillidae and Emberizidae. The facts have been selected from a large body of valuable notes kindly sent me by him. The bullfinch makes his advances in front of the female, and then puffs out his breast, so that many more of the crimson feathers are seen at once than otherwise would be the case. At the same time he twists and bows his black tail from side to side in a ludicrous manner. The male chaffinch also stands in front of the female, thus shewing his red breast and “blue bell,” as the fanciers call his head; the wings at the same time being slightly expanded, with the pure white bands on the shoulders thus rendered conspicuous. The common linnet distends his rosy breast, slightly expands his brown wings and tail, so as to make the best of them by exhibiting their white edgings. We must, however, be cautious in concluding that the wings are spread out solely for display, as some birds do so whose wings are not beautiful. This is the case with the domestic cock, but it is always the wing on the side opposite to the female which is expanded, and at the same time scraped on the ground. The male goldfinch behaves differently from all other finches: his wings are beautiful, the shoulders being black, with the dark-tipped wing-feathers spotted with white and edged with golden yellow. When he courts the female, he sways his body from side to side, and quickly turns his slightly expanded wings first to one side, then to the other, with a golden flashing effect. Mr. Weir informs me that no other British finch turns thus from side to side during his courtship, not even the closely-allied male siskin, for he would not thus add to his beauty.

Most of the British Buntings are plain coloured birds; but in the spring the feathers on the head of the male reed-bunting (Emberiza schoeniculus) acquire a fine black colour by the abrasion of the dusky tips; and these are erected during the act of courtship. Mr. Weir has kept two species of Amadina from Australia: the A. castanotis is a very small and chastely coloured finch, with a dark tail, white rump, and jet-black upper tail-coverts, each of the latter being marked with three large conspicuous oval spots of white. (91. For the description of these birds, see Gould’s ‘Handbook to the Birds of Australia,’ vol. i. 1865, p. 417.) This species, when courting the female, slightly spreads out and vibrates these parti-coloured tail-coverts in a very peculiar manner. The male Amadina Lathami behaves very differently, exhibiting before the female his brilliantly spotted breast, scarlet rump, and scarlet upper tail-coverts. I may here add from Dr. Jerdon that the Indian bulbul (Pycnonotus hoemorrhous) has its under tail-coverts of a crimson colour, and these, it might be thought, could never be well exhibited; but the bird “when excited often spreads them out laterally, so that they can be seen even from above.” (92. ‘Birds of India,’ vol. ii. p. 96.) The crimson under tail-coverts of some other birds, as with one of the woodpeckers, Picus major, can be seen without any such display. The common pigeon has iridescent feathers on the breast, and every one must have seen how the male inflates his breast whilst courting the female, thus shewing them off to the best advantage. One of the beautiful bronze-winged pigeons of Australia (Ocyphaps lophotes) behaves, as described to me by Mr. Weir, very differently: the male, whilst standing before the female, lowers his head almost to the ground, spreads out and raises his tail, and half expands his wings. He then alternately and slowly raises and depresses his body, so that the iridescent metallic feathers are all seen at once, and glitter in the sun.

Sufficient facts have now been given to shew with what care male birds display their various charms, and this they do with the utmost skill. Whilst preening their feathers, they have frequent opportunities for admiring themselves, and of studying how best to exhibit their beauty. But as all the males of the same species display themselves in exactly the same manner, it appears that actions, at first perhaps intentional, have become instinctive. If so, we ought not to accuse birds of conscious vanity; yet when we see a peacock strutting about, with expanded and quivering tail-feathers, he seems the very emblem of pride and vanity.

The various ornaments possessed by the males are certainly of the highest importance to them, for in some cases they have been acquired at the expense of greatly impeded powers of flight or of running. The African night-jar (Cosmetornis), which during the pairing-season has one of its primary wing-feathers developed into a streamer of very great length, is thereby much retarded in its flight, although at other times remarkable for its swiftness. The “unwieldy size” of the secondary wing-feathers of the male Argus pheasant is said “almost entirely to deprive the bird of flight.” The fine plumes of male birds of paradise trouble them during a high wind. The extremely long tail-feathers of the male widow-birds (Vidua) of Southern Africa render “their flight heavy;” but as soon as these are cast off they fly as well as the females. As birds always breed when food is abundant, the males probably do not suffer much inconvenience in searching for food from their impeded powers of movement; but there can hardly be a doubt that they must be much more liable to be struck down by birds of prey. Nor can we doubt that the long train of the peacock and the long tail and wing-feathers of the Argus pheasant must render them an easier prey to any prowling tiger-cat than would otherwise be the case. Even the bright colours of many male birds cannot fail to make them conspicuous to their enemies of all kinds. Hence, as Mr. Gould has remarked, it probably is that such birds are generally of a shy disposition, as if conscious that their beauty was a source of danger, and are much more difficult to discover or approach, than the sombre coloured and comparatively tame females or than the young and as yet unadorned males. (93. On the Cosmetornis, see Livingstone’s ‘Expedition to the Zambesi,’ 1865, p. 66. On the Argus pheasant, Jardine’s ‘Nat. Hist. Lib.: Birds,’ vol. xiv. p. 167. On Birds of Paradise, Lesson, quoted by Brehm, ‘Thierleben,’ B. iii. s. 325. On the widow-bird, Barrow’s ‘Travels in Africa,’ vol. i. p. 243, and ‘Ibis,’ vol. iii. 1861 p. 133. Mr. Gould, on the shyness of male birds, ‘Handbook to Birds of Australia,’ vol. i. 1865, pp. 210, 457.)

It is a more curious fact that the males of some birds which are provided with special weapons for battle, and which in a state of nature are so pugnacious that they often kill each other, suffer from possessing certain ornaments. Cock-fighters trim the hackles and cut off the combs and gills of their cocks; and the birds are then said to be dubbed. An undubbed bird, as Mr. Tegetmeier insists, “is at a fearful disadvantage; the comb and gills offer an easy hold to his adversary’s beak, and as a cock always strikes where he holds, when once he has seized his foe, he has him entirely in his power. Even supposing that the bird is not killed, the loss of blood suffered by an undubbed cock is much greater than that sustained by one that has been trimmed.” (94. Tegetmeier, ‘The Poultry Book,’ 1866, p. 139.) Young turkey-cocks in fighting always seize hold of each other’s wattles; and I presume that the old birds fight in the same manner. It may perhaps be objected that the comb and wattles are not ornamental, and cannot be of service to the birds in this way; but even to our eyes, the beauty of the glossy black Spanish cock is much enhanced by his white face and crimson comb; and no one who has ever seen the splendid blue wattles of the male Tragopan pheasant distended in courtship can for a moment doubt that beauty is the object gained. From the foregoing facts we clearly see that the plumes and other ornaments of the males must be of the highest importance to them; and we further see that beauty is even sometimes more important than success in battle.

Chapter XIV

CHAPTER XIV. BIRDS—continued.

Choice exerted by the female—Length of courtship—Unpaired birds—Mental qualities and taste for the beautiful—Preference or antipathy shewn by the female for particular males—Variability of birds—Variations sometimes abrupt—Laws of variation—Formation of ocelli—Gradations of character—Case of Peacock, Argus pheasant, and Urosticte.

When the sexes differ in beauty or in the power of singing, or in producing what I have called instrumental music, it is almost invariably the male who surpasses the female. These qualities, as we have just seen, are evidently of high importance to the male. When they are gained for only a part of the year it is always before the breeding-season. It is the male alone who elaborately displays his varied attractions, and often performs strange antics on the ground or in the air, in the presence of the female. Each male drives away, or if he can, kills his rivals. Hence we may conclude that it is the object of the male to induce the female to pair with him, and for this purpose he tries to excite or charm her in various ways; and this is the opinion of all those who have carefully studied the habits of living birds. But there remains a question which has an all important bearing on sexual selection, namely, does every male of the same species excite and attract the female equally? Or does she exert a choice, and prefer certain males? This latter question can be answered in the affirmative by much direct and indirect evidence. It is far more difficult to decide what qualities determine the choice of the females; but here again we have some direct and indirect evidence that it is to a large extent the external attractions of the male; though no doubt his vigour, courage, and other mental qualities come into play. We will begin with the indirect evidence.

LENGTH OF COURTSHIP.

The lengthened period during which both sexes of certain birds meet day after day at an appointed place probably depends partly on the courtship being a prolonged affair, and partly on reiteration in the act of pairing. Thus in Germany and Scandinavia the balzen or leks of the black-cocks last from the middle of March, all through April into May. As many as forty or fifty, or even more birds congregate at the leks; and the same place is often frequented during successive years. The lek of the capercailzie lasts from the end of March to the middle or even end of May. In North America “the partridge dances” of the Tetrao phasianellus “last for a month or more.” Other kinds of grouse, both in North America and Eastern Siberia (1. Nordman describes (‘Bull. Soc. Imp. des Nat. Moscou,’ 1861, tom. xxxiv. p. 264) the balzen of Tetrao urogalloides in Amur Land. He estimated the number of birds assembled at above a hundred, not counting the females, which lie hid in the surrounding bushes. The noises uttered differ from those of T. urogallus.), follow nearly the same habits. The fowlers discover the hillocks where the ruffs congregate by the grass being trampled bare, and this shews that the same spot is long frequented. The Indians of Guiana are well acquainted with the cleared arenas, where they expect to find the beautiful cocks of the Rock; and the natives of New Guinea know the trees where from ten to twenty male birds of paradise in full plumage congregate. In this latter case it is not expressly stated that the females meet on the same trees, but the hunters, if not specially asked, would probably not mention their presence, as their skins are valueless. Small parties of an African weaver (Ploceus) congregate, during the breeding-season, and perform for hours their graceful evolutions. Large numbers of the Solitary snipe (Scolopax major) assemble during dusk in a morass; and the same place is frequented for the same purpose during successive years; here they may be seen running about “like so many large rats,” puffing out their feathers, flapping their wings, and uttering the strangest cries. (2. With respect to the assemblages of the above named grouse, see Brehm, ‘Thierleben,’ B. iv. s. 350; also L. Lloyd, ‘Game Birds of Sweden,’ 1867, pp. 19, 78. Richardson, ‘Fauna Bor. Americana: Birds,’ p. 362. References in regard to the assemblages of other birds have already been given. On Paradisea, see Wallace, in ‘Annals and Mag. of Nat. Hist.’ vol. xx. 1857, p. 412. On the snipe, Lloyd, ibid. p. 221.)

Some of the above birds,—the black-cock, capercailzie, pheasant-grouse, ruff, solitary snipe, and perhaps others,—are, as is believed, polygamists. With such birds it might have been thought that the stronger males would simply have driven away the weaker, and then at once have taken possession of as many females as possible; but if it be indispensable for the male to excite or please the female, we can understand the length of the courtship and the congregation of so many individuals of both sexes at the same spot. Certain strictly monogamous species likewise hold nuptial assemblages; this seems to be the case in Scandinavia with one of the ptarmigans, and their leks last from the middle of March to the middle of May. In Australia the lyre-bird (Menura superba) forms “small round hillocks,” and the M. Alberti scratches for itself shallow holes, or, as they are called by the natives, “corroborying places,” where it is believed both sexes assemble. The meetings of the M. superba are sometimes very large; and an account has lately been published (3. Quoted by Mr. T.W. Wood, in the ‘Student,’ April 1870, p. 125.) by a traveller, who heard in a valley beneath him, thickly covered with scrub, “a din which completely astonished” him; on crawling onwards he beheld, to his amazement, about one hundred and fifty of the magnificent lyre-cocks, “ranged in order of battle, and fighting with indescribable fury.” The bowers of the Bower-birds are the resort of both sexes during the breeding-season; and “here the males meet and contend with each other for the favours of the female, and here the latter assemble and coquet with the males.” With two of the genera, the same bower is resorted to during many years. (4. Gould, ‘Handbook to the Birds of Australia,’ vol. i. pp. 300, 308, 448, 451. On the ptarmigan, above alluded to, see Lloyd, ibid. p. 129.)

The common magpie (Corvus pica, Linn.), as I have been informed by the Rev. W. Darwin Fox, used to assemble from all parts of Delamere Forest, in order to celebrate the “great magpie marriage.” Some years ago these birds abounded in extraordinary numbers, so that a gamekeeper killed in one morning nineteen males, and another killed by a single shot seven birds at roost together. They then had the habit of assembling very early in the spring at particular spots, where they could be seen in flocks, chattering, sometimes fighting, bustling and flying about the trees. The whole affair was evidently considered by the birds as one of the highest importance. Shortly after the meeting they all separated, and were then observed by Mr. Fox and others to be paired for the season. In any district in which a species does not exist in large numbers, great assemblages cannot, of course, be held, and the same species may have different habits in different countries. For instance, I have heard of only one instance, from Mr. Wedderburn, of a regular assemblage of black game in Scotland, yet these assemblages are so well known in Germany and Scandinavia that they have received special names.

UNPAIRED BIRDS.

From the facts now given, we may conclude that the courtship of birds belonging to widely different groups, is often a prolonged, delicate, and troublesome affair. There is even reason to suspect, improbable as this will at first appear, that some males and females of the same species, inhabiting the same district, do not always please each other, and consequently do not pair. Many accounts have been published of either the male or female of a pair having been shot, and quickly replaced by another. This has been observed more frequently with the magpie than with any other bird, owing perhaps to its conspicuous appearance and nest. The illustrious Jenner states that in Wiltshire one of a pair was daily shot no less than seven times successively, “but all to no purpose, for the remaining magpie soon found another mate”; and the last pair reared their young. A new partner is generally found on the succeeding day; but Mr. Thompson gives the case of one being replaced on the evening of the same day. Even after the eggs are hatched, if one of the old birds is destroyed a mate will often be found; this occurred after an interval of two days, in a case recently observed by one of Sir J. Lubbock’s keepers. (5. On magpies, Jenner, in ‘Philosophical Transactions,’ 1824, p. 21. Macgillivray, ‘Hist. British Birds,’ vol. i. p. 570. Thompson, in ‘Annals and Magazine of Natural History,’ vol. viii. 1842, p. 494.) The first and most obvious conjecture is that male magpies must be much more numerous than females; and that in the above cases, as well as in many others which could be given, the males alone had been killed. This apparently holds good in some instances, for the gamekeepers in Delamere Forest assured Mr. Fox that the magpies and carrion-crows which they formerly killed in succession in large numbers near their nests, were all males; and they accounted for this fact by the males being easily killed whilst bringing food to the sitting females. Macgillivray, however, gives, on the authority of an excellent observer, an instance of three magpies successively killed on the same nest, which were all females; and another case of six magpies successively killed whilst sitting on the same eggs, which renders it probable that most of them were females; though, as I hear from Mr. Fox, the male will sit on the eggs when the female is killed.

Sir J. Lubbock’s gamekeeper has repeatedly shot, but how often he could not say, one of a pair of jays (Garrulus glandarius), and has never failed shortly afterwards to find the survivor re-matched. Mr. Fox, Mr. F. Bond, and others have shot one of a pair of carrion-crows (Corvus corone), but the nest was soon again tenanted by a pair. These birds are rather common; but the peregrine-falcon (Falco peregrinus) is rare, yet Mr. Thompson states that in Ireland “if either an old male or female be killed in the breeding-season (not an uncommon circumstance), another mate is found within a very few days, so that the eyries, notwithstanding such casualties, are sure to turn out their complement of young.” Mr. Jenner Weir has known the same thing with the peregrine-falcons at Beachy Head. The same observer informs me that three kestrels (Falco tinnunculus), all males, were killed one after the other whilst attending the same nest; two of these were in mature plumage, but the third was in the plumage of the previous year. Even with the rare golden eagle (Aquila chrysaetos), Mr. Birkbeck was assured by a trustworthy gamekeeper in Scotland, that if one is killed, another is soon found. So with the white owl (Strix flammea), “the survivor readily found a mate, and the mischief went on.”

White of Selborne, who gives the case of the owl, adds that he knew a man, who from believing that partridges when paired were disturbed by the males fighting, used to shoot them; and though he had widowed the same female several times, she always soon found a fresh partner. This same naturalist ordered the sparrows, which deprived the house-martins of their nests, to be shot; but the one which was left, “be it cock or hen, presently procured a mate, and so for several times following.” I could add analogous cases relating to the chaffinch, nightingale, and redstart. With respect to the latter bird (Phoenicura ruticilla), a writer expresses much surprise how the sitting female could so soon have given effectual notice that she was a widow, for the species was not common in the neighbourhood. Mr. Jenner Weir has mentioned to me a nearly similar case; at Blackheath he never sees or hears the note of the wild bullfinch, yet when one of his caged males has died, a wild one in the course of a few days has generally come and perched near the widowed female, whose call-note is not loud. I will give only one other fact, on the authority of this same observer; one of a pair of starlings (Sturnus vulgaris) was shot in the morning; by noon a new mate was found; this was again shot, but before night the pair was complete; so that the disconsolate widow or widower was thrice consoled during the same day. Mr. Engleheart also informs me that he used during several years to shoot one of a pair of starlings which built in a hole in a house at Blackheath; but the loss was always immediately repaired. During one season he kept an account, and found that he had shot thirty-five birds from the same nest; these consisted of both males and females, but in what proportion he could not say: nevertheless, after all this destruction, a brood was reared. (6. On the peregrine falcon, see Thompson, ‘Nat. Hist. of Ireland: Birds,’ vol. i. 1849, p. 39. On owls, sparrows, and partridges, see White, ‘Nat. Hist. of Selborne,’ edit. of 1825, vol. i. p. 139. On the Phoenicura, see Loudon’s ‘Mag. of Nat. Hist.’ vol. vii. 1834, p. 245. Brehm (‘Thierleben,’ B. iv. s. 991) also alludes to cases of birds thrice mated during the same day.)

These facts well deserve attention. How is it that there are birds enough ready to replace immediately a lost mate of either sex? Magpies, jays, carrion-crows, partridges, and some other birds, are always seen during the spring in pairs, and never by themselves; and these offer at first sight the most perplexing cases. But birds of the same sex, although of course not truly paired, sometimes live in pairs or in small parties, as is known to be the case with pigeons and partridges. Birds also sometimes live in triplets, as has been observed with starlings, carrion-crows, parrots, and partridges. With partridges two females have been known to live with one male, and two males with one female. In all such cases it is probable that the union would be easily broken; and one of the three would readily pair with a widow or widower. The males of certain birds may occasionally be heard pouring forth their love-song long after the proper time, shewing that they have either lost or never gained a mate. Death from accident or disease of one of a pair would leave the other free and single; and there is reason to believe that female birds during the breeding-season are especially liable to premature death. Again, birds which have had their nests destroyed, or barren pairs, or retarded individuals, would easily be induced to desert their mates, and would probably be glad to take what share they could of the pleasures and duties of rearing offspring although not their own. (7. See White (‘Nat. Hist. of Selborne,’ 1825, vol. i. p. 140) on the existence, early in the season, of small coveys of male partridges, of which fact I have heard other instances. See Jenner, on the retarded state of the generative organs in certain birds, in ‘Phil. Transact.’ 1824. In regard to birds living in triplets, I owe to Mr. Jenner Weir the cases of the starlings and parrots, and to Mr. Fox, of partridges; on carrion-crows, see the ‘Field,’ 1868, p. 415. On various male birds singing after the proper period, see Rev. L. Jenyns, ‘Observations in Natural History,’ 1846, p. 87.) Such contingencies as these probably explain most of the foregoing cases. (8. The following case has been given (‘The Times,’ Aug. 6, 1868) by the Rev. F.O. Morris, on the authority of the Hon. and Rev. O.W. Forester. “The gamekeeper here found a hawk’s nest this year, with five young ones on it. He took four and killed them, but left one with its wings clipped as a decoy to destroy the old ones by. They were both shot next day, in the act of feeding the young one, and the keeper thought it was done with. The next day he came again and found two other charitable hawks, who had come with an adopted feeling to succour the orphan. These two he killed, and then left the nest. On returning afterwards he found two more charitable individuals on the same errand of mercy. One of these he killed; the other he also shot, but could not find. No more came on the like fruitless errand.”) Nevertheless, it is a strange fact that within the same district, during the height of the breeding-season, there should be so many males and females always ready to repair the loss of a mated bird. Why do not such spare birds immediately pair together? Have we not some reason to suspect, and the suspicion has occurred to Mr. Jenner Weir, that as the courtship of birds appears to be in many cases prolonged and tedious, so it occasionally happens that certain males and females do not succeed, during the proper season, in exciting each other’s love, and consequently do not pair? This suspicion will appear somewhat less improbable after we have seen what strong antipathies and preferences female birds occasionally evince towards particular males.

MENTAL QUALITIES OF BIRDS, AND THEIR TASTE FOR THE BEAUTIFUL.

Before we further discuss the question whether the females select the more attractive males or accept the first whom they may encounter, it will be advisable briefly to consider the mental powers of birds. Their reason is generally, and perhaps justly, ranked as low; yet some facts could be given leading to an opposite conclusion. (9. I am indebted to Prof. Newton for the following passage from Mr. Adam’s ‘Travels of a Naturalist,’ 1870, p. 278. Speaking of Japanese nut-hatches in confinement, he says: “Instead of the more yielding fruit of the yew, which is the usual food of the nut-hatch of Japan, at one time I substituted hard hazel-nuts. As the bird was unable to crack them, he placed them one by one in his water-glass, evidently with the notion that they would in time become softer—an interesting proof of intelligence on the part of these birds.”) Low powers of reasoning, however, are compatible, as we see with mankind, with strong affections, acute perception, and a taste for the beautiful; and it is with these latter qualities that we are here concerned. It has often been said that parrots become so deeply attached to each other that when one dies the other pines for a long time; but Mr. Jenner Weir thinks that with most birds the strength of their affection has been much exaggerated. Nevertheless when one of a pair in a state of nature has been shot, the survivor has been heard for days afterwards uttering a plaintive call; and Mr. St. John gives various facts proving the attachment of mated birds. (10. ‘A Tour in Sutherlandshire,’ vol. i. 1849, p. 185. Dr. Buller says (‘Birds of New Zealand,’ 1872, p. 56) that a male King Lory was killed; and the female “fretted and moped, refused her food, and died of a broken heart.”) Mr. Bennett relates (11. ‘Wanderings in New South Wales,’ vol. ii. 1834, p. 62.) that in China after a drake of the beautiful mandarin Teal had been stolen, the duck remained disconsolate, though sedulously courted by another mandarin drake, who displayed before her all his charms. After an interval of three weeks the stolen drake was recovered, and instantly the pair recognised each other with extreme joy. On the other hand, starlings, as we have seen, may be consoled thrice in the same day for the loss of their mates. Pigeons have such excellent local memories, that they have been known to return to their former homes after an interval of nine months, yet, as I hear from Mr. Harrison Weir, if a pair which naturally would remain mated for life be separated for a few weeks during the winter, and afterwards matched with other birds, the two when brought together again, rarely, if ever, recognise each other.

Birds sometimes exhibit benevolent feelings; they will feed the deserted young ones even of distinct species, but this perhaps ought to be considered as a mistaken instinct. They will feed, as shewn in an earlier part of this work, adult birds of their own species which have become blind. Mr. Buxton gives a curious account of a parrot which took care of a frost-bitten and crippled bird of a distinct species, cleansed her feathers, and defended her from the attacks of the other parrots which roamed freely about his garden. It is a still more curious fact that these birds apparently evince some sympathy for the pleasures of their fellows. When a pair of cockatoos made a nest in an acacia tree, “it was ridiculous to see the extravagant interest taken in the matter by the others of the same species.” These parrots, also, evinced unbounded curiosity, and clearly had “the idea of property and possession.” (12. ‘Acclimatization of Parrots,’ by C. Buxton, M.P., ‘Annals and Mag. of Nat. Hist.’ Nov. 1868, p. 381.) They have good memories, for in the Zoological Gardens they have plainly recognised their former masters after an interval of some months.

Birds possess acute powers of observation. Every mated bird, of course, recognises its fellow. Audubon states that a certain number of mocking-thrushes (Mimus polyglottus) remain all the year round in Louisiana, whilst others migrate to the Eastern States; these latter, on their return, are instantly recognised, and always attacked, by their southern brethren. Birds under confinement distinguish different persons, as is proved by the strong and permanent antipathy or affection which they shew, without any apparent cause, towards certain individuals. I have heard of numerous instances with jays, partridges, canaries, and especially bullfinches. Mr. Hussey has described in how extraordinary a manner a tamed partridge recognised everybody: and its likes and dislikes were very strong. This bird seemed “fond of gay colours, and no new gown or cap could be put on without catching his attention.” (13. The ‘Zoologist,’ 1847-48, p. 1602.) Mr. Hewitt has described the habits of some ducks (recently descended from wild birds), which, at the approach of a strange dog or cat, would rush headlong into the water, and exhaust themselves in their attempts to escape; but they knew Mr. Hewitt’s own dogs and cats so well that they would lie down and bask in the sun close to them. They always moved away from a strange man, and so they would from the lady who attended them if she made any great change in her dress. Audubon relates that he reared and tamed a wild turkey which always ran away from any strange dog; this bird escaped into the woods, and some days afterwards Audubon saw, as he thought, a wild turkey, and made his dog chase it; but, to his astonishment, the bird did not run away, and the dog, when he came up, did not attack the bird, for they mutually recognised each other as old friends. (14. Hewitt on wild ducks, ‘Journal of Horticulture,’ Jan. 13, 1863, p. 39. Audubon on the wild turkey, ‘Ornithological Biography,’ vol. i. p. 14. On the mocking-thrush, ibid. vol. i. p. 110.)

Mr. Jenner Weir is convinced that birds pay particular attention to the colours of other birds, sometimes out of jealousy, and sometimes as a sign of kinship. Thus he turned a reed-bunting (Emberiza schoeniculus), which had acquired its black head-dress, into his aviary, and the new-comer was not noticed by any bird, except by a bullfinch, which is likewise black-headed. This bullfinch was a very quiet bird, and had never before quarrelled with any of its comrades, including another reed-bunting, which had not as yet become black-headed: but the reed-bunting with a black head was so unmercifully treated that it had to be removed. Spiza cyanea, during the breeding-season, is of a bright blue colour; and though generally peaceable, it attacked S. ciris, which has only the head blue, and completely scalped the unfortunate bird. Mr. Weir was also obliged to turn out a robin, as it fiercely attacked all the birds in his aviary with any red in their plumage, but no other kinds; it actually killed a red-breasted crossbill, and nearly killed a goldfinch. On the other hand, he has observed that some birds, when first introduced, fly towards the species which resemble them most in colour, and settle by their sides.

As male birds display their fine plumage and other ornaments with so much care before the females, it is obviously probable that these appreciate the beauty of their suitors. It is, however, difficult to obtain direct evidence of their capacity to appreciate beauty. When birds gaze at themselves in a looking-glass (of which many instances have been recorded) we cannot feel sure that it is not from jealousy of a supposed rival, though this is not the conclusion of some observers. In other cases it is difficult to distinguish between mere curiosity and admiration. It is perhaps the former feeling which, as stated by Lord Lilford (15. The ‘Ibis,’ vol. ii. 1860, p. 344.), attracts the ruff towards any bright object, so that, in the Ionian Islands, “it will dart down to a bright-coloured handkerchief, regardless of repeated shots.” The common lark is drawn down from the sky, and is caught in large numbers, by a small mirror made to move and glitter in the sun. Is it admiration or curiosity which leads the magpie, raven, and some other birds to steal and secrete bright objects, such as silver articles or jewels?

Mr. Gould states that certain humming-birds decorate the outsides of their nests “with the utmost taste; they instinctively fasten thereon beautiful pieces of flat lichen, the larger pieces in the middle, and the smaller on the part attached to the branch. Now and then a pretty feather is intertwined or fastened to the outer sides, the stem being always so placed that the feather stands out beyond the surface.” The best evidence, however, of a taste for the beautiful is afforded by the three genera of Australian bower-birds already mentioned. Their bowers (Fig. 46), where the sexes congregate and play strange antics, are variously constructed, but what most concerns us is, that they are decorated by the several species in a different manner. The Satin bower-bird collects gaily-coloured articles, such as the blue tail-feathers of parrakeets, bleached bones and shells, which it sticks between the twigs or arranges at the entrance. Mr. Gould found in one bower a neatly-worked stone tomahawk and a slip of blue cotton, evidently procured from a native encampment. These objects are continually re-arranged, and carried about by the birds whilst at play. The bower of the Spotted bower-bird “is beautifully lined with tall grasses, so disposed that the heads nearly meet, and the decorations are very profuse.” Round stones are used to keep the grass-stems in their proper places, and to make divergent paths leading to the bower. The stones and shells are often brought from a great distance. The Regent bird, as described by Mr. Ramsay, ornaments its short bower with bleached land-shells belonging to five or six species, and with “berries of various colours, blue, red, and black, which give it when fresh a very pretty appearance. Besides these there were several newly-picked leaves and young shoots of a pinkish colour, the whole showing a decided taste for the beautiful.” Well may Mr. Gould say that “these highly decorated halls of assembly must be regarded as the most wonderful instances of bird-architecture yet discovered;” and the taste, as we see, of the several species certainly differs. (16. On the ornamented nests of humming-birds, Gould, ‘Introduction to the Trochilidae,’ 1861, p. 19. On the bower-birds, Gould, ‘Handbook to the Birds of Australia,’ 1865, vol. i. pp. 444-461. Ramsay, in the ‘Ibis,’ 1867, p. 456.)

PREFERENCE FOR A PARTICULAR MALES BY THE FEMALES.

Having made these preliminary remarks on the discrimination and taste of birds, I will give all the facts known to me which bear on the preference shewn by the female for particular males. It is certain that distinct species of birds occasionally pair in a state of nature and produce hybrids. Many instances could be given: thus Macgillivray relates how a male blackbird and female thrush “fell in love with each other,” and produced offspring. (17. ‘History of Brit. Birds,’ vol. ii. p. 92.) Several years ago eighteen cases had been recorded of the occurrence in Great Britain of hybrids between the black grouse and pheasant (18. ‘Zoologist,’ 1853-1854, p. 3946.); but most of these cases may perhaps be accounted for by solitary birds not finding one of their own species to pair with. With other birds, as Mr. Jenner Weir has reason to believe, hybrids are sometimes the result of the casual intercourse of birds building in close proximity. But these remarks do not apply to the many recorded instances of tamed or domestic birds, belonging to distinct species, which have become absolutely fascinated with each other, although living with their own species. Thus Waterton (19. Waterton, ‘Essays on Nat. Hist.’ 2nd series, pp. 42 and 117. For the following statements see on the wigeon, ‘Loudon’s Mag. of Nat. Hist.’ vol. ix. p. 616; L. Lloyd, ‘Scandinavian Adventures,’ vol. i. 1854, p. 452. Dixon, ‘Ornamental and Domestic Poultry,’ p. 137; Hewitt, in ‘Journal of Horticulture,’ Jan. 13, 1863, p. 40; Bechstein, ‘Stubenvögel,’ 1840, s. 230. Mr. J. Jenner Weir has lately given me an analogous case with ducks of two species.) states that out of a flock of twenty-three Canada geese, a female paired with a solitary Bernicle gander, although so different in appearance and size; and they produced hybrid offspring. A male wigeon (Mareca penelope), living with females of the same species, has been known to pair with a pintail duck, Querquedula acuta. Lloyd describes the remarkable attachment between a shield-drake (Tadorna vulpanser) and a common duck. Many additional instances could be given; and the Rev. E.S. Dixon remarks that “those who have kept many different species of geese together well know what unaccountable attachments they are frequently forming, and that they are quite as likely to pair and rear young with individuals of a race (species) apparently the most alien to themselves as with their own stock.”

The Rev. W.D. Fox informs me that he possessed at the same time a pair of Chinese geese (Anser cygnoides), and a common gander with three geese. The two lots kept quite separate, until the Chinese gander seduced one of the common geese to live with him. Moreover, of the young birds hatched from the eggs of the common geese, only four were pure, the other eighteen proving hybrids; so that the Chinese gander seems to have had prepotent charms over the common gander. I will give only one other case; Mr. Hewitt states that a wild duck, reared in captivity, “after breeding a couple of seasons with her own mallard, at once shook him off on my placing a male Pintail on the water. It was evidently a case of love at first sight, for she swam about the new-comer caressingly, though he appeared evidently alarmed and averse to her overtures of affection. From that hour she forgot her old partner. Winter passed by, and the next spring the pintail seemed to have become a convert to her blandishments, for they nested and produced seven or eight young ones.”

What the charm may have been in these several cases, beyond mere novelty, we cannot even conjecture. Colour, however, sometimes comes into play; for in order to raise hybrids from the siskin (Fringilla spinus) and the canary, it is much the best plan, according to Bechstein, to place birds of the same tint together. Mr. Jenner Weir turned a female canary into his aviary, where there were male linnets, goldfinches, siskins, greenfinches, chaffinches, and other birds, in order to see which she would choose; but there never was any doubt, and the greenfinch carried the day. They paired and produced hybrid offspring.

The fact of the female preferring to pair with one male rather than with another of the same species is not so likely to excite attention, as when this occurs, as we have just seen, between distinct species. The former cases can best be observed with domesticated or confined birds; but these are often pampered by high feeding, and sometimes have their instincts vitiated to an extreme degree. Of this latter fact I could give sufficient proofs with pigeons, and especially with fowls, but they cannot be here related. Vitiated instincts may also account for some of the hybrid unions above mentioned; but in many of these cases the birds were allowed to range freely over large ponds, and there is no reason to suppose that they were unnaturally stimulated by high feeding.

With respect to birds in a state of nature, the first and most obvious supposition which will occur to every one is that the female at the proper season accepts the first male whom she may encounter; but she has at least the opportunity for exerting a choice, as she is almost invariably pursued by many males. Audubon—and we must remember that he spent a long life in prowling about the forests of the United States and observing the birds—does not doubt that the female deliberately chooses her mate; thus, speaking of a woodpecker, he says the hen is followed by half-a-dozen gay suitors, who continue performing strange antics, “until a marked preference is shewn for one.” The female of the red-winged starling (Agelaeus phoeniceus) is likewise pursued by several males, “until, becoming fatigued, she alights, receives their addresses, and soon makes a choice.” He describes also how several male night-jars repeatedly plunge through the air with astonishing rapidity, suddenly turning, and thus making a singular noise; “but no sooner has the female made her choice than the other males are driven away.” With one of the vultures (Cathartes aura) of the United States, parties of eight, ten, or more males and females assemble on fallen logs, “exhibiting the strongest desire to please mutually,” and after many caresses, each male leads off his partner on the wing. Audubon likewise carefully observed the wild flocks of Canada geese (Anser canadensis), and gives a graphic description of their love-antics; he says that the birds which had been previously mated “renewed their courtship as early as the month of January, while the others would be contending or coquetting for hours every day, until all seemed satisfied with the choice they had made, after which, although they remained together, any person could easily perceive that they were careful to keep in pairs. I have observed also that the older the birds the shorter were the preliminaries of their courtship. The bachelors and old maids whether in regret, or not caring to be disturbed by the bustle, quietly moved aside and lay down at some distance from the rest.” (20. Audubon, ‘Ornithological Biography,’ vol. i. pp. 191, 349; vol. ii. pp. 42, 275; vol. iii. p. 2.) Many similar statements with respect to other birds could be cited from this same observer.

Turning now to domesticated and confined birds, I will commence by giving what little I have learnt respecting the courtship of fowls. I have received long letters on this subject from Messrs. Hewitt and Tegetmeier, and almost an essay from the late Mr. Brent. It will be admitted by every one that these gentlemen, so well known from their published works, are careful and experienced observers. They do not believe that the females prefer certain males on account of the beauty of their plumage; but some allowance must be made for the artificial state under which these birds have long been kept. Mr. Tegetmeier is convinced that a gamecock, though disfigured by being dubbed and with his hackles trimmed, would be accepted as readily as a male retaining all his natural ornaments. Mr. Brent, however, admits that the beauty of the male probably aids in exciting the female; and her acquiescence is necessary. Mr. Hewitt is convinced that the union is by no means left to mere chance, for the female almost invariably prefers the most vigorous, defiant, and mettlesome male; hence it is almost useless, as he remarks, “to attempt true breeding if a game-cock in good health and condition runs the locality, for almost every hen on leaving the roosting-place will resort to the game-cock, even though that bird may not actually drive away the male of her own variety.” Under ordinary circumstances the males and females of the fowl seem to come to a mutual understanding by means of certain gestures, described to me by Mr. Brent. But hens will often avoid the officious attentions of young males. Old hens, and hens of a pugnacious disposition, as the same writer informs me, dislike strange males, and will not yield until well beaten into compliance. Ferguson, however, describes how a quarrelsome hen was subdued by the gentle courtship of a Shanghai cock. (21. ‘Rare and Prize Poultry,’ 1854, p. 27.)

There is reason to believe that pigeons of both sexes prefer pairing with birds of the same breed; and dovecot-pigeons dislike all the highly improved breeds. (22. ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 103.) Mr. Harrison Weir has lately heard from a trustworthy observer, who keeps blue pigeons, that these drive away all other coloured varieties, such as white, red, and yellow; and from another observer, that a female dun carrier could not, after repeated trials, be matched with a black male, but immediately paired with a dun. Again, Mr. Tegetmeier had a female blue turbit that obstinately refused to pair with two males of the same breed, which were successively shut up with her for weeks; but on being let out she would have immediately accepted the first blue dragon that offered. As she was a valuable bird, she was then shut up for many weeks with a silver (i.e., very pale blue) male, and at last mated with him. Nevertheless, as a general rule, colour appears to have little influence on the pairing of pigeons. Mr. Tegetmeier, at my request, stained some of his birds with magenta, but they were not much noticed by the others.

Female pigeons occasionally feel a strong antipathy towards certain males, without any assignable cause. Thus MM. Boitard and Corbie, whose experience extended over forty-five years, state: “Quand une femelle éprouve de l’antipathie pour un mâle avec lequel on veut l’accoupler, malgré tous les feux de l’amour, malgré l’alpiste et le chenevis dont on la nourrit pour augmenter son ardeur, malgré un emprisonnement de six mois et même d’un an, elle refuse constamment ses caresses; les avances empressées, les agaceries, les tournoiemens, les tendres roucoulemens, rien ne peut lui plaire ni l’émouvoir; gonflée, boudeuse, blottie dans un coin de sa prison, elle n’en sort que pour boire et manger, ou pour repousser avec une espèce de rage des caresses devenues trop pressantes.” (23. Boitard and Corbie, ‘Les Pigeons,’ etc., 1824, p. 12. Prosper Lucas (‘Traité de l’Héréd. Nat.’ tom. ii. 1850, p. 296) has himself observed nearly similar facts with pigeons.) On the other hand, Mr. Harrison Weir has himself observed, and has heard from several breeders, that a female pigeon will occasionally take a strong fancy for a particular male, and will desert her own mate for him. Some females, according to another experienced observer, Riedel (24. Die Taubenzucht, 1824, s. 86.), are of a profligate disposition, and prefer almost any stranger to their own mate. Some amorous males, called by our English fanciers “gay birds,” are so successful in their gallantries, that, as Mr. H. Weir informs me, they must be shut up on account of the mischief which they cause.

Wild turkeys in the United States, according to Audubon, “sometimes pay their addresses to the domesticated females, and are generally received by them with great pleasure.” So that these females apparently prefer the wild to their own males. (25. ‘Ornithological Biography,’ vol. i. p. 13. See to the same effect, Dr. Bryant, in Allen’s ‘Mammals and Birds of Florida,’ p. 344.)

Here is a more curious case. Sir R. Heron during many years kept an account of the habits of the peafowl, which he bred in large numbers. He states that “the hens have frequently great preference to a particular peafowl. They were all so fond of an old pied cock, that one year, when he was confined, though still in view, they were constantly assembled close to the trellice-walls of his prison, and would not suffer a japanned peacock to touch them. On his being let out in the autumn, the oldest of the hens instantly courted him and was successful in her courtship. The next year he was shut up in a stable, and then the hens all courted his rival.” (26. ‘Proceedings, Zoological Society,’ 1835, p. 54. The japanned peacock is considered by Mr. Sclater as a distinct species, and has been named Pavo nigripennis; but the evidence seems to me to show that it is only a variety.) This rival was a japanned or black-winged peacock, to our eyes a more beautiful bird than the common kind.

Lichtenstein, who was a good observer and had excellent opportunities of observation at the Cape of Good Hope, assured Rudolphi that the female widow-bird (Chera progne) disowns the male when robbed of the long tail-feathers with which he is ornamented during the breeding-season. I presume that this observation must have been made on birds under confinement. (27. Rudolphi, ‘Beiträge zur Anthropologie,’ 1812, s. 184.) Here is an analogous case; Dr. Jaeger (28. ‘Die Darwin’sche Theorie, und ihre Stellung zu Moral und Religion,’ 1869, s. 59.), director of the Zoological Gardens of Vienna, states that a male silver-pheasant, who had been triumphant over all other males and was the accepted lover of the females, had his ornamental plumage spoiled. He was then immediately superseded by a rival, who got the upper hand and afterwards led the flock.

It is a remarkable fact, as shewing how important colour is in the courtship of birds, that Mr. Boardman, a well-known collector and observer of birds for many years in the Northern United States, has never in his large experience seen an albino paired with another bird; yet he has had opportunities of observing many albinos belonging to several species. (29. This statement is given by Mr. A. Leith Adams, in his ‘Field and Forest Rambles,’ 1873, p. 76, and accords with his own experience.) It can hardly be maintained that albinos in a state of nature are incapable of breeding, as they can be raised with the greatest facility under confinement. It appears, therefore, that we must attribute the fact that they do not pair to their rejection by their normally coloured comrades.

Female birds not only exert a choice, but in some few cases they court the male, or even fight together for his possession. Sir R. Heron states that with peafowl, the first advances are always made by the female; something of the same kind takes place, according to Audubon, with the older females of the wild turkey. With the capercailzie, the females flit round the male whilst he is parading at one of the places of assemblage, and solicit his attention. (30. In regard to peafowl, see Sir R. Heron, ‘Proc. Zoolog. Soc.’ 1835, p. 54, and the Rev. E.S. Dixon, ‘Ornamental Poultry,’ 1848, p. 8. For the turkey, Audubon, ibid. p. 4. For the capercailzie, Lloyd, ‘Game Birds of Sweden,’ 1867, p. 23.) We have seen that a tame wild-duck seduced an unwilling pintail drake after a long courtship. Mr. Bartlett believes that the Lophophorus, like many other gallinaceous birds, is naturally polygamous, but two females cannot be placed in the same cage with a male, as they fight so much together. The following instance of rivalry is more surprising as it relates to bullfinches, which usually pair for life. Mr. Jenner Weir introduced a dull-coloured and ugly female into his aviary, and she immediately attacked another mated female so unmercifully that the latter had to be separated. The new female did all the courtship, and was at last successful, for she paired with the male; but after a time she met with a just retribution, for, ceasing to be pugnacious, she was replaced by the old female, and the male then deserted his new and returned to his old love.

In all ordinary cases the male is so eager that he will accept any female, and does not, as far as we can judge, prefer one to the other; but, as we shall hereafter see, exceptions to this rule apparently occur in some few groups. With domesticated birds, I have heard of only one case of males shewing any preference for certain females, namely, that of the domestic cock, who, according to the high authority of Mr. Hewitt, prefers the younger to the older hens. On the other hand, in effecting hybrid unions between the male pheasant and common hens, Mr. Hewitt is convinced that the pheasant invariably prefers the older birds. He does not appear to be in the least influenced by their colour; but “is most capricious in his attachments” (31. Mr. Hewitt, quoted in Tegetmeier’s ‘Poultry Book,’ 1866, p. 165.): from some inexplicable cause he shews the most determined aversion to certain hens, which no care on the part of the breeder can overcome. Mr. Hewitt informs me that some hens are quite unattractive even to the males of their own species, so that they may be kept with several cocks during a whole season, and not one egg out of forty or fifty will prove fertile. On the other hand, with the long-tailed duck (Harelda glacialis), “it has been remarked,” says M. Ekstrom, “that certain females are much more courted than the rest. Frequently, indeed, one sees an individual surrounded by six or eight amorous males.” Whether this statement is credible, I know not; but the native sportsmen shoot these females in order to stuff them as decoys. (32. Quoted in Lloyd’s ‘Game Birds of Sweden,’ p. 345.)

With respect to female birds feeling a preference for particular males, we must bear in mind that we can judge of choice being exerted only by analogy. If an inhabitant of another planet were to behold a number of young rustics at a fair courting a pretty girl, and quarrelling about her like birds at one of their places of assemblage, he would, by the eagerness of the wooers to please her and to display their finery, infer that she had the power of choice. Now with birds the evidence stands thus: they have acute powers of observation, and they seem to have some taste for the beautiful both in colour and sound. It is certain that the females occasionally exhibit, from unknown causes, the strongest antipathies and preferences for particular males. When the sexes differ in colour or in other ornaments the males with rare exceptions are the more decorated, either permanently or temporarily during the breeding-season. They sedulously display their various ornaments, exert their voices, and perform strange antics in the presence of the females. Even well-armed males, who, it might be thought, would altogether depend for success on the law of battle, are in most cases highly ornamented; and their ornaments have been acquired at the expense of some loss of power. In other cases ornaments have been acquired, at the cost of increased risk from birds and beasts of prey. With various species many individuals of both sexes congregate at the same spot, and their courtship is a prolonged affair. There is even reason to suspect that the males and females within the same district do not always succeed in pleasing each other and pairing.

What then are we to conclude from these facts and considerations? Does the male parade his charms with so much pomp and rivalry for no purpose? Are we not justified in believing that the female exerts a choice, and that she receives the addresses of the male who pleases her most? It is not probable that she consciously deliberates; but she is most excited or attracted by the most beautiful, or melodious, or gallant males. Nor need it be supposed that the female studies each stripe or spot of colour; that the peahen, for instance, admires each detail in the gorgeous train of the peacock—she is probably struck only by the general effect. Nevertheless, after hearing how carefully the male Argus pheasant displays his elegant primary wing-feathers, and erects his ocellated plumes in the right position for their full effect; or again, how the male goldfinch alternately displays his gold-bespangled wings, we ought not to feel too sure that the female does not attend to each detail of beauty. We can judge, as already remarked, of choice being exerted, only from analogy; and the mental powers of birds do not differ fundamentally from ours. From these various considerations we may conclude that the pairing of birds is not left to chance; but that those males, which are best able by their various charms to please or excite the female, are under ordinary circumstances accepted. If this be admitted, there is not much difficulty in understanding how male birds have gradually acquired their ornamental characters. All animals present individual differences, and as man can modify his domesticated birds by selecting the individuals which appear to him the most beautiful, so the habitual or even occasional preference by the female of the more attractive males would almost certainly lead to their modification; and such modifications might in the course of time be augmented to almost any extent, compatible with the existence of the species.

VARIABILITY OF BIRDS, AND ESPECIALLY OF THEIR SECONDARY SEXUAL CHARACTERS.

Variability and inheritance are the foundations for the work of selection. That domesticated birds have varied greatly, their variations being inherited, is certain. That birds in a state of nature have been modified into distinct races is now universally admitted. (33. According to Dr. Blasius (‘Ibis,’ vol. ii. 1860, p. 297), there are 425 indubitable species of birds which breed in Europe, besides sixty forms, which are frequently regarded as distinct species. Of the latter, Blasius thinks that only ten are really doubtful, and that the other fifty ought to be united with their nearest allies; but this shews that there must be a considerable amount of variation with some of our European birds. It is also an unsettled point with naturalists, whether several North American birds ought to be ranked as specifically distinct from the corresponding European species. So again many North American forms which until lately were named as distinct species, are now considered to be local races.) Variations may be divided into two classes; those which appear to our ignorance to arise spontaneously, and those which are directly related to the surrounding conditions, so that all or nearly all the individuals of the same species are similarly modified. Cases of the latter kind have recently been observed with care by Mr. J.A. Allen (34. ‘Mammals and Birds of East Florida,’ also an ‘Ornithological Reconnaissance of Kansas,’ etc. Notwithstanding the influence of climate on the colours of birds, it is difficult to account for the dull or dark tints of almost all the species inhabiting certain countries, for instance, the Galapagos Islands under the equator, the wide temperate plains of Patagonia, and, as it appears, Egypt (see Mr. Hartshorne in the ‘American Naturalist,’ 1873, p. 747). These countries are open, and afford little shelter to birds; but it seems doubtful whether the absence of brightly coloured species can be explained on the principle of protection, for on the Pampas, which are equally open, though covered by green grass, and where the birds would be equally exposed to danger, many brilliant and conspicuously coloured species are common. I have sometimes speculated whether the prevailing dull tints of the scenery in the above named countries may not have affected the appreciation of bright colours by the birds inhabiting them.), who shews that in the United States many species of birds gradually become more strongly coloured in proceeding southward, and more lightly coloured in proceeding westward to the arid plains of the interior. Both sexes seem generally to be affected in a like manner, but sometimes one sex more than the other. This result is not incompatible with the belief that the colours of birds are mainly due to the accumulation of successive variations through sexual selection; for even after the sexes have been greatly differentiated, climate might produce an equal effect on both sexes, or a greater effect on one sex than on the other, owing to some constitutional difference.

Individual differences between the members of the same species are admitted by every one to occur under a state of nature. Sudden and strongly marked variations are rare; it is also doubtful whether if beneficial they would often be preserved through selection and transmitted to succeeding generations. (35. ‘Origin of Species’ fifth edit. 1869, p.104. I had always perceived, that rare and strongly-marked deviations of structure, deserving to be called monstrosities, could seldom be preserved through natural selection, and that the preservation of even highly-beneficial variations would depend to a certain extent on chance. I had also fully appreciated the importance of mere individual differences, and this led me to insist so strongly on the importance of that unconscious form of selection by man, which follows from the preservation of the most valued individuals of each breed, without any intention on his part to modify the characters of the breed. But until I read an able article in the ‘North British Review’ (March 1867, p. 289, et seq.), which has been of more use to me than any other Review, I did not see how great the chances were against the preservation of variations, whether slight or strongly pronounced, occurring only in single individuals.) Nevertheless, it may be worth while to give the few cases which I have been able to collect, relating chiefly to colour,—simple albinism and melanism being excluded. Mr. Gould is well known to admit the existence of few varieties, for he esteems very slight differences as specific; yet he states (36. ‘Introduction to the Trochlidae,’ p. 102.) that near Bogota certain humming-birds belonging to the genus Cynanthus are divided into two or three races or varieties, which differ from each other in the colouring of the tail—“some having the whole of the feathers blue, while others have the eight central ones tipped with beautiful green.” It does not appear that intermediate gradations have been observed in this or the following cases. In the males alone of one of the Australian parrakeets “the thighs in some are scarlet, in others grass-green.” In another parrakeet of the same country “some individuals have the band across the wing-coverts bright-yellow, while in others the same part is tinged with red.” (37. Gould, ‘Handbook to Birds of Australia,’ vol. ii. pp. 32 and 68.) In the United States some few of the males of the scarlet tanager (Tanagra rubra) have “a beautiful transverse band of glowing red on the smaller wing-coverts” (38. Audubon, ‘Ornithological Biography,’ 1838, vol. iv. p. 389.); but this variation seems to be somewhat rare, so that its preservation through sexual selection would follow only under usually favourable circumstances. In Bengal the Honey buzzard (Pernis cristata) has either a small rudimental crest on its head, or none at all: so slight a difference, however, would not have been worth notice, had not this same species possessed in Southern India a well-marked occipital crest formed of several graduated feathers.” (39. Jerdon, ‘Birds of India,’ vol. i. p. 108; and Mr. Blyth, in ‘Land and Water,’ 1868, p. 381.)

The following case is in some respects more interesting. A pied variety of the raven, with the head, breast, abdomen, and parts of the wings and tail-feathers white, is confined to the Feroe Islands. It is not very rare there, for Graba saw during his visit from eight to ten living specimens. Although the characters of this variety are not quite constant, yet it has been named by several distinguished ornithologists as a distinct species. The fact of the pied birds being pursued and persecuted with much clamour by the other ravens of the island was the chief cause which led Brunnich to conclude that they were specifically distinct; but this is now known to be an error. (40. Graba, ‘Tagebuch Reise nach Faro,’ 1830, ss. 51-54. Macgillivray, ‘History of British Birds,’ vol. iii. p. 745, ‘Ibis,’ vol. v. 1863, p. 469.) This case seems analogous to that lately given of albino birds not pairing from being rejected by their comrades.

In various parts of the northern seas a remarkable variety of the common Guillemot (Uria troile) is found; and in Feroe, one out of every five birds, according to Graba’s estimation, presents this variation. It is characterised (41. Graba, ibid. s. 54. Macgillivray, ibid. vol. v. p. 327.) by a pure white ring round the eye, with a curved narrow white line, an inch and a half in length, extending back from the ring. This conspicuous character has caused the bird to be ranked by several ornithologists as a distinct species under the name of U. lacrymans, but it is now known to be merely a variety. It often pairs with the common kind, yet intermediate gradations have never been seen; nor is this surprising, for variations which appear suddenly, are often, as I have elsewhere shewn (42. ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 92.), transmitted either unaltered or not at all. We thus see that two distinct forms of the same species may co-exist in the same district, and we cannot doubt that if the one had possessed any advantage over the other, it would soon have been multiplied to the exclusion of the latter. If, for instance, the male pied ravens, instead of being persecuted by their comrades, had been highly attractive (like the above pied peacock) to the black female ravens their numbers would have rapidly increased. And this would have been a case of sexual selection.

With respect to the slight individual differences which are common, in a greater or less degree, to all the members of the same species, we have every reason to believe that they are by far the most important for the work of selection. Secondary sexual characters are eminently liable to vary, both with animals in a state of nature and under domestication. (43. On these points see also ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 253; vol ii. pp. 73, 75.) There is also reason to believe, as we have seen in our eighth chapter, that variations are more apt to occur in the male than in the female sex. All these contingencies are highly favourable for sexual selection. Whether characters thus acquired are transmitted to one sex or to both sexes, depends, as we shall see in the following chapter, on the form of inheritance which prevails.

It is sometimes difficult to form an opinion whether certain slight differences between the sexes of birds are simply the result of variability with sexually-limited inheritance, without the aid of sexual selection, or whether they have been augmented through this latter process. I do not here refer to the many instances where the male displays splendid colours or other ornaments, of which the female partakes to a slight degree; for these are almost certainly due to characters primarily acquired by the male having been more or less transferred to the female. But what are we to conclude with respect to certain birds in which, for instance, the eyes differ slightly in colour in the two sexes? (44. See, for instance, on the irides of a Podica and Gallicrex in ‘Ibis,’ vol. ii. 1860, p. 206; and vol. v. 1863, p. 426.) In some cases the eyes differ conspicuously; thus with the storks of the genus Xenorhynchus, those of the male are blackish-hazel, whilst those of the females are gamboge-yellow; with many hornbills (Buceros), as I hear from Mr. Blyth (45. See also Jerdon, ‘Birds of India,’ vol. i. pp. 243-245.), the males have intense crimson eyes, and those of the females are white. In the Buceros bicornis, the hind margin of the casque and a stripe on the crest of the beak are black in the male, but not so in the female. Are we to suppose that these black marks and the crimson colour of the eyes have been preserved or augmented through sexual selection in the males? This is very doubtful; for Mr. Bartlett shewed me in the Zoological Gardens that the inside of the mouth of this Buceros is black in the male and flesh-coloured in the female; and their external appearance or beauty would not be thus affected. I observed in Chile (46. ‘Zoology of the Voyage of H.M.S. “Beagle,”’ 1841, p. 6.) that the iris in the condor, when about a year old, is dark-brown, but changes at maturity into yellowish-brown in the male, and into bright red in the female. The male has also a small, longitudinal, leaden-coloured, fleshy crest or comb. The comb of many gallinaceous birds is highly ornamental, and assumes vivid colours during the act of courtship; but what are we to think of the dull-coloured comb of the condor, which does not appear to us in the least ornamental? The same question may be asked in regard to various other characters, such as the knob on the base of the beak of the Chinese goose (Anser cygnoides), which is much larger in the male than in the female. No certain answer can be given to these questions; but we ought to be cautious in assuming that knobs and various fleshy appendages cannot be attractive to the female, when we remember that with savage races of man various hideous deformities—deep scars on the face with the flesh raised into protuberances, the septum of the nose pierced by sticks or bones, holes in the ears and lips stretched widely open—are all admired as ornamental.

Whether or not unimportant differences between the sexes, such as those just specified, have been preserved through sexual selection, these differences, as well as all others, must primarily depend on the laws of variation. On the principle of correlated development, the plumage often varies on different parts of the body, or over the whole body, in the same manner. We see this well illustrated in certain breeds of the fowl. In all the breeds the feathers on the neck and loins of the males are elongated, and are called hackles; now when both sexes acquire a top-knot, which is a new character in the genus, the feathers on the head of the male become hackle-shaped, evidently on the principle of correlation; whilst those on the head of the female are of the ordinary shape. The colour also of the hackles forming the top-knot of the male, is often correlated with that of the hackles on the neck and loins, as may be seen by comparing these feathers in the golden and silver-spangled Polish, the Houdans, and Creve-coeur breeds. In some natural species we may observe exactly the same correlation in the colours of these same feathers, as in the males of the splendid Gold and Amherst pheasants.

The structure of each individual feather generally causes any change in its colouring to be symmetrical; we see this in the various laced, spangled, and pencilled breeds of the fowl; and on the principle of correlation the feathers over the whole body are often coloured in the same manner. We are thus enabled without much trouble to rear breeds with their plumage marked almost as symmetrically as in natural species. In laced and spangled fowls the coloured margins of the feathers are abruptly defined; but in a mongrel raised by me from a black Spanish cock glossed with green, and a white game-hen, all the feathers were greenish-black, excepting towards their extremities, which were yellowish-white; but between the white extremities and the black bases, there was on each feather a symmetrical, curved zone of dark-brown. In some instances the shaft of the feather determines the distribution of the tints; thus with the body-feathers of a mongrel from the same black Spanish cock and a silver-spangled Polish hen, the shaft, together with a narrow space on each side, was greenish-black, and this was surrounded by a regular zone of dark-brown, edged with brownish-white. In these cases we have feathers symmetrically shaded, like those which give so much elegance to the plumage of many natural species. I have also noticed a variety of the common pigeon with the wing-bars symmetrically zoned with three bright shades, instead of being simply black on a slaty-blue ground, as in the parent-species.

In many groups of birds the plumage is differently coloured in the several species, yet certain spots, marks, or stripes are retained by all. Analogous cases occur with the breeds of the pigeon, which usually retain the two wing-bars, though they may be coloured red, yellow, white, black, or blue, the rest of the plumage being of some wholly different tint. Here is a more curious case, in which certain marks are retained, though coloured in a manner almost exactly the opposite of what is natural; the aboriginal pigeon has a blue tail, with the terminal halves of the outer webs of the two outer tail feathers white; now there is a sub-variety having a white instead of a blue tail, with precisely that part black which is white in the parent-species. (47. Bechstein, ‘Naturgeschichte Deutschlands,’ B. iv. 1795, s. 31, on a sub-variety of the Monck pigeon.)

FORMATION AND VARIABILITY OF THE OCELLI OR EYE-LIKE SPOTS ON THE PLUMAGE OF BIRDS.

[Fig. 53. Cyllo leda, Linn., from a drawing by Mr. Trimen, shewing the extreme range of variation in the ocelli. A. Specimen, from Mauritius, upper surface of fore-wing. A1. Specimen, from Natal, ditto. B. Specimen, from Java, upper surface of hind-wing. B1. Specimen, from Mauritius, ditto.]

As no ornaments are more beautiful than the ocelli on the feathers of various birds, on the hairy coats of some mammals, on the scales of reptiles and fishes, on the skin of amphibians, on the wings of many Lepidoptera and other insects, they deserve to be especially noticed. An ocellus consists of a spot within a ring of another colour, like the pupil within the iris, but the central spot is often surrounded by additional concentric zones. The ocelli on the tail-coverts of the peacock offer a familiar example, as well as those on the wings of the peacock-butterfly (Vanessa). Mr. Trimen has given me a description of a S. African moth (Gynanisa isis), allied to our Emperor moth, in which a magnificent ocellus occupies nearly the whole surface of each hinder wing; it consists of a black centre, including a semi-transparent crescent-shaped mark, surrounded by successive, ochre-yellow, black, ochre-yellow, pink, white, pink, brown, and whitish zones. Although we do not know the steps by which these wonderfully beautiful and complex ornaments have been developed, the process has probably been a simple one, at least with insects; for, as Mr. Trimen writes to me, “no characters of mere marking or coloration are so unstable in the Lepidoptera as the ocelli, both in number and size.” Mr. Wallace, who first called my attention to this subject, shewed me a series of specimens of our common meadow-brown butterfly (Hipparchia janira) exhibiting numerous gradations from a simple minute black spot to an elegantly-shaded ocellus. In a S. African butterfly (Cyllo leda, Linn.), belonging to the same family, the ocelli are even still more variable. In some specimens (A, Fig. 53) large spaces on the upper surface of the wings are coloured black, and include irregular white marks; and from this state a complete gradation can be traced into a tolerably perfect ocellus (A1), and this results from the contraction of the irregular blotches of colour. In another series of specimens a gradation can be followed from excessively minute white dots, surrounded by a scarcely visible black line (B), into perfectly symmetrical and large ocelli (B1). (48. This woodcut has been engraved from a beautiful drawing, most kindly made for me by Mr. Trimen; see also his description of the wonderful amount of variation in the coloration and shape of the wings of this butterfly, in his ‘Rhopalocera Africae Australis,’ p. 186.) In cases like these, the development of a perfect ocellus does not require a long course of variation and selection.

With birds and many other animals, it seems to follow from the comparison of allied species that circular spots are often generated by the breaking up and contraction of stripes. In the Tragopan pheasant faint white lines in the female represent the beautiful white spots in the male (49. Jerdon, ‘Birds of India,’ vol. iii. p. 517.); and something of the same kind may be observed in the two sexes of the Argus pheasant. However this may be, appearances strongly favour the belief that on the one hand, a dark spot is often formed by the colouring matter being drawn towards a central point from a surrounding zone, which latter is thus rendered lighter; and, on the other hand, that a white spot is often formed by the colour being driven away from a central point, so that it accumulates in a surrounding darker zone. In either case an ocellus is the result. The colouring matter seems to be a nearly constant quantity, but is redistributed, either centripetally or centrifugally. The feathers of the common guinea-fowl offer a good instance of white spots surrounded by darker zones; and wherever the white spots are large and stand near each other, the surrounding dark zones become confluent. In the same wing-feather of the Argus pheasant dark spots may be seen surrounded by a pale zone, and white spots by a dark zone. Thus the formation of an ocellus in its most elementary state appears to be a simple affair. By what further steps the more complex ocelli, which are surrounded by many successive zones of colour, have been generated, I will not pretend to say. But the zoned feathers of the mongrels from differently coloured fowls, and the extraordinary variability of the ocelli on many Lepidoptera, lead us to conclude that their formation is not a complex process, but depends on some slight and graduated change in the nature of the adjoining tissues.

GRADATION OF SECONDARY SEXUAL CHARACTERS.

[Fig. 54. Feather of Peacock, about two-thirds of natural size, drawn by Mr. Ford. The transparent zone is represented by the outermost white zone, confined to the upper end of the disc.]

Cases of gradation are important, as shewing us that highly complex ornaments may be acquired by small successive steps. In order to discover the actual steps by which the male of any existing bird has acquired his magnificent colours or other ornaments, we ought to behold the long line of his extinct progenitors; but this is obviously impossible. We may, however, generally gain a clue by comparing all the species of the same group, if it be a large one; for some of them will probably retain, at least partially, traces of their former characters. Instead of entering on tedious details respecting various groups, in which striking instances of gradation could be given, it seems the best plan to take one or two strongly marked cases, for instance that of the peacock, in order to see if light can be thrown on the steps by which this bird has become so splendidly decorated. The peacock is chiefly remarkable from the extraordinary length of his tail-coverts; the tail itself not being much elongated. The barbs along nearly the whole length of these feathers stand separate or are decomposed; but this is the case with the feathers of many species, and with some varieties of the domestic fowl and pigeon. The barbs coalesce towards the extremity of the shaft forming the oval disc or ocellus, which is certainly one of the most beautiful objects in the world. It consists of an iridescent, intensely blue, indented centre, surrounded by a rich green zone, this by a broad coppery-brown zone, and this by five other narrow zones of slightly different iridescent shades. A trifling character in the disc deserves notice; the barbs, for a space along one of the concentric zones are more or less destitute of their barbules, so that a part of the disc is surrounded by an almost transparent zone, which gives it a highly finished aspect. But I have elsewhere described (50. ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 254.) an exactly analogous variation in the hackles of a sub-variety of the game-cock, in which the tips, having a metallic lustre, “are separated from the lower part of the feather by a symmetrically shaped transparent zone, composed of the naked portions of the barbs.” The lower margin or base of the dark-blue centre of the ocellus is deeply indented on the line of the shaft. The surrounding zones likewise shew traces, as may be seen in the drawing (Fig. 54), of indentations, or rather breaks. These indentations are common to the Indian and Javan peacocks (Pavo cristatus and P. muticus); and they seem to deserve particular attention, as probably connected with the development of the ocellus; but for a long time I could not conjecture their meaning.

If we admit the principle of gradual evolution, there must formerly have existed many species which presented every successive step between the wonderfully elongated tail-coverts of the peacock and the short tail-coverts of all ordinary birds; and again between the magnificent ocelli of the former, and the simpler ocelli or mere coloured spots on other birds; and so with all the other characters of the peacock. Let us look to the allied Gallinaceae for any still-existing gradations. The species and sub-species of Polyplectron inhabit countries adjacent to the native land of the peacock; and they so far resemble this bird that they are sometimes called peacock-pheasants. I am also informed by Mr. Bartlett that they resemble the peacock in their voice and in some of their habits. During the spring the males, as previously described, strut about before the comparatively plain-coloured females, expanding and erecting their tail and wing-feathers, which are ornamented with numerous ocelli. I request the reader to turn back to the drawing (Fig. 51) of a Polyplectron; In P. napoleonis the ocelli are confined to the tail, and the back is of a rich metallic blue; in which respects this species approaches the Java peacock. P. hardwickii possesses a peculiar top-knot, which is also somewhat like that of the Java peacock. In all the species the ocelli on the wings and tail are either circular or oval, and consist of a beautiful, iridescent, greenish-blue or greenish-purple disc, with a black border. This border in P. chinquis shades into brown, edged with cream colour, so that the ocellus is here surrounded with variously shaded, though not bright, concentric zones. The unusual length of the tail-coverts is another remarkable character in Polyplectron; for in some of the species they are half, and in others two-thirds as long as the true tail-feathers. The tail-coverts are ocellated as in the peacock. Thus the several species of Polyplectron manifestly make a graduated approach to the peacock in the length of their tail-coverts, in the zoning of the ocelli, and in some other characters.

[Fig. 55. Part of a tail-covert of Polyplectron chinquis, with the two ocelli of natural size.

Fig. 56. Part of a tail-covert of Polyplectron malaccense, with the two ocelli, partially confluent, of natural size.]

Notwithstanding this approach, the first species of Polyplectron which I examined almost made me give up the search; for I found not only that the true tail-feathers, which in the peacock are quite plain, were ornamented with ocelli, but that the ocelli on all the feathers differed fundamentally from those of the peacock, in there being two on the same feather (Fig. 55), one on each side of the shaft. Hence I concluded that the early progenitors of the peacock could not have resembled a Polyplectron. But on continuing my search, I observed that in some of the species the two ocelli stood very near each other; that in the tail-feathers of P. hardwickii they touched each other; and, finally, that on the tail-coverts of this same species as well as of P. malaccense (Fig. 56) they were actually confluent. As the central part alone is confluent, an indentation is left at both the upper and lower ends; and the surrounding coloured zones are likewise indented. A single ocellus is thus formed on each tail-covert, though still plainly betraying its double origin. These confluent ocelli differ from the single ocelli of the peacock in having an indentation at both ends, instead of only at the lower or basal end. The explanation, however, of this difference is not difficult; in some species of Polyplectron the two oval ocelli on the same feather stand parallel to each other; in other species (as in P. chinquis) they converge towards one end; now the partial confluence of two convergent ocelli would manifestly leave a much deeper indentation at the divergent than at the convergent end. It is also manifest that if the convergence were strongly pronounced and the confluence complete, the indentation at the convergent end would tend to disappear.

The tail-feathers in both species of the peacock are entirely destitute of ocelli, and this apparently is related to their being covered up and concealed by the long tail-coverts. In this respect they differ remarkably from the tail-feathers of Polyplectron, which in most of the species are ornamented with larger ocelli than those on the tail-coverts. Hence I was led carefully to examine the tail-feathers of the several species, in order to discover whether their ocelli shewed any tendency to disappear; and to my great satisfaction, this appeared to be so. The central tail-feathers of P. napoleonis have the two ocelli on each side of the shaft perfectly developed; but the inner ocellus becomes less and less conspicuous on the more exterior tail-feathers, until a mere shadow or rudiment is left on the inner side of the outermost feather. Again, in P. malaccense, the ocelli on the tail-coverts are, as we have seen, confluent; and these feathers are of unusual length, being two-thirds of the length of the tail-feathers, so that in both these respects they approach the tail-coverts of the peacock. Now in P. malaccense, the two central tail-feathers alone are ornamented, each with two brightly-coloured ocelli, the inner ocellus having completely disappeared from all the other tail-feathers. Consequently the tail-coverts and tail-feathers of this species of Polyplectron make a near approach in structure and ornamentation to the corresponding feathers of the peacock.

As far, then, as gradation throws light on the steps by which the magnificent train of the peacock has been acquired, hardly anything more is needed. If we picture to ourselves a progenitor of the peacock in an almost exactly intermediate condition between the existing peacock, with his enormously elongated tail-coverts, ornamented with single ocelli, and an ordinary gallinaceous bird with short tail-coverts, merely spotted with some colour, we shall see a bird allied to Polyplectron—that is, with tail-coverts, capable of erection and expansion, ornamented with two partially confluent ocelli, and long enough almost to conceal the tail-feathers, the latter having already partially lost their ocelli. The indentation of the central disc and of the surrounding zones of the ocellus, in both species of peacock, speaks plainly in favour of this view, and is otherwise inexplicable. The males of Polyplectron are no doubt beautiful birds, but their beauty, when viewed from a little distance, cannot be compared with that of the peacock. Many female progenitors of the peacock must, during a long line of descent, have appreciated this superiority; for they have unconsciously, by the continued preference for the most beautiful males, rendered the peacock the most splendid of living birds.

ARGUS PHEASANT.

Another excellent case for investigation is offered by the ocelli on the wing-feathers of the Argus pheasant, which are shaded in so wonderful a manner as to resemble balls lying loose within sockets, and consequently differ from ordinary ocelli. No one, I presume, will attribute the shading, which has excited the admiration of many experienced artists, to chance—to the fortuitous concourse of atoms of colouring matter. That these ornaments should have been formed through the selection of many successive variations, not one of which was originally intended to produce the ball-and-socket effect, seems as incredible as that one of Raphael’s Madonnas should have been formed by the selection of chance daubs of paint made by a long succession of young artists, not one of whom intended at first to draw the human figure. In order to discover how the ocelli have been developed, we cannot look to a long line of progenitors, nor to many closely-allied forms, for such do not now exist. But fortunately the several feathers on the wing suffice to give us a clue to the problem, and they prove to demonstration that a gradation is at least possible from a mere spot to a finished ball-and-socket ocellus.

[Fig. 57. Part of secondary wing-feather of Argus pheasant, shewing two perfect ocelli, a and b. A, B, C, D, etc., are dark stripes running obliquely down, each to an ocellus. [Much of the web on both sides, especially to the left of the shaft, has been cut off.]

Fig.59. Portion of one of the secondary wing-feathers near to the body, shewing the so-called elliptic ornaments. The right-hand figure is given merely as a diagram for the sake of the letters of reference. A, B, C, D, etc. Rows of spots running down to and forming the elliptic ornaments. b. Lowest spot or mark in row B. c. The next succeeding spot or mark in the same row. d. Apparently a broken prolongation of the spot c. in the same row B.]

The wing-feathers, bearing the ocelli, are covered with dark stripes (Fig. 57) or with rows of dark spots (Fig. 59), each stripe or row of spots running obliquely down the outer side of the shaft to one of the ocelli. The spots are generally elongated in a line transverse to the row in which they stand. They often become confluent either in the line of the row—and then they form a longitudinal stripe—or transversely, that is, with the spots in the adjoining rows, and then they form transverse stripes. A spot sometimes breaks up into smaller spots, which still stand in their proper places.

It will be convenient first to describe a perfect ball-and-socket ocellus. This consists of an intensely black circular ring, surrounding a space shaded so as exactly to resemble a ball. The figure here given has been admirably drawn by Mr. Ford and well engraved, but a woodcut cannot exhibit the exquisite shading of the original. The ring is almost always slightly broken or interrupted (Fig. 57) at a point in the upper half, a little to the right of and above the white shade on the enclosed ball; it is also sometimes broken towards the base on the right hand. These little breaks have an important meaning. The ring is always much thickened, with the edges ill-defined towards the left-hand upper corner, the feather being held erect, in the position in which it is here drawn. Beneath this thickened part there is on the surface of the ball an oblique almost pure-white mark, which shades off downwards into a pale-leaden hue, and this into yellowish and brown tints, which insensibly become darker and darker towards the lower part of the ball. It is this shading which gives so admirably the effect of light shining on a convex surface. If one of the balls be examined, it will be seen that the lower part is of a brown tint and is indistinctly separated by a curved oblique line from the upper part, which is yellower and more leaden; this curved oblique line runs at right angles to the longer axis of the white patch of light, and indeed of all the shading; but this difference in colour, which cannot of course be shewn in the woodcut, does not in the least interfere with the perfect shading of the ball. It should be particularly observed that each ocellus stands in obvious connection either with a dark stripe, or with a longitudinal row of dark spots, for both occur indifferently on the same feather. Thus in Fig. 57 stripe A runs to ocellus a; B runs to ocellus b; stripe C is broken in the upper part, and runs down to the next succeeding ocellus, not represented in the woodcut; D to the next lower one, and so with the stripes E and F. Lastly, the several ocelli are separated from each other by a pale surface bearing irregular black marks.

[Fig. 58. Basal part of the secondary wing feather, nearest to the body.]

I will next describe the other extreme of the series, namely, the first trace of an ocellus. The short secondary wing-feather (Fig. 58), nearest to the body, is marked like the other feathers, with oblique, longitudinal, rather irregular, rows of very dark spots. The basal spot, or that nearest the shaft, in the five lower rows (excluding the lowest one) is a little larger than the other spots of the same row, and a little more elongated in a transverse direction. It differs also from the other spots by being bordered on its upper side with some dull fulvous shading. But this spot is not in any way more remarkable than those on the plumage of many birds, and might easily be overlooked. The next higher spot does not differ at all from the upper ones in the same row. The larger basal spots occupy exactly the same relative position on these feathers as do the perfect ocelli on the longer wing-feathers.

By looking to the next two or three succeeding wing-feathers, an absolutely insensible gradation can be traced from one of the last-described basal spots, together with the next higher one in the same row, to a curious ornament, which cannot be called an ocellus, and which I will name, from the want of a better term, an “elliptic ornament.” These are shewn in the accompanying figure (Fig. 59). We here see several oblique rows, A, B, C, D, etc. (see the lettered diagram on the right hand), of dark spots of the usual character. Each row of spots runs down to and is connected with one of the elliptic ornaments, in exactly the same manner as each stripe in Fig. 57 runs down to and is connected with one of the ball-and-socket ocelli. Looking to any one row, for instance, B, in Fig. 59, the lowest mark (b) is thicker and considerably longer than the upper spots, and has its left extremity pointed and curved upwards. This black mark is abruptly bordered on its upper side by a rather broad space of richly shaded tints, beginning with a narrow brown zone, which passes into orange, and this into a pale leaden tint, with the end towards the shaft much paler. These shaded tints together fill up the whole inner space of the elliptic ornament. The mark (b) corresponds in every respect with the basal shaded spot of the simple feather described in the last paragraph (Fig. 58), but is more highly developed and more brightly coloured. Above and to the right of this spot (b, Fig. 59), with its bright shading, there is a long narrow, black mark (c), belonging to the same row, and which is arched a little downwards so as to face (b). This mark is sometimes broken into two portions. It is also narrowly edged on the lower side with a fulvous tint. To the left of and above c, in the same oblique direction, but always more or less distinct from it, there is another black mark (d). This mark is generally sub-triangular and irregular in shape, but in the one lettered in the diagram it is unusually narrow, elongated, and regular. It apparently consists of a lateral and broken prolongation of the mark (c), together with its confluence with a broken and prolonged part of the next spot above; but I do not feel sure of this. These three marks, b, c, and d, with the intervening bright shades, form together the so-called elliptic ornament. These ornaments placed parallel to the shaft, manifestly correspond in position with the ball-and-socket ocelli. Their extremely elegant appearance cannot be appreciated in the drawing, as the orange and leaden tints, contrasting so well with the black marks, cannot be shewn.

[Fig. 60. An ocellus in an intermediate condition between the elliptic ornament and the perfect ball-and-socket ocellus.]

Between one of the elliptic ornaments and a perfect ball-and-socket ocellus, the gradation is so perfect that it is scarcely possible to decide when the latter term ought to be used. The passage from the one into the other is effected by the elongation and greater curvature in opposite directions of the lower black mark (b, Fig. 59), and more especially of the upper one (c), together with the contraction of the elongated sub-triangular or narrow mark (d), so that at last these three marks become confluent, forming an irregular elliptic ring. This ring is gradually rendered more and more circular and regular, increasing at the same time in diameter. I have here given a drawing (Fig. 60) of the natural size of an ocellus not as yet quite perfect. The lower part of the black ring is much more curved than is the lower mark in the elliptic ornament (b, Fig. 59). The upper part of the ring consists of two or three separate portions; and there is only a trace of the thickening of the portion which forms the black mark above the white shade. This white shade itself is not as yet much concentrated; and beneath it the surface is brighter coloured than in a perfect ball-and-socket ocellus. Even in the most perfect ocelli traces of the junction of three or four elongated black marks, by which the ring has been formed, may often be detected. The irregular sub-triangular or narrow mark (d, Fig. 59), manifestly forms, by its contraction and equalisation, the thickened portion of the ring above the white shade on a perfect ball-and-socket ocellus. The lower part of the ring is invariably a little thicker than the other parts (Fig. 57), and this follows from the lower black mark of the elliptic ornament (b, Fig. 59) having originally been thicker than the upper mark (c). Every step can be followed in the process of confluence and modification; and the black ring which surrounds the ball of the ocellus is unquestionably formed by the union and modification of the three black marks, b, c, d, of the elliptic ornament. The irregular zigzag black marks between the successive ocelli (Fig. 57) are plainly due to the breaking up of the somewhat more regular but similar marks between the elliptic ornaments.

The successive steps in the shading of the ball-and-socket ocelli can be followed out with equal clearness. The brown, orange, and pale-leadened narrow zones, which border the lower black mark of the elliptic ornament, can be seen gradually to become more and more softened and shaded into each other, with the upper lighter part towards the left-hand corner rendered still lighter, so as to become almost white, and at the same time more contracted. But even in the most perfect ball-and-socket ocelli a slight difference in the tints, though not in the shading, between the upper and lower parts of the ball can be perceived, as before noticed; and the line of separation is oblique, in the same direction as the bright coloured shades of the elliptic ornaments. Thus almost every minute detail in the shape and colouring of the ball-and-socket ocelli can be shewn to follow from gradual changes in the elliptic ornaments; and the development of the latter can be traced by equally small steps from the union of two almost simple spots, the lower one (Fig. 58) having some dull fulvous shading on its upper side.

[Fig. 61. Portion near summit of one of the secondary wing-feathers, bearing perfect ball-and-socket ocelli. a. Ornamented upper part. b. Uppermost, imperfect ball-and-socket ocellus. (The shading above the white mark on the summit of the ocellus is here a little too dark.) c. Perfect ocellus.]

The extremities of the longer secondary feathers which bear the perfect ball-and-socket ocelli, are peculiarly ornamented (Fig. 61). The oblique longitudinal stripes suddenly cease upwards and become confused; and above this limit the whole upper end of the feather (a) is covered with white dots, surrounded by little black rings, standing on a dark ground. The oblique stripe belonging to the uppermost ocellus (b) is barely represented by a very short irregular black mark with the usual, curved, transverse base. As this stripe is thus abruptly cut off, we can perhaps understand from what has gone before, how it is that the upper thickened part of the ring is here absent; for, as before stated, this thickened part apparently stands in some relation with a broken prolongation from the next higher spot. From the absence of the upper and thickened part of the ring, the uppermost ocellus, though perfect in all other respects, appears as if its top had been obliquely sliced off. It would, I think, perplex any one, who believes that the plumage of the Argus pheasant was created as we now see it, to account for the imperfect condition of the uppermost ocellus. I should add that on the secondary wing-feather farthest from the body all the ocelli are smaller and less perfect than on the other feathers, and have the upper part of the ring deficient, as in the case just mentioned. The imperfection here seems to be connected with the fact that the spots on this feather shew less tendency than usual to become confluent into stripes; they are, on the contrary, often broken up into smaller spots, so that two or three rows run down to the same ocellus.

There still remains another very curious point, first observed by Mr. T.W. Wood (51. The ‘Field,’ May 28, 1870.), which deserves attention. In a photograph, given me by Mr. Ward, of a specimen mounted as in the act of display, it may be seen that on the feathers which are held perpendicularly, the white marks on the ocelli, representing light reflected from a convex surface, are at the upper or further end, that is, are directed upwards; and the bird whilst displaying himself on the ground would naturally be illuminated from above. But here comes the curious point; the outer feathers are held almost horizontally, and their ocelli ought likewise to appear as if illuminated from above, and consequently the white marks ought to be placed on the upper sides of the ocelli; and, wonderful as is the fact, they are thus placed! Hence the ocelli on the several feathers, though occupying very different positions with respect to the light, all appear as if illuminated from above, just as an artist would have shaded them. Nevertheless they are not illuminated from strictly the same point as they ought to be; for the white marks on the ocelli of the feathers which are held almost horizontally, are placed rather too much towards the further end; that is, they are not sufficiently lateral. We have, however, no right to expect absolute perfection in a part rendered ornamental through sexual selection, any more than we have in a part modified through natural selection for real use; for instance, in that wondrous organ the human eye. And we know what Helmholtz, the highest authority in Europe on the subject, has said about the human eye; that if an optician had sold him an instrument so carelessly made, he would have thought himself fully justified in returning it. (52. ‘Popular Lectures on Scientific Subjects,’ Eng. trans. 1873, pp. 219, 227, 269, 390.)

We have now seen that a perfect series can be followed, from simple spots to the wonderful ball-and-socket ornaments. Mr. Gould, who kindly gave me some of these feathers, fully agrees with me in the completeness of the gradation. It is obvious that the stages in development exhibited by the feathers on the same bird do not at all necessarily shew us the steps passed through by the extinct progenitors of the species; but they probably give us the clue to the actual steps, and they at least prove to demonstration that a gradation is possible. Bearing in mind how carefully the male Argus pheasant displays his plumes before the female, as well as the many facts rendering it probable that female birds prefer the more attractive males, no one who admits the agency of sexual selection in any case will deny that a simple dark spot with some fulvous shading might be converted, through the approximation and modification of two adjoining spots, together with some slight increase of colour, into one of the so-called elliptic ornaments. These latter ornaments have been shewn to many persons, and all have admitted that they are beautiful, some thinking them even more so than the ball-and-socket ocelli. As the secondary plumes became lengthened through sexual selection, and as the elliptic ornaments increased in diameter, their colours apparently became less bright; and then the ornamentation of the plumes had to be gained by an improvement in the pattern and shading; and this process was carried on until the wonderful ball-and-socket ocelli were finally developed. Thus we can understand—and in no other way as it seems to me—the present condition and origin of the ornaments on the wing-feathers of the Argus pheasant.

From the light afforded by the principle of gradation—from what we know of the laws of variation—from the changes which have taken place in many of our domesticated birds—and, lastly, from the character (as we shall hereafter see more clearly) of the immature plumage of young birds—we can sometimes indicate, with a certain amount of confidence, the probable steps by which the males have acquired their brilliant plumage and various ornaments; yet in many cases we are involved in complete darkness. Mr. Gould several years ago pointed out to me a humming-bird, the Urosticte benjamini, remarkable for the curious differences between the sexes. The male, besides a splendid gorget, has greenish-black tail-feathers, with the four CENTRAL ones tipped with white; in the female, as with most of the allied species, the three OUTER tail-feathers on each side are tipped with white, so that the male has the four central, whilst the female has the six exterior feathers ornamented with white tips. What makes the case more curious is that, although the colouring of the tail differs remarkably in both sexes of many kinds of humming-birds, Mr. Gould does not know a single species, besides the Urosticte, in which the male has the four central feathers tipped with white.

The Duke of Argyll, in commenting on this case (53. ‘The Reign of Law,’ 1867, p. 247.), passes over sexual selection, and asks, “What explanation does the law of natural selection give of such specific varieties as these?” He answers “none whatever”; and I quite agree with him. But can this be so confidently said of sexual selection? Seeing in how many ways the tail-feathers of humming-birds differ, why should not the four central feathers have varied in this one species alone, so as to have acquired white tips? The variations may have been gradual, or somewhat abrupt as in the case recently given of the humming-birds near Bogota, in which certain individuals alone have the “central tail-feathers tipped with beautiful green.” In the female of the Urosticte I noticed extremely minute or rudimental white tips to the two outer of the four central black tail-feathers; so that here we have an indication of change of some kind in the plumage of this species. If we grant the possibility of the central tail-feathers of the male varying in whiteness, there is nothing strange in such variations having been sexually selected. The white tips, together with the small white ear-tufts, certainly add, as the Duke of Argyll admits, to the beauty of the male; and whiteness is apparently appreciated by other birds, as may be inferred from such cases as the snow-white male of the Bell-bird. The statement made by Sir R. Heron should not be forgotten, namely, that his peahens, when debarred from access to the pied peacock, would not unite with any other male, and during that season produced no offspring. Nor is it strange that variations in the tail-feathers of the Urosticte should have been specially selected for the sake of ornament, for the next succeeding genus in the family takes its name of Metallura from the splendour of these feathers. We have, moreover, good evidence that humming-birds take especial pains in displaying their tail-feathers; Mr. Belt (54. ‘The Naturalist in Nicaragua,’ 1874, p. 112.), after describing the beauty of the Florisuga mellivora, says, “I have seen the female sitting on a branch, and two males displaying their charms in front of her. One would shoot up like a rocket, then suddenly expanding the snow-white tail, like an inverted parachute, slowly descend in front of her, turning round gradually to shew off back and front...The expanded white tail covered more space than all the rest of the bird, and was evidently the grand feature in the performance. Whilst one male was descending, the other would shoot up and come slowly down expanded. The entertainment would end in a fight between the two performers; but whether the most beautiful or the most pugnacious was the accepted suitor, I know not.” Mr. Gould, after describing the peculiar plumage of the Urosticte, adds, “that ornament and variety is the sole object, I have myself but little doubt.” (55. ‘Introduction to the Trochilidae,’ 1861, p. 110.) If this be admitted, we can perceive that the males which during former times were decked in the most elegant and novel manner would have gained an advantage, not in the ordinary struggle for life, but in rivalry with other males, and would have left a larger number of offspring to inherit their newly-acquired beauty.

Chapter XV

CHAPTER XV. BIRDS—continued.

Discussion as to why the males alone of some species, and both sexes of others, are brightly coloured—On sexually-limited inheritance, as applied to various structures and to brightly-coloured plumage—Nidification in relation to colour—Loss of nuptial plumage during the winter.

We have in this chapter to consider why the females of many birds have not acquired the same ornaments as the male; and why, on the other hand, both sexes of many other birds are equally, or almost equally, ornamented? In the following chapter we shall consider the few cases in which the female is more conspicuously coloured than the male.

In my ‘Origin of Species’ (1. Fourth edition, 1866, p. 241.) I briefly suggested that the long tail of the peacock would be inconvenient and the conspicuous black colour of the male capercailzie dangerous, to the female during the period of incubation: and consequently that the transmission of these characters from the male to the female offspring had been checked through natural selection. I still think that this may have occurred in some few instances: but after mature reflection on all the facts which I have been able to collect, I am now inclined to believe that when the sexes differ, the successive variations have generally been from the first limited in their transmission to the same sex in which they first arose. Since my remarks appeared, the subject of sexual coloration has been discussed in some very interesting papers by Mr. Wallace (2. ‘Westminster Review,’ July 1867. ‘Journal of Travel,’ vol. i. 1868, p. 73.), who believes that in almost all cases the successive variations tended at first to be transmitted equally to both sexes; but that the female was saved, through natural selection, from acquiring the conspicuous colours of the male, owing to the danger which she would thus have incurred during incubation.

This view necessitates a tedious discussion on a difficult point, namely, whether the transmission of a character, which is at first inherited by both sexes can be subsequently limited in its transmission to one sex alone by means of natural selection. We must bear in mind, as shewn in the preliminary chapter on sexual selection, that characters which are limited in their development to one sex are always latent in the other. An imaginary illustration will best aid us in seeing the difficulty of the case; we may suppose that a fancier wished to make a breed of pigeons, in which the males alone should be coloured of a pale blue, whilst the females retained their former slaty tint. As with pigeons characters of all kinds are usually transmitted to both sexes equally, the fancier would have to try to convert this latter form of inheritance into sexually-limited transmission. All that he could do would be to persevere in selecting every male pigeon which was in the least degree of a paler blue; and the natural result of this process, if steadily carried on for a long time, and if the pale variations were strongly inherited or often recurred, would be to make his whole stock of a lighter blue. But our fancier would be compelled to match, generation after generation, his pale blue males with slaty females, for he wishes to keep the latter of this colour. The result would generally be the production either of a mongrel piebald lot, or more probably the speedy and complete loss of the pale-blue tint; for the primordial slaty colour would be transmitted with prepotent force. Supposing, however, that some pale-blue males and slaty females were produced during each successive generation, and were always crossed together, then the slaty females would have, if I may use the expression, much blue blood in their veins, for their fathers, grandfathers, etc., will all have been blue birds. Under these circumstances it is conceivable (though I know of no distinct facts rendering it probable) that the slaty females might acquire so strong a latent tendency to pale-blueness, that they would not destroy this colour in their male offspring, their female offspring still inheriting the slaty tint. If so, the desired end of making a breed with the two sexes permanently different in colour might be gained.

The extreme importance, or rather necessity in the above case of the desired character, namely, pale-blueness, being present though in a latent state in the female, so that the male offspring should not be deteriorated, will be best appreciated as follows: the male of Soemmerring’s pheasant has a tail thirty-seven inches in length, whilst that of the female is only eight inches; the tail of the male common pheasant is about twenty inches, and that of the female twelve inches long. Now if the female Soemmerring pheasant with her SHORT tail were crossed with the male common pheasant, there can be no doubt that the male hybrid offspring would have a much LONGER tail than that of the pure offspring of the common pheasant. On the other hand, if the female common pheasant, with a tail much longer than that of the female Soemmerring pheasant, were crossed with the male of the latter, the male hybrid offspring would have a much SHORTER tail than that of the pure offspring of Soemmerring’s pheasant. (3. Temminck says that the tail of the female Phasianus Soemmerringii is only six inches long, ‘Planches coloriees,’ vol. v. 1838, pp. 487 and 488: the measurements above given were made for me by Mr. Sclater. For the common pheasant, see Macgillivray, ‘History of British Birds,’ vol. i. pp. 118-121.)

Our fancier, in order to make his new breed with the males of a pale-blue tint, and the females unchanged, would have to continue selecting the males during many generations; and each stage of paleness would have to be fixed in the males, and rendered latent in the females. The task would be an extremely difficult one, and has never been tried, but might possibly be successfully carried out. The chief obstacle would be the early and complete loss of the pale-blue tint, from the necessity of reiterated crosses with the slaty female, the latter not having at first any LATENT tendency to produce pale-blue offspring.

On the other hand, if one or two males were to vary ever so slightly in paleness, and the variations were from the first limited in their transmission to the male sex, the task of making a new breed of the desired kind would be easy, for such males would simply have to be selected and matched with ordinary females. An analogous case has actually occurred, for there are breeds of the pigeon in Belgium (4. Dr. Chapuis, ‘Le Pigeon Voyageur Belge,’ 1865, p. 87.) in which the males alone are marked with black striae. So again Mr. Tegetmeier has recently shewn (5. The ‘Field,’ Sept. 1872.) that dragons not rarely produce silver-coloured birds, which are almost always hens; and he himself has bred ten such females. It is on the other hand a very unusual event when a silver male is produced; so that nothing would be easier, if desired, than to make a breed of dragons with blue males and silver females. This tendency is indeed so strong that when Mr. Tegetmeier at last got a silver male and matched him with one of the silver females, he expected to get a breed with both sexes thus coloured; he was however disappointed, for the young male reverted to the blue colour of his grandfather, the young female alone being silver. No doubt with patience this tendency to reversion in the males, reared from an occasional silver male matched with a silver hen, might be eliminated, and then both sexes would be coloured alike; and this very process has been followed with success by Mr. Esquilant in the case of silver turbits.

With fowls, variations of colour, limited in their transmission to the male sex, habitually occur. When this form of inheritance prevails, it might well happen that some of the successive variations would be transferred to the female, who would then slightly resemble the male, as actually occurs in some breeds. Or again, the greater number, but not all, of the successive steps might be transferred to both sexes, and the female would then closely resemble the male. There can hardly be a doubt that this is the cause of the male pouter pigeon having a somewhat larger crop, and of the male carrier pigeon having somewhat larger wattles, than their respective females; for fanciers have not selected one sex more than the other, and have had no wish that these characters should be more strongly displayed in the male than in the female, yet this is the case with both breeds.

The same process would have to be followed, and the same difficulties encountered, if it were desired to make a breed with the females alone of some new colour.

Lastly, our fancier might wish to make a breed with the two sexes differing from each other, and both from the parent species. Here the difficulty would be extreme, unless the successive variations were from the first sexually limited on both sides, and then there would be no difficulty. We see this with the fowl; thus the two sexes of the pencilled Hamburghs differ greatly from each other, and from the two sexes of the aboriginal Gallus bankiva; and both are now kept constant to their standard of excellence by continued selection, which would be impossible unless the distinctive characters of both were limited in their transmission.

The Spanish fowl offers a more curious case; the male has an immense comb, but some of the successive variations, by the accumulation of which it was acquired, appear to have been transferred to the female; for she has a comb many times larger than that of the females of the parent species. But the comb of the female differs in one respect from that of the male, for it is apt to lop over; and within a recent period it has been ordered by the fancy that this should always be the case, and success has quickly followed the order. Now the lopping of the comb must be sexually limited in its transmission, otherwise it would prevent the comb of the male from being perfectly upright, which would be abhorrent to every fancier. On the other hand, the uprightness of the comb in the male must likewise be a sexually-limited character, otherwise it would prevent the comb of the female from lopping over.

From the foregoing illustrations, we see that even with almost unlimited time at command, it would be an extremely difficult and complex, perhaps an impossible process, to change one form of transmission into the other through selection. Therefore, without distinct evidence in each case, I am unwilling to admit that this has been effected in natural species. On the other hand, by means of successive variations, which were from the first

sexually limited in their transmission, there would not be the least difficulty in rendering a male bird widely different in colour or in any other character from the female; the latter being left unaltered, or slightly altered, or specially modified for the sake of protection.

As bright colours are of service to the males in their rivalry with other males, such colours would be selected whether or not they were transmitted exclusively to the same sex. Consequently the females might be expected often to partake of the brightness of the males to a greater or less degree; and this occurs with a host of species. If all the successive variations were transmitted equally to both sexes, the females would be indistinguishable from the males; and this likewise occurs with many birds. If, however, dull colours were of high importance for the safety of the female during incubation, as with many ground birds, the females which varied in brightness, or which received through inheritance from the males any marked accession of brightness, would sooner or later be destroyed. But the tendency in the males to continue for an indefinite period transmitting to their female offspring their own brightness, would have to be eliminated by a change in the form of inheritance; and this, as shewn by our previous illustration, would be extremely difficult. The more probable result of the long-continued destruction of the more brightly-coloured females, supposing the equal form of transmission to prevail, would be the lessening or annihilation of the bright colours of the males, owing to their continual crossing with the duller females. It would be tedious to follow out all the other possible results; but I may remind the reader that if sexually-limited variations in brightness occurred in the females, even if they were not in the least injurious to them and consequently were not eliminated, yet they would not be favoured or selected, for the male usually accepts any female, and does not select the more attractive individuals; consequently these variations would be liable to be lost, and would have little influence on the character of the race; and this will aid in accounting for the females being commonly duller-coloured than the males.

In the eighth chapter instances were given, to which many might here be added, of variations occurring at various ages, and inherited at the corresponding age. It was also shewn that variations which occur late in life are commonly transmitted to the same sex in which they first appear; whilst variations occurring early in life are apt to be transmitted to both sexes; not that all the cases of sexually-limited transmission can thus be accounted for. It was further shewn that if a male bird varied by becoming brighter whilst young, such variations would be of no service until the age for reproduction had arrived, and there was competition between rival males. But in the case of birds living on the ground and commonly in need of the protection of dull colours, bright tints would be far more dangerous to the young and inexperienced than to the adult males. Consequently the males which varied in brightness whilst young would suffer much destruction and be eliminated through natural selection; on the other hand, the males which varied in this manner when nearly mature, notwithstanding that they were exposed to some additional danger, might survive, and from being favoured through sexual selection, would procreate their kind. As a relation often exists between the period of variation and the form of transmission, if the bright-coloured young males were destroyed and the mature ones were successful in their courtship, the males alone would acquire brilliant colours and would transmit them exclusively to their male offspring. But I by no means wish to maintain that the influence of age on the form of transmission, is the sole cause of the great difference in brilliancy between the sexes of many birds.

When the sexes of birds differ in colour, it is interesting to determine whether the males alone have been modified by sexual selection, the females having been left unchanged, or only partially and indirectly thus changed; or whether the females have been specially modified through natural selection for the sake of protection. I will therefore discuss this question at some length, even more fully than its intrinsic importance deserves; for various curious collateral points may thus be conveniently considered.

Before we enter on the subject of colour, more especially in reference to Mr. Wallace’s conclusions, it may be useful to discuss some other sexual differences under a similar point of view. A breed of fowls formerly existed in Germany (6. Bechstein, ‘Naturgeschichte Deutschlands,’ 1793, B. iii. 339.) in which the hens were furnished with spurs; they were good layers, but they so greatly disturbed their nests with their spurs that they could not be allowed to sit on their own eggs. Hence at one time it appeared to me probable that with the females of the wild Gallinaceae the development of spurs had been checked through natural selection, from the injury thus caused to their nests. This seemed all the more probable, as wing-spurs, which would not be injurious during incubation, are often as well-developed in the female as in the male; though in not a few cases they are rather larger in the male. When the male is furnished with leg-spurs the female almost always exhibits rudiments of them,—the rudiment sometimes consisting of a mere scale, as in Gallus. Hence it might be argued that the females had aboriginally been furnished with well-developed spurs, but that these had subsequently been lost through disuse or natural selection. But if this view be admitted, it would have to be extended to innumerable other cases; and it implies that the female progenitors of the existing spur-bearing species were once encumbered with an injurious appendage.

In some few genera and species, as in Galloperdix, Acomus, and the Javan peacock (Pavo muticus), the females, as well as the males, possess well-developed leg-spurs. Are we to infer from this fact that they construct a different sort of nest from that made by their nearest allies, and not liable to be injured by their spurs; so that the spurs have not been removed? Or are we to suppose that the females of these several species especially require spurs for their defence? It is a more probable conclusion that both the presence and absence of spurs in the females result from different laws of inheritance having prevailed, independently of natural selection. With the many females in which spurs appear as rudiments, we may conclude that some few of the successive variations, through which they were developed in the males, occurred very early in life, and were consequently transferred to the females. In the other and much rarer cases, in which the females possess fully developed spurs, we may conclude that all the successive variations were transferred to them; and that they gradually acquired and inherited the habit of not disturbing their nests.

The vocal organs and the feathers variously modified for producing sound, as well as the proper instincts for using them, often differ in the two sexes, but are sometimes the same in both. Can such differences be accounted for by the males having acquired these organs and instincts, whilst the females have been saved from inheriting them, on account of the danger to which they would have been exposed by attracting the attention of birds or beasts of prey? This does not seem to me probable, when we think of the multitude of birds which with impunity gladden the country with their voices during the spring. (7. Daines Barrington, however, thought it probable (‘Philosophical Transactions,’ 1773, p. 164) that few female birds sing, because the talent would have been dangerous to them during incubation. He adds, that a similar view may possibly account for the inferiority of the female to the male in plumage.) It is a safer conclusion that, as vocal and instrumental organs are of special service only to the males during their courtship, these organs were developed through sexual selection and their constant use in that sex alone—the successive variations and the effects of use having been from the first more or less limited in transmission to the male offspring.

Many analogous cases could be adduced; those for instance of the plumes on the head being generally longer in the male than in the female, sometimes of equal length in both sexes, and occasionally absent in the female,—these several cases occurring in the same group of birds. It would be difficult to account for such a difference between the sexes by the female having been benefited by possessing a slightly shorter crest than the male, and its consequent diminution or complete suppression through natural selection. But I will take a more favourable case, namely the length of the tail. The long train of the peacock would have been not only inconvenient but dangerous to the peahen during the period of incubation and whilst accompanying her young. Hence there is not the least a priori improbability in the development of her tail having been checked through natural selection. But the females of various pheasants, which apparently are exposed on their open nests to as much danger as the peahen, have tails of considerable length. The females as well as the males of the Menura superba have long tails, and they build a domed nest, which is a great anomaly in so large a bird. Naturalists have wondered how the female Menura could manage her tail during incubation; but it is now known (8. Mr. Ramsay, in ‘Proc. Zoolog. Soc.’ 1868, p. 50.) that she “enters the nest head first, and then turns round with her tail sometimes over her back, but more often bent round by her side. Thus in time the tail becomes quite askew, and is a tolerable guide to the length of time the bird has been sitting.” Both sexes of an Australian kingfisher (Tanysiptera sylvia) have the middle tail-feathers greatly lengthened, and the female makes her nest in a hole; and as I am informed by Mr. R.B. Sharpe these feathers become much crumpled during incubation.

In these two latter cases the great length of the tail-feathers must be in some degree inconvenient to the female; and as in both species the tail-feathers of the female are somewhat shorter than those of the male, it might be argued that their full development had been prevented through natural selection. But if the development of the tail of the peahen had been checked only when it became inconveniently or dangerously great, she would have retained a much longer tail than she actually possesses; for her tail is not nearly so long, relatively to the size of her body, as that of many female pheasants, nor longer than that of the female turkey. It must also be borne in mind that, in accordance with this view, as soon as the tail of the peahen became dangerously long, and its development was consequently checked, she would have continually reacted on her male progeny, and thus have prevented the peacock from acquiring his present magnificent train. We may therefore infer that the length of the tail in the peacock and its shortness in the peahen are the result of the requisite variations in the male having been from the first transmitted to the male offspring alone.

We are led to a nearly similar conclusion with respect to the length of the tail in the various species of pheasants. In the Eared pheasant (Crossoptilon auritum) the tail is of equal length in both sexes, namely sixteen or seventeen inches; in the common pheasant it is about twenty inches long in the male and twelve in the female; in Soemmerring’s pheasant, thirty-seven inches in the male and only eight in the female; and lastly in Reeve’s pheasant it is sometimes actually seventy-two inches long in the male and sixteen in the female. Thus in the several species, the tail of the female differs much in length, irrespectively of that of the male; and this can be accounted for, as it seems to me, with much more probability, by the laws of inheritance,—that is by the successive variations having been from the first more or less closely limited in their transmission to the male sex than by the agency of natural selection, resulting from the length of tail being more or less injurious to the females of these several allied species.

We may now consider Mr. Wallace’s arguments in regard to the sexual coloration of birds. He believes that the bright tints originally acquired through sexual selection by the males would in all, or almost all cases, have been transmitted to the females, unless the transference had been checked through natural selection. I may here remind the reader that various facts opposed to this view have already been given under reptiles, amphibians, fishes and lepidoptera. Mr. Wallace rests his belief chiefly, but not exclusively, as we shall see in the next chapter, on the following statement (9. ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 78.), that when both sexes are coloured in a very conspicuous manner, the nest is of such a nature as to conceal the sitting bird; but when there is a marked contrast of colour between the sexes, the male being gay and the female dull-coloured, the nest is open and exposes the sitting bird to view. This coincidence, as far as it goes, certainly seems to favour the belief that the females which sit on open nests have been specially modified for the sake of protection; but we shall presently see that there is another and more probable explanation, namely, that conspicuous females have acquired the instinct of building domed nests oftener than dull-coloured birds. Mr. Wallace admits that there are, as might have been expected, some exceptions to his two rules, but it is a question whether the exceptions are not so numerous as seriously to invalidate them.

There is in the first place much truth in the Duke of Argyll’s remark (10. ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 281.) that a large domed nest is more conspicuous to an enemy, especially to all tree-haunting carnivorous animals, than a smaller open nest. Nor must we forget that with many birds which build open nests, the male sits on the eggs and aids the female in feeding the young: this is the case, for instance, with Pyranga aestiva (11. Audubon, ‘Ornithological Biography,’ vol. i. p. 233.), one of the most splendid birds in the United States, the male being vermilion, and the female light brownish-green. Now if brilliant colours had been extremely dangerous to birds whilst sitting on their open nests, the males in these cases would have suffered greatly. It might, however, be of such paramount importance to the male to be brilliantly coloured, in order to beat his rivals, that this may have more than compensated some additional danger.

Mr. Wallace admits that with the King-crows (Dicrurus), Orioles, and Pittidae, the females are conspicuously coloured, yet build open nests; but he urges that the birds of the first group are highly pugnacious and could defend themselves; that those of the second group take extreme care in concealing their open nests, but this does not invariably hold good (12. Jerdon, ‘Birds of India,’ vol. ii. p. 108. Gould’s ‘Handbook of the Birds of Australia,’ vol. i. p. 463.); and that with the birds of the third group the females are brightly coloured chiefly on the under surface. Besides these cases, pigeons which are sometimes brightly, and almost always conspicuously coloured, and which are notoriously liable to the attacks of birds of prey, offer a serious exception to the rule, for they almost always build open and exposed nests. In another large family, that of the humming-birds, all the species build open nests, yet with some of the most gorgeous species the sexes are alike; and in the majority, the females, though less brilliant than the males, are brightly coloured. Nor can it be maintained that all female humming-birds, which are brightly coloured, escape detection by their tints being green, for some display on their upper surfaces red, blue, and other colours. (13. For instance, the female Eupetomena macroura has the head and tail dark blue with reddish loins; the female Lampornis porphyrurus is blackish-green on the upper surface, with the lores and sides of the throat crimson; the female Eulampis jugularis has the top of the head and back green, but the loins and the tail are crimson. Many other instances of highly conspicuous females could be given. See Mr. Gould’s magnificent work on this family.)

In regard to birds which build in holes or construct domed nests, other advantages, as Mr. Wallace remarks, besides concealment are gained, such as shelter from the rain, greater warmth, and in hot countries protection from the sun (14. Mr. Salvin noticed in Guatemala (‘Ibis,’ 1864, p. 375) that humming-birds were much more unwilling to leave their nests during very hot weather, when the sun was shining brightly, as if their eggs would be thus injured, than during cool, cloudy, or rainy weather.); so that it is no valid objection to his view that many birds having both sexes obscurely coloured build concealed nests. (15. I may specify, as instances of dull-coloured birds building concealed nests, the species belonging to eight Australian genera described in Gould’s ‘Handbook of the Birds of Australia,’ vol. i. pp. 340, 362, 365, 383, 387, 389, 391, 414.) The female Horn-bill (Buceros), for instance, of India and Africa is protected during incubation with extraordinary care, for she plasters up with her own excrement the orifice of the hole in which she sits on her eggs, leaving only a small orifice through which the male feeds her; she is thus kept a close prisoner during the whole period of incubation (16. Mr. C. Horne, ‘Proc. Zoolog. Soc.’ 1869. p. 243.); yet female horn-bills are not more conspicuously coloured than many other birds of equal size which build open nests. It is a more serious objection to Mr. Wallace’s view, as is admitted by him, that in some few groups the males are brilliantly coloured and the females obscure, and yet the latter hatch their eggs in domed nests. This is the case with the Grallinae of Australia, the Superb Warblers (Maluridae) of the same country, the Sun-birds (Nectariniae), and with several of the Australian Honey-suckers or Meliphagidae. (17. On the nidification and colours of these latter species, see Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 504, 527.)

If we look to the birds of England we shall see that there is no close and general relation between the colours of the female and the nature of the nest which is constructed. About forty of our British birds (excluding those of large size which could defend themselves) build in holes in banks, rocks, or trees, or construct domed nests. If we take the colours of the female goldfinch, bullfinch, or blackbird, as a standard of the degree of conspicuousness, which is not highly dangerous to the sitting female, then out of the above forty birds the females of only twelve can be considered as conspicuous to a dangerous degree, the remaining twenty-eight being inconspicuous. (18. I have consulted, on this subject, Macgillivray’s ‘British Birds,’ and though doubts may be entertained in some cases in regard to the degree of concealment of the nest, and to the degree of conspicuousness of the female, yet the following birds, which all lay their eggs in holes or in domed nests, can hardly be considered, by the above standard, as conspicuous: Passer, 2 species; Sturnus, of which the female is considerably less brilliant than the male; Cinclus; Motallica boarula (?); Erithacus (?); Fruticola, 2 sp.; Saxicola; Ruticilla, 2 sp.; Sylvia, 3 sp.; Parus, 3 sp.; Mecistura; Anorthura; Certhia; Sitta; Yunx; Muscicapa, 2 sp.; Hirundo, 3 sp.; and Cypselus. The females of the following 12 birds may be considered as conspicuous according to the same standard, viz., Pastor, Motacilla alba, Parus major and P. caeruleus, Upupa, Picus, 4 sp., Coracias, Alcedo, and Merops.) Nor is there any close relation within the same genus between a well-pronounced difference in colour between the sexes, and the nature of the nest constructed. Thus the male house sparrow (Passer domesticus) differs much from the female, the male tree-sparrow (P. montanus) hardly at all, and yet both build well-concealed nests. The two sexes of the common fly-catcher (Muscicapa grisola) can hardly be distinguished, whilst the sexes of the pied fly-catcher (M. luctuosa) differ considerably, and both species build in holes or conceal their nests. The female blackbird (Turdus merula) differs much, the female ring-ouzel (T. torquatus) differs less, and the female common thrush (T. musicus) hardly at all from their respective males; yet all build open nests. On the other hand, the not very distantly-allied water-ouzel (Cinclus aquaticus) builds a domed nest, and the sexes differ about as much as in the ring-ouzel. The black and red grouse (Tetrao tetrix and T. scoticus) build open nests in equally well-concealed spots, but in the one species the sexes differ greatly, and in the other very little.

Notwithstanding the foregoing objections, I cannot doubt, after reading Mr. Wallace’s excellent essay, that looking to the birds of the world, a large majority of the species in which the females are conspicuously coloured (and in this case the males with rare exceptions are equally conspicuous), build concealed nests for the sake of protection. Mr. Wallace enumerates (19. ‘Journal of Travel,’ edited by A. Murray, vol. i. p. 78.) a long series of groups in which this rule holds good; but it will suffice here to give, as instances, the more familiar groups of kingfishers, toucans, trogons, puff-birds (Capitonidae), plantain-eaters (Musophagae, woodpeckers, and parrots. Mr. Wallace believes that in these groups, as the males gradually acquired through sexual selection their brilliant colours, these were transferred to the females and were not eliminated by natural selection, owing to the protection which they already enjoyed from their manner of nidification. According to this view, their present manner of nesting was acquired before their present colours. But it seems to me much more probable that in most cases, as the females were gradually rendered more and more brilliant from partaking of the colours of the male, they were gradually led to change their instincts (supposing that they originally built open nests), and to seek protection by building domed or concealed nests. No one who studies, for instance, Audubon’s account of the differences in the nests of the same species in the Northern and Southern United States (20. See many statements in the ‘Ornithological Biography.’ See also some curious observations on the nests of Italian birds by Eugenio Bettoni, in the ‘Atti della Società Italiana,’ vol. xi. 1869, p. 487.), will feel any great difficulty in admitting that birds, either by a change (in the strict sense of the word) of their habits, or through the natural selection of so-called spontaneous variations of instinct, might readily be led to modify their manner of nesting.

This way of viewing the relation, as far as it holds good, between the bright colours of female birds and their manner of nesting, receives some support from certain cases occurring in the Sahara Desert. Here, as in most other deserts, various birds, and many other animals, have had their colours adapted in a wonderful manner to the tints of the surrounding surface. Nevertheless there are, as I am informed by the Rev. Mr. Tristram, some curious exceptions to the rule; thus the male of the Monticola cyanea is conspicuous from his bright blue colour, and the female almost equally conspicuous from her mottled brown and white plumage; both sexes of two species of Dromolaea are of a lustrous black; so that these three species are far from receiving protection from their colours, yet they are able to survive, for they have acquired the habit of taking refuge from danger in holes or crevices in the rocks.

With respect to the above groups in which the females are conspicuously coloured and build concealed nests, it is not necessary to suppose that each separate species had its nidifying instinct specially modified; but only that the early progenitors of each group were gradually led to build domed or concealed nests, and afterwards transmitted this instinct, together with their bright colours, to their modified descendants. As far as it can be trusted, the conclusion is interesting, that sexual selection together with equal or nearly equal inheritance by both sexes, have indirectly determined the manner of nidification of whole groups of birds.

According to Mr. Wallace, even in the groups in which the females, from being protected in domed nests during incubation, have not had their bright colours eliminated through natural selection, the males often differ in a slight, and occasionally in a considerable degree from the females. This is a significant fact, for such differences in colour must be accounted for by some of the variations in the males having been from the first limited in transmission to the same sex; as it can hardly be maintained that these differences, especially when very slight, serve as a protection to the female. Thus all the species in the splendid group of the Trogons build in holes; and Mr. Gould gives figures (21. See his Monograph of the Trogonidae, 1st edition.) of both sexes of twenty-five species, in all of which, with one partial exception, the sexes differ sometimes slightly, sometimes conspicuously, in colour,—the males being always finer than the females, though the latter are likewise beautiful. All the species of kingfishers build in holes, and with most of the species the sexes are equally brilliant, and thus far Mr. Wallace’s rule holds good; but in some of the Australian species the colours of the females are rather less vivid than those of the male; and in one splendidly-coloured species, the sexes differ so much that they were at first thought to be specifically distinct. (22. Namely, Cyanalcyon, Gould’s ‘Handbook to the Birds of Australia,’ vol. i. p. 133; see, also, pp. 130, 136.) Mr. R.B. Sharpe, who has especially studied this group, has shewn me some American species (Ceryle) in which the breast of the male is belted with black. Again, in Carcineutes, the difference between the sexes is conspicuous: in the male the upper surface is dull-blue banded with black, the lower surface being partly fawn-coloured, and there is much red about the head; in the female the upper surface is reddish-brown banded with black, and the lower surface white with black markings. It is an interesting fact, as shewing how the same peculiar style of sexual colouring often characterises allied forms, that in three species of Dacelo the male differs from the female only in the tail being dull-blue banded with black, whilst that of the female is brown with blackish bars; so that here the tail differs in colour in the two sexes in exactly the same manner as the whole upper surface in the two sexes of Carcineutes.

With parrots, which likewise build in holes, we find analogous cases: in most of the species, both sexes are brilliantly coloured and indistinguishable, but in not a few species the males are coloured rather more vividly than the females, or even very differently from them. Thus, besides other strongly-marked differences, the whole under surface of the male King Lory (Aprosmictus scapulatus) is scarlet, whilst the throat and chest of the female is green tinged with red: in the Euphema splendida there is a similar difference, the face and wing coverts moreover of the female being of a paler blue than in the male. (23. Every gradation of difference between the sexes may be followed in the parrots of Australia. See Gould’s ‘Handbook,’ etc., vol. ii. pp. 14-102.) In the family of the tits (Parinae), which build concealed nests, the female of our common blue tomtit (Parus caeruleus), is “much less brightly coloured” than the male: and in the magnificent Sultan yellow tit of India the difference is greater. (24. Macgillivray’s ‘British Birds,’ vol. ii. p. 433. Jerdon, ‘Birds of India,’ vol. ii. p. 282.)

Again, in the great group of the woodpeckers (25. All the following facts are taken from M. Malherbe’s magnificent ‘Monographie des Picidees,’ 1861.), the sexes are generally nearly alike, but in the Megapicus validus all those parts of the head, neck, and breast, which are crimson in the male are pale brown in the female. As in several woodpeckers the head of the male is bright crimson, whilst that of the female is plain, it occurred to me that this colour might possibly make the female dangerously conspicuous, whenever she put her head out of the hole containing her nest, and consequently that this colour, in accordance with Mr. Wallace’s belief, had been eliminated. This view is strengthened by what Malherbe states with respect to Indopicus carlotta; namely, that the young females, like the young males, have some crimson about their heads, but that this colour disappears in the adult female, whilst it is intensified in the adult male. Nevertheless the following considerations render this view extremely doubtful: the male takes a fair share in incubation (26. Audubon’s ‘Ornithological Biography,’ vol. ii. p. 75; see also the ‘Ibis,’ vol. i. p. 268.), and would be thus almost equally exposed to danger; both sexes of many species have their heads of an equally bright crimson; in other species the difference between the sexes in the amount of scarlet is so slight that it can hardly make any appreciable difference in the danger incurred; and lastly, the colouring of the head in the two sexes often differs slightly in other ways.

The cases, as yet given, of slight and graduated differences in colour between the males and females in the groups, in which as a general rule the sexes resemble each other, all relate to species which build domed or concealed nests. But similar gradations may likewise be observed in groups in which the sexes as a general rule resemble each other, but which build open nests.

As I have before instanced the Australian parrots, so I may here instance, without giving any details, the Australian pigeons. (27. Gould’s ‘Handbook to the Birds of Australia,’ vol. ii. pp. 109-149.) It deserves especial notice that in all these cases the slight differences in plumage between the sexes are of the same general nature as the occasionally greater differences. A good illustration of this fact has already been afforded by those kingfishers in which either the tail alone or the whole upper surface of the plumage differs in the same manner in the two sexes. Similar cases may be observed with parrots and pigeons. The differences in colour between the sexes of the same species are, also, of the same general nature as the differences in colour between the distinct species of the same group. For when in a group in which the sexes are usually alike, the male differs considerably from the female, he is not coloured in a quite new style. Hence we may infer that within the same group the special colours of both sexes when they are alike, and the colours of the male, when he differs slightly or even considerably from the female, have been in most cases determined by the same general cause; this being sexual selection.

It is not probable, as has already been remarked, that differences in colour between the sexes, when very slight, can be of service to the female as a protection. Assuming, however, that they are of service, they might be thought to be cases of transition; but we have no reason to believe that many species at any one time are undergoing change. Therefore we can hardly admit that the numerous females which differ very slightly in colour from their males are now all commencing to become obscure for the sake of protection. Even if we consider somewhat more marked sexual differences, is it probable, for instance, that the head of the female chaffinch,—the crimson on the breast of the female bullfinch,—the green of the female greenfinch,—the crest of the female golden-crested wren, have all been rendered less bright by the slow process of selection for the sake of protection? I cannot think so; and still less with the slight differences between the sexes of those birds which build concealed nests. On the other hand, the differences in colour between the sexes, whether great or small, may to a large extent be explained on the principle of the successive variations, acquired by the males through sexual selection, having been from the first more or less limited in their transmission to the females. That the degree of limitation should differ in different species of the same group will not surprise any one who has studied the laws of inheritance, for they are so complex that they appear to us in our ignorance to be capricious in their action. (28. See remarks to this effect in ‘Variation of Animals and Plants under Domestication,’ vol. ii. chap. xii.)

As far as I can discover there are few large groups of birds in which all the species have both sexes alike and brilliantly coloured, but I hear from Mr. Sclater, that this appears to be the case with the Musophagae or plantain-eaters. Nor do I believe that any large group exists in which the sexes of all the species are widely dissimilar in colour: Mr. Wallace informs me that the chatterers of S. America (Cotingidae) offer one of the best instances; but with some of the species, in which the male has a splendid red breast, the female exhibits some red on her breast; and the females of other species shew traces of the green and other colours of the males. Nevertheless we have a near approach to close sexual similarity or dissimilarity throughout several groups: and this, from what has just been said of the fluctuating nature of inheritance, is a somewhat surprising circumstance. But that the same laws should largely prevail with allied animals is not surprising. The domestic fowl has produced a great number of breeds and sub-breeds, and in these the sexes generally differ in plumage; so that it has been noticed as an unusual circumstance when in certain sub-breeds they resemble each other. On the other hand, the domestic pigeon has likewise produced a vast number of distinct breeds and sub-breeds, and in these, with rare exceptions, the two sexes are identically alike.

Therefore if other species of Gallus and Columba were domesticated and varied, it would not be rash to predict that similar rules of sexual similarity and dissimilarity, depending on the form of transmission, would hold good in both cases. In like manner the same form of transmission has generally prevailed under nature throughout the same groups, although marked exceptions to this rule occur. Thus within the same family or even genus, the sexes may be identically alike, or very different in colour. Instances have already been given in the same genus, as with sparrows, fly-catchers, thrushes and grouse. In the family of pheasants the sexes of almost all the species are wonderfully dissimilar, but are quite alike in the eared pheasant or Crossoptilon auritum. In two species of Chloephaga, a genus of geese, the male cannot be distinguished from the females, except by size; whilst in two others, the sexes are so unlike that they might easily be mistaken for distinct species. (29. The ‘Ibis,’ vol. vi. 1864, p. 122.)

The laws of inheritance can alone account for the following cases, in which the female acquires, late in life, certain characters proper to the male, and ultimately comes to resemble him more or less completely. Here protection can hardly have come into play. Mr. Blyth informs me that the females of Oriolus melanocephalus and of some allied species, when sufficiently mature to breed, differ considerably in plumage from the adult males; but after the second or third moults they differ only in their beaks having a slight greenish tinge. In the dwarf bitterns (Ardetta), according to the same authority, “the male acquires his final livery at the first moult, the female not before the third or fourth moult; in the meanwhile she presents an intermediate garb, which is ultimately exchanged for the same livery as that of the male.” So again the female Falco peregrinus acquires her blue plumage more slowly than the male. Mr. Swinhoe states that with one of the Drongo shrikes (Dicrurus macrocercus) the male, whilst almost a nestling, moults his soft brown plumage and becomes of a uniform glossy greenish-black; but the female retains for a long time the white striae and spots on the axillary feathers; and does not completely assume the uniform black colour of the male for three years. The same excellent observer remarks that in the spring of the second year the female spoon-bill (Platalea) of China resembles the male of the first year, and that apparently it is not until the third spring that she acquires the same adult plumage as that possessed by the male at a much earlier age. The female Bombycilla carolinensis differs very little from the male, but the appendages, which like beads of red sealing-wax ornament the wing-feathers (30. When the male courts the female, these ornaments are vibrated, and “are shewn off to great advantage,” on the outstretched wings: A. Leith Adams, ‘Field and Forest Rambles,’ 1873, p. 153.), are not developed in her so early in life as in the male. In the male of an Indian parrakeet (Palaeornis javanicus) the upper mandible is coral-red from his earliest youth, but in the female, as Mr. Blyth has observed with caged and wild birds, it is at first black and does not become red until the bird is at least a year old, at which age the sexes resemble each other in all respects. Both sexes of the wild turkey are ultimately furnished with a tuft of bristles on the breast, but in two-year-old birds the tuft is about four inches long in the male and hardly apparent in the female; when, however, the latter has reached her fourth year, it is from four to five inches in length. (31. On Ardetta, Translation of Cuvier’s ‘Regne Animal,’ by Mr. Blyth, footnote, p. 159. On the Peregrine Falcon, Mr. Blyth, in Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 304. On Dicrurus, ‘Ibis,’ 1863, p. 44. On the Platalea, ‘Ibis,’ vol. vi. 1864, p. 366. On the Bombycilla, Audubon’s ‘Ornitholog. Biography,’ vol. i. p. 229. On the Palaeornis, see, also, Jerdon, ‘Birds of India,’ vol. i. p. 263. On the wild turkey, Audubon, ibid. vol. i. p. 15; but I hear from Judge Caton that in Illinois the female very rarely acquires a tuft. Analogous cases with the females of Petrocossyphus are given by Mr. R. Sharpe, ‘Proceedings of the Zoological Society,’ 1872, p. 496.)

These cases must not be confounded with those where diseased or old females abnormally assume masculine characters, nor with those where fertile females, whilst young, acquire the characters of the male, through variation or some unknown cause. (32. Of these latter cases Mr. Blyth has recorded (Translation of Cuvier’s ‘Regne Animal,’ p. 158) various instances with Lanius, Ruticilla, Linaria, and Anas. Audubon has also recorded a similar case (‘Ornitholog. Biography,’ vol. v. p. 519) with Pyranga aestiva.) But all these cases have so much in common that they depend, according to the hypothesis of pangenesis, on gemmules derived from each part of the male being present, though latent, in the female; their development following on some slight change in the elective affinities of her constituent tissues.

A few words must be added on changes of plumage in relation to the season of the year. From reasons formerly assigned there can be little doubt that the elegant plumes, long pendant feathers, crests, etc., of egrets, herons, and many other birds, which are developed and retained only during the summer, serve for ornamental and nuptial purposes, though common to both sexes. The female is thus rendered more conspicuous during the period of incubation than during the winter; but such birds as herons and egrets would be able to defend themselves. As, however, plumes would probably be inconvenient and certainly of no use during the winter, it is possible that the habit of moulting twice in the year may have been gradually acquired through natural selection for the sake of casting off inconvenient ornaments during the winter. But this view cannot be extended to the many waders, whose summer and winter plumages differ very little in colour. With defenceless species, in which both sexes, or the males alone, become extremely conspicuous during the breeding-season,—or when the males acquire at this season such long wing or tail-feathers as to impede their flight, as with Cosmetornis and Vidua,—it certainly at first appears highly probable that the second moult has been gained for the special purpose of throwing off these ornaments. We must, however, remember that many birds, such as some of the Birds of Paradise, the Argus pheasant and peacock, do not cast their plumes during the winter; and it can hardly be maintained that the constitution of these birds, at least of the Gallinaceae, renders a double moult impossible, for the ptarmigan moults thrice in the year. (33. See Gould’s ‘Birds of Great Britain.’) Hence it must be considered as doubtful whether the many species which moult their ornamental plumes or lose their bright colours during the winter, have acquired this habit on account of the inconvenience or danger which they would otherwise have suffered.

I conclude, therefore, that the habit of moulting twice in the year was in most or all cases first acquired for some distinct purpose, perhaps for gaining a warmer winter covering; and that variations in the plumage occurring during the summer were accumulated through sexual selection, and transmitted to the offspring at the same season of the year; that such variations were inherited either by both sexes or by the males alone, according to the form of inheritance which prevailed. This appears more probable than that the species in all cases originally tended to retain their ornamental plumage during the winter, but were saved from this through natural selection, resulting from the inconvenience or danger thus caused.

I have endeavoured in this chapter to shew that the arguments are not trustworthy in favour of the view that weapons, bright colours, and various ornaments, are now confined to the males owing to the conversion, by natural selection, of the equal transmission of characters to both sexes, into transmission to the male sex alone. It is also doubtful whether the colours of many female birds are due to the preservation, for the sake of protection, of variations which were from the first limited in their transmission to the female sex. But it will be convenient to defer any further discussion on this subject until I treat, in the following chapter, of the differences in plumage between the young and old.

Chapter XVI

CHAPTER XVI. BIRDS—concluded.

The immature plumage in relation to the character of the plumage in both sexes when adult—Six classes of cases—Sexual differences between the males of closely-allied or representative species—The female assuming the characters of the male—Plumage of the young in relation to the summer and winter plumage of the adults—On the increase of beauty in the birds of the world—Protective colouring—Conspicuously coloured birds—Novelty appreciated—Summary of the four chapters on Birds.

We must now consider the transmission of characters, as limited by age, in reference to sexual selection. The truth and importance of the principle of inheritance at corresponding ages need not here be discussed, as enough has already been said on the subject. Before giving the several rather complex rules or classes of cases, under which the differences in plumage between the young and the old, as far as known to me, may be included, it will be well to make a few preliminary remarks.

With animals of all kinds when the adults differ in colour from the young, and the colours of the latter are not, as far as we can see, of any special service, they may generally be attributed, like various embryological structures, to the retention of a former character. But this view can be maintained with confidence, only when the young of several species resemble each other closely, and likewise resemble other adult species belonging to the same group; for the latter are the living proofs that such a state of things was formerly possible. Young lions and pumas are marked with feeble stripes or rows of spots, and as many allied species both young and old are similarly marked, no believer in evolution will doubt that the progenitor of the lion and puma was a striped animal, and that the young have retained vestiges of the stripes, like the kittens of black cats, which are not in the least striped when grown up. Many species of deer, which when mature are not spotted, are whilst young covered with white spots, as are likewise some few species in the adult state. So again the young in the whole family of pigs (Suidae), and in certain rather distantly allied animals, such as the tapir, are marked with dark longitudinal stripes; but here we have a character apparently derived from an extinct progenitor, and now preserved by the young alone. In all such cases the old have had their colours changed in the course of time, whilst the young have remained but little altered, and this has been effected through the principle of inheritance at corresponding ages.

This same principle applies to many birds belonging to various groups, in which the young closely resemble each other, and differ much from their respective adult parents. The young of almost all the Gallinaceae, and of some distantly allied birds such as ostriches, are covered with longitudinally striped down; but this character points back to a state of things so remote that it hardly concerns us. Young cross-bills (Loxia) have at first straight beaks like those of other finches, and in their immature striated plumage they resemble the mature red-pole and female siskin, as well as the young of the goldfinch, greenfinch, and some other allied species. The young of many kinds of buntings (Emberiza) resemble one another, and likewise the adult state of the common bunting, E. miliaria. In almost the whole large group of thrushes the young have their breasts spotted—a character which is retained throughout life by many species, but is quite lost by others, as by the Turdus migratorius. So again with many thrushes, the feathers on the back are mottled before they are moulted for the first time, and this character is retained for life by certain eastern species. The young of many species of shrikes (Lanius), of some woodpeckers, and of an Indian pigeon (Chalcophaps indicus), are transversely striped on the under surface; and certain allied species or whole genera are similarly marked when adult. In some closely-allied and resplendent Indian cuckoos (Chrysococcyx), the mature species differ considerably from one another in colour, but the young cannot be distinguished. The young of an Indian goose (Sarkidiornis melanonotus) closely resemble in plumage an allied genus, Dendrocygna, when mature. (1. In regard to thrushes, shrikes, and woodpeckers, see Mr. Blyth, in Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 304; also footnote to his translation of Cuvier’s ‘Regne Animal,’ p. 159. I give the case of Loxia on Mr. Blyth’s information. On thrushes, see also Audubon, ‘Ornith. Biog.’ vol. ii. p. 195. On Chrysococcyx and Chalcophaps, Blyth, as quoted in Jerdon’s ‘Birds of India,’ vol. iii. p. 485. On Sarkidiornis, Blyth, in ‘Ibis,’ 1867, p. 175.) Similar facts will hereafter be given in regard to certain herons. Young black-grouse (Tetrao tetrix) resemble the young as well as the old of certain other species, for instance the red-grouse or T. scoticus. Finally, as Mr. Blyth, who has attended closely to this subject, has well remarked, the natural affinities of many species are best exhibited in their immature plumage; and as the true affinities of all organic beings depend on their descent from a common progenitor, this remark strongly confirms the belief that the immature plumage approximately shews us the former or ancestral condition of the species.

Although many young birds, belonging to various families, thus give us a glimpse of the plumage of their remote progenitors, yet there are many other birds, both dull-coloured and bright-coloured, in which the young closely resemble their parents. In such cases the young of the different species cannot resemble each other more closely than do the parents; nor can they strikingly resemble allied forms when adult. They give us but little insight into the plumage of their progenitors, excepting in so far that, when the young and the old are coloured in the same general manner throughout a whole group of species, it is probable that their progenitors were similarly coloured.

We may now consider the classes of cases, under which the differences and resemblances between the plumage of the young and the old, in both sexes or in one sex alone, may be grouped. Rules of this kind were first enounced by Cuvier; but with the progress of knowledge they require some modification and amplification. This I have attempted to do, as far as the extreme complexity of the subject permits, from information derived from various sources; but a full essay on this subject by some competent ornithologist is much needed. In order to ascertain to what extent each rule prevails, I have tabulated the facts given in four great works, namely, by Macgillivray on the birds of Britain, Audubon on those of North America, Jerdon on those of India, and Gould on those of Australia. I may here premise, first, that the several cases or rules graduate into each other; and secondly, that when the young are said to resemble their parents, it is not meant that they are identically alike, for their colours are almost always less vivid, and the feathers are softer and often of a different shape.

RULES OR CLASSES OF CASES.

I. When the adult male is more beautiful or conspicuous than the adult female, the young of both sexes in their first plumage closely resemble the adult female, as with the common fowl and peacock; or, as occasionally occurs, they resemble her much more closely than they do the adult male.

II. When the adult female is more conspicuous than the adult male, as sometimes though rarely occurs, the young of both sexes in their first plumage resemble the adult male.

III. When the adult male resembles the adult female, the young of both sexes have a peculiar first plumage of their own, as with the robin.

IV. When the adult male resembles the adult female, the young of both sexes in their first plumage resemble the adults, as with the kingfisher, many parrots, crows, hedge-warblers.

V. When the adults of both sexes have a distinct winter and summer plumage, whether or not the male differs from the female, the young resemble the adults of both sexes in their winter dress, or much more rarely in their summer dress, or they resemble the females alone. Or the young may have an intermediate character; or again they may differ greatly from the adults in both their seasonal plumages.

VI. In some few cases the young in their first plumage differ from each other according to sex; the young males resembling more or less closely the adult males, and the young females more or less closely the adult females.

CLASS I. — In this class, the young of both sexes more or less closely resemble the adult female, whilst the adult male differs from the adult female, often in the most conspicuous manner. Innumerable instances in all Orders could be given; it will suffice to call to mind the common pheasant, duck, and house-sparrow. The cases under this class graduate into others. Thus the two sexes when adult may differ so slightly, and the young so slightly from the adults, that it is doubtful whether such cases ought to come under the present, or under the third or fourth classes. So again the young of the two sexes, instead of being quite alike, may differ in a slight degree from each other, as in our sixth class. These transitional cases, however, are few, or at least are not strongly pronounced, in comparison with those which come strictly under the present class.

The force of the present law is well shewn in those groups, in which, as a general rule, the two sexes and the young are all alike; for when in these groups the male does differ from the female, as with certain parrots, kingfishers, pigeons, etc., the young of both sexes resemble the adult female. (2. See, for instance, Mr. Gould’s account (‘Handbook to the Birds of Australia,’ vol. i. p. 133) of Cyanalcyon (one of the Kingfishers), in which, however, the young male, though resembling the adult female, is less brilliantly coloured. In some species of Dacelo the males have blue tails, and the females brown ones; and Mr. R.B. Sharpe informs me that the tail of the young male of D. gaudichaudi is at first brown. Mr. Gould has described (ibid. vol. ii. pp. 14, 20, 37) the sexes and the young of certain black Cockatoos and of the King Lory, with which the same rule prevails. Also Jerdon (‘Birds of India,’ vol. i. p. 260) on the Palaeornis rosa, in which the young are more like the female than the male. See Audubon (‘Ornithological Biography,’ vol. ii. p. 475) on the two sexes and the young of Columba passerina.) We see the same fact exhibited still more clearly in certain anomalous cases; thus the male of Heliothrix auriculata (one of the humming-birds) differs conspicuously from the female in having a splendid gorget and fine ear-tufts, but the female is remarkable from having a much longer tail than that of the male; now the young of both sexes resemble (with the exception of the breast being spotted with bronze) the adult female in all other respects, including the length of her tail, so that the tail of the male actually becomes shorter as he reaches maturity, which is a most unusual circumstance. (3. I owe this information to Mr. Gould, who shewed me the specimens; see also his ‘Introduction to the Trochilidae,’ 1861, p. 120.) Again, the plumage of the male goosander (Mergus merganser) is more conspicuously coloured than that of the female, with the scapular and secondary wing-feathers much longer; but differently from what occurs, as far as I know, in any other bird, the crest of the adult male, though broader than that of the female, is considerably shorter, being only a little above an inch in length; the crest of the female being two and a half inches long. Now the young of both sexes entirely resemble the adult female, so that their crests are actually of greater length, though narrower, than in the adult male. (4. Macgillivray, ‘Hist. Brit. Birds,’ vol. v. pp. 207-214.)

When the young and the females closely resemble each other and both differ from the males, the most obvious conclusion is that the males alone have been modified. Even in the anomalous cases of the Heliothrix and Mergus, it is probable that originally both adult sexes were furnished—the one species with a much elongated tail, and the other with a much elongated crest—these characters having since been partially lost by the adult males from some unexplained cause, and transmitted in their diminished state to their male offspring alone, when arrived at the corresponding age of maturity. The belief that in the present class the male alone has been modified, as far as the differences between the male and the female together with her young are concerned, is strongly supported by some remarkable facts recorded by Mr. Blyth (5. See his admirable paper in the ‘Journal of the Asiatic Soc. of Bengal,’ vol. xix. 1850, p. 223; see also Jerdon, ‘Birds of India,’ vol. i. introduction, p. xxix. In regard to Tanysiptera, Prof. Schlegel told Mr. Blyth that he could distinguish several distinct races, solely by comparing the adult males.), with respect to closely-allied species which represent each other in distinct countries. For with several of these representative species the adult males have undergone a certain amount of change and can be distinguished; the females and the young from the distinct countries being indistinguishable, and therefore absolutely unchanged. This is the case with certain Indian chats (Thamnobia), with certain honey-suckers (Nectarinia), shrikes (Tephrodornis), certain kingfishers (Tanysiptera), Kalij pheasants (Gallophasis), and tree-partridges (Arboricola).

In some analogous cases, namely with birds having a different summer and winter plumage, but with the two sexes nearly alike, certain closely-allied species can easily be distinguished in their summer or nuptial plumage, yet are indistinguishable in their winter as well as in their immature plumage. This is the case with some of the closely-allied Indian wagtails or Motacillae. Mr. Swinhoe (6. See also Mr. Swinhoe, in ‘Ibis,’ July 1863, p. 131; and a previous paper, with an extract from a note by Mr. Blyth, in ‘Ibis,’ January, 1861, p. 25.) informs me that three species of Ardeola, a genus of herons, which represent one another on separate continents, are “most strikingly different” when ornamented with their summer plumes, but are hardly, if at all, distinguishable during the winter. The young also of these three species in their immature plumage closely resemble the adults in their winter dress. This case is all the more interesting, because with two other species of Ardeola both sexes retain, during the winter and summer, nearly the same plumage as that possessed by the three first species during the winter and in their immature state; and this plumage, which is common to several distinct species at different ages and seasons, probably shews us how the progenitors of the genus were coloured. In all these cases, the nuptial plumage which we may assume was originally acquired by the adult males during the breeding-season, and transmitted to the adults of both sexes at the corresponding season, has been modified, whilst the winter and immature plumages have been left unchanged.

The question naturally arises, how is it that in these latter cases the winter plumage of both sexes, and in the former cases the plumage of the adult females, as well as the immature plumage of the young, have not been at all affected? The species which represent each other in distinct countries will almost always have been exposed to somewhat different conditions, but we can hardly attribute to this action the modification of the plumage in the males alone, seeing that the females and the young, though similarly exposed, have not been affected. Hardly any fact shews us more clearly how subordinate in importance is the direct action of the conditions of life, in comparison with the accumulation through selection of indefinite variations, than the surprising difference between the sexes of many birds; for both will have consumed the same food, and have been exposed to the same climate. Nevertheless we are not precluded from believing that in the course of time new conditions may produce some direct effect either on both sexes, or from their constitutional differences chiefly on one sex. We see only that this is subordinate in importance to the accumulated results of selection. Judging, however, from a wide-spread analogy, when a species migrates into a new country (and this must precede the formation of representative species), the changed conditions to which they will almost always have been exposed will cause them to undergo a certain amount of fluctuating variability. In this case sexual selection, which depends on an element liable to change—the taste or admiration of the female—will have had new shades of colour or other differences to act on and accumulate; and as sexual selection is always at work, it would (from what we know of the results on domestic animals of man’s unintentional selection), be surprising if animals inhabiting separate districts, which can never cross and thus blend their newly-acquired characters, were not, after a sufficient lapse of time, differently modified. These remarks likewise apply to the nuptial or summer plumage, whether confined to the males, or common to both sexes.

Although the females of the above closely-allied or representative species, together with their young, differ hardly at all from one another, so that the males alone can be distinguished, yet the females of most species within the same genus obviously differ from each other. The differences, however, are rarely as great as between the males. We see this clearly in the whole family of the Gallinaceae: the females, for instance, of the common and Japan pheasant, and especially of the gold and Amherst pheasant —of the silver pheasant and the wild fowl—resemble one another very closely in colour, whilst the males differ to an extraordinary degree. So it is with the females of most of the Cotingidae, Fringillidae, and many other families. There can indeed be no doubt that, as a general rule, the females have been less modified than the males. Some few birds, however, offer a singular and inexplicable exception; thus the females of Paradisea apoda and P. papuana differ from each other more than do their respective males (7. Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 394.); the female of the latter species having the under surface pure white, whilst the female P. apoda is deep brown beneath. So, again, as I hear from Professor Newton, the males of two species of Oxynotus (shrikes), which represent each other in the islands of Mauritius and Bourbon (8. These species are described with coloured figures, by M. F. Pollen, in ‘Ibis,’ 1866, p. 275.), differ but little in colour, whilst the females differ much. In the Bourbon species the female appears to have partially retained an immature condition of plumage, for at first sight she “might be taken for the young of the Mauritian species.” These differences may be compared with those inexplicable ones, which occur independently of man’s selection in certain sub-breeds of the game-fowl, in which the females are very different, whilst the males can hardly be distinguished. (9. ‘Variation of Animals,’ etc., vol. i. p. 251.)

As I account so largely by sexual selection for the differences between the males of allied species, how can the differences between the females be accounted for in all ordinary cases? We need not here consider the species which belong to distinct genera; for with these, adaptation to different habits of life, and other agencies, will have come into play. In regard to the differences between the females within the same genus, it appears to me almost certain, after looking through various large groups, that the chief agent has been the greater or less transference to the female of the characters acquired by the males through sexual selection. In the several British finches, the two sexes differ either very slightly or considerably; and if we compare the females of the greenfinch, chaffinch, goldfinch, bullfinch, crossbill, sparrow, etc., we shall see that they differ from one another chiefly in the points in which they partially resemble their respective males; and the colours of the males may safely be attributed to sexual selection. With many gallinaceous species the sexes differ to an extreme degree, as with the peacock, pheasant, and fowl, whilst with other species there has been a partial or even complete transference of character from the male to the female. The females of the several species of Polyplectron exhibit in a dim condition, and chiefly on the tail, the splendid ocelli of their males. The female partridge differs from the male only in the red mark on her breast being smaller; and the female wild turkey only in her colours being much duller. In the guinea-fowl the two sexes are indistinguishable. There is no improbability in the plain, though peculiarly spotted plumage of this latter bird having been acquired through sexual selection by the males, and then transmitted to both sexes; for it is not essentially different from the much more beautifully spotted plumage, characteristic of the males alone of the Tragopan pheasants.

It should be observed that, in some instances, the transference of characters from the male to the female has been effected apparently at a remote period, the male having subsequently undergone great changes, without transferring to the female any of his later-gained characters. For instance, the female and the young of the black-grouse (Tetrao tetrix) resemble pretty closely both sexes and the young of the red-grouse (T. scoticus); and we may consequently infer that the black-grouse is descended from some ancient species, of which both sexes were coloured in nearly the same manner as the red-grouse. As both sexes of this latter species are more distinctly barred during the breeding-season than at any other time, and as the male differs slightly from the female in his more strongly-pronounced red and brown tints (10. Macgillivray, ‘History of British Birds,’ vol. i. pp. 172-174.), we may conclude that his plumage has been influenced by sexual selection, at least to a certain extent. If so, we may further infer that nearly similar plumage of the female black-grouse was similarly produced at some former period. But since this period the male black-grouse has acquired his fine black plumage, with his forked and outwardly-curled tail-feathers; but of these characters there has hardly been any transference to the female, excepting that she shews in her tail a trace of the curved fork.

We may therefore conclude that the females of distinct though allied species have often had their plumage rendered more or less different by the transference in various degrees of characters acquired by the males through sexual selection, both during former and recent times. But it deserves especial attention that brilliant colours have been transferred much more rarely than other tints. For instance, the male of the red-throated blue-breast (Cyanecula suecica) has a rich blue breast, including a sub-triangular red mark; now marks of nearly the same shape have been transferred to the female, but the central space is fulvous instead of red, and is surrounded by mottled instead of blue feathers. The Gallinaceae offer many analogous cases; for none of the species, such as partridges, quails, guinea-fowls, etc., in which the colours of the plumage have been largely transferred from the male to the female, are brilliantly coloured. This is well exemplified with the pheasants, in which the male is generally so much more brilliant than the female; but with the Eared and Cheer pheasants (Crossoptilon auritum and Phasianus wallichii) the sexes closely resemble each other and their colours are dull. We may go so far as to believe that if any part of the plumage in the males of these two pheasants had been brilliantly coloured, it would not have been transferred to the females. These facts strongly support Mr. Wallace’s view that with birds which are exposed to much danger during incubation, the transference of bright colours from the male to the female has been checked through natural selection. We must not, however, forget that another explanation, before given, is possible; namely, that the males which varied and became bright, whilst they were young and inexperienced, would have been exposed to much danger, and would generally have been destroyed; the older and more cautious males, on the other hand, if they varied in a like manner, would not only have been able to survive, but would have been favoured in their rivalry with other males. Now variations occurring late in life tend to be transmitted exclusively to the same sex, so that in this case extremely bright tints would not have been transmitted to the females. On the other hand, ornaments of a less conspicuous kind, such as those possessed by the Eared and Cheer pheasants, would not have been dangerous, and if they appeared during early youth, would generally have been transmitted to both sexes.

In addition to the effects of the partial transference of characters from the males to the females, some of the differences between the females of closely allied species may be attributed to the direct or definite action of the conditions of life. (11. See, on this subject, chap. xxiii. in the ‘Variation of Animals and Plants under Domestication.’) With the males, any such action would generally have been masked by the brilliant colours gained through sexual selection; but not so with the females. Each of the endless diversities in plumage which we see in our domesticated birds is, of course, the result of some definite cause; and under natural and more uniform conditions, some one tint, assuming that it was in no way injurious, would almost certainly sooner or later prevail. The free intercrossing of the many individuals belonging to the same species would ultimately tend to make any change of colour, thus induced, uniform in character.

No one doubts that both sexes of many birds have had their colours adapted for the sake of protection; and it is possible that the females alone of some species may have been modified for this end. Although it would be a difficult, perhaps an impossible process, as shewn in the last chapter, to convert one form of transmission into another through selection, there would not be the least difficulty in adapting the colours of the female, independently of those of the male, to surrounding objects, through the accumulation of variations which were from the first limited in their transmission to the female sex. If the variations were not thus limited, the bright tints of the male would be deteriorated or destroyed. Whether the females alone of many species have been thus specially modified, is at present very doubtful. I wish I could follow Mr. Wallace to the full extent; for the admission would remove some difficulties. Any variations which were of no service to the female as a protection would be at once obliterated, instead of being lost simply by not being selected, or from free intercrossing, or from being eliminated when transferred to the male and in any way injurious to him. Thus the plumage of the female would be kept constant in character. It would also be a relief if we could admit that the obscure tints of both sexes of many birds had been acquired and preserved for the sake of protection,—for example, of the hedge-warbler or kitty-wren (Accentor modularis and Troglodytes vulgaris), with respect to which we have no sufficient evidence of the action of sexual selection. We ought, however, to be cautious in concluding that colours which appear to us dull, are not attractive to the females of certain species; we should bear in mind such cases as that of the common house-sparrow, in which the male differs much from the female, but does not exhibit any bright tints. No one probably will dispute that many gallinaceous birds which live on the open ground, have acquired their present colours, at least in part, for the sake of protection. We know how well they are thus concealed; we know that ptarmigans, whilst changing from their winter to their summer plumage, both of which are protective, suffer greatly from birds of prey. But can we believe that the very slight differences in tints and markings between, for instance, the female black-grouse and red-grouse serve as a protection? Are partridges, as they are now coloured, better protected than if they had resembled quails? Do the slight differences between the females of the common pheasant, the Japan and gold pheasants, serve as a protection, or might not their plumages have been interchanged with impunity? From what Mr. Wallace has observed of the habits of certain gallinaceous birds in the East, he thinks that such slight differences are beneficial. For myself, I will only say that I am not convinced.

Formerly when I was inclined to lay much stress on protection as accounting for the duller colours of female birds, it occurred to me that possibly both sexes and the young might aboriginally have been equally bright coloured; but that subsequently, the females from the danger incurred during incubation, and the young from being inexperienced, had been rendered dull as a protection. But this view is not supported by any evidence, and is not probable; for we thus in imagination expose during past times the females and the young to danger, from which it has subsequently been necessary to shield their modified descendants. We have, also, to reduce, through a gradual process of selection, the females and the young to almost exactly the same tints and markings, and to transmit them to the corresponding sex and period of life. On the supposition that the females and the young have partaken during each stage of the process of modification of a tendency to be as brightly coloured as the males, it is also a somewhat strange fact that the females have never been rendered dull-coloured without the young participating in the same change; for there are no instances, as far as I can discover, of species with the females dull and the young bright coloured. A partial exception, however, is offered by the young of certain woodpeckers, for they have “the whole upper part of the head tinged with red,” which afterwards either decreases into a mere circular red line in the adults of both sexes, or quite disappears in the adult females. (12. Audubon, ‘Ornith. Biography,’ vol. i. p. 193. Macgillivray, ‘History of British Birds,’ vol. iii. p. 85. See also the case before given of Indopicus carlotta.)

Finally, with respect to our present class of cases, the most probable view appears to be that successive variations in brightness or in other ornamental characters, occurring in the males at a rather late period of life have alone been preserved; and that most or all of these variations, owing to the late period of life at which they appeared, have been from the first transmitted only to the adult male offspring. Any variations in brightness occurring in the females or in the young, would have been of no service to them, and would not have been selected; and moreover, if dangerous, would have been eliminated. Thus the females and the young will either have been left unmodified, or (as is much more common) will have been partially modified by receiving through transference from the males some of his successive variations. Both sexes have perhaps been directly acted on by the conditions of life to which they have long been exposed: but the females from not being otherwise much modified, will best exhibit any such effects. These changes and all others will have been kept uniform by the free intercrossing of many individuals. In some cases, especially with ground birds, the females and the young may possibly have been modified, independently of the males, for the sake of protection, so as to have acquired the same dull-coloured plumage.

CLASS II. — WHEN THE ADULT FEMALE IS MORE CONSPICUOUS THAN THE ADULT MALE, THE YOUNG OF BOTH SEXES IN THEIR FIRST PLUMAGE RESEMBLE THE ADULT MALE.

This class is exactly the reverse of the last, for the females are here brighter coloured or more conspicuous than the males; and the young, as far as they are known, resemble the adult males instead of the adult females. But the difference between the sexes is never nearly so great as with many birds in the first class, and the cases are comparatively rare. Mr. Wallace, who first called attention to the singular relation which exists between the less bright colours of the males and their performing the duties of incubation, lays great stress on this point (13. ‘Westminster Review,’ July 1867, and A. Murray, ‘Journal of Travel,’ 1868, p. 83.), as a crucial test that obscure colours have been acquired for the sake of protection during the period of nesting. A different view seems to me more probable. As the cases are curious and not numerous, I will briefly give all that I have been able to find.

In one section of the genus Turnix, quail-like birds, the female is invariably larger than the male (being nearly twice as large in one of the Australian species), and this is an unusual circumstance with the Gallinaceae. In most of the species the female is more distinctly coloured and brighter than the male (14. For the Australian species, see Gould’s ‘Handbook,’ etc., vol. ii. pp. 178, 180, 186, and 188. In the British Museum specimens of the Australian Plain-wanderer (Pedionomus torquatus) may be seen, shewing similar sexual differences.), but in some few species the sexes are alike. In Turnix taigoor of India the male “wants the black on the throat and neck, and the whole tone of the plumage is lighter and less pronounced than that of the female.” The female appears to be noisier, and is certainly much more pugnacious than the male; so that the females and not the males are often kept by the natives for fighting, like game-cocks. As male birds are exposed by the English bird-catchers for a decoy near a trap, in order to catch other males by exciting their rivalry, so the females of this Turnix are employed in India. When thus exposed the females soon begin their “loud purring call, which can be heard a long way off, and any females within ear-shot run rapidly to the spot, and commence fighting with the caged bird.” In this way from twelve to twenty birds, all breeding females, may be caught in the course of a single day. The natives assert that the females after laying their eggs associate in flocks, and leave the males to sit on them. There is no reason to doubt the truth of this assertion, which is supported by some observations made in China by Mr. Swinhoe. (15. Jerdon, ‘Birds of India,’ vol. iii. p. 596. Mr. Swinhoe, in ‘Ibis,’ 1865, p. 542; 1866, pp. 131, 405.) Mr. Blyth believes, that the young of both sexes resemble the adult male.

[Fig. 62. Rhynchaea capensis (from Brehm).]

The females of the three species of Painted Snipes (Rhynchaea, Fig. 62) “are not only larger but much more richly coloured than the males.” (16. Jerdon, ‘Birds of India,’ vol. iii. p. 677.) With all other birds in which the trachea differs in structure in the two sexes it is more developed and complex in the male than in the female; but in the Rhynchaea australis it is simple in the male, whilst in the female it makes four distinct convolutions before entering the lungs. (17. Gould’s ‘Handbook to the Birds of Australia,’ vol. ii. p. 275.) The female therefore of this species has acquired an eminently masculine character. Mr. Blyth ascertained, by examining many specimens, that the trachea is not convoluted in either sex of R. bengalensis, which species resembles R. australis so closely, that it can hardly be distinguished except by its shorter toes. This fact is another striking instance of the law that secondary sexual characters are often widely different in closely-allied forms, though it is a very rare circumstance when such differences relate to the female sex. The young of both sexes of R. bengalensis in their first plumage are said to resemble the mature male. (18. ‘The Indian Field,’ Sept. 1858, p. 3.) There is also reason to believe that the male undertakes the duty of incubation, for Mr. Swinhoe (19. ‘Ibis,’ 1866, p. 298.) found the females before the close of the summer associated in flocks, as occurs with the females of the Turnix.

The females of Phalaropus fulicarius and P. hyperboreus are larger, and in their summer plumage “more gaily attired than the males.” But the difference in colour between the sexes is far from conspicuous. According to Professor Steenstrup, the male alone of P. fulicarius undertakes the duty of incubation; this is likewise shewn by the state of his breast-feathers during the breeding-season. The female of the dotterel plover (Eudromias morinellus) is larger than the male, and has the red and black tints on the lower surface, the white crescent on the breast, and the stripes over the eyes, more strongly pronounced. The male also takes at least a share in hatching the eggs; but the female likewise attends to the young. (20. For these several statements, see Mr. Gould’s ‘Birds of Great Britain.’ Prof. Newton informs me that he has long been convinced, from his own observations and from those of others, that the males of the above-named species take either the whole or a large share of the duties of incubation, and that they “shew much greater devotion towards their young, when in danger, than do the females.” So it is, as he informs me, with Limosa lapponica and some few other Waders, in which the females are larger and have more strongly contrasted colours than the males.) I have not been able to discover whether with these species the young resemble the adult males more closely than the adult females; for the comparison is somewhat difficult to make on account of the double moult.

Turning now to the ostrich Order: the male of the common cassowary (Casuarius galeatus) would be thought by any one to be the female, from his smaller size and from the appendages and naked skin about his head being much less brightly coloured; and I am informed by Mr. Bartlett that in the Zoological Gardens, it is certainly the male alone who sits on the eggs and takes care of the young. (21. The natives of Ceram (Wallace, ‘Malay Archipelago,’ vol. ii. p. 150) assert that the male and female sit alternately on the eggs; but this assertion, as Mr. Bartlett thinks, may be accounted for by the female visiting the nest to lay her eggs.) The female is said by Mr. T.W. Wood (22. The ‘Student,’ April 1870, p. 124.) to exhibit during the breeding-season a most pugnacious disposition; and her wattles then become enlarged and more brilliantly coloured. So again the female of one of the emus (Dromoeus irroratus) is considerably larger than the male, and she possesses a slight top-knot, but is otherwise indistinguishable in plumage. She appears, however, “to have greater power, when angry or otherwise excited, of erecting, like a turkey-cock, the feathers of her neck and breast. She is usually the more courageous and pugilistic. She makes a deep hollow guttural boom especially at night, sounding like a small gong. The male has a slenderer frame and is more docile, with no voice beyond a suppressed hiss when angry, or a croak.” He not only performs the whole duty of incubation, but has to defend the young from their mother; “for as soon as she catches sight of her progeny she becomes violently agitated, and notwithstanding the resistance of the father appears to use her utmost endeavours to destroy them. For months afterwards it is unsafe to put the parents together, violent quarrels being the inevitable result, in which the female generally comes off conqueror.” (23. See the excellent account of the habits of this bird under confinement, by Mr. A.W. Bennett, in ‘Land and Water,’ May 1868, p. 233.) So that with this emu we have a complete reversal not only of the parental and incubating instincts, but of the usual moral qualities of the two sexes; the females being savage, quarrelsome, and noisy, the males gentle and good. The case is very different with the African ostrich, for the male is somewhat larger than the female and has finer plumes with more strongly contrasted colours; nevertheless he undertakes the whole duty of incubation. (24. Mr. Sclater, on the incubation of the Struthiones, ‘Proc. Zool. Soc.’ June 9, 1863. So it is with the Rhea darwinii: Captain Musters says (‘At Home with the Patagonians,’ 1871, p. 128), that the male is larger, stronger and swifter than the female, and of slightly darker colours; yet he takes sole charge of the eggs and of the young, just as does the male of the common species of Rhea.)

I will specify the few other cases known to me, in which the female is more conspicuously coloured than the male, although nothing is known about the manner of incubation. With the carrion-hawk of the Falkland Islands (Milvago leucurus) I was much surprised to find by dissection that the individuals, which had all their tints strongly pronounced, with the cere and legs orange-coloured, were the adult females; whilst those with duller plumage and grey legs were the males or the young. In an Australian tree-creeper (Climacteris erythrops) the female differs from the male in “being adorned with beautiful, radiated, rufous markings on the throat, the male having this part quite plain.” Lastly, in an Australian night-jar “the female always exceeds the male in size and in the brilliance of her tints; the males, on the other hand, have two white spots on the primaries more conspicuous than in the female.” (25. For the Milvago, see ‘Zoology of the Voyage of the “Beagle,” Birds,’ 1841, p. 16. For the Climacteris and night-jar (Eurostopodus), see Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 602 and 97. The New Zealand shieldrake (Tadorna variegata) offers a quite anomalous case; the head of the female is pure white, and her back is redder than that of the male; the head of the male is of a rich dark bronzed colour, and his back is clothed with finely pencilled slate-coloured feathers, so that altogether he may be considered as the more beautiful of the two. He is larger and more pugnacious than the female, and does not sit on the eggs. So that in all these respects this species comes under our first class of cases; but Mr. Sclater (‘Proceedings of the Zoological Society,’ 1866, p. 150) was much surprised to observe that the young of both sexes, when about three months old, resembled in their dark heads and necks the adult males, instead of the adult females; so that it would appear in this case that the females have been modified, whilst the males and the young have retained a former state of plumage.)

We thus see that the cases in which female birds are more conspicuously coloured than the males, with the young in their immature plumage resembling the adult males instead of the adult females, as in the previous class, are not numerous, though they are distributed in various Orders. The amount of difference, also, between the sexes is incomparably less than that which frequently occurs in the last class; so that the cause of the difference, whatever it may have been, has here acted on the females either less energetically or less persistently than on the males in the last class. Mr. Wallace believes that the males have had their colours rendered less conspicuous for the sake of protection during the period of incubation; but the difference between the sexes in hardly any of the foregoing cases appears sufficiently great for this view to be safely accepted. In some of the cases, the brighter tints of the female are almost confined to the lower surface, and the males, if thus coloured, would not have been exposed to danger whilst sitting on the eggs. It should also be borne in mind that the males are not only in a slight degree less conspicuously coloured than the females, but are smaller and weaker. They have, moreover, not only acquired the maternal instinct of incubation, but are less pugnacious and vociferous than the females, and in one instance have simpler vocal organs. Thus an almost complete transposition of the instincts, habits, disposition, colour, size, and of some points of structure, has been effected between the two sexes.

Now if we might assume that the males in the present class have lost some of that ardour which is usual to their sex, so that they no longer search eagerly for the females; or, if we might assume that the females have become much more numerous than the males—and in the case of one Indian Turnix the females are said to be “much more commonly met with than the males” (26. Jerdon, ‘Birds of India,’ vol. iii. p. 598.)—then it is not improbable that the females would have been led to court the males, instead of being courted by them. This indeed is the case to a certain extent with some birds, as we have seen with the peahen, wild turkey, and certain kinds of grouse. Taking as our guide the habits of most male birds, the greater size and strength as well as the extraordinary pugnacity of the females of the Turnix and emu, must mean that they endeavour to drive away rival females, in order to gain possession of the male; and on this view all the facts become clear; for the males would probably be most charmed or excited by the females which were the most attractive to them by their bright colours, other ornaments, or vocal powers. Sexual selection would then do its work, steadily adding to the attractions of the females; the males and the young being left not at all, or but little modified.

CLASS III. — WHEN THE ADULT MALE RESEMBLES THE ADULT FEMALE, THE YOUNG OF BOTH SEXES HAVE A PECULIAR FIRST PLUMAGE OF THEIR OWN.

In this class the sexes when adult resemble each other, and differ from the young. This occurs with many birds of many kinds. The male robin can hardly be distinguished from the female, but the young are widely different, with their mottled dusky-olive and brown plumage. The male and female of the splendid scarlet ibis are alike, whilst the young are brown; and the scarlet colour, though common to both sexes, is apparently a sexual character, for it is not well developed in either sex under confinement; and a loss of colour often occurs with brilliant males when they are confined. With many species of herons the young differ greatly from the adults; and the summer plumage of the latter, though common to both sexes, clearly has a nuptial character. Young swans are slate-coloured, whilst the mature birds are pure white; but it would be superfluous to give additional instances. These differences between the young and the old apparently depend, as in the last two classes, on the young having retained a former or ancient state of plumage, whilst the old of both sexes have acquired a new one. When the adults are bright coloured, we may conclude from the remarks just made in relation to the scarlet ibis and to many herons, and from the analogy of the species in the first class, that such colours have been acquired through sexual selection by the nearly mature males; but that, differently from what occurs in the first two classes, the transmission, though limited to the same age, has not been limited to the same sex. Consequently, the sexes when mature resemble each other and differ from the young.

CLASS IV. — WHEN THE ADULT MALE RESEMBLES THE ADULT FEMALE, THE YOUNG OF BOTH SEXES IN THEIR FIRST PLUMAGE RESEMBLE THE ADULTS.

In this class the young and the adults of both sexes, whether brilliantly or obscurely coloured, resemble each other. Such cases are, I think, more common than those in the last class. We have in England instances in the kingfisher, some woodpeckers, the jay, magpie, crow, and many small dull-coloured birds, such as the hedge-warbler or kitty-wren. But the similarity in plumage between the young and the old is never complete, and graduates away into dissimilarity. Thus the young of some members of the kingfisher family are not only less vividly coloured than the adults, but many of the feathers on the lower surface are edged with brown (27. Jerdon, ‘Birds of India,’ vol. i. pp. 222, 228. Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 124, 130.),—a vestige probably of a former state of the plumage. Frequently in the same group of birds, even within the same genus, for instance in an Australian genus of parrakeets (Platycercus), the young of some species closely resemble, whilst the young of other species differ considerably, from their parents of both sexes, which are alike. (28. Gould, ibid. vol. ii. pp. 37, 46, 56.) Both sexes and the young of the common jay are closely similar; but in the Canada jay (Perisoreus canadensis) the young differ so much from their parents that they were formerly described as distinct species. (29. Audubon, ‘Ornith. Biography,’ vol. ii. p. 55.)

I may remark before proceeding that, under the present and next two classes of cases, the facts are so complex and the conclusions so doubtful, that any one who feels no especial interest in the subject had better pass them over.

The brilliant or conspicuous colours which characterise many birds in the present class, can rarely or never be of service to them as a protection; so that they have probably been gained by the males through sexual selection, and then transferred to the females and the young. It is, however, possible that the males may have selected the more attractive females; and if these transmitted their characters to their offspring of both sexes, the same results would follow as from the selection of the more attractive males by the females. But there is evidence that this contingency has rarely, if ever, occurred in any of those groups of birds in which the sexes are generally alike; for, if even a few of the successive variations had failed to be transmitted to both sexes, the females would have slightly exceeded the males in beauty. Exactly the reverse occurs under nature; for, in almost every large group in which the sexes generally resemble each other, the males of some few species are in a slight degree more brightly coloured than the females. It is again possible that the females may have selected the more beautiful males, these males having reciprocally selected the more beautiful females; but it is doubtful whether this double process of selection would be likely to occur, owing to the greater eagerness of one sex than the other, and whether it would be more efficient than selection on one side alone. It is, therefore, the most probable view that sexual selection has acted, in the present class, as far as ornamental characters are concerned, in accordance with the general rule throughout the animal kingdom, that is, on the males; and that these have transmitted their gradually-acquired colours, either equally or almost equally, to their offspring of both sexes.

Another point is more doubtful, namely, whether the successive variations first appeared in the males after they had become nearly mature, or whilst quite young. In either case sexual selection must have acted on the male when he had to compete with rivals for the possession of the female; and in both cases the characters thus acquired have been transmitted to both sexes and all ages. But these characters if acquired by the males when adult, may have been transmitted at first to the adults alone, and at some subsequent period transferred to the young. For it is known that, when the law of inheritance at corresponding ages fails, the offspring often inherit characters at an earlier age than that at which they first appeared in their parents. (30. ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 79.) Cases apparently of this kind have been observed with birds in a state of nature. For instance Mr. Blyth has seen specimens of Lanius rufus and of Colymbus glacialis which had assumed whilst young, in a quite anomalous manner, the adult plumage of their parents. (31. ‘Charlesworth’s Magazine of Natural History,’ vol. i. 1837, pp. 305, 306.) Again, the young of the common swan (Cygnus olor) do not cast off their dark feathers and become white until eighteen months or two years old; but Dr. F. Forel has described the case of three vigorous young birds, out of a brood of four, which were born pure white. These young birds were not albinos, as shewn by the colour of their beaks and legs, which nearly resembled the same parts in the adults. (32. ‘Bulletin de la Soc. Vaudoise des Sc. Nat.’ vol. x. 1869, p. 132. The young of the Polish swan, Cygnus immutabilis of Yarrell, are always white; but this species, as Mr. Sclater informs me, is believed to be nothing more than a variety of the domestic swan (Cygnus olor).)

It may be worth while to illustrate the above three modes by which, in the present class, the two sexes and the young may have come to resemble each other, by the curious case of the genus Passer. (33. I am indebted to Mr. Blyth for information in regard to this genus. The sparrow of Palestine belongs to the sub-genus Petronia.) In the house-sparrow (P. domesticus) the male differs much from the female and from the young. The young and the females are alike, and resemble to a large extent both sexes and the young of the sparrow of Palestine (P. brachydactylus), as well as of some allied species. We may therefore assume that the female and young of the house-sparrow approximately shew us the plumage of the progenitor of the genus. Now with the tree-sparrow (P. montanus) both sexes and the young closely resemble the male of the house-sparrow; so that they have all been modified in the same manner, and all depart from the typical colouring of their early progenitor. This may have been effected by a male ancestor of the tree-sparrow having varied, firstly, when nearly mature; or, secondly, whilst quite young, and by having in either case transmitted his modified plumage to the females and the young; or, thirdly, he may have varied when adult and transmitted his plumage to both adult sexes, and, owing to the failure of the law of inheritance at corresponding ages, at some subsequent period to his young.

It is impossible to decide which of these three modes has generally prevailed throughout the present class of cases. That the males varied whilst young, and transmitted their variations to their offspring of both sexes, is the most probable. I may here add that I have, with little success, endeavoured, by consulting various works, to decide how far the period of variation in birds has generally determined the transmission of characters to one sex or to both. The two rules, often referred to (namely, that variations occurring late in life are transmitted to one and the same sex, whilst those which occur early in life are transmitted to both sexes), apparently hold good in the first (34. For instance, the males of Tanagra aestiva and Fringilla cyanea require three years, the male of Fringilla ciris four years, to complete their beautiful plumage. (See Audubon, ‘Ornith. Biography,’ vol. i. pp. 233, 280, 378). The Harlequin duck takes three years (ibid. vol. iii. p. 614). The male of the Gold pheasant, as I hear from Mr. Jenner Weir, can be distinguished from the female when about three months old, but he does not acquire his full splendour until the end of the September in the following year.), second, and fourth classes of cases; but they fail in the third, often in the fifth (35. Thus the Ibis tantalus and Grus americanus take four years, the Flamingo several years, and the Ardea ludovicana two years, before they acquire their perfect plumage. See Audubon, ibid. vol. i. p. 221; vol. iii. pp. 133, 139, 211.), and in the sixth small class. They apply, however, as far as I can judge, to a considerable majority of the species; and we must not forget the striking generalisation by Dr. W. Marshall with respect to the protuberances on the heads of birds. Whether or not the two rules generally hold good, we may conclude from the facts given in the eighth chapter, that the period of variation is one important element in determining the form of transmission.

With birds it is difficult to decide by what standard we ought to judge of the earliness or lateness of the period of variation, whether by the age in reference to the duration of life, or to the power of reproduction, or to the number of moults through which the species passes. The moulting of birds, even within the same family, sometimes differs much without any assignable cause. Some birds moult so early, that nearly all the body feathers are cast off before the first wing-feathers are fully grown; and we cannot believe that this was the primordial state of things. When the period of moulting has been accelerated, the age at which the colours of the adult plumage are first developed will falsely appear to us to be earlier than it really is. This may be illustrated by the practice followed by some bird-fanciers, who pull out a few feathers from the breast of nestling bullfinches, and from the head or neck of young gold-pheasants, in order to ascertain their sex; for in the males, these feathers are immediately replaced by coloured ones. (36. Mr. Blyth, in Charlesworth’s ‘Magazine of Natural History,’ vol. i. 1837, p. 300. Mr. Bartlett has informed me in regard to gold pheasants.) The actual duration of life is known in but few birds, so that we can hardly judge by this standard. And, with reference to the period at which the power of reproduction is gained, it is a remarkable fact that various birds occasionally breed whilst retaining their immature plumage. (37. I have noticed the following cases in Audubon’s ‘Ornith. Biography.’ The redstart of America (Muscapica ruticilla, vol. i. p. 203). The Ibis tantalus takes four years to come to full maturity, but sometimes breeds in the second year (vol. iii. p. 133). The Grus americanus takes the same time, but breeds before acquiring its full plumage (vol. iii. p. 211). The adults of Ardea caerulea are blue, and the young white; and white, mottled, and mature blue birds may all be seen breeding together (vol. iv. p. 58): but Mr. Blyth informs me that certain herons apparently are dimorphic, for white and coloured individuals of the same age may be observed. The Harlequin duck (Anas histrionica, Linn.) takes three years to acquire its full plumage, though many birds breed in the second year (vol. iii. p. 614). The White-headed Eagle (Falco leucocephalus, vol. iii. p. 210) is likewise known to breed in its immature state. Some species of Oriolus (according to Mr. Blyth and Mr. Swinhoe, in ‘Ibis,’ July 1863, p. 68) likewise breed before they attain their full plumage.)

The fact of birds breeding in their immature plumage seems opposed to the belief that sexual selection has played as important a part, as I believe it has, in giving ornamental colours, plumes, etc., to the males, and, by means of equal transmission, to the females of many species. The objection would be a valid one, if the younger and less ornamented males were as successful in winning females and propagating their kind, as the older and more beautiful males. But we have no reason to suppose that this is the case. Audubon speaks of the breeding of the immature males of Ibis tantalus as a rare event, as does Mr. Swinhoe, in regard to the immature males of Oriolus. (38. See footnote 37 above.) If the young of any species in their immature plumage were more successful in winning partners than the adults, the adult plumage would probably soon be lost, as the males would prevail, which retained their immature dress for the longest period, and thus the character of the species would ultimately be modified. (39. Other animals, belonging to quite distinct classes, are either habitually or occasionally capable of breeding before they have fully acquired their adult characters. This is the case with the young males of the salmon. Several amphibians have been known to breed whilst retaining their larval structure. Fritz Müller has shewn (‘Facts and arguments for Darwin,’ Eng. trans. 1869, p. 79) that the males of several amphipod crustaceans become sexually mature whilst young; and I infer that this is a case of premature breeding, because they have not as yet acquired their fully-developed claspers. All such facts are highly interesting, as bearing on one means by which species may undergo great modifications of character.) If, on the other hand, the young never succeeded in obtaining a female, the habit of early reproduction would perhaps be sooner or later eliminated, from being superfluous and entailing waste of power.

The plumage of certain birds goes on increasing in beauty during many years after they are fully mature; this is the case with the train of the peacock, with some of the birds of paradise, and with the crest and plumes of certain herons, for instance, the Ardea ludovicana. (40. Jerdon, ‘Birds of India,’ vol. iii. p. 507, on the peacock. Dr. Marshall thinks that the older and more brilliant males of birds of paradise, have an advantage over the younger males; see ‘Archives Neerlandaises,’ tom. vi. 1871.—On Ardea, Audubon, ibid. vol. iii. p. 139.) But it is doubtful whether the continued development of such feathers is the result of the selection of successive beneficial variations (though this is the most probable view with birds of paradise) or merely of continuous growth. Most fishes continue increasing in size, as long as they are in good health and have plenty of food; and a somewhat similar law may prevail with the plumes of birds.

CLASS V. — WHEN THE ADULTS OF BOTH SEXES HAVE A DISTINCT WINTER AND SUMMER PLUMAGE, WHETHER OR NOT THE MALE DIFFERS FROM THE FEMALE, THE YOUNG RESEMBLE THE ADULTS OF BOTH SEXES IN THEIR WINTER DRESS, OR MUCH MORE RARELY IN THEIR SUMMER DRESS, OR THEY RESEMBLE THE FEMALES ALONE. OR THE YOUNG MAY HAVE AN INTERMEDIATE CHARACTER; OR, AGAIN, THEY MAY DIFFER GREATLY FROM THE ADULTS IN BOTH THEIR SEASONAL PLUMAGES.

The cases in this class are singularly complex; nor is this surprising, as they depend on inheritance, limited in a greater or less degree in three different ways, namely, by sex, age, and the season of the year. In some cases the individuals of the same species pass through at least five distinct states of plumage. With the species, in which the male differs from the female during the summer season alone, or, which is rarer, during both seasons (41. For illustrative cases, see vol. iv. of Macgillivray’s ‘History of British Birds;’ on Tringa, etc., pp. 229, 271; on the Machetes, p. 172; on the Charadrius hiaticula, p. 118; on the Charadrius pluvialis, p. 94.), the young generally resemble the females,—as with the so-called goldfinch of North America, and apparently with the splendid Maluri of Australia. (42. For the goldfinch of N. America, Fringilla tristis, Linn., see Audubon, ‘Ornithological Biography,’ vol. i. p. 172. For the Maluri, Gould’s ‘Handbook of the Birds of Australia,’ vol. i. p. 318.) With those species, the sexes of which are alike during both the summer and winter, the young may resemble the adults, firstly, in their winter dress; secondly, and this is of much rarer occurrence, in their summer dress; thirdly, they may be intermediate between these two states; and, fourthly, they may differ greatly from the adults at all seasons. We have an instance of the first of these four cases in one of the egrets of India (Buphus coromandus), in which the young and the adults of both sexes are white during the winter, the adults becoming golden-buff during the summer.

With the gaper (Anastomus oscitans) of India we have a similar case, but the colours are reversed: for the young and the adults of both sexes are grey and black during the winter, the adults becoming white during the summer. (43. I am indebted to Mr. Blyth for information as to the Buphus; see also Jerdon, ‘Birds of India,’ vol. iii. p. 749. On the Anastomus, see Blyth, in ‘Ibis,’ 1867, p. 173.) As an instance of the second case, the young of the razor-bill (Alca torda, Linn.), in an early state of plumage, are coloured like the adults during the summer; and the young of the white-crowned sparrow of North America (Fringilla leucophrys), as soon as fledged, have elegant white stripes on their heads, which are lost by the young and the old during the winter. (44. On the Alca, see Macgillivray, ‘Hist. Brit. Birds,’ vol. v. p. 347. On the Fringilla leucophrys, Audubon, ibid. vol. ii. p. 89. I shall have hereafter to refer to the young of certain herons and egrets being white.) With respect to the third case, namely, that of the young having an intermediate character between the summer and winter adult plumages, Yarrell (45. ‘History of British Birds,’ vol. i. 1839, p. 159.) insists that this occurs with many waders. Lastly, in regard to the young differing greatly from both sexes in their adult summer and winter plumages, this occurs with some herons and egrets of North America and India,—the young alone being white.

I will make only a few remarks on these complicated cases. When the young resemble the females in their summer dress, or the adults of both sexes in their winter dress, the cases differ from those given under Classes I. and III. only in the characters originally acquired by the males during the breeding-season, having been limited in their transmission to the corresponding season. When the adults have a distinct summer and winter plumage, and the young differ from both, the case is more difficult to understand. We may admit as probable that the young have retained an ancient state of plumage; we can account by sexual selection for the summer or nuptial plumage of the adults, but how are we to account for their distinct winter plumage? If we could admit that this plumage serves in all cases as a protection, its acquirement would be a simple affair; but there seems no good reason for this admission. It may be suggested that the widely different conditions of life during the winter and summer have acted in a direct manner on the plumage; this may have had some effect, but I have not much confidence in so great a difference as we sometimes see between the two plumages, having been thus caused. A more probable explanation is, that an ancient style of plumage, partially modified through the transference of some characters from the summer plumage, has been retained by the adults during the winter. Finally, all the cases in our present class apparently depend on characters acquired by the adult males, having been variously limited in their transmission according to age, season, and sex; but it would not be worth while to attempt to follow out these complex relations.

CLASS VI. — THE YOUNG IN THEIR FIRST PLUMAGE DIFFER FROM EACH OTHER ACCORDING TO SEX; THE YOUNG MALES RESEMBLING MORE OR LESS CLOSELY THE ADULT MALES, AND THE YOUNG FEMALES MORE OR LESS CLOSELY THE ADULT FEMALES.

The cases in the present class, though occurring in various groups, are not numerous; yet it seems the most natural thing that the young should at first somewhat resemble the adults of the same sex, and gradually become more and more like them. The adult male blackcap (Sylvia atricapilla) has a black head, that of the female being reddish-brown; and I am informed by Mr. Blyth, that the young of both sexes can be distinguished by this character even as nestlings. In the family of thrushes an unusual number of similar cases have been noticed; thus, the male blackbird (Turdus merula) can be distinguished in the nest from the female. The two sexes of the mocking bird (Turdus polyglottus, Linn.) differ very little from each other, yet the males can easily be distinguished at a very early age from the females by showing more pure white. (46. Audubon, ‘Ornith. Biography,’ vol. i. p. 113.) The males of a forest-thrush and of a rock-thrush (Orocetes erythrogastra and Petrocincla cyanea) have much of their plumage of a fine blue, whilst the females are brown; and the nestling males of both species have their main wing and tail-feathers edged with blue whilst those of the female are edged with brown. (47. Mr. C.A. Wright, in ‘Ibis,’ vol. vi. 1864, p. 65. Jerdon, ‘Birds of India,’ vol. i. p. 515. See also on the blackbird, Blyth in Charlesworth’s ‘Magazine of Natural History,’ vol. i. 1837, p. 113.) In the young blackbird the wing-feathers assume their mature character and become black after the others; on the other hand, in the two species just named the wing-feathers become blue before the others. The most probable view with reference to the cases in the present class is that the males, differently from what occurs in Class I., have transmitted their colours to their male offspring at an earlier age than that at which they were first acquired; for, if the males had varied whilst quite young, their characters would probably have been transmitted to both sexes. (48. The following additional cases may be mentioned; the young males of Tanagra rubra can be distinguished from the young females (Audubon, ‘Ornith. Biography,’ vol. iv. p. 392), and so it is within the nestlings of a blue nuthatch, Dendrophila frontalis of India (Jerdon, ‘Birds of India,’ vol. i. p. 389). Mr. Blyth also informs me that the sexes of the stonechat, Saxicola rubicola, are distinguishable at a very early age. Mr. Salvin gives (‘Proc. Zoolog. Soc.’ 1870, p. 206) the case of a humming-bird, like the following one of Eustephanus.)

In Aithurus polytmus, a humming-bird, the male is splendidly coloured black and green, and two of the tail-feathers are immensely lengthened; the female has an ordinary tail and inconspicuous colours; now the young males, instead of resembling the adult female, in accordance with the common rule, begin from the first to assume the colours proper to their sex, and their tail-feathers soon become elongated. I owe this information to Mr. Gould, who has given me the following more striking and as yet unpublished case. Two humming-birds belonging to the genus Eustephanus, both beautifully coloured, inhabit the small island of Juan Fernandez, and have always been ranked as specifically distinct. But it has lately been ascertained that the one which is of a rich chestnut-brown colour with a golden-red head, is the male, whilst the other which is elegantly variegated with green and white with a metallic green head is the female. Now the young from the first somewhat resemble the adults of the corresponding sex, the resemblance gradually becoming more and more complete.

In considering this last case, if as before we take the plumage of the young as our guide, it would appear that both sexes have been rendered beautiful independently; and not that one sex has partially transferred its beauty to the other. The male apparently has acquired his bright colours through sexual selection in the same manner as, for instance, the peacock or pheasant in our first class of cases; and the female in the same manner as the female Rhynchaea or Turnix in our second class of cases. But there is much difficulty in understanding how this could have been effected at the same time with the two sexes of the same species. Mr. Salvin states, as we have seen in the eighth chapter, that with certain humming-birds the males greatly exceed the females in number, whilst with other species inhabiting the same country the females greatly exceed the males. If, then, we might assume that during some former lengthened period the males of the Juan Fernandez species had greatly exceeded the females in number, but that during another lengthened period the females had far exceeded the males, we could understand how the males at one time, and the females at another, might have been rendered beautiful by the selection of the brighter coloured individuals of either sex; both sexes transmitting their characters to their young at a rather earlier age than usual. Whether this is the true explanation I will not pretend to say; but the case is too remarkable to be passed over without notice.

We have now seen in all six classes, that an intimate relation exists between the plumage of the young and the adults, either of one sex or both. These relations are fairly well explained on the principle that one sex—this being in the great majority of cases the male—first acquired through variation and sexual selection bright colours or other ornaments, and transmitted them in various ways, in accordance with the recognised laws of inheritance. Why variations have occurred at different periods of life, even sometimes with species of the same group, we do not know, but with respect to the form of transmission, one important determining cause seems to be the age at which the variations first appear.

From the principle of inheritance at corresponding ages, and from any variations in colour which occurred in the males at an early age not being then selected—on the contrary being often eliminated as dangerous—whilst similar variations occurring at or near the period of reproduction have been preserved, it follows that the plumage of the young will often have been left unmodified, or but little modified. We thus get some insight into the colouring of the progenitors of our existing species. In a vast number of species in five out of our six classes of cases, the adults of one sex or of both are bright coloured, at least during the breeding-season, whilst the young are invariably less brightly coloured than the adults, or are quite dull coloured; for no instance is known, as far as I can discover, of the young of dull-coloured species displaying bright colours, or of the young of bright-coloured species being more brilliant than their parents. In the fourth class, however, in which the young and the old resemble each other, there are many species (though by no means all), of which the young are bright-coloured, and as these form old groups, we may infer that their early progenitors were likewise bright. With this exception, if we look to the birds of the world, it appears that their beauty has been much increased since that period, of which their immature plumage gives us a partial record.

ON THE COLOUR OF THE PLUMAGE IN RELATION TO PROTECTION.

It will have been seen that I cannot follow Mr. Wallace in the belief that dull colours, when confined to the females, have been in most cases specially gained for the sake of protection. There can, however, be no doubt, as formerly remarked, that both sexes of many birds have had their colours modified, so as to escape the notice of their enemies; or in some instances, so as to approach their prey unobserved, just as owls have had their plumage rendered soft, that their flight may not be overheard. Mr. Wallace remarks (49. ‘Westminster Review,’ July 1867, p. 5.) that “it is only in the tropics, among forests which never lose their foliage, that we find whole groups of birds, whose chief colour is green.” It will be admitted by every one, who has ever tried, how difficult it is to distinguish parrots in a leaf-covered tree. Nevertheless, we must remember that many parrots are ornamented with crimson, blue, and orange tints, which can hardly be protective. Woodpeckers are eminently arboreal, but besides green species, there are many black, and black-and-white kinds—all the species being apparently exposed to nearly the same dangers. It is therefore probable that with tree-haunting birds, strongly-pronounced colours have been acquired through sexual selection, but that a green tint has been acquired oftener than any other, from the additional advantage of protection.

In regard to birds which live on the ground, every one admits that they are coloured so as to imitate the surrounding surface. How difficult it is to see a partridge, snipe, woodcock, certain plovers, larks, and night-jars when crouched on ground. Animals inhabiting deserts offer the most striking cases, for the bare surface affords no concealment, and nearly all the smaller quadrupeds, reptiles, and birds depend for safety on their colours. Mr. Tristram has remarked in regard to the inhabitants of the Sahara, that all are protected by their “isabelline or sand-colour.” (50. ‘Ibis,’ 1859, vol. i. p. 429, et seq. Dr. Rohlfs, however, remarks to me in a letter that according to his experience of the Sahara, this statement is too strong.) Calling to my recollection the desert-birds of South America, as well as most of the ground-birds of Great Britain, it appeared to me that both sexes in such cases are generally coloured nearly alike. Accordingly, I applied to Mr. Tristram with respect to the birds of the Sahara, and he has kindly given me the following information. There are twenty-six species belonging to fifteen genera, which manifestly have their plumage coloured in a protective manner; and this colouring is all the more striking, as with most of these birds it differs from that of their congeners. Both sexes of thirteen out of the twenty-six species are coloured in the same manner; but these belong to genera in which this rule commonly prevails, so that they tell us nothing about the protective colours being the same in both sexes of desert-birds. Of the other thirteen species, three belong to genera in which the sexes usually differ from each other, yet here they have the sexes alike. In the remaining ten species, the male differs from the female; but the difference is confined chiefly to the under surface of the plumage, which is concealed when the bird crouches on the ground; the head and back being of the same sand-coloured hue in the two sexes. So that in these ten species the upper surfaces of both sexes have been acted on and rendered alike, through natural selection, for the sake of protection; whilst the lower surfaces of the males alone have been diversified, through sexual selection, for the sake of ornament. Here, as both sexes are equally well protected, we clearly see that the females have not been prevented by natural selection from inheriting the colours of their male parents; so that we must look to the law of sexually-limited transmission.

In all parts of the world both sexes of many soft-billed birds, especially those which frequent reeds or sedges, are obscurely coloured. No doubt if their colours had been brilliant, they would have been much more conspicuous to their enemies; but whether their dull tints have been specially gained for the sake of protection seems, as far as I can judge, rather doubtful. It is still more doubtful whether such dull tints can have been gained for the sake of ornament. We must, however, bear in mind that male birds, though dull-coloured, often differ much from their females (as with the common sparrow), and this leads to the belief that such colours have been gained through sexual selection, from being attractive. Many of the soft-billed birds are songsters; and a discussion in a former chapter should not be forgotten, in which it was shewn that the best songsters are rarely ornamented with bright tints. It would appear that female birds, as a general rule, have selected their mates either for their sweet voices or gay colours, but not for both charms combined. Some species, which are manifestly coloured for the sake of protection, such as the jack-snipe, woodcock, and night-jar, are likewise marked and shaded, according to our standard of taste, with extreme elegance. In such cases we may conclude that both natural and sexual selection have acted conjointly for protection and ornament. Whether any bird exists which does not possess some special attraction, by which to charm the opposite sex, may be doubted. When both sexes are so obscurely coloured that it would be rash to assume the agency of sexual selection, and when no direct evidence can be advanced shewing that such colours serve as a protection, it is best to own complete ignorance of the cause, or, which comes to nearly the same thing, to attribute the result to the direct action of the conditions of life.

Both sexes of many birds are conspicuously, though not brilliantly coloured, such as the numerous black, white, or piebald species; and these colours are probably the result of sexual selection. With the common blackbird, capercailzie, blackcock, black scoter-duck (Oidemia), and even with one of the birds of paradise (Lophorina atra), the males alone are black, whilst the females are brown or mottled; and there can hardly be a doubt that blackness in these cases has been a sexually selected character. Therefore it is in some degree probable that the complete or partial blackness of both sexes in such birds as crows, certain cockatoos, storks, and swans, and many marine birds, is likewise the result of sexual selection, accompanied by equal transmission to both sexes; for blackness can hardly serve in any case as a protection. With several birds, in which the male alone is black, and in others in which both sexes are black, the beak or skin about the head is brightly coloured, and the contrast thus afforded adds much to their beauty; we see this in the bright yellow beak of the male blackbird, in the crimson skin over the eyes of the blackcock and capercailzie, in the brightly and variously coloured beak of the scoter-drake (Oidemia), in the red beak of the chough (Corvus graculus, Linn.), of the black swan, and the black stork. This leads me to remark that it is not incredible that toucans may owe the enormous size of their beaks to sexual selection, for the sake of displaying the diversified and vivid stripes of colour, with which these organs are ornamented. (51. No satisfactory explanation has ever been offered of the immense size, and still less of the bright colours, of the toucan’s beak. Mr. Bates (‘The Naturalist on the Amazons,’ vol. ii. 1863, p. 341) states that they use their beaks for reaching fruit at the extreme tips of the branches; and likewise, as stated by other authors, for extracting eggs and young birds from the nests of other birds. But, as Mr. Bates admits, the beak “can scarcely be considered a very perfectly-formed instrument for the end to which it is applied.” The great bulk of the beak, as shewn by its breadth, depth, as well as length, is not intelligible on the view, that it serves merely as an organ of prehension. Mr. Belt believes (‘The Naturalist in Nicaragua,’ p. 197) that the principal use of the beak is as a defence against enemies, especially to the female whilst nesting in a hole in a tree.) The naked skin, also, at the base of the beak and round the eyes is likewise often brilliantly coloured; and Mr. Gould, in speaking of one species (52. Rhamphastos carinatus, Gould’s ‘Monograph of Ramphastidae.’), says that the colours of the beak “are doubtless in the finest and most brilliant state during the time of pairing.” There is no greater improbability that toucans should be encumbered with immense beaks, though rendered as light as possible by their cancellated structure, for the display of fine colours (an object falsely appearing to us unimportant), than that the male Argus pheasant and some other birds should be encumbered with plumes so long as to impede their flight.

In the same manner, as the males alone of various species are black, the females being dull-coloured; so in a few cases the males alone are either wholly or partially white, as with the several bell-birds of South America (Chasmorhynchus), the Antarctic goose (Bernicla antarctica), the silver pheasant, etc., whilst the females are brown or obscurely mottled. Therefore, on the same principle as before, it is probable that both sexes of many birds, such as white cockatoos, several egrets with their beautiful plumes, certain ibises, gulls, terns, etc., have acquired their more or less completely white plumage through sexual selection. In some of these cases the plumage becomes white only at maturity. This is the case with certain gannets, tropic-birds, etc., and with the snow-goose (Anser hyperboreus). As the latter breeds on the “barren grounds,” when not covered with snow, and as it migrates southward during the winter, there is no reason to suppose that its snow-white adult plumage serves as a protection. In the Anastomus oscitans, we have still better evidence that the white plumage is a nuptial character, for it is developed only during the summer; the young in their immature state, and the adults in their winter dress, being grey and black. With many kinds of gulls (Larus), the head and neck become pure white during the summer, being grey or mottled during the winter and in the young state. On the other hand, with the smaller gulls, or sea-mews (Gavia), and with some terns (Sterna), exactly the reverse occurs; for the heads of the young birds during the first year, and of the adults during the winter, are either pure white, or much paler coloured than during the breeding-season. These latter cases offer another instance of the capricious manner in which sexual selection appears often to have acted. (53. On Larus, Gavia, and Sterna, see Macgillivray, ‘History of British Birds,’ vol. v. pp. 515, 584, 626. On the Anser hyperboreus, Audubon, ‘Ornithological Biography,’ vol. iv. p. 562. On the Anastomus, Mr. Blyth, in ‘Ibis,’ 1867, p. 173.)

That aquatic birds have acquired a white plumage so much oftener than terrestrial birds, probably depends on their large size and strong powers of flight, so that they can easily defend themselves or escape from birds of prey, to which moreover they are not much exposed. Consequently, sexual selection has not here been interfered with or guided for the sake of protection. No doubt with birds which roam over the open ocean, the males and females could find each other much more easily, when made conspicuous either by being perfectly white or intensely black; so that these colours may possibly serve the same end as the call-notes of many land-birds. (54. It may be noticed that with vultures, which roam far and wide high in the air, like marine birds over the ocean, three or four species are almost wholly or largely white, and that many others are black. So that here again conspicuous colours may possibly aid the sexes in finding each other during the breeding-season.) A white or black bird when it discovers and flies down to a carcase floating on the sea or cast up on the beach, will be seen from a great distance, and will guide other birds of the same and other species, to the prey; but as this would be a disadvantage to the first finders, the individuals which were the whitest or blackest would not thus procure more food than the less strongly coloured individuals. Hence conspicuous colours cannot have been gradually acquired for this purpose through natural selection.

As sexual selection depends on so fluctuating an element as taste, we can understand how it is that, within the same group of birds having nearly the same habits, there should exist white or nearly white, as well as black, or nearly black species,—for instance, both white and black cockatoos, storks, ibises, swans, terns, and petrels. Piebald birds likewise sometimes occur in the same groups together with black and white species; for instance, the black-necked swan, certain terns, and the common magpie. That a strong contrast in colour is agreeable to birds, we may conclude by looking through any large collection, for the sexes often differ from each other in the male having the pale parts of a purer white, and the variously coloured dark parts of still darker tints than the female.

It would even appear that mere novelty, or slight changes for the sake of change, have sometimes acted on female birds as a charm, like changes of fashion with us. Thus the males of some parrots can hardly be said to be more beautiful than the females, at least according to our taste, but they differ in such points, as in having a rose-coloured collar instead of “a bright emeraldine narrow green collar”; or in the male having a black collar instead of “a yellow demi-collar in front,” with a pale roseate instead of a plum-blue head. (55. See Jerdon on the genus Palaeornis, ‘Birds of India,’ vol. i. pp. 258-260.) As so many male birds have elongated tail-feathers or elongated crests for their chief ornament, the shortened tail, formerly described in the male of a humming-bird, and the shortened crest of the male goosander, seem like one of the many changes of fashion which we admire in our own dresses.

Some members of the heron family offer a still more curious case of novelty in colouring having, as it appears, been appreciated for the sake of novelty. The young of the Ardea asha are white, the adults being dark slate-coloured; and not only the young, but the adults in their winter plumage, of the allied Buphus coromandus are white, this colour changing into a rich golden-buff during the breeding-season. It is incredible that the young of these two species, as well as of some other members of the same family (56. The young of Ardea rufescens and A. caerulea of the United States are likewise white, the adults being coloured in accordance with their specific names. Audubon (‘Ornithological Biography,’ vol. iii. p. 416; vol. iv. p. 58) seems rather pleased at the thought that this remarkable change of plumage will greatly “disconcert the systematists.”), should for any special purpose have been rendered pure white and thus made conspicuous to their enemies; or that the adults of one of these two species should have been specially rendered white during the winter in a country which is never covered with snow. On the other hand we have good reason to believe that whiteness has been gained by many birds as a sexual ornament. We may therefore conclude that some early progenitor of the Ardea asha and the Buphus acquired a white plumage for nuptial purposes, and transmitted this colour to their young; so that the young and the old became white like certain existing egrets; and that the whiteness was afterwards retained by the young, whilst it was exchanged by the adults for more strongly-pronounced tints. But if we could look still further back to the still earlier progenitors of these two species, we should probably see the adults dark-coloured. I infer that this would be the case, from the analogy of many other birds, which are dark whilst young, and when adult are white; and more especially from the case of the Ardea gularis, the colours of which are the reverse of those of A. asha, for the young are dark-coloured and the adults white, the young having retained a former state of plumage. It appears therefore that, during a long line of descent, the adult progenitors of the Ardea asha, the Buphus, and of some allies, have undergone the following changes of colour: first, a dark 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.

A 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. Générale,’ 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 heads 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 Aethiopian 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. ‘Traité de l’Héréd. Nat.’ tom. ii. 1850, p. 296.) quotes various statements from French authorities, and remarks, “On voit des étalons 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

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 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.

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.

A 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.

PART III. SEXUAL SELECTION IN RELATION TO MAN, AND CONCLUSION.

Chapter XIX: Secondary Sexual Characters of Man

Differences between man and woman—Causes of such differences and of certain characters common to both sexes—Law of battle—Differences in mental powers, and voice—On the influence of beauty in determining the marriages of mankind—Attention paid by savages to ornaments—Their ideas of beauty in woman—The tendency to exaggerate each natural peculiarity.

With mankind the differences between the sexes are greater than in most of the Quadrumana, but not so great as in some, for instance, the mandrill. Man on an average is considerably taller, heavier, and stronger than woman, with squarer shoulders and more plainly-pronounced muscles. Owing to the relation which exists between muscular development and the projection of the brows (1. Schaaffhausen, translation in ‘Anthropological Review,’ Oct. 1868, pp. 419, 420, 427.), the superciliary ridge is generally more marked in man than in woman. His body, and especially his face, is more hairy, and his voice has a different and more powerful tone. In certain races the women are said to differ slightly in tint from the men. For instance, Schweinfurth, in speaking of a negress belonging to the Monbuttoos, who inhabit the interior of Africa a few degrees north of the equator, says, “Like all her race, she had a skin several shades lighter than her husband’s, being something of the colour of half-roasted coffee.” (2. ‘The Heart of Africa,’ English transl. 1873, vol i. p. 544.) As the women labour in the fields and are quite unclothed, it is not likely that they differ in colour from the men owing to less exposure to the weather. European women are perhaps the brighter coloured of the two sexes, as may be seen when both have been equally exposed.

Man is more courageous, pugnacious and energetic than woman, and has a more inventive genius. His brain is absolutely larger, but whether or not proportionately to his larger body, has not, I believe, been fully ascertained. In woman the face is rounder; the jaws and the base of the skull smaller; the outlines of the body rounder, in parts more prominent; and her pelvis is broader than in man (3. Ecker, translation, in ‘Anthropological Review,’ Oct. 1868, pp. 351-356. The comparison of the form of the skull in men and women has been followed out with much care by Welcker.); but this latter character may perhaps be considered rather as a primary than a secondary sexual character. She comes to maturity at an earlier age than man.

As with animals of all classes, so with man, the distinctive characters of the male sex are not fully developed until he is nearly mature; and if emasculated they never appear. The beard, for instance, is a secondary

sexual character, and male children are beardless, though at an early age they have abundant hair on the head. It is probably due to the rather late appearance in life of the successive variations whereby man has acquired his masculine characters, that they are transmitted to the male sex alone. Male and female children resemble each other closely, like the young of so many other animals in which the adult sexes differ widely; they likewise resemble the mature female much more closely than the mature male. The female, however, ultimately assumes certain distinctive characters, and in the formation of her skull, is said to be intermediate between the child and the man. (4. Ecker and Welcker, ibid. pp. 352, 355; Vogt, ‘Lectures on Man,’ Eng. translat. p. 81.) Again, as the young of closely allied though distinct species do not differ nearly so much from each other as do the adults, so it is with the children of the different races of man. Some have even maintained that race-differences cannot be detected in the infantile skull. (5. Schaaffhausen, ‘Anthropolog. Review,’ ibid. p. 429.) In regard to colour, the new-born negro child is reddish nut-brown, which soon becomes slaty-grey; the black colour being fully developed within a year in the Soudan, but not until three years in Egypt. The eyes of the negro are at first blue, and the hair chestnut-brown rather than black, being curled only at the ends. The children of the Australians immediately after birth are yellowish-brown, and become dark at a later age. Those of the Guaranys of Paraguay are whitish-yellow, but they acquire in the course of a few weeks the yellowish-brown tint of their parents. Similar observations have been made in other parts of America. (6. Pruner-Bey, on negro infants as quoted by Vogt, ‘Lectures on Man,’ Eng. translat. 1864, p. 189: for further facts on negro infants, as quoted from Winterbottom and Camper, see Lawrence, ‘Lectures on Physiology,’ etc. 1822, p. 451. For the infants of the Guaranys, see Rengger, ‘Säugethiere,’ etc. s. 3. See also Godron, ‘De l’Espèce,’ tom. ii. 1859, p. 253. For the Australians, Waitz, ‘Introduction to Anthropology,’ Eng. translat. 1863, p. 99.)

I have specified the foregoing differences between the male and female sex in mankind, because they are curiously like those of the Quadrumana. With these animals the female is mature at an earlier age than the male; at least this is certainly the case in Cebus azarae. (7. Rengger, ‘Säugethiere,’ etc., 1830, s. 49.) The males of most species are larger and stronger than the females, of which fact the gorilla affords a well-known instance. Even in so trifling a character as the greater prominence of the superciliary ridge, the males of certain monkeys differ from the females (8. As in Macacus cynomolgus (Desmarest, ‘Mammalogie,’ p. 65), and in Hylobates agilis (Geoffroy St.-Hilaire and F. Cuvier, ‘Histoire Nat. des Mammifères,’ 1824, tom. i. p. 2)., and agree in this respect with mankind. In the gorilla and certain other monkeys, the cranium of the adult male presents a strongly-marked sagittal crest, which is absent in the female; and Ecker found a trace of a similar difference between the two sexes in the Australians. (9. ‘Anthropological Review,’ Oct. 1868, p. 353.) With monkeys when there is any difference in the voice, that of the male is the more powerful. We have seen that certain male monkeys have a well-developed beard, which is quite deficient, or much less developed in the female. No instance is known of the beard, whiskers, or moustache being larger in the female than in the male monkey. Even in the colour of the beard there is a curious parallelism between man and the Quadrumana, for with man when the beard differs in colour from the hair of the head, as is commonly the case, it is, I believe, almost always of a lighter tint, being often reddish. I have repeatedly observed this fact in England; but two gentlemen have lately written to me, saying that they form an exception to the rule. One of these gentlemen accounts for the fact by the wide difference in colour of the hair on the paternal and maternal sides of his family. Both had been long aware of this peculiarity (one of them having often been accused of dyeing his beard), and had been thus led to observe other men, and were convinced that the exceptions were very rare. Dr. Hooker attended to this little point for me in Russia, and found no exception to the rule. In Calcutta, Mr. J. Scott, of the Botanic Gardens, was so kind as to observe the many races of men to be seen there, as well as in some other parts of India, namely, two races of Sikhim, the Bhoteas, Hindoos, Burmese, and Chinese, most of which races have very little hair on the face; and he always found that when there was any difference in colour between the hair of the head and the beard, the latter was invariably lighter. Now with monkeys, as has already been stated, the beard frequently differs strikingly in colour from the hair of the head, and in such cases it is always of a lighter hue, being often pure white, sometimes yellow or reddish. (10. Mr. Blyth informs me that he has only seen one instance of the beard, whiskers, etc., in a monkey becoming white with old age, as is so commonly the case with us. This, however, occurred in an aged Macacus cynomolgus, kept in confinement whose moustaches were “remarkably long and human-like.” Altogether this old monkey presented a ludicrous resemblance to one of the reigning monarchs of Europe, after whom he was universally nick-named. In certain races of man the hair on the head hardly ever becomes grey; thus Mr. D. Forbes has never, as he informs me, seen an instance with the Aymaras and Quichuas of South America.)

In regard to the general hairiness of the body, the women in all races are less hairy than the men; and in some few Quadrumana the under side of the body of the female is less hairy than that of the male. (11. This is the case with the females of several species of Hylobates; see Geoffroy St.-Hilaire and F. Cuvier, ‘Hist. Nat. des Mamm.’ tom. i. See also, on H. lar, ‘Penny Cyclopedia,’ vol. ii. pp. 149, 150.) Lastly, male monkeys, like men, are bolder and fiercer than the females. They lead the troop, and when there is danger, come to the front. We thus see how close is the parallelism between the sexual differences of man and the Quadrumana. With some few species, however, as with certain baboons, the orang and the gorilla, there is a considerably greater difference between the sexes, as in the size of the canine teeth, in the development and colour of the hair, and especially in the colour of the naked parts of the skin, than in mankind.

All the secondary sexual characters of man are highly variable, even within the limits of the same race; and they differ much in the several races. These two rules hold good generally throughout the animal kingdom. In the excellent observations made on board the Novara (12. The results were deduced by Dr. Weisbach from the measurements made by Drs. K. Scherzer and Schwarz, see ‘Reise der Novara: Anthropolog. Theil,’ 1867, ss. 216, 231, 234, 236, 239, 269.), the male Australians were found to exceed the females by only 65 millim. in height, whilst with the Javans the average excess was 218 millim.; so that in this latter race the difference in height between the sexes is more than thrice as great as with the Australians. Numerous measurements were carefully made of the stature, the circumference of the neck and chest, the length of the back-bone and of the arms, in various races; and nearly all these measurements shew that the males differ much more from one another than do the females. This fact indicates that, as far as these characters are concerned, it is the male which has been chiefly modified, since the several races diverged from their common stock.

The development of the beard and the hairiness of the body differ remarkably in the men of distinct races, and even in different tribes or families of the same race. We Europeans see this amongst ourselves. In the Island of St. Kilda, according to Martin (13. ‘Voyage to St. Kilda’ (3rd ed. 1753), p. 37.), the men do not acquire beards until the age of thirty or upwards, and even then the beards are very thin. On the Europaeo-Asiatic continent, beards prevail until we pass beyond India; though with the natives of Ceylon they are often absent, as was noticed in ancient times by Diodorus. (14. Sir J.E. Tennent, ‘Ceylon,’ vol. ii. 1859, p. 107.) Eastward of India beards disappear, as with the Siamese, Malays, Kalmucks, Chinese, and Japanese; nevertheless, the Ainos (15. Quatrefages, ‘Revue des Cours Scientifiques,’ Aug. 29, 1868, p. 630; Vogt, ‘Lectures on Man,’ Eng. trans. p. 127.), who inhabit the northernmost islands of the Japan Archipelago, are the hairiest men in the world. With negroes the beard is scanty or wanting, and they rarely have whiskers; in both sexes the body is frequently almost destitute of fine down. (16. On the beards of negroes, Vogt, ‘Lectures,’ etc. p. 127; Waitz, ‘Introduct. to Anthropology,’ Engl. translat. 1863, vol. i. p. 96. It is remarkable that in the United States (‘Investigations in Military and Anthropological Statistics of American Soldiers,’ 1869, p. 569) the pure negroes and their crossed offspring seem to have bodies almost as hairy as Europeans.) On the other hand, the Papuans of the Malay Archipelago, who are nearly as black as negroes, possess well-developed beards. (17. Wallace, ‘The Malay Arch.’ vol. ii. 1869, p. 178.) In the Pacific Ocean the inhabitants of the Fiji Archipelago have large bushy beards, whilst those of the not distant archipelagoes of Tonga and Samoa are beardless; but these men belong to distinct races. In the Ellice group all the inhabitants belong to the same race; yet on one island alone, namely Nunemaya, “the men have splendid beards”; whilst on the other islands “they have, as a rule, a dozen straggling hairs for a beard.” (18. Dr. J. Barnard Davis on Oceanic Races, in ‘Anthropological Review,’ April 1870, pp. 185, 191.)

Throughout the great American continent the men may be said to be beardless; but in almost all the tribes a few short hairs are apt to appear on the face, especially in old age. With the tribes of North America, Catlin estimates that eighteen out of twenty men are completely destitute by nature of a beard; but occasionally there may be seen a man, who has neglected to pluck out the hairs at puberty, with a soft beard an inch or two in length. The Guaranys of Paraguay differ from all the surrounding tribes in having a small beard, and even some hair on the body, but no whiskers. (19. Catlin, ‘North American Indians,’ 3rd. ed. 1842, vol. ii. p. 227. On the Guaranys, see Azara, ‘Voyages dans l’Amérique Merid.’ tom. ii. 1809, p. 85; also Rengger, ‘Säugethiere von Paraguay,’ s. 3.) I am informed by Mr. D. Forbes, who particularly attended to this point, that the Aymaras and Quichuas of the Cordillera are remarkably hairless, yet in old age a few straggling hairs occasionally appear on the chin. The men of these two tribes have very little hair on the various parts of the body where hair grows abundantly in Europeans, and the women have none on the corresponding parts. The hair on the head, however, attains an extraordinary length in both sexes, often reaching almost to the ground; and this is likewise the case with some of the N. American tribes. In the amount of hair, and in the general shape of the body, the sexes of the American aborigines do not differ so much from each other, as in most other races. (20. Prof. and Mrs. Agassiz (‘Journey in Brazil,’ p. 530) remark that the sexes of the American Indians differ less than those of the negroes and of the higher races. See also Rengger, ibid. p. 3, on the Guaranys.) This fact is analogous with what occurs with some closely allied monkeys; thus the sexes of the chimpanzee are not as different as those of the orang or gorilla. (21. Rutimeyer, ‘Die Grenzen der Thierwelt; eine Betrachtung zu Darwin’s Lehre,’ 1868, s. 54.)

In the previous chapters we have seen that with mammals, birds, fishes, insects, etc., many characters, which there is every reason to believe were primarily gained through sexual selection by one sex, have been transferred to the other. As this same form of transmission has apparently prevailed much with mankind, it will save useless repetition if we discuss the origin of characters peculiar to the male sex together with certain other characters common to both sexes.

LAW OF BATTLE.

With savages, for instance, the Australians, the women are the constant cause of war both between members of the same tribe and between distinct tribes. So no doubt it was in ancient times; “nam fuit ante Helenam mulier teterrima belli causa.” With some of the North American Indians, the contest is reduced to a system. That excellent observer, Hearne (22. ‘A Journey from Prince of Wales Fort,’ 8vo. ed. Dublin, 1796, p. 104. Sir J. Lubbock (‘Origin of Civilisation,’ 1870, p. 69) gives other and similar cases in North America. For the Guanas of South America see Azara, ‘Voyages,’ etc. tom. ii. p. 94.), says:—“It has ever been the custom among these people for the men to wrestle for any woman to whom they are attached; and, of course, the strongest party always carries off the prize. A weak man, unless he be a good hunter, and well-beloved, is seldom permitted to keep a wife that a stronger man thinks worth his notice. This custom prevails throughout all the tribes, and causes a great spirit of emulation among their youth, who are upon all occasions, from their childhood, trying their strength and skill in wrestling.” With the Guanas of South America, Azara states that the men rarely marry till twenty years old or more, as before that age they cannot conquer their rivals.

Other similar facts could be given; but even if we had no evidence on this head, we might feel almost sure, from the analogy of the higher Quadrumana (23. On the fighting of the male gorillas, see Dr. Savage, in ‘Boston Journal of Natural History,’ vol. v. 1847, p. 423. On Presbytis entellus, see the ‘Indian Field,’ 1859, p. 146.), that the law of battle had prevailed with man during the early stages of his development. The occasional appearance at the present day of canine teeth which project above the others, with traces of a diastema or open space for the reception of the opposite canines, is in all probability a case of reversion to a former state, when the progenitors of man were provided with these weapons, like so many existing male Quadrumana. It was remarked in a former chapter that as man gradually became erect, and continually used his hands and arms for fighting with sticks and stones, as well as for the other purposes of life, he would have used his jaws and teeth less and less. The jaws, together with their muscles, would then have been reduced through disuse, as would the teeth through the not well understood principles of correlation and economy of growth; for we everywhere see that parts, which are no longer of service, are reduced in size. By such steps the original inequality between the jaws and teeth in the two sexes of mankind would ultimately have been obliterated. The case is almost parallel with that of many male Ruminants, in which the canine teeth have been reduced to mere rudiments, or have disappeared, apparently in consequence of the development of horns. As the prodigious difference between the skulls of the two sexes in the orang and gorilla stands in close relation with the development of the immense canine teeth in the males, we may infer that the reduction of the jaws and teeth in the early male progenitors of man must have led to a most striking and favourable change in his appearance.

There can be little doubt that the greater size and strength of man, in comparison with woman, together with his broader shoulders, more developed muscles, rugged outline of body, his greater courage and pugnacity, are all due in chief part to inheritance from his half-human male ancestors. These characters would, however, have been preserved or even augmented during the long ages of man’s savagery, by the success of the strongest and boldest men, both in the general struggle for life and in their contests for wives; a success which would have ensured their leaving a more numerous progeny than their less favoured brethren. It is not probable that the greater strength of man was primarily acquired through the inherited effects of his having worked harder than woman for his own subsistence and that of his family; for the women in all barbarous nations are compelled to work at least as hard as the men. With civilised people the arbitrament of battle for the possession of the women has long ceased; on the other hand, the men, as a general rule, have to work harder than the women for their joint subsistence, and thus their greater strength will have been kept up.

DIFFERENCE IN THE MENTAL POWERS OF THE TWO SEXES.

With respect to differences of this nature between man and woman, it is probable that sexual selection has played a highly important part. I am aware that some writers doubt whether there is any such inherent difference; but this is at least probable from the analogy of the lower animals which present other secondary sexual characters. No one disputes that the bull differs in disposition from the cow, the wild-boar from the sow, the stallion from the mare, and, as is well known to the keepers of menageries, the males of the larger apes from the females. Woman seems to differ from man in mental disposition, chiefly in her greater tenderness and less selfishness; and this holds good even with savages, as shewn by a well-known passage in Mungo Park’s Travels, and by statements made by many other travellers. Woman, owing to her maternal instincts, displays these qualities towards her infants in an eminent degree; therefore it is likely that she would often extend them towards her fellow-creatures. Man is the rival of other men; he delights in competition, and this leads to ambition which passes too easily into selfishness. These latter qualities seem to be his natural and unfortunate birthright. It is generally admitted that with woman the powers of intuition, of rapid perception, and perhaps of imitation, are more strongly marked than in man; but some, at least, of these faculties are characteristic of the lower races, and therefore of a past and lower state of civilisation.

The chief distinction in the intellectual powers of the two sexes is shewn by man’s attaining to a higher eminence, in whatever he takes up, than can woman—whether requiring deep thought, reason, or imagination, or merely the use of the senses and hands. If two lists were made of the most eminent men and women in poetry, painting, sculpture, music (inclusive both of composition and performance), history, science, and philosophy, with half-a-dozen names under each subject, the two lists would not bear comparison. We may also infer, from the law of the deviation from averages, so well illustrated by Mr. Galton, in his work on ‘Hereditary Genius,’ that if men are capable of a decided pre-eminence over women in many subjects, the average of mental power in man must be above that of woman.

Amongst the half-human progenitors of man, and amongst savages, there have been struggles between the males during many generations for the possession of the females. But mere bodily strength and size would do little for victory, unless associated with courage, perseverance, and determined energy. With social animals, the young males have to pass through many a contest before they win a female, and the older males have to retain their females by renewed battles. They have, also, in the case of mankind, to defend their females, as well as their young, from enemies of all kinds, and to hunt for their joint subsistence. But to avoid enemies or to attack them with success, to capture wild animals, and to fashion weapons, requires the aid of the higher mental faculties, namely, observation, reason, invention, or imagination. These various faculties will thus have been continually put to the test and selected during manhood; they will, moreover, have been strengthened by use during this same period of life. Consequently in accordance with the principle often alluded to, we might expect that they would at least tend to be transmitted chiefly to the male offspring at the corresponding period of manhood.

Now, when two men are put into competition, or a man with a woman, both possessed of every mental quality in equal perfection, save that one has higher energy, perseverance, and courage, the latter will generally become more eminent in every pursuit, and will gain the ascendancy. (24. J. Stuart Mill remarks (‘The Subjection of Women,’ 1869, p. 122), “The things in which man most excels woman are those which require most plodding, and long hammering at single thoughts.” What is this but energy and perseverance?) He may be said to possess genius—for genius has been declared by a great authority to be patience; and patience, in this sense, means unflinching, undaunted perseverance. But this view of genius is perhaps deficient; for without the higher powers of the imagination and reason, no eminent success can be gained in many subjects. These latter faculties, as well as the former, will have been developed in man, partly through sexual selection,—that is, through the contest of rival males, and partly through natural selection, that is, from success in the general struggle for life; and as in both cases the struggle will have been during maturity, the characters gained will have been transmitted more fully to the male than to the female offspring. It accords in a striking manner with this view of the modification and re-inforcement of many of our mental faculties by sexual selection, that, firstly, they notoriously undergo a considerable change at puberty (25. Maudsley, ‘Mind and Body,’ p. 31.), and, secondly, that eunuchs remain throughout life inferior in these same qualities. Thus, man has ultimately become superior to woman. It is, indeed, fortunate that the law of the equal transmission of characters to both sexes prevails with mammals; otherwise, it is probable that man would have become as superior in mental endowment to woman, as the peacock is in ornamental plumage to the peahen.

It must be borne in mind that the tendency in characters acquired by either sex late in life, to be transmitted to the same sex at the same age, and of early acquired characters to be transmitted to both sexes, are rules which, though general, do not always hold. If they always held good, we might conclude (but I here exceed my proper bounds) that the inherited effects of the early education of boys and girls would be transmitted equally to both sexes; so that the present inequality in mental power between the sexes would not be effaced by a similar course of early training; nor can it have been caused by their dissimilar early training. In order that woman should reach the same standard as man, she ought, when nearly adult, to be trained to energy and perseverance, and to have her reason and imagination exercised to the highest point; and then she would probably transmit these qualities chiefly to her adult daughters. All women, however, could not be thus raised, unless during many generations those who excelled in the above robust virtues were married, and produced offspring in larger numbers than other women. As before remarked of bodily strength, although men do not now fight for their wives, and this form of selection has passed away, yet during manhood, they generally undergo a severe struggle in order to maintain themselves and their families; and this will tend to keep up or even increase their mental powers, and, as a consequence, the present inequality between the sexes. (26. An observation by Vogt bears on this subject: he says, “It is a remarkable circumstance, that the difference between the sexes, as regards the cranial cavity, increases with the development of the race, so that the male European excels much more the female, than the negro the negress. Welcker confirms this statement of Huschke from his measurements of negro and German skulls.” But Vogt admits (‘Lectures on Man,’ Eng. translat. 1864, p. 81) that more observations are requisite on this point.

VOICE AND MUSICAL POWERS.

In some species of Quadrumana there is a great difference between the adult sexes, in the power of their voices and in the development of the vocal organs; and man appears to have inherited this difference from his early progenitors. His vocal cords are about one-third longer than in woman, or than in boys; and emasculation produces the same effect on him as on the lower animals, for it “arrests that prominent growth of the thyroid, etc., which accompanies the elongation of the cords.” (27. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 603.) With respect to the cause of this difference between the sexes, I have nothing to add to the remarks in the last chapter on the probable effects of the long-continued use of the vocal organs by the male under the excitement of love, rage and jealousy. According to Sir Duncan Gibb (28. ‘Journal of the Anthropological Society,’ April 1869, p. lvii. and lxvi.), the voice and the form of the larynx differ in the different races of mankind; but with the Tartars, Chinese, etc., the voice of the male is said not to differ so much from that of the female, as in most other races.

The capacity and love for singing or music, though not a sexual character in man, must not here be passed over. Although the sounds emitted by animals of all kinds serve many purposes, a strong case can be made out, that the vocal organs were primarily used and perfected in relation to the propagation of the species. Insects and some few spiders are the lowest animals which voluntarily produce any sound; and this is generally effected by the aid of beautifully constructed stridulating organs, which are often confined to the males. The sounds thus produced consist, I believe in all cases, of the same note, repeated rhythmically (29. Dr. Scudder, ‘Notes on Stridulation,’ in ‘Proc. Boston Soc. of Nat. Hist.’ vol. xi. April 1868.); and this is sometimes pleasing even to the ears of man. The chief and, in some cases, exclusive purpose appears to be either to call or charm the opposite sex.

The sounds produced by fishes are said in some cases to be made only by the males during the breeding-season. All the air-breathing Vertebrata necessarily possess an apparatus for inhaling and expelling air, with a pipe capable of being closed at one end. Hence when the primeval members of this class were strongly excited and their muscles violently contracted, purposeless sounds would almost certainly have been produced; and these, if they proved in any way serviceable, might readily have been modified or intensified by the preservation of properly adapted variations. The lowest Vertebrates which breathe air are Amphibians; and of these, frogs and toads possess vocal organs, which are incessantly used during the breeding-season, and which are often more highly developed in the male than in the female. The male alone of the tortoise utters a noise, and this only during the season of love. Male alligators roar or bellow during the same season. Every one knows how much birds use their vocal organs as a means of courtship; and some species likewise perform what may be called instrumental music.

In the class of Mammals, with which we are here more particularly concerned, the males of almost all the species use their voices during the breeding-season much more than at any other time; and some are absolutely mute excepting at this season. With other species both sexes, or only the females, use their voices as a love-call. Considering these facts, and that the vocal organs of some quadrupeds are much more largely developed in the male than in the female, either permanently or temporarily during the breeding-season; and considering that in most of the lower classes the sounds produced by the males, serve not only to call but to excite or allure the female, it is a surprising fact that we have not as yet any good evidence that these organs are used by male mammals to charm the females. The American Mycetes caraya perhaps forms an exception, as does the Hylobates agilis, an ape allied to man. This gibbon has an extremely loud but musical voice. Mr. Waterhouse states (30. Given in W.C.L. Martin’s ‘General Introduction to Natural History of Mamm. Animals,’ 1841, p. 432; Owen, ‘Anatomy of Vertebrates,’ vol. iii, p. 600.), “It appeared to me that in ascending and descending the scale, the intervals were always exactly half-tones; and I am sure that the highest note was the exact octave to the lowest. The quality of the notes is very musical; and I do not doubt that a good violinist would be able to give a correct idea of the gibbon’s composition, excepting as regards its loudness.” Mr. Waterhouse then gives the notes. Professor Owen, who is a musician, confirms the foregoing statement, and remarks, though erroneously, that this gibbon “alone of brute mammals may be said to sing.” It appears to be much excited after its performance. Unfortunately, its habits have never been closely observed in a state of nature; but from the analogy of other animals, it is probable that it uses its musical powers more especially during the season of courtship.

This gibbon is not the only species in the genus which sings, for my son, Francis Darwin, attentively listened in the Zoological Gardens to H. leuciscus whilst singing a cadence of three notes, in true musical intervals and with a clear musical tone. It is a more surprising fact that certain rodents utter musical sounds. Singing mice have often been mentioned and exhibited, but imposture has commonly been suspected. We have, however, at last a clear account by a well-known observer, the Rev. S. Lockwood (31. The ‘American Naturalist,’ 1871, p. 761.), of the musical powers of an American species, the Hesperomys cognatus, belonging to a genus distinct from that of the English mouse. This little animal was kept in confinement, and the performance was repeatedly heard. In one of the two chief songs, “the last bar would frequently be prolonged to two or three; and she would sometimes change from C sharp and D, to C natural and D, then warble on these two notes awhile, and wind up with a quick chirp on C sharp and D. The distinctness between the semitones was very marked, and easily appreciable to a good ear.” Mr. Lockwood gives both songs in musical notation; and adds that though this little mouse “had no ear for time, yet she would keep to the key of B (two flats) and strictly in a major key.”...”Her soft clear voice falls an octave with all the precision possible; then at the wind up, it rises again into a very quick trill on C sharp and D.”

A critic has asked how the ears of man, and he ought to have added of other animals, could have been adapted by selection so as to distinguish musical notes. But this question shews some confusion on the subject; a noise is the sensation resulting from the co-existence of several aerial “simple vibrations” of various periods, each of which intermits so frequently that its separate existence cannot be perceived. It is only in the want of continuity of such vibrations, and in their want of harmony inter se, that a noise differs from a musical note. Thus an ear to be capable of discriminating noises—and the high importance of this power to all animals is admitted by every one—must be sensitive to musical notes. We have evidence of this capacity even low down in the animal scale: thus Crustaceans are provided with auditory hairs of different lengths, which have been seen to vibrate when the proper musical notes are struck. (32. Helmholtz, ‘Theorie Phys. de la Musique,’ 1868, p. 187.) As stated in a previous chapter, similar observations have been made on the hairs of the antennae of gnats. It has been positively asserted by good observers that spiders are attracted by music. It is also well known that some dogs howl when hearing particular tones. (33. Several accounts have been published to this effect. Mr. Peach writes to me that an old dog of his howls when B flat is sounded on the flute, and to no other note. I may add another instance of a dog always whining, when one note on a concertina, which was out of tune, was played.) Seals apparently appreciate music, and their fondness for it “was well known to the ancients, and is often taken advantage of by the hunters at the present day.” (34. Mr. R. Brown, in ‘Proc. Zool. Soc.’ 1868, p. 410.)

Therefore, as far as the mere perception of musical notes is concerned, there seems no special difficulty in the case of man or of any other animal. Helmholtz has explained on physiological principles why concords are agreeable, and discords disagreeable to the human ear; but we are little concerned with these, as music in harmony is a late invention. We are more concerned with melody, and here again, according to Helmholtz, it is intelligible why the notes of our musical scale are used. The ear analyses all sounds into their component “simple vibrations,” although we are not conscious of this analysis. In a musical note the lowest in pitch of these is generally predominant, and the others which are less marked are the octave, the twelfth, the second octave, etc., all harmonies of the fundamental predominant note; any two notes of our scale have many of these harmonic over-tones in common. It seems pretty clear then, that if an animal always wished to sing precisely the same song, he would guide himself by sounding those notes in succession, which possess many over-tones in common—that is, he would choose for his song, notes which belong to our musical scale.

But if it be further asked why musical tones in a certain order and rhythm give man and other animals pleasure, we can no more give the reason than for the pleasantness of certain tastes and smells. That they do give pleasure of some kind to animals, we may infer from their being produced during the season of courtship by many insects, spiders, fishes, amphibians, and birds; for unless the females were able to appreciate such sounds and were excited or charmed by them, the persevering efforts of the males, and the complex structures often possessed by them alone, would be useless; and this it is impossible to believe.

Human song is generally admitted to be the basis or origin of instrumental music. As neither the enjoyment nor the capacity of producing musical notes are faculties of the least use to man in reference to his daily habits of life, they must be ranked amongst the most mysterious with which he is endowed. They are present, though in a very rude condition, in men of all races, even the most savage; but so different is the taste of the several races, that our music gives no pleasure to savages, and their music is to us in most cases hideous and unmeaning. Dr. Seemann, in some interesting remarks on this subject (35. ‘Journal of Anthropological Society,’ Oct. 1870, p. clv. See also the several later chapters in Sir John Lubbock’s ‘Prehistoric Times,’ 2nd ed. 1869, which contain an admirable account of the habits of savages.), “doubts whether even amongst the nations of Western Europe, intimately connected as they are by close and frequent intercourse, the music of the one is interpreted in the same sense by the others. By travelling eastwards we find that there is certainly a different language of music. Songs of joy and dance-accompaniments are no longer, as with us, in the major keys, but always in the minor.” Whether or not the half-human progenitors of man possessed, like the singing gibbons, the capacity of producing, and therefore no doubt of appreciating, musical notes, we know that man possessed these faculties at a very remote period. M. Lartet has described two flutes made out of the bones and horns of the reindeer, found in caves together with flint tools and the remains of extinct animals. The arts of singing and of dancing are also very ancient, and are now practised by all or nearly all the lowest races of man. Poetry, which may be considered as the offspring of song, is likewise so ancient, that many persons have felt astonished that it should have arisen during the earliest ages of which we have any record.

We see that the musical faculties, which are not wholly deficient in any race, are capable of prompt and high development, for Hottentots and Negroes have become excellent musicians, although in their native countries they rarely practise anything that we should consider music. Schweinfurth, however, was pleased with some of the simple melodies which he heard in the interior of Africa. But there is nothing anomalous in the musical faculties lying dormant in man: some species of birds which never naturally sing, can without much difficulty be taught to do so; thus a house-sparrow has learnt the song of a linnet. As these two species are closely allied, and belong to the order of Insessores, which includes nearly all the singing-birds in the world, it is possible that a progenitor of the sparrow may have been a songster. It is more remarkable that parrots, belonging to a group distinct from the Insessores, and having differently constructed vocal organs, can be taught not only to speak, but to pipe or whistle tunes invented by man, so that they must have some musical capacity. Nevertheless it would be very rash to assume that parrots are descended from some ancient form which was a songster. Many cases could be advanced of organs and instincts originally adapted for one purpose, having been utilised for some distinct purpose. (36. Since this chapter was printed, I have seen a valuable article by Mr. Chauncey Wright (‘North American Review,’ Oct. 1870, page 293), who, in discussing the above subject, remarks, “There are many consequences of the ultimate laws or uniformities of nature, through which the acquisition of one useful power will bring with it many resulting advantages as well as limiting disadvantages, actual or possible, which the principle of utility may not have comprehended in its action.” As I have attempted to shew in an early chapter of this work, this principle has an important bearing on the acquisition by man of some of his mental characteristics.) Hence the capacity for high musical development which the savage races of man possess, may be due either to the practice by our semi-human progenitors of some rude form of music, or simply to their having acquired the proper vocal organs for a different purpose. But in this latter case we must assume, as in the above instance of parrots, and as seems to occur with many animals, that they already possessed some sense of melody.

Music arouses in us various emotions, but not the more terrible ones of horror, fear, rage, etc. It awakens the gentler feelings of tenderness and love, which readily pass into devotion. In the Chinese annals it is said, “Music hath the power of making heaven descend upon earth.” It likewise stirs up in us the sense of triumph and the glorious ardour for war. These powerful and mingled feelings may well give rise to the sense of sublimity. We can concentrate, as Dr. Seemann observes, greater intensity of feeling in a single musical note than in pages of writing. It is probable that nearly the same emotions, but much weaker and far less complex, are felt by birds when the male pours forth his full volume of song, in rivalry with other males, to captivate the female. Love is still the commonest theme of our songs. As Herbert Spencer remarks, “music arouses dormant sentiments of which we had not conceived the possibility, and do not know the meaning; or, as Richter says, tells us of things we have not seen and shall not see.” Conversely, when vivid emotions are felt and expressed by the orator, or even in common speech, musical cadences and rhythm are instinctively used. The negro in Africa when excited often bursts forth in song; “another will reply in song, whilst the company, as if touched by a musical wave, murmur a chorus in perfect unison.” (37. Winwood Reade, ‘The Martyrdom of Man,’ 1872, p. 441, and ‘African Sketch Book,’ 1873, vol. ii. p. 313.) Even monkeys express strong feelings in different tones—anger and impatience by low,—fear and pain by high notes. (38. Rengger, ‘Säugethiere von Paraguay,’ s. 49.) The sensations and ideas thus excited in us by music, or expressed by the cadences of oratory, appear from their vagueness, yet depth, like mental reversions to the emotions and thoughts of a long-past age.

All these facts with respect to music and impassioned speech become intelligible to a certain extent, if we may assume that musical tones and rhythm were used by our half-human ancestors, during the season of courtship, when animals of all kinds are excited not only by love, but by the strong passions of jealousy, rivalry, and triumph. From the deeply-laid principle of inherited associations, musical tones in this case would be likely to call up vaguely and indefinitely the strong emotions of a long-past age. As we have every reason to suppose that articulate speech is one of the latest, as it certainly is the highest, of the arts acquired by man, and as the instinctive power of producing musical notes and rhythms is developed low down in the animal series, it would be altogether opposed to the principle of evolution, if we were to admit that man’s musical capacity has been developed from the tones used in impassioned speech. We must suppose that the rhythms and cadences of oratory are derived from previously developed musical powers. (39. See the very interesting discussion on the ‘Origin and Function of Music,’ by Mr. Herbert Spencer, in his collected ‘Essays,’ 1858, p. 359. Mr. Spencer comes to an exactly opposite conclusion to that at which I have arrived. He concludes, as did Diderot formerly, that the cadences used in emotional speech afford the foundation from which music has been developed; whilst I conclude that musical notes and rhythm were first acquired by the male or female progenitors of mankind for the sake of charming the opposite sex. Thus musical tones became firmly associated with some of the strongest passions an animal is capable of feeling, and are consequently used instinctively, or through association when strong emotions are expressed in speech. Mr. Spencer does not offer any satisfactory explanation, nor can I, why high or deep notes should be expressive, both with man and the lower animals, of certain emotions. Mr. Spencer gives also an interesting discussion on the relations between poetry, recitative and song.) We can thus understand how it is that music, dancing, song, and poetry are such very ancient arts. We may go even further than this, and, as remarked in a former chapter, believe that musical sounds afforded one of the bases for the development of language. (40. I find in Lord Monboddo’s ‘Origin of Language,’ vol. i. 1774, p. 469, that Dr. Blacklock likewise thought “that the first language among men was music, and that before our ideas were expressed by articulate sounds, they were communicated by tones varied according to different degrees of gravity and acuteness.”)

As the males of several quadrumanous animals have their vocal organs much more developed than in the females, and as a gibbon, one of the anthropomorphous apes, pours forth a whole octave of musical notes and may be said to sing, it appears probable that the progenitors of man, either the males or females or both sexes, before acquiring the power of expressing their mutual love in articulate language, endeavoured to charm each other with musical notes and rhythm. So little is known about the use of the voice by the Quadrumana during the season of love, that we have no means of judging whether the habit of singing was first acquired by our male or female ancestors. Women are generally thought to possess sweeter voices than men, and as far as this serves as any guide, we may infer that they first acquired musical powers in order to attract the other sex. (41. See an interesting discussion on this subject by Haeckel, ‘Generelle Morphologie,’ B. ii. 1866, s. 246.) But if so, this must have occurred long ago, before our ancestors had become sufficiently human to treat and value their women merely as useful slaves. The impassioned orator, bard, or musician, when with his varied tones and cadences he excites the strongest emotions in his hearers, little suspects that he uses the same means by which his half-human ancestors long ago aroused each other’s ardent passions, during their courtship and rivalry.

THE INFLUENCE OF BEAUTY IN DETERMINING THE MARRIAGES OF MANKIND.

In civilised life man is largely, but by no means exclusively, influenced in the choice of his wife by external appearance; but we are chiefly concerned with primeval times, and our only means of forming a judgment on this subject is to study the habits of existing semi-civilised and savage nations. If it can be shewn that the men of different races prefer women having various characteristics, or conversely with the women, we have then to enquire whether such choice, continued during many generations, would produce any sensible effect on the race, either on one sex or both according to the form of inheritance which has prevailed.

It will be well first to shew in some detail that savages pay the greatest attention to their personal appearance. (42. A full and excellent account of the manner in which savages in all parts of the world ornament themselves, is given by the Italian traveller, Professor Mantegazza, ‘Rio de la Plata, Viaggi e Studi,’ 1867, pp. 525-545; all the following statements, when other references are not given, are taken from this work. See, also, Waitz, ‘Introduction to Anthropology,’ Eng. translat. vol. i. 1863, p. 275, et passim. Lawrence also gives very full details in his ‘Lectures on Physiology,’ 1822. Since this chapter was written Sir J. Lubbock has published his ‘Origin of Civilisation,’ 1870, in which there is an interesting chapter on the present subject, and from which (pp. 42, 48) I have taken some facts about savages dyeing their teeth and hair, and piercing their teeth.) That they have a passion for ornament is notorious; and an English philosopher goes so far as to maintain, that clothes were first made for ornament and not for warmth. As Professor Waitz remarks, “however poor and miserable man is, he finds a pleasure in adorning himself.” The extravagance of the naked Indians of South America in decorating themselves is shewn “by a man of large stature gaining with difficulty enough by the labour of a fortnight to procure in exchange the chica necessary to paint himself red.” (43. Humboldt, ‘Personal Narrative,’ Eng. translat. vol. iv. p. 515; on the imagination shewn in painting the body, p. 522; on modifying the form of the calf of the leg, p. 466.) The ancient barbarians of Europe during the Reindeer period brought to their caves any brilliant or singular objects which they happened to find. Savages at the present day everywhere deck themselves with plumes, necklaces, armlets, ear-rings, etc. They paint themselves in the most diversified manner. “If painted nations,” as Humboldt observes, “had been examined with the same attention as clothed nations, it would have been perceived that the most fertile imagination and the most mutable caprice have created the fashions of painting, as well as those of garments.”

In one part of Africa the eyelids are coloured black; in another the nails are coloured yellow or purple. In many places the hair is dyed of various tints. In different countries the teeth are stained black, red, blue, etc., and in the Malay Archipelago it is thought shameful to have white teeth “like those of a dog.” Not one great country can be named, from the polar regions in the north to New Zealand in the south, in which the aborigines do not tattoo themselves. This practice was followed by the Jews of old, and by the ancient Britons. In Africa some of the natives tattoo themselves, but it is a much more common practice to raise protuberances by rubbing salt into incisions made in various parts of the body; and these are considered by the inhabitants of Kordofan and Darfur “to be great personal attractions.” In the Arab countries no beauty can be perfect until the cheeks “or temples have been gashed.” (44. ‘The Nile Tributaries,’ 1867; ‘The Albert N’yanza,’ 1866, vol. i. p. 218.) In South America, as Humboldt remarks, “a mother would be accused of culpable indifference towards her children, if she did not employ artificial means to shape the calf of the leg after the fashion of the country.” In the Old and New Worlds the shape of the skull was formerly modified during infancy in the most extraordinary manner, as is still the case in many places, and such deformities are considered ornamental. For instance, the savages of Colombia (45. Quoted by Prichard, ‘Physical History of Mankind,’ 4th ed. vol. i. 1851, p. 321.) deem a much flattened head “an essential point of beauty.”

The hair is treated with especial care in various countries; it is allowed to grow to full length, so as to reach to the ground, or is combed into “a compact frizzled mop, which is the Papuan’s pride and glory.” (46. On the Papuans, Wallace, ‘The Malay Archipelago,’ vol. ii. p. 445. On the coiffure of the Africans, Sir S. Baker, ‘The Albert N’yanza,’ vol. i. p. 210.) In northern Africa “a man requires a period of from eight to ten years to perfect his coiffure.” With other nations the head is shaved, and in parts of South America and Africa even the eyebrows and eyelashes are eradicated. The natives of the Upper Nile knock out the four front teeth, saying that they do not wish to resemble brutes. Further south, the Batokas knock out only the two upper incisors, which, as Livingstone (47. ‘Travels,’ p. 533.) remarks, gives the face a hideous appearance, owing to the prominence of the lower jaw; but these people think the presence of the incisors most unsightly, and on beholding some Europeans, cried out, “Look at the great teeth!” The chief Sebituani tried in vain to alter this fashion. In various parts of Africa and in the Malay Archipelago the natives file the incisors into points like those of a saw, or pierce them with holes, into which they insert studs.

As the face with us is chiefly admired for its beauty, so with savages it is the chief seat of mutilation. In all quarters of the world the septum, and more rarely the wings of the nose are pierced; rings, sticks, feathers, and other ornaments being inserted into the holes. The ears are everywhere pierced and similarly ornamented, and with the Botocudos and Lenguas of South America the hole is gradually so much enlarged that the lower edge touches the shoulder. In North and South America and in Africa either the upper or lower lip is pierced; and with the Botocudos the hole in the lower lip is so large that a disc of wood, four inches in diameter, is placed in it. Mantegazza gives a curious account of the shame felt by a South American native, and of the ridicule which he excited, when he sold his tembeta,—the large coloured piece of wood which is passed through the hole. In Central Africa the women perforate the lower lip and wear a crystal, which, from the movement of the tongue, has “a wriggling motion, indescribably ludicrous during conversation.” The wife of the chief of Latooka told Sir S. Baker (49. ‘The Albert N’yanza,’ 1866, vol. i. p. 217.) that Lady Baker “would be much improved if she would extract her four front teeth from the lower jaw, and wear the long pointed polished crystal in her under lip.” Further south with the Makalolo, the upper lip is perforated, and a large metal and bamboo ring, called a pelele, is worn in the hole. “This caused the lip in one case to project two inches beyond the tip of the nose; and when the lady smiled, the contraction of the muscles elevated it over the eyes. ‘Why do the women wear these things?’ the venerable chief, Chinsurdi, was asked. Evidently surprised at such a stupid question, he replied, ‘For beauty! They are the only beautiful things women have; men have beards, women have none. What kind of a person would she be without the pelele? She would not be a woman at all with a mouth like a man, but no beard.’” (49. Livingstone, ‘British Association,’ 1860; report given in the ‘Athenaeum,’ July 7, 1860, p. 29.)

Hardly any part of the body, which can be unnaturally modified, has escaped. The amount of suffering thus caused must have been extreme, for many of the operations require several years for their completion, so that the idea of their necessity must be imperative. The motives are various; the men paint their bodies to make themselves appear terrible in battle; certain mutilations are connected with religious rites, or they mark the age of puberty, or the rank of the man, or they serve to distinguish the tribes. Amongst savages the same fashions prevail for long periods (50. Sir S. Baker (ibid. vol. i. p. 210) speaking of the natives of Central Africa says, “every tribe has a distinct and unchanging fashion for dressing the hair.” See Agassiz (‘Journey in Brazil,’ 1868, p. 318) on invariability of the tattooing of Amazonian Indians.), and thus mutilations, from whatever cause first made, soon come to be valued as distinctive marks. But self-adornment, vanity, and the admiration of others, seem to be the commonest motives. In regard to tattooing, I was told by the missionaries in New Zealand that when they tried to persuade some girls to give up the practice, they answered, “We must just have a few lines on our lips; else when we grow old we shall be so very ugly.” With the men of New Zealand, a most capable judge (51. Rev. R. Taylor, ‘New Zealand and its Inhabitants,’ 1855, p. 152.) says, “to have fine tattooed faces was the great ambition of the young, both to render themselves attractive to the ladies, and conspicuous in war.” A star tattooed on the forehead and a spot on the chin are thought by the women in one part of Africa to be irresistible attractions. (52. Mantegazza, ‘Viaggi e Studi,’ p. 542.) In most, but not all parts of the world, the men are more ornamented than the women, and often in a different manner; sometimes, though rarely, the women are hardly at all ornamented. As the women are made by savages to perform the greatest share of the work, and as they are not allowed to eat the best kinds of food, so it accords with the characteristic selfishness of man that they should not be allowed to obtain, or use the finest ornaments. Lastly, it is a remarkable fact, as proved by the foregoing quotations, that the same fashions in modifying the shape of the head, in ornamenting the hair, in painting, tattooing, in perforating the nose, lips, or ears, in removing or filing the teeth, etc., now prevail, and have long prevailed, in the most distant quarters of the world. It is extremely improbable that these practices, followed by so many distinct nations, should be due to tradition from any common source. They indicate the close similarity of the mind of man, to whatever race he may belong, just as do the almost universal habits of dancing, masquerading, and making rude pictures.

Having made these preliminary remarks on the admiration felt by savages for various ornaments, and for deformities most unsightly in our eyes, let us see how far the men are attracted by the appearance of their women, and what are their ideas of beauty. I have heard it maintained that savages are quite indifferent about the beauty of their women, valuing them solely as slaves; it may therefore be well to observe that this conclusion does not at all agree with the care which the women take in ornamenting themselves, or with their vanity. Burchell (53. ‘Travels in South Africa,’ 1824, vol. i. p. 414.) gives an amusing account of a Bush-woman who used as much grease, red ochre, and shining powder “as would have ruined any but a very rich husband.” She displayed also “much vanity and too evident a consciousness of her superiority.” Mr. Winwood Reade informs me that the negroes of the West Coast often discuss the beauty of their women. Some competent observers have attributed the fearfully common practice of infanticide partly to the desire felt by the women to retain their good looks. (54. See, for references, Gerland, ‘Ueber das Aussterben der Naturvölker,’ 1868, ss. 51, 53, 55; also Azara, ‘Voyages,’ etc., tom. ii. p. 116.) In several regions the women wear charms and use love-philters to gain the affections of the men; and Mr. Brown enumerates four plants used for this purpose by the women of North-Western America. (55. On the vegetable productions used by the North-Western American Indians, see ‘Pharmaceutical Journal,’ vol. x.)

Hearne (56. ‘A Journey from Prince of Wales Fort,’ 8vo. ed. 1796, p. 89.), an excellent observer, who lived many years with the American Indians, says, in speaking of the women, “Ask a Northern Indian what is beauty, and he will answer, a broad flat face, small eyes, high cheek-bones, three or four broad black lines across each cheek, a low forehead, a large broad chin, a clumsy hook nose, a tawny hide, and breasts hanging down to the belt.” Pallas, who visited the northern parts of the Chinese empire, says, “those women are preferred who have the Mandschu form; that is to say, a broad face, high cheek-bones, very broad noses, and enormous ears”(57. Quoted by Prichard, ‘Physical History of Mankind,’ 3rd ed. vol. iv. 1844, p. 519; Vogt, ‘Lectures on Man,’ Eng. translat. p. 129. On the opinion of the Chinese on the Cingalese, E. Tennent, ‘Ceylon,’ 1859, vol. ii. p. 107.); and Vogt remarks that the obliquity of the eye, which is proper to the Chinese and Japanese, is exaggerated in their pictures for the purpose, as it “seems, of exhibiting its beauty, as contrasted with the eye of the red-haired barbarians.” It is well known, as Huc repeatedly remarks, that the Chinese of the interior think Europeans hideous, with their white skins and prominent noses. The nose is far from being too prominent, according to our ideas, in the natives of Ceylon; yet “the Chinese in the seventh century, accustomed to the flat features of the Mongol races, were surprised at the prominent noses of the Cingalese; and Thsang described them as having ‘the beak of a bird, with the body of a man.’”

Finlayson, after minutely describing the people of Cochin China, says that their rounded heads and faces are their chief characteristics; and, he adds, “the roundness of the whole countenance is more striking in the women, who are reckoned beautiful in proportion as they display this form of face.” The Siamese have small noses with divergent nostrils, a wide mouth, rather thick lips, a remarkably large face, with very high and broad cheek-bones. It is, therefore, not wonderful that “beauty, according to our notion, is a stranger to them. Yet they consider their own females to be much more beautiful than those of Europe.” (58. Prichard, as taken from Crawfurd and Finlayson, ‘Phys. Hist. of Mankind,’ vol. iv. pp. 534, 535.)

It is well known that with many Hottentot women the posterior part of the body projects in a wonderful manner; they are steatopygous; and Sir Andrew Smith is certain that this peculiarity is greatly admired by the men. (59. Idem illustrissimus viator dixit mihi praecinctorium vel tabulam foeminae, quod nobis teterrimum est, quondam permagno aestimari ab hominibus in hac gente. Nunc res mutata est, et censent talem conformationem minime optandam esse.) He once saw a woman who was considered a beauty, and she was so immensely developed behind, that when seated on level ground she could not rise, and had to push herself along until she came to a slope. Some of the women in various negro tribes have the same peculiarity; and, according to Burton, the Somal men are said to choose their wives by ranging them in a line, and by picking her out who projects farthest a tergo. Nothing can be more hateful to a negro than the opposite form.” (60. The ‘Anthropological Review,’ November 1864, p. 237. For additional references, see Waitz, ‘Introduction to Anthropology,’ Eng. translat., 1863, vol. i. p. 105.)

With respect to colour, the negroes rallied Mungo Park on the whiteness of his skin and the prominence of his nose, both of which they considered as “unsightly and unnatural conformations.” He in return praised the glossy jet of their skins and the lovely depression of their noses; this they said was “honeymouth,” nevertheless they gave him food. The African Moors, also, “knitted their brows and seemed to shudder” at the whiteness of his skin. On the eastern coast, the negro boys when they saw Burton, cried out, “Look at the white man; does he not look like a white ape?” On the western coast, as Mr. Winwood Reade informs me, the negroes admire a very black skin more than one of a lighter tint. But their horror of whiteness may be attributed, according to this same traveller, partly to the belief held by most negroes that demons and spirits are white, and partly to their thinking it a sign of ill-health.

The Banyai of the more southern part of the continent are negroes, but “a great many of them are of a light coffee-and-milk colour, and, indeed, this colour is considered handsome throughout the whole country”; so that here we have a different standard of taste. With the Kaffirs, who differ much from negroes, “the skin, except among the tribes near Delagoa Bay, is not usually black, the prevailing colour being a mixture of black and red, the most common shade being chocolate. Dark complexions, as being most common, are naturally held in the highest esteem. To be told that he is light-coloured, or like a white man, would be deemed a very poor compliment by a Kaffir. I have heard of one unfortunate man who was so very fair that no girl would marry him.” One of the titles of the Zulu king is, “You who are black.” (61. Mungo Park’s ‘Travels in Africa,’ 4to. 1816, pp. 53, 131. Burton’s statement is quoted by Schaaffhausen, ‘Archiv. fur Anthropologie,’ 1866, s. 163. On the Banyai, Livingstone, ‘Travels,’ p. 64. On the Kaffirs, the Rev. J. Shooter, ‘The Kafirs of Natal and the Zulu Country,’ 1857, p. 1.) Mr. Galton, in speaking to me about the natives of S. Africa, remarked that their ideas of beauty seem very different from ours; for in one tribe two slim, slight, and pretty girls were not admired by the natives.

Turning to other quarters of the world; in Java, a yellow, not a white girl, is considered, according to Madame Pfeiffer, a beauty. A man of Cochin China “spoke with contempt of the wife of the English Ambassador, that she had white teeth like a dog, and a rosy colour like that of potato-flowers.” We have seen that the Chinese dislike our white skin, and that the N. Americans admire “a tawny hide.” In S. America, the Yuracaras, who inhabit the wooded, damp slopes of the eastern Cordillera, are remarkably pale-coloured, as their name in their own language expresses; nevertheless they consider European women as very inferior to their own. (62. For the Javans and Cochin-Chinese, see Waitz, ‘Introduct. to Anthropology,’ Eng. translat. vol. i. p. 305. On the Yuracaras, A. d’Orbigny, as quoted in Prichard, ‘Physical History of Mankind,’ vol. v. 3rd ed. p. 476.)

In several of the tribes of North America the hair on the head grows to a wonderful length; and Catlin gives a curious proof how much this is esteemed, for the chief of the Crows was elected to this office from having the longest hair of any man in the tribe, namely ten feet and seven inches. The Aymaras and Quichuas of S. America, likewise have very long hair; and this, as Mr. D. Forbes informs me, is so much valued as a beauty, that cutting it off was the severest punishment which he could inflict on them. In both the Northern and Southern halves of the continent the natives sometimes increase the apparent length of their hair by weaving into it fibrous substances. Although the hair on the head is thus cherished, that on the face is considered by the North American Indians “as very vulgar,” and every hair is carefully eradicated. This practice prevails throughout the American continent from Vancouver’s Island in the north to Tierra del Fuego in the south. When York Minster, a Fuegian on board the “Beagle,” was taken back to his country, the natives told him he ought to pull out the few short hairs on his face. They also threatened a young missionary, who was left for a time with them, to strip him naked, and pluck the hair from his face and body, yet he was far from being a hairy man. This fashion is carried so far that the Indians of Paraguay eradicate their eyebrows and eyelashes, saying that they do not wish to be like horses. (63. ‘North American Indians,’ by G. Catlin, 3rd ed., 1842, vol. i. p. 49; vol. ii, p. 227. On the natives of Vancouver’s Island, see Sproat, ‘Scenes and Studies of Savage Life,’ 1868, p. 25. On the Indians of Paraguay, Azara, ‘Voyages,’ tom. ii. p. 105.)

It is remarkable that throughout the world the races which are almost completely destitute of a beard dislike hairs on the face and body, and take pains to eradicate them. The Kalmucks are beardless, and they are well known, like the Americans, to pluck out all straggling hairs; and so it is with the Polynesians, some of the Malays, and the Siamese. Mr. Veitch states that the Japanese ladies “all objected to our whiskers, considering them very ugly, and told us to cut them off, and be like Japanese men.” The New Zealanders have short, curled beards; yet they formerly plucked out the hairs on the face. They had a saying that “there is no woman for a hairy man;” but it would appear that the fashion has changed in New Zealand, perhaps owing to the presence of Europeans, and I am assured that beards are now admired by the Maories. (64. On the Siamese, Prichard, ibid. vol. iv. p. 533. On the Japanese, Veitch in ‘Gardeners’ Chronicle,’ 1860, p. 1104. On the New Zealanders, Mantegazza, ‘Viaggi e Studi,’ 1867, p. 526. For the other nations mentioned, see references in Lawrence, ‘Lectures on Physiology,’ etc., 1822, p. 272.)

On the other hand, bearded races admire and greatly value their beards; among the Anglo-Saxons every part of the body had a recognised value; “the loss of the beard being estimated at twenty shillings, while the breaking of a thigh was fixed at only twelve.” (65. Lubbock, ‘Origin of Civilisation,’ 1870, p. 321.) In the East men swear solemnly by their beards. We have seen that Chinsurdi, the chief of the Makalolo in Africa, thought that beards were a great ornament. In the Pacific the Fijian’s beard is “profuse and bushy, and is his greatest pride”; whilst the inhabitants of the adjacent archipelagoes of Tonga and Samoa are “beardless, and abhor a rough chin.” In one island alone of the Ellice group “the men are heavily bearded, and not a little proud thereof.” (66. Dr. Barnard Davis quotes Mr. Prichard and others for these facts in regard to the Polynesians, in ‘Anthropolog. Review,’ April 1870, pp. 185, 191.)

We thus see how widely the different races of man differ in their taste for the beautiful. In every nation sufficiently advanced to have made effigies of their gods or of their deified rulers, the sculptors no doubt have endeavoured to express their highest ideal of beauty and grandeur. (67. Ch. Comte has remarks to this effect in his ‘Traité de Législation,’ 3rd ed. 1837, p. 136.) Under this point of view it is well to compare in our mind the Jupiter or Apollo of the Greeks with the Egyptian or Assyrian statues; and these with the hideous bas-reliefs on the ruined buildings of Central America.

I have met with very few statements opposed to this conclusion. Mr. Winwood Reade, however, who has had ample opportunities for observation, not only with the negroes of the West Coast of Africa, but with those of the interior who have never associated with Europeans, is convinced that their ideas of beauty are ON THE WHOLE the same as ours; and Dr. Rohlfs writes to me to the same effect with respect to Bornu and the countries inhabited by the Pullo tribes. Mr. Reade found that he agreed with the negroes in their estimation of the beauty of the native girls; and that their appreciation of the beauty of European women corresponded with ours. They admire long hair, and use artificial means to make it appear abundant; they admire also a beard, though themselves very scantily provided. Mr. Reade feels doubtful what kind of nose is most appreciated; a girl has been heard to say, “I do not want to marry him, he has got no nose”; and this shews that a very flat nose is not admired. We should, however, bear in mind that the depressed, broad noses and projecting jaws of the negroes of the West Coast are exceptional types with the inhabitants of Africa. Notwithstanding the foregoing statements, Mr. Reade admits that negroes “do not like the colour of our skin; they look on blue eyes with aversion, and they think our noses too long and our lips too thin.” He does not think it probable that negroes would ever prefer the most beautiful European woman, on the mere grounds of physical admiration, to a good-looking negress. (68. The ‘African Sketch Book,’ vol. ii. 1873, pp. 253, 394, 521. The Fuegians, as I have been informed by a missionary who long resided with them, consider European women as extremely beautiful; but from what we have seen of the judgment of the other aborigines of America, I cannot but think that this must be a mistake, unless indeed the statement refers to the few Fuegians who have lived for some time with Europeans, and who must consider us as superior beings. I should add that a most experienced observer, Capt. Burton, believes that a woman whom we consider beautiful is admired throughout the world. ‘Anthropological Review,’ March, 1864, p. 245.)

The general truth of the principle, long ago insisted on by Humboldt (69. ‘Personal Narrative,’ Eng. translat. vol. iv. p. 518, and elsewhere. Mantegazza, in his ‘Viaggi e Studi,’ strongly insists on this same principle.), that man admires and often tries to exaggerate whatever characters nature may have given him, is shewn in many ways. The practice of beardless races extirpating every trace of a beard, and often all the hairs on the body affords one illustration. The skull has been greatly modified during ancient and modern times by many nations; and there can be little doubt that this has been practised, especially in N. and S. America, in order to exaggerate some natural and admired peculiarity. Many American Indians are known to admire a head so extremely flattened as to appear to us idiotic. The natives on the north-western coast compress the head into a pointed cone; and it is their constant practice to gather the hair into a knot on the top of the head, for the sake, as Dr. Wilson remarks, “of increasing the apparent elevation of the favourite conoid form.” The inhabitants of Arakhan admire a broad, smooth forehead, and in order to produce it, they fasten a plate of lead on the heads of the new-born children. On the other hand, “a broad, well-rounded occiput is considered a great beauty” by the natives of the Fiji Islands. (70. On the skulls of the American tribes, see Nott and Gliddon, ‘Types of Mankind,’ 1854, p. 440; Prichard, ‘Physical History of Mankind,’ vol. i. 3rd ed. p. 321; on the natives of Arakhan, ibid. vol. iv. p. 537. Wilson, ‘Physical Ethnology,’ Smithsonian Institution, 1863, p. 288; on the Fijians, p. 290. Sir J. Lubbock (‘Prehistoric Times,’ 2nd ed. 1869, p. 506) gives an excellent resume on this subject.)

As with the skull, so with the nose; the ancient Huns during the age of Attila were accustomed to flatten the noses of their infants with bandages, “for the sake of exaggerating a natural conformation.” With the Tahitians, to be called LONG-NOSE is considered as an insult, and they compress the noses and foreheads of their children for the sake of beauty. The same holds with the Malays of Sumatra, the Hottentots, certain Negroes, and the natives of Brazil. (71. On the Huns, Godron, ‘De l’Espèce,’ tom. ii. 1859, p. 300. On the Tahitians, Waitz, ‘Anthropology,’ Eng. translat. vol. i. p. 305. Marsden, quoted by Prichard, ‘Phys. Hist. of Mankind,’ 3rd edit. vol. v. p. 67. Lawrence, ‘Lectures on Physiology,’ p. 337.) The Chinese have by nature unusually small feet (72. This fact was ascertained in the ‘Reise der Novara: Anthropolog. Theil.’ Dr. Weisbach, 1867, s. 265.); and it is well known that the women of the upper classes distort their feet to make them still smaller. Lastly, Humboldt thinks that the American Indians prefer colouring their bodies with red paint in order to exaggerate their natural tint; and until recently European women added to their naturally bright colours by rouge and white cosmetics; but it may be doubted whether barbarous nations have generally had any such intention in painting themselves.

In the fashions of our own dress we see exactly the same principle and the same desire to carry every point to an extreme; we exhibit, also, the same spirit of emulation. But the fashions of savages are far more permanent than ours; and whenever their bodies are artificially modified, this is necessarily the case. The Arab women of the Upper Nile occupy about three days in dressing their hair; they never imitate other tribes, “but simply vie with each other in the superlativeness of their own style.” Dr. Wilson, in speaking of the compressed skulls of various American races, adds, “such usages are among the least eradicable, and long survive the shock of revolutions that change dynasties and efface more important national peculiarities.” (73. ‘Smithsonian Institution,’ 1863, p. 289. On the fashions of Arab women, Sir S. Baker, ‘The Nile Tributaries,’ 1867, p. 121.) The same principle comes into play in the art of breeding; and we can thus understand, as I have elsewhere explained (74. The ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 214; vol. ii. p. 240.), the wonderful development of the many races of animals and plants, which have been kept merely for ornament. Fanciers always wish each character to be somewhat increased; they do not admire a medium standard; they certainly do not desire any great and abrupt change in the character of their breeds; they admire solely what they are accustomed to, but they ardently desire to see each characteristic feature a little more developed.

The senses of man and of the lower animals seem to be so constituted that brilliant colours and certain forms, as well as harmonious and rhythmical sounds, give pleasure and are called beautiful; but why this should be so we know not. It is certainly not true that there is in the mind of man any universal standard of beauty with respect to the human body. It is, however, possible that certain tastes may in the course of time become inherited, though there is no evidence in favour of this belief: and if so, each race would possess its own innate ideal standard of beauty. It has been argued (75. Schaaffhausen, ‘Archiv. für Anthropologie,’ 1866, s. 164.) that ugliness consists in an approach to the structure of the lower animals, and no doubt this is partly true with the more civilised nations, in which intellect is highly appreciated; but this explanation will hardly apply to all forms of ugliness. The men of each race prefer what they are accustomed to; they cannot endure any great change; but they like variety, and admire each characteristic carried to a moderate extreme. (76. Mr. Bain has collected (‘Mental and Moral Science,’ 1868, pp. 304-314) about a dozen more or less different theories of the idea of beauty; but none is quite the same as that here given.) Men accustomed to a nearly oval face, to straight and regular features, and to bright colours, admire, as we Europeans know, these points when strongly developed. On the other hand, men accustomed to a broad face, with high cheek-bones, a depressed nose, and a black skin, admire these peculiarities when strongly marked. No doubt characters of all kinds may be too much developed for beauty. Hence a perfect beauty, which implies many characters modified in a particular manner, will be in every race a prodigy. As the great anatomist Bichat long ago said, if every one were cast in the same mould, there would be no such thing as beauty. If all our women were to become as beautiful as the Venus de’ Medici, we should for a time be charmed; but we should soon wish for variety; and as soon as we had obtained variety, we should wish to see certain characters a little exaggerated beyond the then existing common standard.

Chapter XX

CHAPTER XX. SECONDARY SEXUAL CHARACTERS OF MAN—continued.

On the effects of the continued selection of women according to a different standard of beauty in each race—On the causes which interfere with sexual selection in civilised and savage nations—Conditions favourable to sexual selection during primeval times—On the manner of action of sexual selection with mankind—On the women in savage tribes having some power to choose their husbands—Absence of hair on the body, and development of the beard—Colour of the skin—Summary.

We have seen in the last chapter that with all barbarous races ornaments, dress, and external appearance are highly valued; and that the men judge of the beauty of their women by widely different standards. We must next inquire whether this preference and the consequent selection during many generations of those women, which appear to the men of each race the most attractive, has altered the character either of the females alone, or of both sexes. With mammals the general rule appears to be that characters of all kinds are inherited equally by the males and females; we might therefore expect that with mankind any characters gained by the females or by the males through sexual selection would commonly be transferred to the offspring of both sexes. If any change has thus been effected, it is almost certain that the different races would be differently modified, as each has its own standard of beauty.

With mankind, especially with savages, many causes interfere with the action of sexual selection as far as the bodily frame is concerned. Civilised men are largely attracted by the mental charms of women, by their wealth, and especially by their social position; for men rarely marry into a much lower rank. The men who succeed in obtaining the more beautiful women will not have a better chance of leaving a long line of descendants than other men with plainer wives, save the few who bequeath their fortunes according to primogeniture. With respect to the opposite form of selection, namely, of the more attractive men by the women, although in civilised nations women have free or almost free choice, which is not the case with barbarous races, yet their choice is largely influenced by the social position and wealth of the men; and the success of the latter in life depends much on their intellectual powers and energy, or on the fruits of these same powers in their forefathers. No excuse is needed for treating this subject in some detail; for, as the German philosopher Schopenhauer remarks, “the final aim of all love intrigues, be they comic or tragic, is really of more importance than all other ends in human life. What it all turns upon is nothing less than the composition of the next generation...It is not the weal or woe of any one individual, but that of the human race to come, which is here at stake.” (1. ‘Schopenhauer and Darwinism,’ in ‘Journal of Anthropology,’ Jan. 1871, p. 323.

There is, however, reason to believe that in certain civilised and semi-civilised nations sexual selection has effected something in modifying the bodily frame of some of the members. Many persons are convinced, as it appears to me with justice, that our aristocracy, including under this term all wealthy families in which primogeniture has long prevailed, from having chosen during many generations from all classes the more beautiful women as their wives, have become handsomer, according to the European standard, than the middle classes; yet the middle classes are placed under equally favourable conditions of life for the perfect development of the body. Cook remarks that the superiority in personal appearance “which is observable in the erees or nobles in all the other islands (of the Pacific) is found in the Sandwich Islands”; but this may be chiefly due to their better food and manner of life.

The old traveller Chardin, in describing the Persians, says their “blood is now highly refined by frequent intermixtures with the Georgians and Circassians, two nations which surpass all the world in personal beauty. There is hardly a man of rank in Persia who is not born of a Georgian or Circassian mother.” He adds that they inherit their beauty, “not from their ancestors, for without the above mixture, the men of rank in Persia, who are descendants of the Tartars, would be extremely ugly.” (2. These quotations are taken from Lawrence (‘Lectures on Physiology,’ etc., 1822, p. 393), who attributes the beauty of the upper classes in England to the men having long selected the more beautiful women.) Here is a more curious case; the priestesses who attended the temple of Venus Erycina at San-Giuliano in Sicily, were selected for their beauty out of the whole of Greece; they were not vestal virgins, and Quatrefages (3. ‘Anthropologie,’ ‘Revue des Cours Scientifiques,’ Oct. 1868, p. 721.), who states the foregoing fact, says that the women of San-Giuliano are now famous as the most beautiful in the island, and are sought by artists as models. But it is obvious that the evidence in all the above cases is doubtful.

The following case, though relating to savages, is well worth giving for its curiosity. Mr. Winwood Reade informs me that the Jollofs, a tribe of negroes on the west coast of Africa, “are remarkable for their uniformly fine appearance.” A friend of his asked one of these men, “How is it that every one whom I meet is so fine looking, not only your men but your women?” The Jollof answered, “It is very easily explained: it has always been our custom to pick out our worst-looking slaves and to sell them.” It need hardly be added that with all savages, female slaves serve as concubines. That this negro should have attributed, whether rightly or wrongly, the fine appearance of his tribe to the long-continued elimination of the ugly women is not so surprising as it may at first appear; for I have elsewhere shewn (4. ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 207.) that negroes fully appreciate the importance of selection in the breeding of their domestic animals, and I could give from Mr. Reade additional evidence on this head.

THE CAUSES WHICH PREVENT OR CHECK THE ACTION OF SEXUAL SELECTION WITH SAVAGES.

The chief causes are, first, so-called communal marriages or promiscuous intercourse; secondly, the consequences of female infanticide; thirdly, early betrothals; and lastly, the low estimation in which women are held, as mere slaves. These four points must be considered in some detail.

It is obvious that as long as the pairing of man, or of any other animal, is left to mere chance, with no choice exerted by either sex, there can be no sexual selection; and no effect will be produced on the offspring by certain individuals having had an advantage over others in their courtship. Now it is asserted that there exist at the present day tribes which practise what Sir J. Lubbock by courtesy calls communal marriages; that is, all the men and women in the tribe are husbands and wives to one another. The licentiousness of many savages is no doubt astonishing, but it seems to me that more evidence is requisite, before we fully admit that their intercourse is in any case promiscuous. Nevertheless all those who have most closely studied the subject (5. Sir J. Lubbock, ‘The Origin of Civilisation,’ 1870, chap. iii. especially pp. 60-67. Mr. M’Lennan, in his extremely valuable work on ‘Primitive Marriage,’ 1865, p. 163, speaks of the union of the sexes “in the earliest times as loose, transitory, and in some degree promiscuous.” Mr. M’Lennan and Sir J. Lubbock have collected much evidence on the extreme licentiousness of savages at the present time. Mr. L.H. Morgan, in his interesting memoir of the classificatory system of relationship. (‘Proceedings of the American Academy of Sciences,’ vol. vii. Feb. 1868, p. 475), concludes that polygamy and all forms of marriage during primeval times were essentially unknown. It appears also, from Sir J. Lubbock’s work, that Bachofen likewise believes that communal intercourse originally prevailed.), and whose judgment is worth much more than mine, believe that communal marriage (this expression being variously guarded) was the original and universal form throughout the world, including therein the intermarriage of brothers and sisters. The late Sir A. Smith, who had travelled widely in S. Africa, and knew much about the habits of savages there and elsewhere, expressed to me the strongest opinion that no race exists in which woman is considered as the property of the community. I believe that his judgment was largely determined by what is implied by the term marriage. Throughout the following discussion I use the term in the same sense as when naturalists speak of animals as monogamous, meaning thereby that the male is accepted by or chooses a single female, and lives with her either during the breeding-season or for the whole year, keeping possession of her by the law of might; or, as when they speak of a polygamous species, meaning that the male lives with several females. This kind of marriage is all that concerns us here, as it suffices for the work of sexual selection. But I know that some of the writers above referred to imply by the term marriage a recognised right protected by the tribe.

The indirect evidence in favour of the belief of the former prevalence of communal marriages is strong, and rests chiefly on the terms of relationship which are employed between the members of the same tribe, implying a connection with the tribe, and not with either parent. But the subject is too large and complex for even an abstract to be here given, and I will confine myself to a few remarks. It is evident in the case of such marriages, or where the marriage tie is very loose, that the relationship of the child to its father cannot be known. But it seems almost incredible that the relationship of the child to its mother should ever be completely ignored, especially as the women in most savage tribes nurse their infants for a long time. Accordingly, in many cases the lines of descent are traced through the mother alone, to the exclusion of the father. But in other cases the terms employed express a connection with the tribe alone, to the exclusion even of the mother. It seems possible that the connection between the related members of the same barbarous tribe, exposed to all sorts of danger, might be so much more important, owing to the need of mutual protection and aid, than that between the mother and her child, as to lead to the sole use of terms expressive of the former relationships; but Mr. Morgan is convinced that this view is by no means sufficient.

The terms of relationship used in different parts of the world may be divided, according to the author just quoted, into two great classes, the classificatory and descriptive, the latter being employed by us. It is the classificatory system which so strongly leads to the belief that communal and other extremely loose forms of marriage were originally universal. But as far as I can see, there is no necessity on this ground for believing in absolutely promiscuous intercourse; and I am glad to find that this is Sir J. Lubbock’s view. Men and women, like many of the lower animals, might formerly have entered into strict though temporary unions for each birth, and in this case nearly as much confusion would have arisen in the terms of relationship as in the case of promiscuous intercourse. As far as sexual selection is concerned, all that is required is that choice should be exerted before the parents unite, and it signifies little whether the unions last for life or only for a season.

Besides the evidence derived from the terms of relationship, other lines of reasoning indicate the former wide prevalence of communal marriage. Sir J. Lubbock accounts for the strange and widely-extended habit of exogamy—that is, the men of one tribe taking wives from a distinct tribe,—by communism having been the original form of intercourse; so that a man never obtained a wife for himself unless he captured her from a neighbouring and hostile tribe, and then she would naturally have become his sole and valuable property. Thus the practice of capturing wives might have arisen; and from the honour so gained it might ultimately have become the universal habit. According to Sir J. Lubbock (6. ‘Address to British Association On the Social and Religious Condition of the Lower Races of Man,’ 1870, p. 20.), we can also thus understand “the necessity of expiation for marriage as an infringement of tribal rites, since according to old ideas, a man had no right to appropriate to himself that which belonged to the whole tribe.” Sir J. Lubbock further gives a curious body of facts shewing that in old times high honour was bestowed on women who were utterly licentious; and this, as he explains, is intelligible, if we admit that promiscuous intercourse was the aboriginal, and therefore long revered custom of the tribe. (7. ‘Origin of Civilisation,’ 1870, p. 86. In the several works above quoted, there will be found copious evidence on relationship through the females alone, or with the tribe alone.)

Although the manner of development of the marriage tie is an obscure subject, as we may infer from the divergent opinions on several points between the three authors who have studied it most closely, namely, Mr. Morgan, Mr. M’Lennan, and Sir J. Lubbock, yet from the foregoing and several other lines of evidence it seems probable (8. Mr. C. Staniland Wake argues strongly (‘Anthropologia,’ March, 1874, p. 197) against the views held by these three writers on the former prevalence of almost promiscuous intercourse; and he thinks that the classificatory system of relationship can be otherwise explained.) that the habit of marriage, in any strict sense of the word, has been gradually developed; and that almost promiscuous or very loose intercourse was once extremely common throughout the world. Nevertheless, from the strength of the feeling of jealousy all through the animal kingdom, as well as from the analogy of the lower animals, more particularly of those which come nearest to man, I cannot believe that absolutely promiscuous intercourse prevailed in times past, shortly before man attained to his present rank in the zoological scale. Man, as I have attempted to shew, is certainly descended from some ape-like creature. With the existing Quadrumana, as far as their habits are known, the males of some species are monogamous, but live during only a part of the year with the females: of this the orang seems to afford an instance. Several kinds, for example some of the Indian and American monkeys, are strictly monogamous, and associate all the year round with their wives. Others are polygamous, for example the gorilla and several American species, and each family lives separate. Even when this occurs, the families inhabiting the same district are probably somewhat social; the chimpanzee, for instance, is occasionally met with in large bands. Again, other species are polygamous, but several males, each with his own females, live associated in a body, as with several species of baboons. (9. Brehm (‘Thierleben,’ B. i. p. 77) says Cynocephalus hamadryas lives in great troops containing twice as many adult females as adult males. See Rengger on American polygamous species, and Owen (‘Anatomy of Vertebrates,’ vol. iii. p. 746) on American monogamous species. Other references might be added.) We may indeed conclude from what we know of the jealousy of all male quadrupeds, armed, as many of them are, with special weapons for battling with their rivals, that promiscuous intercourse in a state of nature is extremely improbable. The pairing may not last for life, but only for each birth; yet if the males which are the strongest and best able to defend or otherwise assist their females and young, were to select the more attractive females, this would suffice for sexual selection.

Therefore, looking far enough back in the stream of time, and judging from the social habits of man as he now exists, the most probable view is that he aboriginally lived in small communities, each with a single wife, or if powerful with several, whom he jealously guarded against all other men. Or he may not have been a social animal, and yet have lived with several wives, like the gorilla; for all the natives “agree that but one adult male is seen in a band; when the young male grows up, a contest takes place for mastery, and the strongest, by killing and driving out the others, establishes himself as the head of the community.” (10. Dr. Savage, in ‘Boston Journal of Natural History,’ vol. v. 1845-47, p. 423.) The younger males, being thus expelled and wandering about, would, when at last successful in finding a partner, prevent too close interbreeding within the limits of the same family.

Although savages are now extremely licentious, and although communal marriages may formerly have largely prevailed, yet many tribes practise some form of marriage, but of a far more lax nature than that of civilised nations. Polygamy, as just stated, is almost universally followed by the leading men in every tribe. Nevertheless there are tribes, standing almost at the bottom of the scale, which are strictly monogamous. This is the case with the Veddahs of Ceylon: they have a saying, according to Sir J. Lubbock (11. ‘Prehistoric Times,’ 1869, p. 424.), “that death alone can separate husband and wife.” An intelligent Kandyan chief, of course a polygamist, “was perfectly scandalised at the utter barbarism of living with only one wife, and never parting until separated by death.” It was, he said, “just like the Wanderoo monkeys.” Whether savages who now enter into some form of marriage, either polygamous or monogamous, have retained this habit from primeval times, or whether they have returned to some form of marriage, after passing through a stage of promiscuous intercourse, I will not pretend to conjecture.

INFANTICIDE.

This practice is now very common throughout the world, and there is reason to believe that it prevailed much more extensively during former times. (12. Mr. M’Lennan, ‘Primitive Marriage,’ 1865. See especially on exogamy and infanticide, pp. 130, 138, 165.) Barbarians find it difficult to support themselves and their children, and it is a simple plan to kill their infants. In South America some tribes, according to Azara, formerly destroyed so many infants of both sexes that they were on the point of extinction. In the Polynesian Islands women have been known to kill from four or five, to even ten of their children; and Ellis could not find a single woman who had not killed at least one. In a village on the eastern frontier of India Colonel MacCulloch found not a single female child. Wherever infanticide (13. Dr. Gerland (‘Ueber das Aussterben der Naturvölker,’ 1868) has collected much information on infanticide, see especially ss. 27, 51, 54. Azara (‘Voyages,’ etc., tom. ii. pp. 94, 116) enters in detail on the motives. See also M’Lennan (ibid. p. 139) for cases in India. In the former reprints of the 2nd edition of this book an incorrect quotation from Sir G. Grey was unfortunately given in the above passage and has now been removed from the text.) prevails the struggle for existence will be in so far less severe, and all the members of the tribe will have an almost equally good chance of rearing their few surviving children. In most cases a larger number of female than of male infants are destroyed, for it is obvious that the latter are of more value to the tribe, as they will, when grown up, aid in defending it, and can support themselves. But the trouble experienced by the women in rearing children, their consequent loss of beauty, the higher estimation set on them when few, and their happier fate, are assigned by the women themselves, and by various observers, as additional motives for infanticide.

When, owing to female infanticide, the women of a tribe were few, the habit of capturing wives from neighbouring tribes would naturally arise. Sir J. Lubbock, however, as we have seen, attributes the practice in chief part to the former existence of communal marriage, and to the men having consequently captured women from other tribes to hold as their sole property. Additional causes might be assigned, such as the communities being very small, in which case, marriageable women would often be deficient. That the habit was most extensively practised during former times, even by the ancestors of civilised nations, is clearly shewn by the preservation of many curious customs and ceremonies, of which Mr. M’Lennan has given an interesting account. In our own marriages the “best man” seems originally to have been the chief abettor of the bridegroom in the act of capture. Now as long as men habitually procured their wives through violence and craft, they would have been glad to seize on any woman, and would not have selected the more attractive ones. But as soon as the practice of procuring wives from a distinct tribe was effected through barter, as now occurs in many places, the more attractive women would generally have been purchased. The incessant crossing, however, between tribe and tribe, which necessarily follows from any form of this habit, would tend to keep all the people inhabiting the same country nearly uniform in character; and this would interfere with the power of sexual selection in differentiating the tribes.

The scarcity of women, consequent on female infanticide, leads, also, to another practice, that of polyandry, still common in several parts of the world, and which formerly, as Mr. M’Lennan believes, prevailed almost universally: but this latter conclusion is doubted by Mr. Morgan and Sir J. Lubbock. (14. ‘Primitive Marriage,’ p. 208; Sir J. Lubbock, ‘Origin of Civilisation,’ p. 100. See also Mr. Morgan, loc. cit., on the former prevalence of polyandry.) Whenever two or more men are compelled to marry one woman, it is certain that all the women of the tribe will get married, and there will be no selection by the men of the more attractive women. But under these circumstances the women no doubt will have the power of choice, and will prefer the more attractive men. Azara, for instance, describes how carefully a Guana woman bargains for all sorts of privileges, before accepting some one or more husbands; and the men in consequence take unusual care of their personal appearance. So amongst the Todas of India, who practise polyandry, the girls can accept or refuse any man. (15. Azara, ‘Voyages,’ etc., tom. ii. pp. 92-95; Colonel Marshall, ‘Amongst the Todas,’ p. 212.) A very ugly man in these cases would perhaps altogether fail in getting a wife, or get one later in life; but the handsomer men, although more successful in obtaining wives, would not, as far as we can see, leave more offspring to inherit their beauty than the less handsome husbands of the same women.

EARLY BETROTHALS AND SLAVERY OF WOMEN.

With many savages it is the custom to betroth the females whilst mere infants; and this would effectually prevent preference being exerted on either side according to personal appearance. But it would not prevent the more attractive women from being afterwards stolen or taken by force from their husbands by the more powerful men; and this often happens in Australia, America, and elsewhere. The same consequences with reference to sexual selection would to a certain extent follow, when women are valued almost solely as slaves or beasts of burden, as is the case with many savages. The men, however, at all times would prefer the handsomest slaves according to their standard of beauty.

We thus see that several customs prevail with savages which must greatly interfere with, or completely stop, the action of sexual selection. On the other hand, the conditions of life to which savages are exposed, and some of their habits, are favourable to natural selection; and this comes into play at the same time with sexual selection. Savages are known to suffer severely from recurrent famines; they do not increase their food by artificial means; they rarely refrain from marriage (16. Burchell says (‘Travels in S. Africa,’ vol. ii. 1824, p. 58), that among the wild nations of Southern Africa, neither men nor women ever pass their lives in a state of celibacy. Azara (‘Voyages dans l’Amérique Merid.’ tom. ii. 1809, p. 21) makes precisely the same remark in regard to the wild Indians of South America.), and generally marry whilst young. Consequently they must be subjected to occasional hard struggles for existence, and the favoured individuals will alone survive.

At a very early period, before man attained to his present rank in the scale, many of his conditions would be different from what now obtains amongst savages. Judging from the analogy of the lower animals, he would then either live with a single female, or be a polygamist. The most powerful and able males would succeed best in obtaining attractive females. They would also succeed best in the general struggle for life, and in defending their females, as well as their offspring, from enemies of all kinds. At this early period the ancestors of man would not be sufficiently advanced in intellect to look forward to distant contingencies; they would not foresee that the rearing of all their children, especially their female children, would make the struggle for life severer for the tribe. They would be governed more by their instincts and less by their reason than are savages at the present day. They would not at that period have partially lost one of the strongest of all instincts, common to all the lower animals, namely the love of their young offspring; and consequently they would not have practised female infanticide. Women would not have been thus rendered scarce, and polyandry would not have been practised; for hardly any other cause, except the scarcity of women seems sufficient to break down the natural and widely prevalent feeling of jealousy, and the desire of each male to possess a female for himself. Polyandry would be a natural stepping-stone to communal marriages or almost promiscuous intercourse; though the best authorities believe that this latter habit preceded polyandry. During primordial times there would be no early betrothals, for this implies foresight. Nor would women be valued merely as useful slaves or beasts of burthen. Both sexes, if the females as well as the males were permitted to exert any choice, would choose their partners not for mental charms, or property, or social position, but almost solely from external appearance. All the adults would marry or pair, and all the offspring, as far as that was possible, would be reared; so that the struggle for existence would be periodically excessively severe. Thus during these times all the conditions for sexual selection would have been more favourable than at a later period, when man had advanced in his intellectual powers but had retrograded in his instincts. Therefore, whatever influence sexual selection may have had in producing the differences between the races of man, and between man and the higher Quadrumana, this influence would have been more powerful at a remote period than at the present day, though probably not yet wholly lost.

THE MANNER OF ACTION OF SEXUAL SELECTION WITH MANKIND.

With primeval man under the favourable conditions just stated, and with those savages who at the present time enter into any marriage tie, sexual selection has probably acted in the following manner, subject to greater or less interference from female infanticide, early betrothals, etc. The strongest and most vigorous men—those who could best defend and hunt for their families, who were provided with the best weapons and possessed the most property, such as a large number of dogs or other animals,—would succeed in rearing a greater average number of offspring than the weaker and poorer members of the same tribes. There can, also, be no doubt that such men would generally be able to select the more attractive women. At present the chiefs of nearly every tribe throughout the world succeed in obtaining more than one wife. I hear from Mr. Mantell that, until recently, almost every girl in New Zealand who was pretty, or promised to be pretty, was tapu to some chief. With the Kafirs, as Mr. C. Hamilton states (17. ‘Anthropological Review,’ Jan. 1870, p. xvi.), “the chiefs generally have the pick of the women for many miles round, and are most persevering in establishing or confirming their privilege.” We have seen that each race has its own style of beauty, and we know that it is natural to man to admire each characteristic point in his domestic animals, dress, ornaments, and personal appearance, when carried a little beyond the average. If then the several foregoing propositions be admitted, and I cannot see that they are doubtful, it would be an inexplicable circumstance if the selection of the more attractive women by the more powerful men of each tribe, who would rear on an average a greater number of children, did not after the lapse of many generations somewhat modify the character of the tribe.

When a foreign breed of our domestic animals is introduced into a new country, or when a native breed is long and carefully attended to, either for use or ornament, it is found after several generations to have undergone a greater or less amount of change whenever the means of comparison exist. This follows from unconscious selection during a long series of generations—that is, the preservation of the most approved individuals—without any wish or expectation of such a result on the part of the breeder. So again, if during many years two careful breeders rear animals of the same family, and do not compare them together or with a common standard, the animals are found to have become, to the surprise of their owners, slightly different. (18. The ‘Variation of Animals and Plants under Domestication,’ vol. ii. pp. 210-217.) Each breeder has impressed, as von Nathusius well expresses it, the character of his own mind—his own taste and judgment—on his animals. What reason, then, can be assigned why similar results should not follow from the long-continued selection of the most admired women by those men of each tribe who were able to rear the greatest number of children? This would be unconscious selection, for an effect would be produced, independently of any wish or expectation on the part of the men who preferred certain women to others.

Let us suppose the members of a tribe, practising some form of marriage, to spread over an unoccupied continent, they would soon split up into distinct hordes, separated from each other by various barriers, and still more effectually by the incessant wars between all barbarous nations. The hordes would thus be exposed to slightly different conditions and habits of life, and would sooner or later come to differ in some small degree. As soon as this occurred, each isolated tribe would form for itself a slightly different standard of beauty (19. An ingenious writer argues, from a comparison of the pictures of Raphael, Rubens, and modern French artists, that the idea of beauty is not absolutely the same even throughout Europe: see the ‘Lives of Haydn and Mozart,’ by Bombet (otherwise M. Beyle), English translation, p. 278.); and then unconscious selection would come into action through the more powerful and leading men preferring certain women to others. Thus the differences between the tribes, at first very slight, would gradually and inevitably be more or less increased.

With animals in a state of nature, many characters proper to the males, such as size, strength, special weapons, courage and pugnacity, have been acquired through the law of battle. The semi-human progenitors of man, like their allies the Quadrumana, will almost certainly have been thus modified; and, as savages still fight for the possession of their women, a similar process of selection has probably gone on in a greater or less degree to the present day. Other characters proper to the males of the lower animals, such as bright colours and various ornaments, have been acquired by the more attractive males having been preferred by the females. There are, however, exceptional cases in which the males are the selectors, instead of having been the selected. We recognise such cases by the females being more highly ornamented than the males,—their ornamental characters having been transmitted exclusively or chiefly to their female offspring. One such case has been described in the order to which man belongs, that of the Rhesus monkey.

Man is more powerful in body and mind than woman, and in the savage state he keeps her in a far more abject state of bondage than does the male of any other animal; therefore it is not surprising that he should have gained the power of selection. Women are everywhere conscious of the value of their own beauty; and when they have the means, they take more delight in decorating themselves with all sorts of ornaments than do men. They borrow the plumes of male birds, with which nature has decked this sex, in order to charm the females. As women have long been selected for beauty, it is not surprising that some of their successive variations should have been transmitted exclusively to the same sex; consequently that they should have transmitted beauty in a somewhat higher degree to their female than to their male offspring, and thus have become more beautiful, according to general opinion, than men. Women, however, certainly transmit most of their characters, including some beauty, to their offspring of both sexes; so that the continued preference by the men of each race for the more attractive women, according to their standard of taste, will have tended to modify in the same manner all the individuals of both sexes belonging to the race.

With respect to the other form of sexual selection (which with the lower animals is much the more common), namely, when the females are the selectors, and accept only those males which excite or charm them most, we have reason to believe that it formerly acted on our progenitors. Man in all probability owes his beard, and perhaps some other characters, to inheritance from an ancient progenitor who thus gained his ornaments. But this form of selection may have occasionally acted during later times; for in utterly barbarous tribes the women have more power in choosing, rejecting, and tempting their lovers, or of afterwards changing their husbands, than might have been expected. As this is a point of some importance, I will give in detail such evidence as I have been able to collect.

Hearne describes how a woman in one of the tribes of Arctic America repeatedly ran away from her husband and joined her lover; and with the Charruas of S. America, according to Azara, divorce is quite optional. Amongst the Abipones, a man on choosing a wife bargains with the parents about the price. But “it frequently happens that the girl rescinds what has been agreed upon between the parents and the bridegroom, obstinately rejecting the very mention of marriage.” She often runs away, hides herself, and thus eludes the bridegroom. Captain Musters who lived with the Patagonians, says that their marriages are always settled by inclination; “if the parents make a match contrary to the daughter’s will, she refuses and is never compelled to comply.” In Tierra del Fuego a young man first obtains the consent of the parents by doing them some service, and then he attempts to carry off the girl; “but if she is unwilling, she hides herself in the woods until her admirer is heartily tired of looking for her, and gives up the pursuit; but this seldom happens.” In the Fiji Islands the man seizes on the woman whom he wishes for his wife by actual or pretended force; but “on reaching the home of her abductor, should she not approve of the match, she runs to some one who can protect her; if, however, she is satisfied, the matter is settled forthwith.” With the Kalmucks there is a regular race between the bride and bridegroom, the former having a fair start; and Clarke “was assured that no instance occurs of a girl being caught, unless she has a partiality to the pursuer.” Amongst the wild tribes of the Malay Archipelago there is also a racing match; and it appears from M. Bourien’s account, as Sir J. Lubbock remarks, that “the race, ‘is not to the swift, nor the battle to the strong,’ but to the young man who has the good fortune to please his intended bride.” A similar custom, with the same result, prevails with the Koraks of North-Eastern Asia.

Turning to Africa: the Kafirs buy their wives, and girls are severely beaten by their fathers if they will not accept a chosen husband; but it is manifest from many facts given by the Rev. Mr. Shooter, that they have considerable power of choice. Thus very ugly, though rich men, have been known to fail in getting wives. The girls, before consenting to be betrothed, compel the men to shew themselves off first in front and then behind, and “exhibit their paces.” They have been known to propose to a man, and they not rarely run away with a favoured lover. So again, Mr. Leslie, who was intimately acquainted with the Kafirs, says, “it is a mistake to imagine that a girl is sold by her father in the same manner, and with the same authority, with which he would dispose of a cow.” Amongst the degraded Bushmen of S. Africa, “when a girl has grown up to womanhood without having been betrothed, which, however, does not often happen, her lover must gain her approbation, as well as that of the parents.” (20. Azara, ‘Voyages,’ etc., tom. ii. p. 23. Dobrizhoffer, ‘An Account of the Abipones,’ vol. ii. 1822, p. 207. Capt. Musters, in ‘Proc. R. Geograph. Soc.’ vol. xv. p. 47. Williams on the Fiji Islanders, as quoted by Lubbock, ‘Origin of Civilisation,’ 1870, p. 79. On the Fuegians, King and Fitzroy, ‘Voyages of the “Adventure” and “Beagle,”’ vol. ii. 1839, p. 182. On the Kalmucks, quoted by M’Lennan, ‘Primitive Marriage,’ 1865, p. 32. On the Malays, Lubbock, ibid. p. 76. The Rev. J. Shooter, ‘On the Kafirs of Natal,’ 1857, pp. 52-60. Mr. D. Leslie, ‘Kafir Character and Customs,’ 1871, p. 4. On the Bush-men, Burchell, ‘Travels in S. Africa,’ ii. 1824, p. 59. On the Koraks by McKennan, as quoted by Mr. Wake, in ‘Anthropologia,’ Oct. 1873, p. 75.) Mr. Winwood Reade made inquiries for me with respect to the negroes of Western Africa, and he informs me that “the women, at least among the more intelligent Pagan tribes, have no difficulty in getting the husbands whom they may desire, although it is considered unwomanly to ask a man to marry them. They are quite capable of falling in love, and of forming tender, passionate, and faithful attachments.” Additional cases could be given.

We thus see that with savages the women are not in quite so abject a state in relation to marriage as has often been supposed. They can tempt the men whom they prefer, and can sometimes reject those whom they dislike, either before or after marriage. Preference on the part of the women, steadily acting in any one direction, would ultimately affect the character of the tribe; for the women would generally choose not merely the handsomest men, according to their standard of taste, but those who were at the same time best able to defend and support them. Such well-endowed pairs would commonly rear a larger number of offspring than the less favoured. The same result would obviously follow in a still more marked manner if there was selection on both sides; that is, if the more attractive, and at the same time more powerful men were to prefer, and were preferred by, the more attractive women. And this double form of selection seems actually to have occurred, especially during the earlier periods of our long history.

We will now examine a little more closely some of the characters which distinguish the several races of man from one another and from the lower animals, namely, the greater or less deficiency of hair on the body, and the colour of the skin. We need say nothing about the great diversity in the shape of the features and of the skull between the different races, as we have seen in the last chapter how different is the standard of beauty in these respects. These characters will therefore probably have been acted on through sexual selection; but we have no means of judging whether they have been acted on chiefly from the male or female side. The musical faculties of man have likewise been already discussed.

ABSENCE OF HAIR ON THE BODY, AND ITS DEVELOPMENT ON THE FACE AND HEAD.

From the presence of the woolly hair or lanugo on the human foetus, and of rudimentary hairs scattered over the body during maturity, we may infer that man is descended from some animal which was born hairy and remained so during life. The loss of hair is an inconvenience and probably an injury to man, even in a hot climate, for he is thus exposed to the scorching of the sun, and to sudden chills, especially during wet weather. As Mr. Wallace remarks, the natives in all countries are glad to protect their naked backs and shoulders with some slight covering. No one supposes that the nakedness of the skin is any direct advantage to man; his body therefore cannot have been divested of hair through natural selection. (21. ‘Contributions to the Theory of Natural Selection,’ 1870, p. 346. Mr. Wallace believes (p. 350) “that some intelligent power has guided or determined the development of man”; and he considers the hairless condition of the skin as coming under this head. The Rev. T.R. Stebbing, in commenting on this view (‘Transactions of Devonshire Association for Science,’ 1870) remarks, that had Mr. Wallace “employed his usual ingenuity on the question of man’s hairless skin, he might have seen the possibility of its selection through its superior beauty or the health attaching to superior cleanliness.”) Nor, as shewn in a former chapter, have we any evidence that this can be due to the direct action of climate, or that it is the result of correlated development.

The absence of hair on the body is to a certain extent a secondary sexual character; for in all parts of the world women are less hairy than men. Therefore we may reasonably suspect that this character has been gained through sexual selection. We know that the faces of several species of monkeys, and large surfaces at the posterior end of the body of other species, have been denuded of hair; and this we may safely attribute to sexual selection, for these surfaces are not only vividly coloured, but sometimes, as with the male mandrill and female rhesus, much more vividly in the one sex than in the other, especially during the breeding-season. I am informed by Mr. Bartlett that, as these animals gradually reach maturity, the naked surfaces grow larger compared with the size of their bodies. The hair, however, appears to have been removed, not for the sake of nudity, but that the colour of the skin may be more fully displayed. So again with many birds, it appears as if the head and neck had been divested of feathers through sexual selection, to exhibit the brightly-coloured skin.

As the body in woman is less hairy than in man, and as this character is common to all races, we may conclude that it was our female semi-human ancestors who were first divested of hair, and that this occurred at an extremely remote period before the several races had diverged from a common stock. Whilst our female ancestors were gradually acquiring this new character of nudity, they must have transmitted it almost equally to their offspring of both sexes whilst young; so that its transmission, as with the ornaments of many mammals and birds, has not been limited either by sex or age. There is nothing surprising in a partial loss of hair having been esteemed as an ornament by our ape-like progenitors, for we have seen that innumerable strange characters have been thus esteemed by animals of all kinds, and have consequently been gained through sexual selection. Nor is it surprising that a slightly injurious character should have been thus acquired; for we know that this is the case with the plumes of certain birds, and with the horns of certain stags.

The females of some of the anthropoid apes, as stated in a former chapter, are somewhat less hairy on the under surface than the males; and here we have what might have afforded a commencement for the process of denudation. With respect to the completion of the process through sexual selection, it is well to bear in mind the New Zealand proverb, “There is no woman for a hairy man.” All who have seen photographs of the Siamese hairy family will admit how ludicrously hideous is the opposite extreme of excessive hairiness. And the king of Siam had to bribe a man to marry the first hairy woman in the family; and she transmitted this character to her young offspring of both sexes. (22. The ‘Variation of Animals and Plants under Domestication,’ vol. ii. 1868, p. 237.)

Some races are much more hairy than others, especially the males; but it must not be assumed that the more hairy races, such as the European, have retained their primordial condition more completely than the naked races, such as the Kalmucks or Americans. It is more probable that the hairiness of the former is due to partial reversion; for characters which have been at some former period long inherited are always apt to return. We have seen that idiots are often very hairy, and they are apt to revert in other characters to a lower animal type. It does not appear that a cold climate has been influential in leading to this kind of reversion; excepting perhaps with the negroes, who have been reared during several generations in the United States (23. ‘Investigations into Military and Anthropological Statistics of American Soldiers,’ by B.A. Gould, 1869, p. 568:—Observations were carefully made on the hairiness of 2129 black and coloured soldiers, whilst they were bathing; and by looking to the published table, “it is manifest at a glance that there is but little, if any, difference between the white and the black races in this respect.” It is, however, certain that negroes in their native and much hotter land of Africa, have remarkably smooth bodies. It should be particularly observed, that both pure blacks and mulattoes were included in the above enumeration; and this is an unfortunate circumstance, as in accordance with a principle, the truth of which I have elsewhere proved, crossed races of man would be eminently liable to revert to the primordial hairy character of their early ape-like progenitors.), and possibly with the Ainos, who inhabit the northern islands of the Japan archipelago. But the laws of inheritance are so complex that we can seldom understand their action. If the greater hairiness of certain races be the result of reversion, unchecked by any form of selection, its extreme variability, even within the limits of the same race, ceases to be remarkable. (24. Hardly any view advanced in this work has met with so much disfavour (see for instance, Sprengel, ‘Die Fortschritte des Darwinismus,’ 1874, p. 80) as the above explanation of the loss of hair in mankind through sexual selection; but none of the opposed arguments seem to me of much weight, in comparison with the facts shewing that the nudity of the skin is to a certain extent a secondary sexual character in man and in some of the Quadrumana.)

With respect to the beard in man, if we turn to our best guide, the Quadrumana, we find beards equally developed in both sexes of many species, but in some, either confined to the males, or more developed in them than in the females. From this fact and from the curious arrangement, as well as the bright colours of the hair about the heads of many monkeys, it is highly probable, as before explained, that the males first acquired their beards through sexual selection as an ornament, transmitting them in most cases, equally or nearly so, to their offspring of both sexes. We know from Eschricht (25. ‘Ueber die Richtung der Haare am Menschlichen Körper,’ in Müller’s ‘Archiv. für Anat. und Phys.’ 1837, s. 40.) that with mankind the female as well as the male foetus is furnished with much hair on the face, especially round the mouth; and this indicates that we are descended from progenitors of whom both sexes were bearded. It appears therefore at first sight probable that man has retained his beard from a very early period, whilst woman lost her beard at the same time that her body became almost completely divested of hair. Even the colour of our beards seems to have been inherited from an ape-like progenitor; for when there is any difference in tint between the hair of the head and the beard, the latter is lighter coloured in all monkeys and in man. In those Quadrumana in which the male has a larger beard than that of the female, it is fully developed only at maturity, just as with mankind; and it is possible that only the later stages of development have been retained by man. In opposition to this view of the retention of the beard from an early period is the fact of its great variability in different races, and even within the same race; for this indicates reversion,—long lost characters being very apt to vary on re-appearance.

Nor must we overlook the part which sexual selection may have played in later times; for we know that with savages the men of the beardless races take infinite pains in eradicating every hair from their faces as something odious, whilst the men of the bearded races feel the greatest pride in their beards. The women, no doubt, participate in these feelings, and if so sexual selection can hardly have failed to have effected something in the course of later times. It is also possible that the long-continued habit of eradicating the hair may have produced an inherited effect. Dr. Brown-Sequard has shewn that if certain animals are operated on in a particular manner, their offspring are affected. Further evidence could be given of the inheritance of the effects of mutilations; but a fact lately ascertained by Mr. Salvin (26. On the tail-feathers of Motmots, ‘Proceedings of the Zoological Society,’ 1873, p. 429.) has a more direct bearing on the present question; for he has shewn that the motmots, which are known habitually to bite off the barbs of the two central tail-feathers, have the barbs of these feathers naturally somewhat reduced. (27. Mr. Sproat has suggested (‘Scenes and Studies of Savage Life,’ 1868, p. 25) this same view. Some distinguished ethnologists, amongst others M. Gosse of Geneva, believe that artificial modifications of the skull tend to be inherited.) Nevertheless, with mankind the habit of eradicating the beard and the hairs on the body would probably not have arisen until these had already become by some means reduced.

It is difficult to form any judgment as to how the hair on the head became developed to its present great length in many races. Eschricht (28. ‘Ueber die Richtung,’ ibid. s. 40.) states that in the human foetus the hair on the face during the fifth month is longer than that on the head; and this indicates that our semi-human progenitors were not furnished with long tresses, which must therefore have been a late acquisition. This is likewise indicated by the extraordinary difference in the length of the hair in the different races; in the negro the hair forms a mere curly mat; with us it is of great length, and with the American natives it not rarely reaches to the ground. Some species of Semnopithecus have their heads covered with moderately long hair, and this probably serves as an ornament and was acquired through sexual selection. The same view may perhaps be extended to mankind, for we know that long tresses are now and were formerly much admired, as may be observed in the works of almost every poet; St. Paul says, “if a woman have long hair, it is a glory to her;” and we have seen that in North America a chief was elected solely from the length of his hair.

COLOUR OF THE SKIN.

The best kind of evidence that in man the colour of the skin has been modified through sexual selection is scanty; for in most races the sexes do not differ in this respect, and only slightly, as we have seen, in others. We know, however, from the many facts already given that the colour of the skin is regarded by the men of all races as a highly important element in their beauty; so that it is a character which would be likely to have been modified through selection, as has occurred in innumerable instances with the lower animals. It seems at first sight a monstrous supposition that the jet-blackness of the negro should have been gained through sexual selection; but this view is supported by various analogies, and we know that negroes admire their own colour. With mammals, when the sexes differ in colour, the male is often black or much darker than the female; and it depends merely on the form of inheritance whether this or any other tint is transmitted to both sexes or to one alone. The resemblance to a negro in miniature of Pithecia satanas with his jet black skin, white rolling eyeballs, and hair parted on the top of the head, is almost ludicrous.

The colour of the face differs much more widely in the various kinds of monkeys than it does in the races of man; and we have some reason to believe that the red, blue, orange, almost white and black tints of their skin, even when common to both sexes, as well as the bright colours of their fur, and the ornamental tufts about the head, have all been acquired through sexual selection. As the order of development during growth, generally indicates the order in which the characters of a species have been developed and modified during previous generations; and as the newly-born infants of the various races of man do not differ nearly as much in colour as do the adults, although their bodies are as completely destitute of hair, we have some slight evidence that the tints of the different races were acquired at a period subsequent to the removal of the hair, which must have occurred at a very early period in the history of man.

A SUMMARY.

We may conclude that the greater size, strength, courage, pugnacity, and energy of man, in comparison with woman, were acquired during primeval times, and have subsequently been augmented, chiefly through the contests of rival males for the possession of the females. The greater intellectual vigour and power of invention in man is probably due to natural selection, combined with the inherited effects of habit, for the most able men will have succeeded best in defending and providing for themselves and for their wives and offspring. As far as the extreme intricacy of the subject permits us to judge, it appears that our male ape-like progenitors acquired their beards as an ornament to charm or excite the opposite sex, and transmitted them only to their male offspring. The females apparently first had their bodies denuded of hair, also as a sexual ornament; but they transmitted this character almost equally to both sexes. It is not improbable that the females were modified in other respects for the same purpose and by the same means; so that women have acquired sweeter voices and become more beautiful than men.

It deserves attention that with mankind the conditions were in many respects much more favourable for sexual selection, during a very early period, when man had only just attained to the rank of manhood, than during later times. For he would then, as we may safely conclude, have been guided more by his instinctive passions, and less by foresight or reason. He would have jealously guarded his wife or wives. He would not have practised infanticide; nor valued his wives merely as useful slaves; nor have been betrothed to them during infancy. Hence we may infer that the races of men were differentiated, as far as sexual selection is concerned, in chief part at a very remote epoch; and this conclusion throws light on the remarkable fact that at the most ancient period, of which we have not as yet any record, the races of man had already come to differ nearly or quite as much as they do at the present day.

The views here advanced, on the part which sexual selection has played in the history of man, want scientific precision. He who does not admit this agency in the case of the lower animals, will disregard all that I have written in the later chapters on man. We cannot positively say that this character, but not that, has been thus modified; it has, however, been shewn that the races of man differ from each other and from their nearest allies, in certain characters which are of no service to them in their daily habits of life, and which it is extremely probable would have been modified through sexual selection. We have seen that with the lowest savages the people of each tribe admire their own characteristic qualities,—the shape of the head and face, the squareness of the cheek-bones, the prominence or depression of the nose, the colour of the skin, the length of the hair on the head, the absence of hair on the face and body, or the presence of a great beard, and so forth. Hence these and other such points could hardly fail to be slowly and gradually exaggerated, from the more powerful and able men in each tribe, who would succeed in rearing the largest number of offspring, having selected during many generations for their wives the most strongly characterised and therefore most attractive women. For my own part I conclude that of all the causes which have led to the differences in external appearance between the races of man, and to a certain extent between man and the lower animals, sexual selection has been the most efficient.

Chapter XXI: General a Summary and Conclusion

Main conclusion that man is descended from some lower form—Manner of development—Genealogy of man—Intellectual and moral faculties—Sexual Selection—Concluding remarks.

A brief summary will be sufficient to recall to the reader’s mind the more salient points in this work. Many of the views which have been advanced are highly speculative, and some no doubt will prove erroneous; but I have in every case given the reasons which have led me to one view rather than to another. It seemed worth while to try how far the principle of evolution would throw light on some of the more complex problems in the natural history of man. False facts are highly injurious to the progress of science, for they often endure long; but false views, if supported by some evidence, do little harm, for every one takes a salutary pleasure in proving their falseness: and when this is done, one path towards error is closed and the road to truth is often at the same time opened.

The main conclusion here arrived at, and now held by many naturalists who are well competent to form a sound judgment, is that man is descended from some less highly organised form. The grounds upon which this conclusion rests will never be shaken, for the close similarity between man and the lower animals in embryonic development, as well as in innumerable points of structure and constitution, both of high and of the most trifling importance,—the rudiments which he retains, and the abnormal reversions to which he is occasionally liable,—are facts which cannot be disputed. They have long been known, but until recently they told us nothing with respect to the origin of man. Now when viewed by the light of our knowledge of the whole organic world, their meaning is unmistakable. The great principle of evolution stands up clear and firm, when these groups or facts are considered in connection with others, such as the mutual affinities of the members of the same group, their geographical distribution in past and present times, and their geological succession. It is incredible that all these facts should speak falsely. He who is not content to look, like a savage, at the phenomena of nature as disconnected, cannot any longer believe that man is the work of a separate act of creation. He will be forced to admit that the close resemblance of the embryo of man to that, for instance, of a dog—the construction of his skull, limbs and whole frame on the same plan with that of other mammals, independently of the uses to which the parts may be put—the occasional re-appearance of various structures, for instance of several muscles, which man does not normally possess, but which are common to the Quadrumana—and a crowd of analogous facts—all point in the plainest manner to the conclusion that man is the co-descendant with other mammals of a common progenitor.

We have seen that man incessantly presents individual differences in all parts of his body and in his mental faculties. These differences or variations seem to be induced by the same general causes, and to obey the same laws as with the lower animals. In both cases similar laws of inheritance prevail. Man tends to increase at a greater rate than his means of subsistence; consequently he is occasionally subjected to a severe struggle for existence, and natural selection will have effected whatever lies within its scope. A succession of strongly-marked variations of a similar nature is by no means requisite; slight fluctuating differences in the individual suffice for the work of natural selection; not that we have any reason to suppose that in the same species, all parts of the organisation tend to vary to the same degree. We may feel assured that the inherited effects of the long-continued use or disuse of parts will have done much in the same direction with natural selection. Modifications formerly of importance, though no longer of any special use, are long-inherited. When one part is modified, other parts change through the principle of correlation, of which we have instances in many curious cases of correlated monstrosities. Something may be attributed to the direct and definite action of the surrounding conditions of life, such as abundant food, heat or moisture; and lastly, many characters of slight physiological importance, some indeed of considerable importance, have been gained through sexual selection.

No doubt man, as well as every other animal, presents structures, which seem to our limited knowledge, not to be now of any service to him, nor to have been so formerly, either for the general conditions of life, or in the relations of one sex to the other. Such structures cannot be accounted for by any form of selection, or by the inherited effects of the use and disuse of parts. We know, however, that many strange and strongly-marked peculiarities of structure occasionally appear in our domesticated productions, and if their unknown causes were to act more uniformly, they would probably become common to all the individuals of the species. We may hope hereafter to understand something about the causes of such occasional modifications, especially through the study of monstrosities: hence the labours of experimentalists, such as those of M. Camille Dareste, are full of promise for the future. In general we can only say that the cause of each slight variation and of each monstrosity lies much more in the constitution of the organism, than in the nature of the surrounding conditions; though new and changed conditions certainly play an important part in exciting organic changes of many kinds.

Through the means just specified, aided perhaps by others as yet undiscovered, man has been raised to his present state. But since he attained to the rank of manhood, he has diverged into distinct races, or as they may be more fitly called, sub-species. Some of these, such as the Negro and European, are so distinct that, if specimens had been brought to a naturalist without any further information, they would undoubtedly have been considered by him as good and true species. Nevertheless all the races agree in so many unimportant details of structure and in so many mental peculiarities that these can be accounted for only by inheritance from a common progenitor; and a progenitor thus characterised would probably deserve to rank as man.

It must not be supposed that the divergence of each race from the other races, and of all from a common stock, can be traced back to any one pair of progenitors. On the contrary, at every stage in the process of modification, all the individuals which were in any way better fitted for their conditions of life, though in different degrees, would have survived in greater numbers than the less well-fitted. The process would have been like that followed by man, when he does not intentionally select particular individuals, but breeds from all the superior individuals, and neglects the inferior. He thus slowly but surely modifies his stock, and unconsciously forms a new strain. So with respect to modifications acquired independently of selection, and due to variations arising from the nature of the organism and the action of the surrounding conditions, or from changed habits of life, no single pair will have been modified much more than the other pairs inhabiting the same country, for all will have been continually blended through free intercrossing.

By considering the embryological structure of man,—the homologies which he presents with the lower animals,—the rudiments which he retains,—and the reversions to which he is liable, we can partly recall in imagination the former condition of our early progenitors; and can approximately place them in their proper place in the zoological series. We thus learn that man is descended from a hairy, tailed quadruped, probably arboreal in its habits, and an inhabitant of the Old World. This creature, if its whole structure had been examined by a naturalist, would have been classed amongst the Quadrumana, as surely as the still more ancient progenitor of the Old and New World monkeys. The Quadrumana and all the higher mammals are probably derived from an ancient marsupial animal, and this through a long line of diversified forms, from some amphibian-like creature, and this again from some fish-like animal. In the dim obscurity of the past we can see that the early progenitor of all the Vertebrata must have been an aquatic animal, provided with branchiae, with the two sexes united in the same individual, and with the most important organs of the body (such as the brain and heart) imperfectly or not at all developed. This animal seems to have been more like the larvae of the existing marine Ascidians than any other known form.

The high standard of our intellectual powers and moral disposition is the greatest difficulty which presents itself, after we have been driven to this conclusion on the origin of man. But every one who admits the principle of evolution, must see that the mental powers of the higher animals, which are the same in kind with those of man, though so different in degree, are capable of advancement. Thus the interval between the mental powers of one of the higher apes and of a fish, or between those of an ant and scale-insect, is immense; yet their development does not offer any special difficulty; for with our domesticated animals, the mental faculties are certainly variable, and the variations are inherited. No one doubts that they are of the utmost importance to animals in a state of nature. Therefore the conditions are favourable for their development through natural selection. The same conclusion may be extended to man; the intellect must have been all-important to him, even at a very remote period, as enabling him to invent and use language, to make weapons, tools, traps, etc., whereby with the aid of his social habits, he long ago became the most dominant of all living creatures.

A great stride in the development of the intellect will have followed, as soon as the half-art and half-instinct of language came into use; for the continued use of language will have reacted on the brain and produced an inherited effect; and this again will have reacted on the improvement of language. As Mr. Chauncey Wright (1. ‘On the Limits of Natural Selection,’ in the ‘North American Review,’ Oct. 1870, p. 295.) has well remarked, the largeness of the brain in man relatively to his body, compared with the lower animals, may be attributed in chief part to the early use of some simple form of language,—that wonderful engine which affixes signs to all sorts of objects and qualities, and excites trains of thought which would never arise from the mere impression of the senses, or if they did arise could not be followed out. The higher intellectual powers of man, such as those of ratiocination, abstraction, self-consciousness, etc., probably follow from the continued improvement and exercise of the other mental faculties.

The development of the moral qualities is a more interesting problem. The foundation lies in the social instincts, including under this term the family ties. These instincts are highly complex, and in the case of the lower animals give special tendencies towards certain definite actions; but the more important elements are love, and the distinct emotion of sympathy. Animals endowed with the social instincts take pleasure in one another’s company, warn one another of danger, defend and aid one another in many ways. These instincts do not extend to all the individuals of the species, but only to those of the same community. As they are highly beneficial to the species, they have in all probability been acquired through natural selection.

A moral being is one who is capable of reflecting on his past actions and their motives—of approving of some and disapproving of others; and the fact that man is the one being who certainly deserves this designation, is the greatest of all distinctions between him and the lower animals. But in the fourth chapter I have endeavoured to shew that the moral sense follows, firstly, from the enduring and ever-present nature of the social instincts; secondly, from man’s appreciation of the approbation and disapprobation of his fellows; and thirdly, from the high activity of his mental faculties, with past impressions extremely vivid; and in these latter respects he differs from the lower animals. Owing to this condition of mind, man cannot avoid looking both backwards and forwards, and comparing past impressions. Hence after some temporary desire or passion has mastered his social instincts, he reflects and compares the now weakened impression of such past impulses with the ever-present social instincts; and he then feels that sense of dissatisfaction which all unsatisfied instincts leave behind them, he therefore resolves to act differently for the future,—and this is conscience. Any instinct, permanently stronger or more enduring than another, gives rise to a feeling which we express by saying that it ought to be obeyed. A pointer dog, if able to reflect on his past conduct, would say to himself, I ought (as indeed we say of him) to have pointed at that hare and not have yielded to the passing temptation of hunting it.

Social animals are impelled partly by a wish to aid the members of their community in a general manner, but more commonly to perform certain definite actions. Man is impelled by the same general wish to aid his fellows; but has few or no special instincts. He differs also from the lower animals in the power of expressing his desires by words, which thus become a guide to the aid required and bestowed. The motive to give aid is likewise much modified in man: it no longer consists solely of a blind instinctive impulse, but is much influenced by the praise or blame of his fellows. The appreciation and the bestowal of praise and blame both rest on sympathy; and this emotion, as we have seen, is one of the most important elements of the social instincts. Sympathy, though gained as an instinct, is also much strengthened by exercise or habit. As all men desire their own happiness, praise or blame is bestowed on actions and motives, according as they lead to this end; and as happiness is an essential part of the general good, the greatest-happiness principle indirectly serves as a nearly safe standard of right and wrong. As the reasoning powers advance and experience is gained, the remoter effects of certain lines of conduct on the character of the individual, and on the general good, are perceived; and then the self-regarding virtues come within the scope of public opinion, and receive praise, and their opposites blame. But with the less civilised nations reason often errs, and many bad customs and base superstitions come within the same scope, and are then esteemed as high virtues, and their breach as heavy crimes.

The moral faculties are generally and justly esteemed as of higher value than the intellectual powers. But we should bear in mind that the activity of the mind in vividly recalling past impressions is one of the fundamental though secondary bases of conscience. This affords the strongest argument for educating and stimulating in all possible ways the intellectual faculties of every human being. No doubt a man with a torpid mind, if his social affections and sympathies are well developed, will be led to good actions, and may have a fairly sensitive conscience. But whatever renders the imagination more vivid and strengthens the habit of recalling and comparing past impressions, will make the conscience more sensitive, and may even somewhat compensate for weak social affections and sympathies.

The moral nature of man has reached its present standard, partly through the advancement of his reasoning powers and consequently of a just public opinion, but especially from his sympathies having been rendered more tender and widely diffused through the effects of habit, example, instruction, and reflection. It is not improbable that after long practice virtuous tendencies may be inherited. With the more civilised races, the conviction of the existence of an all-seeing Deity has had a potent influence on the advance of morality. Ultimately man does not accept the praise or blame of his fellows as his sole guide, though few escape this influence, but his habitual convictions, controlled by reason, afford him the safest rule. His conscience then becomes the supreme judge and monitor. Nevertheless the first foundation or origin of the moral sense lies in the social instincts, including sympathy; and these instincts no doubt were primarily gained, as in the case of the lower animals, through natural selection.

The belief in God has often been advanced as not only the greatest, but the most complete of all the distinctions between man and the lower animals. It is however impossible, as we have seen, to maintain that this belief is innate or instinctive in man. On the other hand a belief in all-pervading spiritual agencies seems to be universal; and apparently follows from a considerable advance in man’s reason, and from a still greater advance in his faculties of imagination, curiosity and wonder. I am aware that the assumed instinctive belief in God has been used by many persons as an argument for His existence. But this is a rash argument, as we should thus be compelled to believe in the existence of many cruel and malignant spirits, only a little more powerful than man; for the belief in them is far more general than in a beneficent Deity. The idea of a universal and beneficent Creator does not seem to arise in the mind of man, until he has been elevated by long-continued culture.

He who believes in the advancement of man from some low organised form, will naturally ask how does this bear on the belief in the immortality of the soul. The barbarous races of man, as Sir J. Lubbock has shewn, possess no clear belief of this kind; but arguments derived from the primeval beliefs of savages are, as we have just seen, of little or no avail. Few persons feel any anxiety from the impossibility of determining at what precise period in the development of the individual, from the first trace of a minute germinal vesicle, man becomes an immortal being; and there is no greater cause for anxiety because the period cannot possibly be determined in the gradually ascending organic scale. (2. The Rev. J.A. Picton gives a discussion to this effect in his ‘New Theories and the Old Faith,’ 1870.)

I am aware that the conclusions arrived at in this work will be denounced by some as highly irreligious; but he who denounces them is bound to shew why it is more irreligious to explain the origin of man as a distinct species by descent from some lower form, through the laws of variation and natural selection, than to explain the birth of the individual through the laws of ordinary reproduction. The birth both of the species and of the individual are equally parts of that grand sequence of events, which our minds refuse to accept as the result of blind chance. The understanding revolts at such a conclusion, whether or not we are able to believe that every slight variation of structure,—the union of each pair in marriage, the dissemination of each seed,—and other such events, have all been ordained for some special purpose.

Sexual selection has been treated at great length in this work; for, as I have attempted to shew, it has played an important part in the history of the organic world. I am aware that much remains doubtful, but I have endeavoured to give a fair view of the whole case. In the lower divisions of the animal kingdom, sexual selection seems to have done nothing: such animals are often affixed for life to the same spot, or have the sexes combined in the same individual, or what is still more important, their perceptive and intellectual faculties are not sufficiently advanced to allow of the feelings of love and jealousy, or of the exertion of choice. When, however, we come to the Arthropoda and Vertebrata, even to the lowest classes in these two great Sub-Kingdoms, sexual selection has effected much.

In the several great classes of the animal kingdom,—in mammals, birds, reptiles, fishes, insects, and even crustaceans,—the differences between the sexes follow nearly the same rules. The males are almost always the wooers; and they alone are armed with special weapons for fighting with their rivals. They are generally stronger and larger than the females, and are endowed with the requisite qualities of courage and pugnacity. They are provided, either exclusively or in a much higher degree than the females, with organs for vocal or instrumental music, and with odoriferous glands. They are ornamented with infinitely diversified appendages, and with the most brilliant or conspicuous colours, often arranged in elegant patterns, whilst the females are unadorned. When the sexes differ in more important structures, it is the male which is provided with special sense-organs for discovering the female, with locomotive organs for reaching her, and often with prehensile organs for holding her. These various structures for charming or securing the female are often developed in the male during only part of the year, namely the breeding-season. They have in many cases been more or less transferred to the females; and in the latter case they often appear in her as mere rudiments. They are lost or never gained by the males after emasculation. Generally they are not developed in the male during early youth, but appear a short time before the age for reproduction. Hence in most cases the young of both sexes resemble each other; and the female somewhat resembles her young offspring throughout life. In almost every great class a few anomalous cases occur, where there has been an almost complete transposition of the characters proper to the two sexes; the females assuming characters which properly belong to the males. This surprising uniformity in the laws regulating the differences between the sexes in so many and such widely separated classes, is intelligible if we admit the action of one common cause, namely sexual selection.

Sexual selection depends on the success of certain individuals over others of the same sex, in relation to the propagation of the species; whilst natural selection depends on the success of both sexes, at all ages, in relation to the general conditions of life. The sexual struggle is of two kinds; in the one it is between individuals of the same sex, generally the males, in order to drive away or kill their rivals, the females remaining passive; whilst in the other, the struggle is likewise between the individuals of the same sex, in order to excite or charm those of the opposite sex, generally the females, which no longer remain passive, but select the more agreeable partners. This latter kind of selection is closely analogous to that which man unintentionally, yet effectually, brings to bear on his domesticated productions, when he preserves during a long period the most pleasing or useful individuals, without any wish to modify the breed.

The laws of inheritance determine whether characters gained through sexual selection by either sex shall be transmitted to the same sex, or to both; as well as the age at which they shall be developed. It appears that variations arising late in life are commonly transmitted to one and the same sex. Variability is the necessary basis for the action of selection, and is wholly independent of it. It follows from this, that variations of the same general nature have often been taken advantage of and accumulated through sexual selection in relation to the propagation of the species, as well as through natural selection in relation to the general purposes of life. Hence secondary sexual characters, when equally transmitted to both sexes can be distinguished from ordinary specific characters only by the light of analogy. The modifications acquired through sexual selection are often so strongly pronounced that the two sexes have frequently been ranked as distinct species, or even as distinct genera. Such strongly-marked differences must be in some manner highly important; and we know that they have been acquired in some instances at the cost not only of inconvenience, but of exposure to actual danger.

The belief in the power of sexual selection rests chiefly on the following considerations. Certain characters are confined to one sex; and this alone renders it probable that in most cases they are connected with the act of reproduction. In innumerable instances these characters are fully developed only at maturity, and often during only a part of the year, which is always the breeding-season. The males (passing over a few exceptional cases) are the more active in courtship; they are the better armed, and are rendered the more attractive in various ways. It is to be especially observed that the males display their attractions with elaborate care in the presence of the females; and that they rarely or never display them excepting during the season of love. It is incredible that all this should be purposeless. Lastly we have distinct evidence with some quadrupeds and birds, that the individuals of one sex are capable of feeling a strong antipathy or preference for certain individuals of the other sex.

Bearing in mind these facts, and the marked results of man’s unconscious selection, when applied to domesticated animals and cultivated plants, it seems to me almost certain that if the individuals of one sex were during a long series of generations to prefer pairing with certain individuals of the other sex, characterised in some peculiar manner, the offspring would slowly but surely become modified in this same manner. I have not attempted to conceal that, excepting when the males are more numerous than the females, or when polygamy prevails, it is doubtful how the more attractive males succeed in leaving a large number of offspring to inherit their superiority in ornaments or other charms than the less attractive males; but I have shewn that this would probably follow from the females,—especially the more vigorous ones, which would be the first to breed,—preferring not only the more attractive but at the same time the more vigorous and victorious males.

Although we have some positive evidence that birds appreciate bright and beautiful objects, as with the bower-birds of Australia, and although they certainly appreciate the power of song, yet I fully admit that it is astonishing that the females of many birds and some mammals should be endowed with sufficient taste to appreciate ornaments, which we have reason to attribute to sexual selection; and this is even more astonishing in the case of reptiles, fish, and insects. But we really know little about the minds of the lower animals. It cannot be supposed, for instance, that male birds of paradise or peacocks should take such pains in erecting, spreading, and vibrating their beautiful plumes before the females for no purpose. We should remember the fact given on excellent authority in a former chapter, that several peahens, when debarred from an admired male, remained widows during a whole season rather than pair with another bird.

Nevertheless I know of no fact in natural history more wonderful than that the female Argus pheasant should appreciate the exquisite shading of the ball-and-socket ornaments and the elegant patterns on the wing-feather of the male. He who thinks that the male was created as he now exists must admit that the great plumes, which prevent the wings from being used for flight, and which are displayed during courtship and at no other time in a manner quite peculiar to this one species, were given to him as an ornament. If so, he must likewise admit that the female was created and endowed with the capacity of appreciating such ornaments. I differ only in the conviction that the male Argus pheasant acquired his beauty gradually, through the preference of the females during many generations for the more highly ornamented males; the aesthetic capacity of the females having been advanced through exercise or habit, just as our own taste is gradually improved. In the male through the fortunate chance of a few feathers being left unchanged, we can distinctly trace how simple spots with a little fulvous shading on one side may have been developed by small steps into the wonderful ball-and-socket ornaments; and it is probable that they were actually thus developed.

Everyone who admits the principle of evolution, and yet feels great difficulty in admitting that female mammals, birds, reptiles, and fish, could have acquired the high taste implied by the beauty of the males, and which generally coincides with our own standard, should reflect that the nerve-cells of the brain in the highest as well as in the lowest members of the Vertebrate series, are derived from those of the common progenitor of this great Kingdom. For we can thus see how it has come to pass that certain mental faculties, in various and widely distinct groups of animals, have been developed in nearly the same manner and to nearly the same degree.

The reader who has taken the trouble to go through the several chapters devoted to sexual selection, will be able to judge how far the conclusions at which I have arrived are supported by sufficient evidence. If he accepts these conclusions he may, I think, safely extend them to mankind; but it would be superfluous here to repeat what I have so lately said on the manner in which sexual selection apparently has acted on man, both on the male and female side, causing the two sexes to differ in body and mind, and the several races to differ from each other in various characters, as well as from their ancient and lowly-organised progenitors.

He who admits the principle of sexual selection will be led to the remarkable conclusion that the nervous system not only regulates most of the existing functions of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities. Courage, pugnacity, perseverance, strength and size of body, weapons of all kinds, musical organs, both vocal and instrumental, bright colours and ornamental appendages, have all been indirectly gained by the one sex or the other, through the exertion of choice, the influence of love and jealousy, and the appreciation of the beautiful in sound, colour or form; and these powers of the mind manifestly depend on the development of the brain.

Man scans with scrupulous care the character and pedigree of his horses, cattle, and dogs before he matches them; but when he comes to his own marriage he rarely, or never, takes any such care. He is impelled by nearly the same motives as the lower animals, when they are left to their own free choice, though he is in so far superior to them that he highly values mental charms and virtues. On the other hand he is strongly attracted by mere wealth or rank. Yet he might by selection do something not only for the bodily constitution and frame of his offspring, but for their intellectual and moral qualities. Both sexes ought to refrain from marriage if they are in any marked degree inferior in body or mind; but such hopes are Utopian and will never be even partially realised until the laws of inheritance are thoroughly known. Everyone does good service, who aids towards this end. When the principles of breeding and inheritance are better understood, we shall not hear ignorant members of our legislature rejecting with scorn a plan for ascertaining whether or not consanguineous marriages are injurious to man.

The advancement of the welfare of mankind is a most intricate problem: all ought to refrain from marriage who cannot avoid abject poverty for their children; for poverty is not only a great evil, but tends to its own increase by leading to recklessness in marriage. On the other hand, as Mr. Galton has remarked, if the prudent avoid marriage, whilst the reckless marry, the inferior members tend to supplant the better members of society. Man, like every other animal, has no doubt advanced to his present high condition through a struggle for existence consequent on his rapid multiplication; and if he is to advance still higher, it is to be feared that he must remain subject to a severe struggle. Otherwise he would sink into indolence, and the more gifted men would not be more successful in the battle of life than the less gifted. Hence our natural rate of increase, though leading to many and obvious evils, must not be greatly diminished by any means. There should be open competition for all men; and the most able should not be prevented by laws or customs from succeeding best and rearing the largest number of offspring. Important as the struggle for existence has been and even still is, yet as far as the highest part of man’s nature is concerned there are other agencies more important. For the moral qualities are advanced, either directly or indirectly, much more through the effects of habit, the reasoning powers, instruction, religion, etc., than through natural selection; though to this latter agency may be safely attributed the social instincts, which afforded the basis for the development of the moral sense.

The main conclusion arrived at in this work, namely, that man is descended from some lowly organised form, will, I regret to think, be highly distasteful to many. But there can hardly be a doubt that we are descended from barbarians. The astonishment which I felt on first seeing a party of Fuegians on a wild and broken shore will never be forgotten by me, for the reflection at once rushed into my mind—such were our ancestors. These men were absolutely naked and bedaubed with paint, their long hair was tangled, their mouths frothed with excitement, and their expression was wild, startled, and distrustful. They possessed hardly any arts, and like wild animals lived on what they could catch; they had no government, and were merciless to every one not of their own small tribe. He who has seen a savage in his native land will not feel much shame, if forced to acknowledge that the blood of some more humble creature flows in his veins. For my own part I would as soon be descended from that heroic little monkey, who braved his dreaded enemy in order to save the life of his keeper, or from that old baboon, who descending from the mountains, carried away in triumph his young comrade from a crowd of astonished dogs—as from a savage who delights to torture his enemies, offers up bloody sacrifices, practices infanticide without remorse, treats his wives like slaves, knows no decency, and is haunted by the grossest superstitions.

Man may be excused for feeling some pride at having risen, though not through his own exertions, to the very summit of the organic scale; and the fact of his having thus risen, instead of having been aboriginally placed there, may give him hope for a still higher destiny in the distant future. But we are not here concerned with hopes or fears, only with the truth as far as our reason permits us to discover it; and I have given the evidence to the best of my ability. We must, however, acknowledge, as it seems to me, that man with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his god-like intellect which has penetrated into the movements and constitution of the solar system—with all these exalted powers—Man still bears in his bodily frame the indelible stamp of his lowly origin.

SUPPLEMENTAL NOTE. ON SEXUAL SELECTION IN RELATION TO MONKEYS.

Reprinted from NATURE, November 2, 1876, p. 18.

In the discussion on Sexual Selection in my ‘Descent of Man,’ no case interested and perplexed me so much as the brightly-coloured hinder ends and adjoining parts of certain monkeys. As these parts are more brightly coloured in one sex than the other, and as they become more brilliant during the season of love, I concluded that the colours had been gained as a sexual attraction. I was well aware that I thus laid myself open to ridicule; though in fact it is not more surprising that a monkey should display his bright-red hinder end than that a peacock should display his magnificent tail. I had, however, at that time no evidence of monkeys exhibiting this part of their bodies during their courtship; and such display in the case of birds affords the best evidence that the ornaments of the males are of service to them by attracting or exciting the females. I have lately read an article by Joh. von Fischer, of Gotha, published in ‘Der Zoologische Garten,’ April 1876, on the expression of monkeys under various emotions, which is well worthy of study by any one interested in the subject, and which shews that the author is a careful and acute observer. In this article there is an account of the behaviour of a young male mandrill when he first beheld himself in a looking-glass, and it is added, that after a time he turned round and presented his red hinder end to the glass. Accordingly I wrote to Herr J. von Fischer to ask what he supposed was the meaning of this strange action, and he has sent me two long letters full of new and curious details, which will, I hope, be hereafter published. He says that he was himself at first perplexed by the above action, and was thus led carefully to observe several individuals of various other species of monkeys, which he has long kept in his house. He finds that not only the mandrill (Cynocephalus mormon) but the drill (C. leucophaeus) and three other kinds of baboons (C. hamadryas, sphinx, and babouin), also Cynopithecus niger, and Macacus rhesus and nemestrinus, turn this part of their bodies, which in all these species is more or less brightly coloured, to him when they are pleased, and to other persons as a sort of greeting. He took pains to cure a Macacus rhesus, which he had kept for five years, of this indecorous habit, and at last succeeded. These monkeys are particularly apt to act in this manner, grinning at the same time, when first introduced to a new monkey, but often also to their old monkey friends; and after this mutual display they begin to play together. The young mandrill ceased spontaneously after a time to act in this manner towards his master, von Fischer, but continued to do so towards persons who were strangers and to new monkeys. A young Cynopithecus niger never acted, excepting on one occasion, in this way towards his master, but frequently towards strangers, and continues to do so up to the present time. From these facts Von Fischer concludes that the monkeys which behaved in this manner before a looking-glass (viz., the mandrill, drill, Cynopithecus niger, Macacus rhesus and nemestrinus) acted as if their reflection were a new acquaintance. The mandrill and drill, which have their hinder ends especially ornamented, display it even whilst quite young, more frequently and more ostentatiously than do the other kinds. Next in order comes Cynocephalus hamadryas, whilst the other species act in this manner seldomer. The individuals, however, of the same species vary in this respect, and some which were very shy never displayed their hinder ends. It deserves especial attention that Von Fischer has never seen any species purposely exhibit the hinder part of its body, if not at all coloured. This remark applies to many individuals of Macacus cynomolgus and Cercocebus radiatus (which is closely allied to M. rhesus), to three species of Cercopithecus and several American monkeys. The habit of turning the hinder ends as a greeting to an old friend or new acquaintance, which seems to us so odd, is not really more so than the habits of many savages, for instance that of rubbing their bellies with their hands, or rubbing noses together. The habit with the mandrill and drill seems to be instinctive or inherited, as it was followed by very young animals; but it is modified or guided, like so many other instincts, by observation, for Von Fischer says that they take pains to make their display fully; and if made before two observers, they turn to him who seems to pay the most attention.

With respect to the origin of the habit, Von Fischer remarks that his monkeys like to have their naked hinder ends patted or stroked, and that they then grunt with pleasure. They often also turn this part of their bodies to other monkeys to have bits of dirt picked off, and so no doubt it would be with respect to thorns. But the habit with adult animals is connected to a certain extent with sexual feelings, for Von Fischer watched through a glass door a female Cynopithecus niger, and she during several days, “umdrehte und dem Männchen mit gurgelnden Tönen die stark geröthete Sitzflache zeigte, was ich früher nie an diesem Thier bemerkt hatte. Beim Anblick dieses Gegenstandes erregte sich das Männchen sichtlich, denn es polterte heftig an den Stäben, ebenfalls gurgelnde Laute ausstossend.” As all the monkeys which have the hinder parts of their bodies more or less brightly coloured live, according to Von Fischer, in open rocky places, he thinks that these colours serve to render one sex conspicuous at a distance to the other; but, as monkeys are such gregarious animals, I should have thought that there was no need for the sexes to recognise each other at a distance. It seems to me more probable that the bright colours, whether on the face or hinder end, or, as in the mandrill, on both, serve as a sexual ornament and attraction. Anyhow, as we now know that monkeys have the habit of turning their hinder ends towards other monkeys, it ceases to be at all surprising that it should have been this part of their bodies which has been more or less decorated. The fact that it is only the monkeys thus characterised which, as far as at present known, act in this manner as a greeting towards other monkeys renders it doubtful whether the habit was first acquired from some independent cause, and that afterwards the parts in question were coloured as a sexual ornament; or whether the colouring and the habit of turning round were first acquired through variation and sexual selection, and that afterwards the habit was retained as a sign of pleasure or as a greeting, through the principle of inherited association. This principle apparently comes into play on many occasions: thus it is generally admitted that the songs of birds serve mainly as an attraction during the season of love, and that the leks, or great congregations of the black-grouse, are connected with their courtship; but the habit of singing has been retained by some birds when they feel happy, for instance by the common robin, and the habit of congregating has been retained by the black-grouse during other seasons of the year.

I beg leave to refer to one other point in relation to sexual selection. It has been objected that this form of selection, as far as the ornaments of the males are concerned, implies that all females within the same district must possess and exercise exactly the same taste. It should, however, be observed, in the first place, that although the range of variation of a species may be very large, it is by no means indefinite. I have elsewhere given a good instance of this fact in the pigeon, of which there are at least a hundred varieties differing widely in their colours, and at least a score of varieties of the fowl differing in the same kind of way; but the range of colour in these two species is extremely distinct. Therefore the females of natural species cannot have an unlimited scope for their taste. In the second place, I presume that no supporter of the principle of sexual selection believes that the females select particular points of beauty in the males; they are merely excited or attracted in a greater degree by one male than by another, and this seems often to depend, especially with birds, on brilliant colouring. Even man, excepting perhaps an artist, does not analyse the slight differences in the features of the woman whom he may admire, on which her beauty depends. The male mandrill has not only the hinder end of his body, but his face gorgeously coloured and marked with oblique ridges, a yellow beard, and other ornaments. We may infer from what we see of the variation of animals under domestication, that the above several ornaments of the mandrill were gradually acquired by one individual varying a little in one way, and another individual in another way. The males which were the handsomest or the most attractive in any manner to the females would pair oftenest, and would leave rather more offspring than other males. The offspring of the former, although variously intercrossed, would either inherit the peculiarities of their fathers or transmit an increased tendency to vary in the same manner. Consequently the whole body of males inhabiting the same country would tend from the effects of constant intercrossing to become modified almost uniformly, but sometimes a little more in one character and sometimes in another, though at an extremely slow rate; all ultimately being thus rendered more attractive to the females. The process is like that which I have called unconscious selection by man, and of which I have given several instances. In one country the inhabitants value a fleet or light dog or horse, and in another country a heavier and more powerful one; in neither country is there any selection of individual animals with lighter or stronger bodies and limbs; nevertheless after a considerable lapse of time the individuals are found to have been modified in the desired manner almost uniformly, though differently in each country. In two absolutely distinct countries inhabited by the same species, the individuals of which can never during long ages have intermigrated and intercrossed, and where, moreover, the variations will probably not have been identically the same, sexual selection might cause the males to differ. Nor does the belief appear to me altogether fanciful that two sets of females, surrounded by a very different environment, would be apt to acquire somewhat different tastes with respect to form, sound, or colour. However this may be, I have given in my ‘Descent of Man’ instances of closely-allied birds inhabiting distinct countries, of which the young and the females cannot be distinguished, whilst the adult males differ considerably, and this may be attributed with much probability to the action of sexual selection.

INDEX. — Abbot, C., on the battles of seals.

Abductor of the fifth metatarsal, presence of, in man.

Abercrombie, Dr., on disease of the brain affecting speech.

Abipones, marriage customs of the.

Abortion, prevalence of the practice of.

Abou-Simbel, caves of.

Abramis brama.

Abstraction, power of, in animals.

Acalles, stridulation of.

Acanthodactylus capensis, sexual differences of colour in.

Accentor Modularis.

Acclimatisation, difference of, in different races of men.

Achetidae, stridulation of the; rudimentary stridulating organs in female.

Acilius sulcatus, elytra of the female.

Acomus, development of spurs in the female of.

Acridiidae, stridulation of the; rudimentary stridulating organs in female.

Acromio-basilar muscle, and quadrupedal gait.

Acting.

Actiniae, bright colours of.

Adams, Mr., migration of birds; intelligence of nut-hatch; on the Bombycilla carolinensis.

Admiral butterfly.

Adoption of the young of other animals by female monkeys.

Advancement in the organic scale, Von Baer’s definition of.

Aeby, on the difference between the skulls of man and the quadrumana.

Aesthetic faculty, not highly developed in savages.

Affection, maternal; manifestation of, by animals; parental and filial, partly the result of natural selection; mutual, of birds; shewn by birds in confinement, for certain persons.

Africa, probably the birthplace of man; South, crossed population of; South, retention of colour by the Dutch in; South, proportion of the sexes in the butterflies of; tattooing practised in; Northern, coiffure of natives of.

Agassiz, L., on conscience in dogs; on the coincidence of the races of man with zoological provinces; on the number of species of man; on the courtship of the land-snails; on the brightness of the colours of male fishes during the breeding season; on the frontal protuberance of the males of Geophagus and Cichla; male fishes hatching ova in their mouths; sexual differences in colour of chromids; on the slight sexual differences of the South Americans; on the tattooing of the Amazonian Indians.

Age, in relation to the transmission of characters in birds; variation in accordance with, in birds.

Agelaeus phoeniceus.

Ageronia feronia, noise produced by.

Agrion, dimorphism in.

Agrion Ramburii, sexes of.

Agrionidae, difference in the sexes of.

Agrotis exclamationis.

Ague, tertian, dog suffering from.

Ainos, hairiness of the.

Aitchison, Mr., on sheep.

Aithurus polytmus, young of.

Albino birds.

Alca torda, young of.

Alces palmata.

Alder and Hancock, MM., on the nudi-branch mollusca.

Allen, J.A., vigour of birds earliest hatched; effect of difference of temperature, light, etc., on birds; colours of birds; on the relative size of the sexes of Callorhinus ursinus; on the name of Otaria jubata; on the pairing of seals; on sexual differences in the colour of bats.

Allen, S., on the habits of Hoplopterus; on the plumes of herons; on the vernal moult of Herodius bubulcus.

Alligator, courtship of the male; roaring of the male.

Amadavat, pugnacity of male.

Amadina Lathami, display of plumage by the male.

Amadina castanotis, display of plumage by the male.

Amazons, butterflies of the; fishes of the.

America, variation in the skulls of aborigines of; wide range of aborigines of; lice of the natives of; general beardlessness of the natives of.

America, North, butterflies of; Indians of, women a cause of strife among the; Indians of, their notions of female beauty.

America, South, character of the natives of; population of parts of; piles of stones in; extinction of the fossil horse of; desert-birds of; slight sexual difference of the aborigines of; prevalence of infanticide in.

American languages, often highly artificial.

Americans, wide geographical range of; native, variability of; and negroes, difference of; aversion of, to hair on the face.

Ammophila, on the jaws of.

Ammotragus tragelaphus, hairy forelegs of.

Amphibia, affinity of, to the ganoid fishes; vocal organs of the.

Amphibians, breeding whilst immature.

Amphioxus.

Amphipoda, males sexually mature while young.

Amunoph III., negro character of, features of.

Anal appendages of insects.

Analogous variation in the plumage of birds.

Anas.

Anas acuta, male plumage of.

Anas boschas, male plumage of.

Anas histrionica.

Anas punctata.

Anastomus oscitans, sexes and young of; white nuptial plumage of.

Anatidae, voices of.

Anax junius, differences in the sexes of.

Andaman islanders, susceptible to change of climate.

Anderson, Dr., on the tail of Macacus brunneus; the Bufo sikimmensis; sounds of Echis carinata.

Andreana fulva.

Anglo-Saxons, estimation of the beard among the.

Animals, domesticated, more fertile than wild; cruelty of savages to; characters common to man and; domestic, change of breeds of.

Annelida, colours of.

Anobium tessellatum, sounds produced by.

Anolis cristatellus, male, crest of; pugnacity of the male; throat-pouch of.

Anser canadensis.

Anset cygnoides; knob at the base of the beak of.

Anser hyperboreus, whiteness of.

Antelope, prong-horned, horns of.

Antelopes, generally polygamous; horns of; canine teeth of some male; use of horns of; dorsal crests in; dewlaps of; winter change of two species of; peculiar markings of.

Antennae, furnished with cushions in the male of Penthe.

Anthidium manicatum, large male of.

Anthocharis cardamines; sexual difference of colour in.

Anthocharis genutia.

Anthocharis sara.

Anthophora acervorum, large male of.

Anthophora retusa, difference of the sexes in.

Anthropidae.

Anthus, moulting of.

Antics of birds.

Antigua, Dr. Nicholson’s observations on yellow fever in.

Antilocapra americana, horns of.

Antilope bezoartica, horned females of; sexual difference in the colour of.

Antilope Dorcas and euchore.

Antilope euchore, horns of.

Antilope montana, rudimentary canines in the young male of.

Antilope niger, sing-sing, caama, and gorgon, sexual differences in the colours of.

Antilope oreas, horns of.

Antilope saiga, polygamous habits of.

Antilope strepsiceros, horns of.

Antilope subgutturosa, absence of suborbital pits in.

Antipathy, shewn by birds in confinement, to certain persons.

Ants, large size of the cerebral ganglia in; soldier, large jaws of; playing together; memory in; intercommunication of, by means of the antennae; habits of; difference of the sexes in; recognition of each other by, after separation.

Ants White, habits of.

Anura.

Apatania muliebris, male unknown.

Apathus, difference of the sexes in.

Apatura Iris.

Apes, difference of the young, from the adult; semi-erect attitude of some; mastoid processes of; influences of the jaw-muscles on the physiognomy of; female, destitute of large canines; building platforms; imitative faculties of; anthropomorphous; probable speedy extermination of the; Gratiolet on the evolution of; canine teeth of male; females of some, less hairy beneath than the males.

Apes, long-armed, their mode of progression.

Aphasia, Dr. Bateman on.

Apis mellifica, large male of.

Apollo, Greek statues of.

Apoplexy in Cebus Azarae.

Appendages, anal, of insects.

Approbation, influence of the love of.

Aprosmictus scapulatus.

Apus, proportion of sexes.

Aquatic birds, frequency of white plumage in.

Aquila chrysaetos.

Arab women, elaborate and peculiar coiffure of.

Arabs, fertility of crosses with other races; gashing of cheeks and temples among the.

Arachnida.

Arakhan, artificial widening of the forehead by the natives of.

Arboricola, young of.

Archeopteryx.

Arctiidae, coloration of the.

Ardea asha, rufescens, and coerulea, change of colour in.

Ardea coerulea, breeding in immature plumage.

Ardea gularis, change of plumage in.

Ardea herodias, love-gestures of the male.

Ardea ludoviciana, age of mature plumage in; continued growth of crest and plumes in the male of.

Ardea nycticorax, cries of.

Ardeola, young of.

Ardetta, changes of plumage in.

Argenteuil.

Argus pheasant, display of plumage by the male; ocellated spots of the; gradation of characters in the.

Argyll, Duke of, on the physical weakness of man; the fashioning of implements peculiar to man; on the contest in man between right and wrong; on the primitive civilisation of man; on the plumage of the male Argus pheasant; on Urosticte Benjamini; on the nests of birds.

Argynnis, colouring of the lower surface of.

Aricoris epitus, sexual differences in the wings of.

Aristocracy, increased beauty of the.

Arms, proportions of, in soldiers and sailors; direction of the hair on the.

Arms and hands, free use of, indirectly correlated with diminution of canines.

Arrest of development.

Arrow-heads, stone, general resemblance of.

Arrows, use of.

Arteries, variations in the course of the.

Artery, effect of tying, upon the lateral channels.

Arthropoda.

Arts practised by savages.

Ascension, coloured incrustation on the rocks of.

Ascidia, affinity of the lancelet to; tad-pole like larvae of.

Ascidians, bright colours of some.

Asinus, Asiatic and African species of.

Asinus taeniopus.

Ass, colour-variations of the.

Ateles, effects of brandy on an; absence of the thumb in.

Ateles beelzebuth, ears of.

Ateles marginatus, colour of the ruff of; hair on the head of.

Ateuchus cicatricosus, habits of.

Ateuchus, stridulation of.

Athalia, proportions of the sexes in.

Atropus pulsatorius.

Attention, manifestations of, in animals.

Audouin, V., on a hymenopterous parasite with a sedentary male.

Audubon, J.J., on the pinioned goose; on the speculum of Mergus cucullatus; on the pugnacity of male birds; on courtship of Caprimulgus; on Tetrao cupido; on Ardea nycticorax; on Sturnella ludoviciana; on the vocal organs of Tetra cupido; on the drumming of the male Tetrao umbellus; on sounds produced by the nightjar; on Ardea herodias and Cathartes jota; on Mimus polyglottus; on display in male birds; on the spring change of colour in some finches; on migration of mocking thrushes; recognition of a dog by a turkey; selection of mate by female birds; on the turkey; on variation in the male scarlet tanager; on the musk-rat; on the habits of Pyranga aestiva; on local differences in the nests of the same species of birds; on the habits of woodpeckers; on Bombycilla carolinensis; on young females of Pyranga aestiva acquiring male characters; on the immature plumage of thrushes; on the immature plumage of birds; on birds breeding in immature plumage; on the growth of the crest and plume in the male Ardea ludoviciana; on the change of colour in some species of Ardea.

Audobon and Bachman, MM., on squirrels fighting; on the Canadian lynx.

Aughey, Prof., on rattlesnakes.

Austen, N.L., on Anolis cristatellus.

Australia, not the birthplace of man; half-castes killed by the natives of; lice of the natives of.

Australia, South, variation in the skulls of aborigines of.

Australians, colour of new-born children of; relative height of the sexes of; women a cause of war among the.

Axis deer, sexual difference in the colour of the.

Aymaras, measurements of the; no grey hair among the; hairlessness of the face in the; long hair of the.

Azara, on the proportion of men and women among the Guaranys; on Palamedea cornuta; on the beards of the Guaranys; on strife for women among the Guanas; on infanticide; on the eradication of the eyebrows and eyelashes by the Indians of Paraguay; on polyandry among the Guanas; celibacy unknown among the savages of South America; on the freedom of divorce among the Charruas.

Babbage C., on the greater proportion of illegitimate female births.

Babirusa, tusks of the.

Baboon, revenge in a; rage excited in, by reading; manifestation of memory by a; employing a mat for shelter against the sun; protected from punishment by its companions.

Baboon, Cape, mane of the male; Hamadryas, mane of the male.

Baboon, effects of intoxicating liquors on; ears of; diversity of the mental faculties in; hands of; habits of; variability of the tail in; manifestation of maternal affection by; using stones and sticks as weapons; co-operation of; silence of, on plundering expeditions; apparent polygamy of; polygamous and social habits of.

Baboons, courtship of.

Bachman, Dr., on the fertility of mulattoes.

Baer, K.E. von, on embryonic development; definition of advancement in the organic scale.

Bagehot, W., on the social virtues among primitive men; slavery formerly beneficial; on the value of obedience; on human progress; on the persistence of savage tribes in classical times.

Bailly, E.M., on the mode of fighting of the Italian buffalo; on the fighting of stags.

Bain, A., on the sense of duty; aid springing from sympathy; on the basis of sympathy; on the love of approbation etc.; on the idea of beauty.

Baird, W., on a difference in colour between the males and females of some Entozoa.

Baker, Mr., observation on the proportion of the sexes in pheasant-chicks.

Baker, Sir S., on the fondness of the Arabs for discordant music; on sexual difference in the colours of an antelope; on the elephant and rhinoceros attacking white or grey horses; on the disfigurements practised by the negroes; on the gashing of the cheeks and temples practised in Arab countries; on the coiffure of the North Africans; on the perforation of the lower lip by the women of Latooka; on the distinctive characters of the coiffure of central African tribes; on the coiffure of Arab women.

“Balz” of the Black-cock.

Bantam, Sebright.

Banteng, horns of; sexual differences in the colours of the.

Banyai, colour of the.

Barbarism, primitive, of civilised nations.

Barbs, filamentous, of the feathers, in certain birds.

Barr, Mr., on sexual preference in dogs.

Barrago, F., on the Simian resemblances of man.

Barrington, Daines, on the language of birds; on the clucking of the hen; on the object of the song of birds; on the singing of female birds; on birds acquiring the songs of other birds; on the muscles of the larynx in song-birds; on the want of the power of song by female birds.

Barrow, on the widow-bird.

Bartels, Dr., supernumerary mammae in men.

Bartlett, A.D., period of hatching of bird’s eggs; on the tragopan; on the development of the spurs in Crossoptilon auritum; on the fighting of the males of Plectopterus gambensis; on the Knot; on display in male birds; on the display of plumage by the male Polyplectron; on Crossoptilon auritum and Phasianus Wallichii; on the habits of Lophophorus; on the colour of the mouth in Buceros bicornis; on the incubation of the cassowary; on the Cape Buffalo; on the use of the horns of antelopes; on the fighting of male wart-hogs; on Ammotragus tragelaphus; on the colours of Cercopithecus cephus; on the colours of the faces of monkeys; on the naked surfaces of monkeys.

Bartlett, on courting of Argus pheasant.

Bartram, on the courtship of the male alligator.

Basque language, highly artificial.

Bate, C.S., on the superior activity of male crustacea; on the proportions of the sexes in crabs; on the chelae of crustacea; on the relative size of the sexes in crustacea; on the colours of crustacea.

Bateman, Dr., tendency to imitation in certain diseased states; on Aphasia.

Bates, H.W., on variation in the form of the head of Amazonian Indians; on the proportion of the sexes among Amazonian butterflies; on sexual differences in the wings of butterflies; on the field-cricket; on Pyrodes pulcherrimus; on the horns of Lamellicorn beetles; on the colours of Epicaliae, etc.; on the coloration of tropical butterflies; on the variability of Papilio Sesostris and Childrenae; on male and female butterflies inhabiting different stations; on mimicry; on the caterpillar of a Sphinx; on the vocal organs of the umbrella-bird; on the toucans; on Brackyurus calvus.

Batokas, knocking out two upper incisors.

Batrachia, eagerness of male.

Bats, scent-glands; sexual differences in the colour of; fur of male frugivorous.

Battle, law of; among beetles; among birds; among mammals; in man.

Beak, sexual difference in the forms of the; in the colour of the.

Beaks, of birds, bright colours of.

Beard, development of, in man; analogy of the, in man and the quadrumana; variation of the development of the, in different races of men; estimation of, among bearded nations; probable origin of the.

Beard, in monkeys; of mammals.

Beautiful, taste for the, in birds; in the quadrumana.

Beauty, sense of, in animals; appreciation of, by birds; influence of; variability of the standard of.

Beauty, sense of, sufficiently permanent for action of sexual selection.

Beaven, Lieut., on the development of the horns in Cervus Eldi.

Beaver, instinct and intelligence of the; voice of the; castoreum of the.

Beavers, battles of male.

Bechstein, on female birds choosing the best singers among the males; on rivalry in song-birds; on the singing of female birds; on birds acquiring the songs of other birds; on pairing the canary and siskin; on a sub-variety of the monk pigeon; on spurred hens.

Beddoe, Dr., on causes of difference in stature.

Bee-eater.

Bees, pollen-baskets and stings of; destruction of drones and queens by; female, secondary sexual characters of; proportion of sexes; difference of the sexes in colour and sexual selection.

Beetle, luminous larva of a.

Beetles, size of the cerebral ganglia in; dilatation of the foretarsi in male; blind; stridulation of.

Belgium, ancient inhabitants of.

Bell, Sir C., on emotional muscles in man; “snarling muscles;” on the hand.

Bell, T., on the numerical proportion of the sexes in moles; on the newts; on the croaking of the frog; on the difference in the coloration of the sexes in Zootoca vivipara; on moles fighting.

Bell-bird, sexual difference in the colour of the.

Bell-birds, colours of.

Belt, Mr., on the nakedness of tropical mankind; on a spider-monkey and eagle; habits of ants; Lampridae distasteful to mammals; mimicry of Leptalides; colours of Nicaraguan frogs; display of humming-birds; on the toucans; protective colouring of skunk.

Benevolence, manifested by birds.

Bennett, A.W., attachment of mated birds; on the habits of Dromaeus irroratus.

Bennett, Dr., on birds of paradise.

Berbers, fertility of crosses with other races.

Bernicla antarctica, colours of.

Bernicle gander pairing with a Canada goose.

Bert, M., crustaceans distinguish colours.

Bettoni, E., on local differences in the nests of Italian birds.

Beyle, M., see Bombet.

Bhoteas, colour of the beard in.

Bhringa, disc-formed tail-feathers of.

Bianconi, Prof., on structures as explained through mechanical principles.

Bibio, sexual differences in the genus.

Bichat, on beauty.

Bickes, proportion of sexes in man.

Bile, coloured, in many animals.

Bimana.

Birds, imitations of the songs of other birds by; dreaming; killed by telegraph wires; language of; sense of beauty in; pleasure of, in incubation; male, incubation by; and reptiles, alliance of; sexual differences in the beak of some; migratory, arrival of the male before the female; apparent relation between polygamy and marked sexual differences in; monogamous, becoming polygamous under domestication; eagerness of male in pursuit of the female; wild, numerical proportion of the sexes in; secondary sexual characters of; difference of size in the sexes of; fights of male, witnessed by females; display of male, to captivate the females; close attention of, to the songs of others; acquiring the song of their foster-parents; brilliant, rarely good songsters; love-antics and dances of; coloration of; moulting of; unpaired; male, singing out of season; mutual affection of; in confinement, distinguish persons; hybrid, production of; Albino; European, number of species of; variability of; geographical distribution of colouring; gradation of secondary sexual characters in; obscurely coloured, building concealed nests; young female, acquiring male characters; breeding in immature plumage; moulting of; aquatic, frequency of white plumage in; vocal courtship of; naked skin of the head and neck in.

Birgus latro, habits of.

Birkbeck, Mr., on the finding of new mates by golden eagles.

Birthplace of man.

Births, numerical proportions of the sexes in, in animals and man; male and female, numerical proportion of, in England.

Bischoff, Prof., on the agreement between the brains of man and of the orang; figure of the embryo of the dog; on the convolutions of the brain in the human foetus; on the difference between the skulls of man and the quadrumana; resemblance between the ape’s and man’s.

Bishop, J., on the vocal organs of frogs; on the vocal organs of cervine birds; on the trachea of the Merganser.

Bison, American, co-operation of; mane of the male.

Bitterns, dwarf, coloration of the sexes of.

Biziura lobata, musky odour of the male; large size of male.

Blackbird, sexual differences in the; proportion of the sexes in the; acquisition of a song by; colour of the beak in the sexes of the; pairing with a thrush; colours and nidification of the; young of the; sexual difference in coloration of the.

Black-buck, Indian, sexual difference in the colour of the.

Blackcap, arrival of the male, before the female; young of the.

Black-cock, polygamous; proportion of the sexes in the; pugnacity and love-dance of the; call of the; moulting of the; duration of the courtship of the; and pheasant, hybrids of; sexual difference in coloration of the; crimson eye-cere of the.

Black-grouse, characters of young.

Blacklock, Dr., on music.

Blackwall, J., on the speaking of the magpie; on the desertion of their young by swallows; on the superior activity of male spiders; on the proportion of the sexes in spiders; on sexual variation of colour in spiders; on male spiders.

Bladder-nose Seal, hood of the.

Blaine, on the affections of dogs.

Blair, Dr., on the relative liability of Europeans to yellow fever.

Blake, C.C., on the jaw from La Naulette.

Blakiston, Captain, on the American snipe; on the dances of Tetrao phasianellus.

Blasius, Dr., on the species of European birds.

Bledius taurus, hornlike processes of male.

Bleeding, tendency to profuse.

Blenkiron, Mr., on sexual preference in horses.

Blennies, crest developed on the head of male, during the breeding season.

Blethisa multipunctata, stridulation of.

Bloch, on the proportions of the sexes in fishes.

Blood, arterial, red colour of.

Blood pheasant, number of spurs in.

Blow-fly, sounds made by.

Bluebreast, red-throated, sexual differences of the.

Blumenbach, on Man; on the large size of the nasal cavities in American aborigines; on the position of man; on the number of species of man.

Blyth, E., on the structure of the hand in the species of Hylobates; observations on Indian crows; on the development of the horns in the Koodoo and Eland antelopes; on the pugnacity of the males of Gallicrex cristatus; on the presence of spurs in the female Euplocamus erythrophthalmus; on the pugnacity of the amadavat; on the spoonbill; on the moulting of Anthus; on the moulting of bustards, plovers, and Gallus bankiva; on the Indian honey-buzzard; on sexual differences in the colour of the eyes of hornbills; on Oriolus melanocephalus; on Palaeornis javanicus; on the genus Ardetta; on the peregrine falcon; on young female birds acquiring male characters; on the immature plumage of birds; on representative species of birds; on the young of Turnix; on anomalous young of Lanius rufus and Colymbus glacialis; on the sexes and young of the sparrows; on dimorphism in some herons; on the ascertainment of the sex of nestling bullfinches by pulling out breast-feathers; on orioles breeding in immature plumage; on the sexes and young of Buphus and Anastomus; on the young of the blackcap and blackbird; on the young of the stonechat; on the white plumage of Anastomus; on the horns of Bovine animals; on the horns of Antilope bezoartica; on the mode of fighting of Ovis cycloceros; on the voice of the Gibbons; on the crest of the male wild goat; on the colours of Portax picta; on the colours of Antilope bezoartica; on the colour of the Axis deer; on sexual difference of colour in Hylobates hoolock; on the hog-deer; on the beard and whiskers in a monkey, becoming white with age.

Boar, wild, polygamous in India; use of the tusks by the; fighting of.

Boardman, Mr., Albino birds in U.S.

Boitard and Corbie, MM., on the transmission of sexual peculiarities in pigeons; on the antipathy shewn by some female pigeons to certain males.

Bold, Mr., on the singing of a sterile hybrid canary.

Bombet, on the variability of the standard of beauty in Europe.

Bombus, difference of the sexes in.

Bombycidae, coloration of; pairing of the; colours of.

Bombycilla carolinensis, red appendages of.

Bombyx cynthia, proportion of the sexes in; pairing of.

Bombyx mori, difference of size of the male and female cocoons of; pairing of.

Bombyx Pernyi, proportion of sexes of.

Bombyx Yamamai, M. Personnat on; proportion of sexes of.

Bonaparte, C.L., on the call-notes of the wild turkey.

Bond, F., on the finding of new mates by crows.

Bone, implements of, skill displayed in making.

Boner, C., on the transfer of male characters to an old female chamois; on the habits of stags; on the pairing of red deer.

Bones, increase of, in length and thickness, when carrying a greater weight.

Bonizzi, P., difference of colour in sexes of pigeons.

Bonnet monkey.

Bonwick, J., extinction of Tasmanians.

Boomerang.

Boreus hyemalis, scarcity of the male.

Bory St. Vincent, on the number of species of man; on the colours of Labrus pavo.

Bos etruscus.

Bos gaurus, horns of.

Bos moschatus.

Bos primigenius.

Bos sondaicus, horns of, colours of.

Botocudos, mode of life of; disfigurement of the ears and lower lip of the.

Boucher de Perthes, J.C. de, on the antiquity of man.

Bourbon, proportion of the sexes in a species of Papilio from.

Bourien on the marriage-customs of the savages of the Malay Archipelago.

Bovidae, dewlaps of.

Bower-birds, habits of the; ornamented playing-places of.

Bows, use of.

Brachycephalic structure, possible explanation of.

Brachyura.

Brachyurus calvus, scarlet face of.

Bradley, Mr., abductor ossis metatarsi quinti in man.

Brain, of man, agreement of the, with that of lower animals; convolutions of, in the human foetus; influence of development of mental faculties upon the size of the; influence of the development of on the spinal column and skull; larger in some existing mammals than in their tertiary prototypes; relation of the development of the, to the progress of language; disease of the, affecting speech; difference in the convolutions of, in different races of men; supplement on, by Prof. Huxley; development of the gyri and sulci.

Brakenridge, Dr., on the influence of climate.

Brandt, A., on hairy men.

Braubach, Prof., on the quasi-religious feeling of a dog towards his master; on the self-restraint of dogs.

Brauer, F., on dimorphism in Neurothemis.

Brazil, skulls found in caves of; population of; compression of the nose by the natives of.

Break between man and the apes.

Bream, proportion of the sexes in the.

Breeding, age of, in birds.

Breeding season, sexual characters making their appearance in the, in birds.

Brehm, on the effects of intoxicating liquors on monkeys; on the recognition of women by male Cynocephali; on the diversity of the mental faculties of monkeys; on the habits of baboons; on revenge taken by monkeys; on manifestations of maternal affection by monkeys and baboons; on the instinctive dread of monkeys for serpents; on the use of stones as missiles by baboons; on a baboon using a mat for shelter from the sun; on the signal-cries of monkeys; on sentinels posted by monkeys; on co-operation of animals; on an eagle attacking a young Cercopithecus; on baboons in confinement protecting one of their number from punishment; on the habits of baboons when plundering; on polygamy in Cynocephalus and Cebus; on the numerical proportion of the sexes in birds; on the love-dance of the blackcock; Palamedea cornuta; on the habits of the Black-grouse; on sounds produced by birds of paradise; on assemblages of grouse; on the finding of new mates by birds; on the fighting of wild boars; on sexual differences in Mycetes; on the habits of Cynocephalus hamadryas.

Brent, Mr., on the courtship of fowls.

Breslau, numerical proportion of male and female births in.

Bridgeman, Laura.

Brimstone butterfly, sexual difference of colour in the.

British, ancient, tattooing practised by.

Broca, Prof., on the occurrence of the supra-condyloid foramen in the human humerus; anthropomorphous apes more bipedal than quadrupedal; on the capacity of Parisian skulls at different periods; comparison of modern and mediaeval skulls; on tails of quadrupeds; on the influence of natural selection; on hybridity in man; on human remains from Les Eyzies; on the cause of the difference between Europeans and Hindoos.

Brodie, Sir B., on the origin of the moral sense in man.

Bronn, H.G., on the copulation of insects of distinct species.

Bronze period, men of, in Europe.

Brown, R., sentinels of seals generally females; on the battles of seals; on the narwhal; on the occasional absence of the tusks in the female walrus; on the bladder-nose seal; on the colours of the sexes in Phoca Groenlandica; on the appreciation of music by seals; on plants used as love-philters, by North American women.

Browne, Dr. Crichton, injury to infants during parturition.

Brown-Sequard, Dr., on the inheritance of the effects of operations by guinea-pig.

Bruce, on the use of the elephant’s tusks.

Brulerie, P. de la, on the habits of Ateuchus cicatricosus; on the stridulation of Ateuchus.

Brunnich, on the pied ravens of the Feroe islands.

Bryant, Dr., preference of tame pigeon for wild mate.

Bryant, Captain, on the courtship of Callorhinus ursinus.

Bubas bison, thoracic projection of.

Bubalus caffer, use of horns.

Bucephalus capensis, difference of the sexes of, in colour.

Buceros, nidification and incubation of.

Buceros bicornis, sexual differences in the colouring of the casque, beak, and mouth in.

Buceros corrugatus, sexual differences in the beak of.

Buchner, L., on the origin of man; on the use of the human foot as a prehensile organ; on the mode of progression of the apes; on want of self-consciousness, etc., in savages.

Bucholz, Dr., quarrels of chamaeleons.

Buckinghamshire, numerical proportion of male and female births in.

Buckland, F., on the numerical proportion of the sexes in rats; on the proportion of the sexes in the trout; on Chimaera monstrosa.

Buckland, W., on the complexity of crinoids.

Buckler, W., proportion of sexes of Lepidoptera reared by.

Bucorax abyssinicus, inflation of the neck-wattle of the male during courtship.

Budytes Raii.

Buffalo, Cape.

Buffalo, Indian, horns of the.

Buffalo, Italian, mode of fighting of the.

Buffon, on the number of species of man.

Bufo sikimmensis.

Bugs.

Buist, R., on the proportion of the sexes in salmon; on the pugnacity of the male salmon.

Bulbul, pugnacity of the male; display of under tail-coverts by the male.

Bull, mode of fighting of the; curled frontal hair of the.

Buller, Dr., on the Huia; the attachment of birds.

Bullfinch, sexual differences in the; piping; female, singing of the; courtship of the; widowed, finding a new mate; attacking a reed-bunting; nestling, sex ascertained by pulling out breast feathers.

Bullfinches, distinguishing persons; rivalry of female.

Bulls, two young, attacking an old one; wild, battles of.

Bull-trout, male, colouring of, during the breeding season.

Bunting, reed, head feathers of the male; attacked by a bullfinch.

Buntings, characters of young.

Buphus coromandus, sexes and young of; change of colour in.

Burchell, Dr., on the zebra; on the extravagance of a Bushwoman in adorning herself; celibacy unknown among the savages of South Africa; on the marriage-customs of the Bushwomen.

Burke, on the number of species of man.

Burmese, colour of the beard in.

Burton, Captain, on negro ideas of female beauty; on a universal ideal of beauty.

Bushmen, marriage among.

Bushwoman, extravagant ornamentation of a.

Bushwomen, hair of; marriage-customs of.

Bustard, throat-pouch of the male; humming noise produced by a male; Indian, ear-tufts of.

Bustards, occurrence of sexual differences and of polygamy among the; love-gestures of the male; double moult in.

Butler, A.G., on sexual differences in the wings of Aricoris epitus; courtship of butterflies; on the colouring of the sexes in species of Thecla; on the resemblance of Iphias glaucippe to a leaf; on the rejection of certain moths and caterpillars by lizards and frogs.

Butterfly, noise produced by a; Emperor; meadow brown, instability of the ocellated spots of.

Butterflies, proportion of the sexes in; forelegs atrophied in some males; sexual difference in the neuration of the wings of; pugnacity of male; protective resemblances of the lower surface of; display of the wings by; white, alighting upon bits of paper; attracted by a dead specimen of the same species; courtship of; male and female, inhabiting different stations.

Buxton, C., observations on macaws; on an instance of benevolence in a parrot.

Buzzard, Indian honey-; variation in the crest of.

Cabbage butterflies.

Cachalot, large head of the male.

Cadences, musical, perception of, by animals.

Caecum, large, in the early progenitors of man.

Cairina moschata, pugnacity of the male.

Californian Indians, decrease of.

Callianassa, chelae of, figured.

Callidryas, colours of sexes.

Callionymus lyra, characters of the male.

Callorhinus ursinus, relative size of the sexes of; courtship of.

Calotes maria.

Calotes nigrilabris, sexual difference in the colour of.

Cambridge, O. Pickard, on the sexes of spiders; on the size of male Nephila.

Camel, canine teeth of male.

Campbell, J., on the Indian elephant; on the proportion of male and female births in the harems of Siam.

Campylopterus hemileucurus.

Canaries distinguishing persons.

Canary, polygamy of the; change of plumage in, after moulting; female, selecting the best singing male; sterile hybrid, singing of a; female, singing of the; selecting a greenfinch; and siskin, pairing of.

Cancer pagurus.

Canestrini, G., on rudimentary characters and the origin of man; on rudimentary characters; on the movement of the ear in man; of the variability of the vermiform appendage in man; on the abnormal division of the malar bone in man; on abnormal conditions of the human uterus; on the persistence of the frontal suture in man; on the proportion of the sexes in silk-moths; secondary sexual characters of spiders.

Canfield, Dr., on the horns of the Antilocapra.

Canine teeth in man, diminution of, in man; diminution of, in horses; disappearance of, in male ruminants; large in the early progenitors of man.

Canines, and horns, inverse development of.

Canoes, use of.

Cantharis, difference of colour in the sexes of a species of.

Cantharus lineatus.

Capercailzie, polygamous; proportion of the sexes in the; pugnacity of the male; pairing of the; autumn meetings of the; call of the; duration of the courtship of; behaviour of the female; inconvenience of black colour to the female; sexual difference in the coloration of the; crimson eye-cere of the male.

Capitonidae, colours and nidification of the.

Capra aegagrus, crest of the male; sexual difference in the colour of.

Capreolus Sibiricus subecaudatus.

Caprice, common to man and animals.

Caprimulgus, noise made by the males of some species of, with their wings.

Caprimulgus virginianus, pairing of.

Carabidae.

Carbonnier, on the natural history of the pike; on the relative size of the sexes in fishes; courtship of Chinese Macropus.

Carcineutes, sexual difference of colour in.

Carcinus moenas.

Cardinalis virginianus.

Carduelis elegans, sexual differences of the beak in.

Carnivora, marine, polygamous habits of; sexual differences in the colours of.

Carp, numerical proportion of the sexes in the.

Carr, R., on the peewit.

Carrier pigeon, late development of the wattle in the.

Carrion beetles, stridulation of.

Carrion-hawk, bright coloured female of.

Carus, Prof. V., on the development of the horns in merino sheep; on antlers of red deer.

Cassowary, sexes and incubation of the.

Castnia, mode of holding wings.

Castoreum.

Castration, effects of.

Casuarius galeatus.

Cat, convoluted body in the extremity of the tail of a; sick, sympathy of a dog with a.

Cataract in Cebus Azarae.

Catarrh, liability of Cebus Azarae to.

Catarrhine monkeys.

Caterpillars, bright colours of.

Cathartes aura.

Cathartes jota, love-gestures of the male.

Catlin, G., correlation of colour and texture of hair in the Mandans; on the development of the beard among the North American Indians; on the great length of the hair in some North American tribes.

Caton, J.D., on the development of the horns in Cervus virginianus and strongyloceros; on the wild turkey; on the presence of traces of horns in the female wapiti; on the fighting of deer; on the crest of the male wapiti; on the colours of the Virginian deer; on sexual differences of colour in the wapiti; on the spots of the Virginian deer.

Cats, dreaming; tortoise-shell; enticed by valerian; colours of.

Cattle, rapid increase of, in South America; domestic, lighter in winter in Siberia; horns of; domestic, sexual differences of, late developed; numerical proportion of the sexes in.

Caudal vertebrae, number of, in macaques and baboons; basal, of monkeys, imbedded in the body.

Cavolini, observations on Serranus.

Cebus, maternal affection in a; gradation of species of.

Cebus Apella.

Cebus Azarae, liability of, to the same diseases as man; distinct sounds produced by; early maturity of the female.

Cebus capucinus, polygamous; sexual differences of colour in; hair on the head of.

Cebus vellerosus, hair on the head of.

Cecidomyiidae, proportions of the sexes in.

Celibacy, unknown among the savages of South Africa and South America.

Centipedes.

Cephalopoda, absence of secondary sexual characters in.

Cephalopterus ornatus.

Cephalopterus penduliger.

Cerambyx heros, stridulant organ of.

Ceratodus, paddle of.

Ceratophora aspera, nasal appendages of.

Ceratophora Stoddartii, nasal horn of.

Cerceris, habits of.

Cercocebus aethiops, whiskers, etc., of.

Cercopithecus, young, seized by an eagle and rescued by the troop; definition of species of.

Cercopithecus cephus, sexual difference of colour in.

Cercopithecus cynosurus and griseo-viridis, colour of the scrotum in.

Cercopithecus Diana, sexual differences of colour in.

Cercopithecus griseo-viridis.

Cercopithecus petaurista, whiskers, etc., of.

Ceres, of birds, bright colours of.

Ceriornis Temminckii, swelling of the wattles of the male during courtship.

Cervulus, weapons of.

Cervulus moschatus, rudimentary horns of the female.

Cervus alces.

Cervus campestris, odour of.

Cervus canadensis, traces of horns in the female; attacking a man; sexual difference in the colour of.

Cervus elaphus, battles of male; horns of, with numerous points; long hairs on the throat of.

Cervus Eldi.

Cervus mantchuricus.

Cervus paludosus, colours of.

Cervus strongyloceros.

Cervus virginianus, horns of, in course of modification.

Ceryle, male black-belted in some species of.

Cetacea, nakedness of.

Ceylon, frequent absence of beard in the natives of.

Chaffinch, proportion of the sexes in the; courtship of the.

Chaffinches, new mates found by.

Chalcophaps indicus, characters of young.

Chalcosoma atlas, sexual differences of.

Chamaeleo, sexual differences in the genus; combats of.

Chamaeleo bifurcus.

Chamaeleo Owenii.

Chamaeleo pumilus.

Chamaepetes unicolor, modified wing-feather in the male.

Chameleons.

Chamois, danger-signals of; transfer of male characters to an old female.

Champneys, Mr., acromio-basilar muscle and quadrupedal gait.

Chapman, Dr., on stridulation in Scolytus.

Chapuis, Dr., on the transmission of sexual peculiarities in pigeons; on streaked Belgian pigeons.

Char, male, colouring of, during the breeding season.

Characters, male, developed in females; secondary sexual, transmitted through both sexes; natural, artificial, exaggeration of, by man.

Charadrus hiaticula and pluvialis, sexes and young of.

Chardin on the Persians.

Charms, worn by women.

Charruas, freedom of divorce among the.

Chasmorhynchus, difference of colour in the sexes of; colours of.

Chasmorhynchus niveus.

Chasmorhynchus nudicollis.

Chasmorhynchus tricarunculatus.

Chastity, early estimation of.

Chatterers, sexual differences in.

Cheever, Rev. H.T., census of the Sandwich Islands.

Cheiroptera, absence of secondary sexual characters in.

Chelae of crustacea.

Chelonia, sexual differences in.

Chenalopex aegyptiacus, wing-knobs of.

Chera progne.

Chest, proportions of, in soldiers and sailors; large, of the Quechua and Aymara Indians.

Chevrotains, canine teeth of.

Chiasognathus, stridulation of.

Chiasognathus Grantii, mandibles of.

Children, legitimate and illegitimate, proportion of the sexes in.

Chiloe, lice of the natives of; population of.

Chimaera monstrosa, bony process on the head of the male.

Chimaeroid fishes, prehensile organs of male.

Chimpanzee, ears of the; representatives of the eyebrows in the; hands of the; absence of mastoid processes in the; platforms built by the; cracking nuts with a stone; direction of the hair on the arms of the; supposed evolution of the; polygamous and social habits of the.

China, North, idea of female beauty in.

China, Southern, inhabitants of.

Chinese, use of flint tools by the; difficulty of distinguishing the races of the; colour of the beard in; general beardlessness of the; opinions of the, on the appearance of Europeans and Cingalese; compression of the feet of.

Chinsurdi, his opinion of beards.

Chlamydera maculata.

Chloeon, pedunculated eyes of the male of.

Chloephaga, coloration of the sexes in.

Chlorocoelus Tanana.

Chorda dorsalis.

Chough, red beak of the.

Chromidae, frontal protuberance in male; sexual differences in colour of.

Chrysemys picta, long claws of the male.

Chrysococcyx, characters of young of.

Chrysomelidae, stridulation of.

Cicada pruinosa.

Cicada septendecim.

Cicadae, songs of the; rudimentary sound-organs in females of.

Cicatrix of a burn, causing modification of the facial bones.

Cichla, frontal protuberance of male.

Cimetiere du Sud, Paris.

Cincloramphus cruralis, large size of male.

Cinclus aquaticus.

Cingalese, Chinese opinion of the appearance of the.

Cirripedes, complemental males of.

Civilisation, effects of, upon natural selection; influence of, in the competition of nations.

Clanging of geese, etc.

Claparede, E., on natural selection applied to man.

Clarke, on the marriage-customs of Kalmucks.

Classification.

Claus, C., on the sexes of Saphirina.

Cleft-palate, inherited.

Climacteris erythrops, sexes of.

Climate, cool, favourable to human progress; power of supporting extremes of, by man; want of connexion of, with colour; direct action of, on colours of birds.

Cloaca, existence of a, in the early progenitors of man.

Cloacal passage existing in the human embryo.

Chapter XXI: General a Summary and Conclusion (part 2)

Clubs, used as weapons before dispersion of mankind.

Clucking of fowls.

Clythra 4-punctata, stridulation of.

Coan, Mr., Sandwich-islanders.

Cobbe, Miss, on morality in hypothetical bee-community.

Cobra, ingenuity of a.

Coccus.

Coccyx, in the human embryo; convoluted body at the extremity of the; imbedded in the body.

Cochin-China, notions of beauty of the inhabitants of.

Cock, blind, fed by its companion; game, killing a kite; comb and wattles of the; preference shewn by the, for young hens; game, transparent zone in the hackles of a.

Cock of the rock.

Cockatoos, nestling; black, immature plumage of.

Coelenterata, absence of secondary sexual characters in.

Coffee, fondness of monkeys for.

Cold, supposed effects of; power of supporting, by man.

Coleoptera, stridulation of; stridulant organs of, discussed.

Colias edusa and hyale.

Collingwood, C., on the pugnacity of the butterflies of Borneo; on butterflies being attracted by a dead specimen of the same species.

Colobus, absence of the thumb.

Colombia, flattened heads of savages of.

Colonists, success of the English as.

Coloration, protective, in birds.

Colour, supposed to be dependent on light and heat; correlation of, with immunity from certain poisons and parasites; purpose of, in lepidoptera; relation of, to sexual functions, in fishes; difference of, in the sexes of snakes; sexual differences of, in lizards; influence of, in the pairing of birds of different species; relation of, to nidification; sexual differences of, in mammals; recognition of, by quadrupeds; of children, in different races of man; of the skin in man.

Colours, admired alike by man and animals; bright, due to sexual selection; bright, among the lower animals; bright, protective to butterflies and moths; bright, in male fishes; transmission of, in birds.

Colquhoun, example of reasoning in a retriever.

Columba passerina, young of.

Colymbus glacialis, anomalous young of.

Comb, development of, in fowls.

Combs and wattles in male birds.

Community, preservation of variations useful to the, by natural selection.

Complexion, different in men and women, in an African tribe.

Compositae, gradation of species among the.

Comte, C., on the expression of the ideal of beauty by sculpture.

Conditions of life, action of changed, upon man; influence of, on plumage of birds.

Condor, eyes and comb of the.

Conjugations, origin of.

Conscience, absence of, in some criminals.

Constitution, difference of, in different races of men.

Consumption, liability of Cebus Azarae to; connection between complexion and.

Convergence of characters.

Cooing of pigeons and doves.

Cook, Captain, on the nobles of the Sandwich Islands.

Cope, E.D., on the Dinosauria.

Cophotis ceylanica, sexual differences of.

Copris.

Copris Isidis, sexual differences of.

Copris lunaris, stridulation of.

Corals, bright colours of.

Coral-snakes.

Cordylus, sexual difference of colour in a species of.

Corfu, habits of the Chaffinch in.

Cornelius, on the proportions of the sexes in Lucanus Cervus.

Corpora Wolffiana, agreement of, with the kidneys of fishes.

Correlated variation.

Correlation, influence of, in the production of races.

Corse, on the mode of fighting of the elephant.

Corvus corone.

Corvus graculus, red beak of.

Corvus pica, nuptial assembly of.

Corydalis cornutus, large jaws of the male.

Cosmetornis.

Cosmetornis vexillarius, elongation of wing-feathers in.

Cotingidae, sexual differences in; coloration of the sexes of; resemblance of the females of distinct species of.

Cottus scorpius, sexual differences in.

Coulter, Dr., on the Californian Indians.

Counting, origin of; limited power of, in primeval man.

Courage, variability of, in the same species; universal high appreciation of; importance of; characteristic of men.

Courtship, greater eagerness of males in; of fishes; of birds.

Cow, winter change of colour.

Crab, devil.

Crab, shore, habits of.

Crabro cribrarius, dilated tibiae of the male.

Crabs, proportions of the sexes in.

Cranz, on the inheritance of dexterity in seal-catching.

Crawfurd, on the number of species of man.

Crenilabrus massa and C. melops, nests, built by.

Crest, origin of, in Polish fowls.

Crests, of birds, difference of, in the sexes; dorsal hairy, of mammals.

Cricket, field-, stridulation of the; pugnacity of male.

Cricket, house-, stridulation of the.

Crickets, sexual differences in.

Crinoids, complexity of.

Crioceridae, stridulation of the.

Croaking of frogs.

Crocodiles, musky odour of, during the breeding season.

Crocodilia.

Crossbills, characters of young.

Crosses in man.

Crossing of races, effects of the.

Crossoptilon auritum, adornment of both sexes of; sexes alike in.

Crotch, G.R., on the stridulation of beetles; on the stridulation of Heliopathes; on the stridulation of Acalles; habit of female deer at breeding time.

Crow, Indians, long hair of the.

Crow, young of the.

Crows, vocal organs of the; living in triplets.

Crows, carrion, new mates found by.

Crows, Indian, feeding their blind companions.

Cruelty of savages to animals.

Crustacea, parasitic, loss of limbs by female; prehensile feet and antennae of; male, more active than female; parthenogenesis in; secondary sexual characters of; amphipod, males sexually mature while young; auditory hairs of.

Crystal worn in the lower lip by some Central African women.

Cuckoo fowls.

Culicidae, attracted by each other’s humming.

Cullen, Dr., on the throat-pouch of the male bustard.

Cultivation of plants, probable origin of.

Cupples, Mr., on the numerical proportion of the sexes in dogs, sheep, and cattle; on the Scotch deerhound; on sexual preference in dogs.

Curculionidae, sexual difference in length of snout in some; hornlike processes in male; musical.

Curiosity, manifestations of, by animals.

Curlews, double moult in.

Cursores, comparative absence of sexual differences among the.

Curtis, J., on the proportion of the sexes in Athalia.

Cuvier, F., on the recognition of women by male quadrumana.

Cuvier, G., on the number of caudal vertebrae in the mandrill; on instinct and intelligence; views of, as to the position of man; on the position of the seals; on Hectocotyle.

Cyanalcyon, sexual difference in colours of; immature plumage of.

Cyanecula suecica, sexual differences of.

Cychrus, sounds produced by.

Cycnia mendica, sexual difference of, in colour.

Cygnus ferus, trachea of.

Cygnus immutabilis.

Cygnus olor, white young of.

Cyllo Leda, instability of the ocellated spots of.

Cynanthus, variation in the genus.

Cynipidae, proportion of the sexes in.

Cynocephalus, difference of the young from the adult; male, recognition of women by; polygamous habits of species of.

Cynocephalus babouin.

Cynocephalus chacma.

Cynocephalus gelada.

Cynocephalus hamadryas, sexual difference of colour in.

Cynocephalus leucophaeus, colours of the sexes of.

Cynocephalus mormon, colours of the male.

Cynocephalus porcarius, mane of the male.

Cynocephalus sphinx.

Cynopithecus niger, ear of.

Cypridina, proportions of the sexes in.

Cyprinidae, proportion of the sexes in the.

Cyprinidae, Indian.

Cyprinodontidae, sexual differences in the.

Cyprinus auratus.

Cypris, relation of the sexes in.

Cyrtodactylus rubidus.

Cystophora cristata, hood of.

Dacelo, sexual difference of colour in.

Dacelo Gaudichaudi, young male of.

Dal-ripa, a kind of ptarmigan.

Damalis albifrons, peculiar markings of.

Damalis pygarga, peculiar markings of.

Dampness of climate, supposed influence of, on the colour of the skin.

Danaidae.

Dances of birds.

Dancing, universality of.

Danger-signals of animals.

Daniell, Dr., his experience of residence in West Africa.

Darfur, protuberances artificially produced by natives of.

Darwin, F., on the stridulation of Dermestes murinus.

Dasychira pudibunda, sexual difference of colour in.

Davis, A.H., on the pugnacity of the male stag-beetle.

Davis, J.B., on the capacity of the skull in various races of men; on the beards of the Polynesians.

Death’s Head Sphinx.

Death-rate higher in towns than in rural districts.

Death-tick.

De Candolle, Alph., on a case of inherited power of moving the scalp.

Declensions, origin of.

Decoration in birds.

Decticus.

Deer, development of the horns in; spots of young; horns of; use of horns of; horns of a, in course of modification; size of the horns of; female, pairing with one male whilst others are fighting for her; male, attracted by the voice of the female; male, odour emitted by.

Deer, Axis, sexual difference in the colour of the.

Deer, fallow, different coloured herds of.

Deer, Mantchurian.

Deer, Virginian, colour of the, not affected by castration; colours of.

Deerhound, Scotch, greater size of the male.

Defensive orders of mammals.

De Geer, C., on a female spider destroying a male.

Dekay, Dr., on the bladder-nose seal.

Delorenzi, G., division of malar bone.

Demerara, yellow fever in.

Dendrocygna.

Dendrophila frontalis, young of.

Denison, Sir W., manner of ridding themselves of vermin among the Australians; extinction of Tasmanians.

Denny, H., on the lice of domestic animals.

Dermestes murinus, stridulation of.

Descent traced through the mother alone.

Deserts, protective colouring of animals inhabiting.

Desmarest, on the absence of suborbital pits in Antilope subgutturosa; on the whiskers of Macacus; on the colour of the opossum; on the colours of the sexes of Mus minutus; on the colouring of the ocelot; on the colours of seals; on Antilope caama; on the colours of goats; on sexual difference of colour in Ateles marginatus; on the mandrill; on Macacus cynomolgus.

Desmoulins, on the number of species of man; on the muskdeer.

Desor, on the imitation of man by monkeys.

Despine, P., on criminals destitute of conscience.

Development, embryonic of man; correlated.

Devil, not believed in by the Fuegians.

Devil-crab.

Devonian, fossil-insect from the.

Dewlaps, of Cattle and antelopes.

Diadema, sexual differences of colouring in the species of.

Diamond-beetles, bright colours of.

Diastema, occurrence of, in man.

Diastylidae, proportion of the sexes in.

Dicrurus, racket-shaped feathers in; nidification of.

Dicrurus macrocercus, change of plumage in.

Didelphis opossum, sexual difference in the colour of.

Differences, comparative, between different species of birds of the same sex.

Digits, supernumerary, more frequent in men than in women; supernumerary, inheritance of; supernumerary, early development of.

Dimorphism, in females of water-beetles; in Neurothemis and Agrion.

Diodorus, on the absence of beard in the natives of Ceylon.

Dipelicus Cantori, sexual differences of.

Diplopoda, prehensile limbs of the male.

Dipsas cynodon, sexual difference in the colour of.

Diptera.

Disease, generated by the contact of distinct peoples.

Diseases, common to man and the lower animals; difference of liability to, in different races of men; new, effects of, upon savages; sexually limited.

Display, coloration of Lepidoptera for; of plumage by male birds.

Distribution, wide, of man; geographical, as evidence of specific distinctness in man.

Disuse, effects of, in producing rudimentary organs; and use of parts, effects of; of parts, influence of, on the races of men.

Divorce, freedom of, among the Charruas.

Dixon, E.S., on the pairing of different species of geese; on the courtship of peafowl.

Dobrizhoffer, on the marriage-customs of the Abipones.

Dobson, Dr., on the Cheiroptera; scent-glands of bats; frugivorous bats.

Dogs, suffering from tertian ague; memory of; dreaming; diverging when drawing sledges over thin ice; exercise of reasoning faculties by; domestic, progress of, in moral qualities; distinct tones uttered by; parallelism between his affection for his master and religious feeling; sociability of the; sympathy of, with a sick cat; sympathy of, with his master; their possession of conscience; possible use of the hair on the fore-legs of the; races of the; numerical proportion of male and female births in; sexual affection between individuals of; howling at certain notes; rolling in carrion.

Dolichocephalic structure, possible cause of.

Dolphins, nakedness of.

Domestic animals, races of; change of breeds of.

Domestication, influence of, in removing the sterility of hybrids.

D’Orbigny, A., on the influence of dampness and dryness on the colour of the skin; on the Yuracaras.

Dotterel.

Doubleday, E., on sexual differences in the wings of butterflies.

Doubleday, H., on the proportion of the sexes in the smaller moths; males of Lasiocampa quercus and on the attraction of the Saturnia carpini by the female; on the proportion of the sexes in the Lepidoptera; on the ticking of Anobium tesselatum; on the structure of Ageronia feronia; on white butterflies alighting upon paper.

Douglas, J.W., on the sexual differences of the Hemiptera; colours of British Homoptera.

Down, of birds.

Draco, gular appendages of.

Dragonet, Gemmeous.

Dragon-flies, caudal appendages of male; relative size of the sexes of; difference in the sexes of; want of pugnacity by the male.

Drake, breeding plumage of the.

Dreams, possible source of the belief in spiritual agencies.

Drill, sexual difference of colour in the.

Dromaeus irroratus.

Dromolaea, Saharan species of.

Drongo shrike.

Drongos, racket-shaped feathers in the tails of.

Dryness of climate, supposed influence of, on the colour of the skin.

Dryopithecus.

Duck, harlequin, age of mature plumage in the; breeding in immature plumage.

Duck, long-tailed, preference of male, for certain females.

Duck, pintail, pairing with a widgeon.

Duck, voice of the; pairing with a shield-drake; immature plumage of the.

Duck, wild, sexual differences in the; speculum and male characters of; pairing with a pin-tail drake.

Ducks, wild, becoming polygamous under partial domestication; dogs and cats recognised by.

Dufosse, Dr., sounds produced by fish.

Dugong, nakedness of; tusks of.

Dujardin, on the relative size of the cerebral ganglia, in insects.

Duncan, Dr., on the fertility of early marriages; comparative health of married and single.

Dupont, M., on the occurrence of the supra-condyloid foramen in the humerus of man.

Durand, J.P., on causes of variation.

Dureau de la Malle, on the songs of birds; on the acquisition of an air by blackbirds.

Dutch, retention of their colour by the, in South Africa.

Duty, sense of.

Duvaucel, female Hylobates washing her young.

Dyaks, pride of, in mere homicide.

Dynastes, large size of males of.

Dynastini, stridulation of.

Dytiscus, dimorphism of females of; grooved elytra of the female.

Eagle, young Cercopithecus rescued from, by the troop.

Eagle, white-headed, breeding in immature plumage.

Eagles, golden, new mates found by.

Ear, motion of the; external shell of the, useless in man; rudimentary point of the, in man.

Ears, more variable in men than women; piercing and ornamentation of the.

Earwigs, parental feeling in.

Echidna.

Echini, bright colours of some.

Echinodermata, absence of secondary sexual characters in.

Echis carinata.

Ecker, figure of the human embryo; on the development of the gyri and sulci of the brain; on the sexual differences in the pelvis in man; on the presence of a sagittal crest in Australians.

Edentata, former wide range of, in America; absence of secondary sexual characters in.

Edolius, racket-shaped feathers in.

Edwards, Mr., on the proportion of the sexes in North American species of Papilio.

Eels, hermaphroditism of.

Egerton, Sir P., on the use of the antlers of deer; on the pairing of red deer; on the bellowing of stags.

Eggs, hatched by male fishes.

Egret, Indian, sexes and young of.

Egrets, breeding plumage of; white.

Ehrenberg, on the mane of the male Hamadryas baboon.

Ekstrom, M., on Harelda glacialis.

Elachista rufocinerea, habits of male.

Eland, development of the horns of the.

Elands, sexual differences of colour in.

Elaphomyia, sexual differences in.

Elaphrus uliginosus, stridulation of.

Elaps.

Elateridae, proportion of the sexes in.

Elaters, luminous.

Elephant, rate of increase of the; nakedness of the; using a fan; Indian, forbearance to his keeper; polygamous habits of the; pugnacity of the male; tusks of; Indian, mode of fighting of the; male, odour emitted by the; attacking white or grey horses.

Elevation of abode, modifying influence of.

Elimination of inferior individuals.

Elk, winter change of the.

Elk, Irish, horns of the.

Ellice Islands, beards of the natives.

Elliot, D.G., on Pelecanus erythrorhynchus.

Elliot, R., on the numerical proportion of the sexes in young rats; on the proportion of the sexes in sheep.

Elliot, Sir W., on the polygamous habits of the Indian wild boar.

Ellis, on the prevalence of infanticide in Polynesia.

Elphinstone, Mr., on local difference of stature among the Hindoos; on the difficulty of distinguishing the native races of India.

Elytra, of the females of Dytiscus Acilius, Hydroporus.

Emberiza, characters of young.

Emberiza miliaria.

Emberiza schoeniclus, head-feathers of the male.

Embryo of man; of the dog.

Embryos of mammals, resemblance of the.

Emigration.

Emotions experienced by the lower animals in common with man; manifested by animals.

Emperor butterfly.

Emperor moth.

Emu, sexes and incubation of.

Emulation of singing birds.

Endurance, estimation of.

Energy, a characteristic of men.

England, numerical proportion of male and female births in.

Engleheart, Mr., on the finding of new mates by starlings.

English, success of, as colonists.

Engravers, short-sighted.

Entomostraca.

Entozoa, difference of colour between the males and females of some.

Environment, direct action of the, in causing differences between the sexes.

Envy, persistence of.

Eocene period, possible divergence of men during the.

Eolidae, colours of, produced by the biliary glands.

Epeira nigra, small size of the male of.

Ephemerae.

Ephemeridae.

Ephippiger vitium, stridulating organs of.

Epicalia, sexual differences of colouring in the species of.

Equus hemionus, winter change of.

Erateina, coloration of.

Ercolani, Prof., hermaphroditism in eels.

Erect attitude of man.

Eristalis, courting of.

Eschricht, on the development of hair in man; on a languinous moustache in a female foetus; on the want of definition between the scalp and the forehead in some children; on the arrangement of the hair in the human foetus; on the hairiness of the face in the human foetus of both sexes.

Esmeralda, difference of colour in the sexes of.

Esox lucius.

Esox reticulatus.

Esquimaux, their belief in the inheritance of dexterity in seal-catching; mode of life of.

Estrelda amandava, pugnacity of the male.

Eubagis, sexual differences of colouring in the species of.

Euchirus longimanus, sound produced by.

Eudromias morinellus.

Eulampis jugularis, colours of the female.

Euler, on the rate of increase in the United States.

Eunomota superciliaris, racket-shaped feathers in the tail of.

Eupetomena macroura, colours of the female.

Euphema splendida.

Euplocamus erythrophthalmus, possession of spurs by the female.

Europe, ancient inhabitants of.

Europeans, difference of, from Hindoos; hairiness of, probably due to reversion.

Eurostopodus, sexes of.

Eurygnathus, different proportions of the head in the sexes of.

Eustephanus, sexual differences of species of; young of.

Exaggeration of natural characters by man.

Exogamy.

Experience, acquisition of, by animals.

Expression, resemblances in, between man and the apes.

Extinction of races, causes of.

Eye, destruction of the; change of position in; obliquity of, regarded as a beauty by the Chinese and Japanese.

Eyebrows, elevation of; development of long hairs in; in monkeys; eradicated in parts of South America and Africa; eradication of, by the Indians of Paraguay.

Eyelashes, eradication of, by the Indians of Paraguay.

Eyelids, coloured black, in part of Africa.

Eyes, pillared, of the male of Chloeon; difference in the colour of, in the sexes of birds.

Eyton, T.C., observations on the development of the horns in the fallow deer.

Eyzies, Les, human remains from.

Fabre, M., on the habits of Cerceris.

Facial bones, causes of modification of the.

Faculties, diversity of, in the same race of men; inheritance of; diversity of, in animals of the same species; mental variation of, in the same species; of birds.

Fakirs, Indian, tortures undergone by.

Falco leucocephalus.

Falco peregrinus.

Falco tinnunclus.

Falcon, peregrine, new mate found by.

Falconer, H., on the mode of fighting of the Indian elephant; on canines in a female deer; on Hyomoschus aquaticus.

Falkland Islands, horses of.

Fallow-deer, different coloured herds of.

Famines, frequency of, among savages.

Farr, Dr., on the effects of profligacy; on the influence of marriage on mortality.

Farrar, F.W., on the origin of language; on the crossing or blending of languages; on the absence of the idea of God in certain races of men; on early marriages of the poor; on the middle ages.

Farre, Dr., on the structure of the uterus.

Fashions, long prevalence of, among savages.

Faye, Prof., on the numerical proportion of male and female births in Norway and Russia; on the greater mortality of male children at and before birth.

Feathers, modified, producing sounds; elongated, in male birds; racket-shaped; barbless and with filamentous barbs in certain birds; shedding of margins of.

Feeding, high, probable influence of, in the pairing of birds of different species.

Feet, thickening of the skin on the soles of the; modification of, in man.

Felis canadensis, throat-ruff of.

Felis pardalis and F. mitis, sexual difference in the colouring of.

Female, behaviour of the, during courtship.

Female birds, differences of.

Females, presence of rudimentary male organs in; preference of, for certain males; pursuit of, by males; occurrence of secondary sexual characters in; development of male character by.

Females and males, comparative numbers of; comparative mortality of, while young.

Femur and tibia, proportions of, in the Aymara Indians.

Fenton, Mr., decrease of Maories; infanticide amongst the Maories.

Ferguson, Mr., on the courtship of fowls.

Fertilisation, phenomena of, in plants; in the lower animals.

Fertility lessened under changed conditions.

Fevers, immunity of Negroes and Mulattoes from.

Fiber zibethicus, protective colouring of it.

Fick, H., effect of conscription for military service.

Fidelity, in the elephant; of savages to one another; importance of.

Field-slaves, difference of, from house-slaves.

Fiji Archipelago, population of the.

Fiji Islands, beards of the natives; marriage-customs of the.

Fijians, burying their old and sick parents alive; estimation of the beard among the; admiration of, for a broad occiput.

Filial affection, partly the result of natural selection.

Filum terminale.

Finch, racket-shaped feathers in the tail of a.

Finches, spring change of colour in; British, females of the.

Fingers, partially coherent, in species of Hylobates.

Finlayson, on the Cochin Chinese.

Fire, use of.

Fischer, on the pugnacity of the male of Lethrus cephalotes.

Fischer, F. Von, on display of brightly coloured parts by monkeys in courtship.

Fish, eagerness of male; proportion of the sexes in; sounds produced by.

Fishes, kidneys of, represented by Corpora Wolffiana in the human embryo; male, hatching ova in their mouths; receptacles for ova possessed by; relative size of the sexes in; fresh-water, of the tropics; protective resemblances in; change of colour in; nest-building; spawning of; sounds produced by; continued growth of.

Flamingo, age of mature plumage.

Flexor pollicis longus, similar variation of, in man.

Flies, humming of.

Flint tools.

Flints, difficulty of chipping into form.

Florida, Quiscalus major in.

Florisuga mellivora.

Flounder, coloration of the.

Flower, W.H., on the abductor of the fifth metatarsal in apes; on the position of the Seals; on the Pithecia monachu; on the throat-pouch of the male bustard.

Fly-catchers, colours and nidification of.

Foetus, human, woolly covering of the; arrangement of the hair on.

Food, influence of, upon stature.

Foot, prehensile power of the, retained in some savages; prehensile, in the early progenitors of man.

Foramen, supra-condyloid, exceptional occurrence of in the humerus of man; in the early progenitors of man.

Forbes, D., on the Aymara Indians; on local variation of colour in the Quichuas; on the hairlessness of the Aymaras and Quichuas; on the long hair of the Aymaras and Quichaus.

Forel, F., on white young swans.

Forester, Hon. O.W., on an orphan hawk.

Formica rufa, size of the cerebral ganglia in.

Fossils, absence of, connecting man with the apes.

Fowl, occurrence of spurs in the female; game, early pugnacity of; Polish, early development of cranial peculiarities of; variations in plumage of; examples of correlated development in the; domestic, breeds and sub-breeds of.

Fowls, spangled Hamburg; inheritance of changes of plumage by; sexual peculiarities in, transmitted only to the same sex; loss of secondary sexual characters by male; Polish, origin of the crest in; period of inheritance of characters by; cuckoo-; development of the comb in; numerical proportion of the sexes in; courtship of; mongrel, between a black Spanish cock and different hens; pencilled Hamburg, difference of the sexes in; Spanish, sexual differences of the comb in; spurred, in both sexes.

Fox, W.D., on some half-tamed wild ducks becoming polygamous, and on polygamy in the guinea-fowl and canary-bird; on the proportion of the sexes in cattle; on the pugnacity of the peacock; on a nuptial assembly of magpies; on the finding of new mates by crows; on partridges living in triplets; on the pairing of a goose with a Chinese gander.

Foxes, wariness of young, in hunting districts; black.

Fraser, C., on the different colours of the sexes in a species of Squilla.

Fraser, G., colours of Thecla.

Frere, Hookham, quoting Theognis on selection in mankind.

Fringilla cannabina.

Fringilla ciris, age of mature plumage in.

Fringilla cyanea, age of mature plumage in.

Fringilla leucophrys, young of.

Fringilla spinus.

Fringilla tristis, change of colour in, in spring; young of.

Fringillidae, resemblance of the females of distinct species of.

Frog, bright coloured and distasteful to birds.

Frogs, male; temporary receptacles for ova possessed by; ready to breed before the females; fighting of; vocal organs of.

Frontal bone, persistence of the suture in.

Fruits, poisonous, avoided by animals.

Fuegians, difference of stature among the; power of sight in the; skill of, in stone-throwing; resistance of the, to their severe climate; mental capacity of the; quasi-religious sentiments of the; resemblance of, in mental characters, to Europeans; mode of life of the; aversion of, to hair on the face; said to admire European women.

Fulgoridae, songs of the.

Fur, whiteness of, in Arctic animals in winter.

Fur-bearing animals, acquired sagacity of.

Gallicrex, sexual difference in the colour of the irides in.

Gallicrex cristatus, pugnacity of male; red carbuncle occurring in the male during the breeding-season.

Gallinaceae, frequency of polygamous habits and of sexual differences in the; love-gestures of; decomposed feathers in; stripes of young; comparative sexual differences between the species of; plumage of.

Gallinaceous birds, weapons of the male; racket-shaped feathers on the heads of.

Gallinula chloropus, pugnacity of the male.

Galloperdix, spurs of; development of spurs in the female.

Gallophasis, young of.

Galls.

Gallus bankiva, neck-hackles of.

Gallus Stanleyi, pugnacity of the male.

Galton, Mr., on hereditary genius; gregariousness and independence in animals; on the struggle between the social and personal impulses; on the effects of natural selection on civilised nations; on the sterility of sole daughters; on the degree of fertility of people of genius; on the early marriages of the poor; on the ancient Greeks; on the Middle Ages; on the progress of the United States; on South African notions of beauty.

Gammarus, use of the chelae of.

Gammarus marinus.

Gannets, white only when mature.

Ganoid fishes.

Gaour, horns of the.

Gap between man and the apes.

Gaper, sexes and young of.

Gardner, on an example of rationality in a Gelasimus.

Garrulus glandarius.

Gartner, on sterility of hybrid plants.

Gasteropoda, pulmoniferous, courtship of.

Gasterosteus, nidification of.

Gasterosteus leiurus.

Gasterosteus trachurus.

Gastrophora, wings of, brightly coloured beneath.

Gauchos, want of humanity among the.

Gaudry, M., on a fossil monkey.

Gavia, seasonal change of plumage in.

Geese, clanging noise made by; pairing of different species of; Canada, selection of mates by.

Gegenbaur, C., on the number of digits in the Ichthyopterygia; on the hermaphroditism of the remote progenitors of the vertebrata; two types of nipple in mammals.

Gelasimus, proportions of the sexes in a species of; use of the enlarged chelae of the male; pugnacity of males of; rational actions of a; difference of colour in the sexes of a species of.

Gemmules, dormant in one sex.

Genius, hereditary.

Genius, fertility of men and women of.

Geoffroy St.-Hilaire, Isid., on the recognition of women by male quadrumana; on monstrosities; coincidences of arrested development with polydactylism; on animal-like anomalies in the human structure; on the correlation of monstrosities; on the distribution of hair in man and monkeys; on the caudal vertebrae of monkeys; on correlated variability; on the classification of man; on the long hair on the heads of species of Semnopithecus; on the hair in monkeys; on the development of horns in female deer; and F. Cuvier, on the mandrill; on Hylobates.

Geographical distribution, as evidence of specific distinctions in man.

Geometrae, brightly coloured beneath.

Geophagus, frontal protuberance of, male; eggs hatched by the male, in the mouth or branchial cavity.

Georgia, change of colour in Germans settled in.

Geotrupes, stridulation of.

Gerbe, M., on the nest-building of Crenilabus massa and C. Melops.

Gerland, Dr., on the prevalence of infanticide; on the extinction of races.

Gervais, P., on the hairiness of the gorilla; on the mandrill.

Gesture-language.

Ghost-moth, sexual difference of colour in the.

Giard, M., disputes descent of vertebrates from Ascidians; colour of sponges and Ascidians; musky odour of Sphinx.

Gibbon, voice of.

Gibbon, Hoolock, nose of.

Gibbs, Sir D., on differences of the voice in different races of men.

Gill, Dr., male seals larger than females; sexual differences in seals.

Giraffe, its mode of using the horns; mute, except in the rutting season.

Giraud-Teulon, on the cause of short sight.

Glanders, communicable to man from the lower animals.

Glands, odoriferous, in mammals.

Glareola, double moult in.

Glomeris limbata, difference of colour in the sexes of.

Glow-worm, female, apterous; luminosity of the.

Gnats, dances of; auditory powers of.

Gnu, skeletons of, found locked together; sexual differences in colour of the.

Goat, male, wild, falling on his horns; male, odour emitted by; male, wild, crest of the; Berbura, mane, dewlap, etc., of the male; Kemas, sexual difference in the colour of the.

Goats, sexual differences in the horns of; horns of; mode of fighting of; domestic, sexual differences of, late developed; beards of.

Goatsucker, Virginian, pairing of the.

Gobies, nidification of.

God, want of the idea of, in some races of men.

Godron, M., on variability; on difference of stature; on the want of connexion between climate and the colour of the skin; on the colour of the skin; on the colour of infants.

Goldfinch, proportion of the sexes in the; sexual differences of the beak in the; courtship of the.

Goldfinch, North American, young of.

Goldfish.

Gomphus, proportions of the sexes in; difference in the sexes of.

Gonepteryx Rhamni, sexual difference of colour in.

Goodsir, Prof., on the affinity of the lancelet to the ascidians.

Goosander, young of.

Goose, Antarctic, colours of the.

Goose, Canada, pairing with a Bernicle gander.

Goose, Chinese, knob on the beak of the.

Goose, Egyptian.

Goose, Sebastopol, plumage of.

Goose, Snow-, whiteness of the.

Goose, Spur-winged.

Gorilla, semi-erect attitude of the; mastoid processes of the; protecting himself from rain with his hands; manner of sitting; supposed to be a kind of mandrill; polygamy of the; voice of the; cranium of; fighting of male.

Gosse, P.H., on the pugnacity of the male Humming-bird.

Gosse, M., on the inheritance of artificial modifications of the skull.

Gould, B.A., on variation in the length of the legs in man; measurements of American soldiers; on the proportions of the body and capacity of the lungs in different races of men; on the inferior vitality of mulattoes.

Gould, J., on migration of swifts; on the arrival of male snipes before the females; on the numerical proportion of the sexes in birds; on Neomorpha Grypus; on the species of Eustephanus; on the Australian musk-duck; on the relative size of the sexes in Briziura lobata and Cincloramphus cruralis; on Lobivanellus lobatus; on habits of Menura Alberti; on the rarity of song in brilliant birds; on Selasphorus platycerus; on the Bower-birds; on the ornamental plumage of the Humming-birds; on the moulting of the ptarmigan; on the display of plumage by the male Humming-birds; on the shyness of adorned male birds; on the decoration of the bowers of Bower-birds; on the decoration of their nest by Humming-birds; on variation in the genus Cynanthus; on the colour of the thighs in a male parrakeet; on Urosticte Benjamini; on the nidification of the Orioles; on obscurely-coloured birds building concealed nests; on trogons and king-fishers; on Australian parrots; on Australian pigeons; on the moulting of the ptarmigan; on the immature plumage of birds; on the Australian species of Turnix; on the young of Aithurus polytmus; on the colours of the bills of toucans; on the relative size of the sexes in the marsupials of Australia; on the colours of the Marsupials.

Goureaux, on the stridulation of Mutilla europaea.

Gout, sexually transmitted.

Graba, on the Pied Ravens of the Feroe Islands; variety of the Guillemot.

Gradation of secondary sexual characters in birds.

Grallatores, absence of secondary sexual characters in; double moult in some.

Grallina, nidification of.

Grasshoppers, stridulation of the.

Gratiolet, Prof., on the anthropomorphous apes; on the evolution of the anthropomorphous apes; on the difference in the development of the brains of apes and of man.

Gray, Asa, on the gradation of species among the Compositae.

Gray, J.E., on the caudal vertebrae of monkeys; on the presence of rudiments of horns in the female of Cervulus moschatus; on the horns of goats and sheep; on crests of male antelopes; on the beard of the ibex; on the Berbura goat; on sexual differences in the coloration of Rodents; ornaments of male sloth; on the colours of the Elands; on the Sing-sing antelope; on the colours of goats; on Lemur Macaco; on the hog-deer.

“Greatest happiness principle.”

Greeks, ancient.

Green, A.H., on beavers fighting; on the voice of the beaver.

Greenfinch, selected by a female canary.

Greg, W.R., on the effects of natural selection on civilised nations; on the early marriages of the poor; on the Ancient Greeks.

Grenadiers, Prussian.

Greyhounds, numerical proportion of the sexes in; numerical proportion of male and female births in.

Grouse, red, monogamous; pugnacity of young male; producing a sound by beating their wings together; duration of courtship of; colours and nidification of.

Gruber, Dr., on the occurrence of the supra-condyloid foramen in the humerus of man; on division of malar bone; stridulation of locust; on ephippiger.

Grus americanus, age of mature plumage in; breeding in immature plumage.

Grus virgo, trachea of.

Gryllus campestris, pugnacity of male.

Gryllus domesticus.

Grypus, sexual differences in the beak in.

Guanacoes, battles of; canine teeth of.

Guanas, strife for women among the; polyandry among the.

Guanche skeletons, occurrence of the supra-condyloid foramen in the humerus of.

Guaranys, proportion of men and women among; colour of new-born children of the; beards of the.

Guenee, A., on the sexes of Hyperythra.

Guilding, L., on the stridulation of the Locustidae.

Guillemot, variety of the.

Guinea, sheep of, with males only horned.

Guinea-fowl, monogamous; occasional polygamy of the; markings of the.

Guinea-pigs, inheritance of the effects of operations by.

Gulls, seasonal change of plumage in; white.

Gunther, Dr., on paddle of Ceradotus; on hermaphroditism in Serranus; on male fishes hatching ova in their mouths; on mistaking infertile female fishes for males; on the prehensile organs of male Plagiostomous fishes; spines and brushes on fishes; on the pugnacity of the male salmon and trout; on the relative size of the sexes in fishes; on sexual differences in fishes; on the genus Callionymus; on a protective resemblance of a pipe-fish; on the genus Solenostoma; on the coloration of frogs and toads; combat of Testudo elegans; on the sexual differences in the Ophidia; on differences of the sexes of lizards.

Gynanisa Isis, ocellated spots of.

Gypsies, uniformity of, in various parts of the world.

Habits, bad, facilitated by familiarity; variability of the force of.

Haeckel, E., on the origin of man; on rudimentary characters; on death caused by inflammation of the vermiform appendage; on the canine teeth in man; on the steps by which man became a biped; on man as a member of the Catarrhine group; on the position of the Lemuridae; on the genealogy of the Mammalia; on the lancelet; on the transparency of pelagic animals; on the musical powers of women.

Hagen, H., and Walsh, B.D., on American Neuroptera.

Hair, development of, in man; character of, supposed to be determined by light and heat; distribution of, in man; possibly removed for ornamental purposes; arrangement and direction of; of the early progenitors of man; different texture of, in distinct races; and skin, correlation of colour of; development of, in mammals; management of, among different peoples; great length of, in some North American tribes; elongation of the, on the human head; possible inherited effect of plucking out.

Hairiness, difference of, in the sexes in man; variation of, in races of men.

Hairs and excretory pores, numerical relation of, in sheep.

Hairy family, Siamese.

Halbertsma, Prof., hermaphroditism in Serranus.

Hamadryas baboon, turning over stones; mane of the male.

Hamilton, C., on the cruelty of the Kaffirs to animals; on the engrossment of the women by the Kaffir chiefs.

Hammering, difficulty of.

Hancock, A., on the colours of the nudibranch Mollusca.

Hands, larger at birth, in the children of labourers; structure of, in the quadrumana; and arms, freedom of, indirectly correlated with diminution of canines.

Handwriting, inherited.

Handyside, Dr., supernumerary mammae in men.

Harcourt, E. Vernon, on Fringilla cannabina.

Hare, protective colouring of the.

Harelda glacialis.

Hares, battles of male.

Harlan, Dr., on the difference between field-and house-slaves.

Harris, J.M., on the relation of complexion to climate.

Harris, T.W., on the Katy-did locust; on the stridulation of the grasshoppers; on Oecanthus nivalis; on the colouring of Lepidoptera; on the colouring of Saturnia Io.

Harting, spur of the Ornithorhynchus.

Hartman, Dr., on the singing of Cicada septendecim.

Hatred, persistence of.

Haughton, S., on a variation of the flexor pollicis longus in man.

Hawks, feeding orphan nestling.

Hayes, Dr., on the diverging of sledge-dogs on thin ice.

Haymond, R., on the drumming of the male Tetrao umbellus; on the drumming of birds.

Head, altered position of, to suit the erect attitude of man; hairiness of, in man; processes of, in male beetles; artificial alterations of the form of the.

Hearne, on strife for women among the North American Indians; on the North American Indians’ notion of female beauty; repeated elopements of a North American woman.

Heart, in the human embryo.

Heat, supposed effects of.

Hectocotyle.

Hedge-warbler, young of the.

Heel, small projection of, in the Aymara Indians.

Hegt, M., on the development of the spurs in peacocks.

Heliconidae, mimicry of, by other butterflies.

Heliopathes, stridulation peculiar to the male.

Heliothrix auriculata, young of.

Helix pomatia, example of individual attachment in.

Hellins, J., proportions of sexes of Lepidoptera reared by.

Helmholtz, on pleasure derived from harmonies; on the human eye; on the vibration of the auditory hairs of crustacea; the physiology of harmony.

Hemiptera.

Hemitragus, beardless in both sexes.

Hemsbach, M. von, on medial mamma in man.

Hen, clucking of.

Hepburn, Mr., on the autumn song of the water-ouzel.

Hepialus humuli, sexual difference of colour in the.

Herbs, poisonous, avoided by animals.

Hermaphroditism, of embryos; in fishes.

Herodias bubulcus, vernal moult of.

Heron, Sir R., on the habits of peafowl.

Herons, love-gestures of; decomposed feathers in; breeding plumage of; young of the; sometimes dimorphic; continued growth of crest and plumes in the males of some; change of colour in some.

Hesperomys cognatus.

Hetaerina, proportion of the sexes in; difference in the sexes of.

Heterocerus, stridulation of.

Hewitt, Mr., on a game-cock killing a kite; on the recognition of dogs and cats by ducks; on the pairing of a wild duck with a pintail drake; on the courtship of fowls; on the coupling of pheasants with common hens.

Hilgendorf, sounds produced by crustaceans.

Hindoo, his horror of breaking his caste.

Hindoos, local difference of stature among; difference of, from Europeans; colour of the beard in.

Hipparchia Janira, instability of the ocellated spots of.

Hippocampus, development of; marsupial receptacles of the male.

Hippocampus minor.

Hippopotamus, nakedness of.

Hips, proportions of, in soldiers and sailors.

Hodgson, S., on the sense of duty.

Hoffberg, on the horns of the reindeer; on sexual preferences shewn by reindeer.

Hoffman, Prof., protective colours; fighting of frogs.

Hog, wart-; river-.

Hog-deer.

Holland, Sir H., on the effects of new diseases.

Homologous structures, correlated variation of.

Homoptera, stridulation of the, and Orthoptera, discussed.

Honduras, Quiscalus major in.

Honey-buzzard of India, variation in the crest of.

Honey-sucker, females and young of.

Honey-suckers, moulting of the; Australian, nidification of.

Honour, law of.

Hooker, Dr., forbearance of elephant to his keeper; on the colour of the beard in man.

Hookham, Mr., on mental concepts in animals.

Hoolock Gibbon, nose of.

Hoopoe, sounds produced by male.

Hoplopterus armatus, wing-spurs of.

Hornbill, African, inflation of the neck-wattle of the male during courtship.

Hornbills, sexual difference in the colour of the eyes in; nidification and incubation of.

Horne, C., on the rejection of a brightly-coloured locust by lizards and birds.

Horns, sexual differences of, in sheep and goats; loss of, in female merino sheep; development of, in deer; development in antelopes; from the head and thorax, in male beetles; of deer; originally a masculine character in sheep; and canine teeth, inverse development of.

Horse, fossil, extinction of the, in South America; polygamous; canine teeth of male; winter change of colour.

Horses, rapid increase of, in South America; diminution of canine teeth in; dreaming; of the Falkland Islands and Pampas; numerical proportion of the sexes, in; lighter in winter in Siberia; sexual preferences in; pairing preferently with those of the same colour; numerical proportion of male and female births in; formerly striped.

Hottentot women, peculiarities of.

Hottentots, lice of; readily become musicians; notions of female beauty of the; compression of nose by.

Hough, Dr. S., men’s temperature more variable than women’s; proportion of sexes in man.

House-slaves, difference of, from field-slaves.

Houzeau, on the baying of the dog; on reason in dogs; birds killed by telegraph wires; on the cries of domestic fowls and parrots; animals feel no pity; suicide in the Aleutian Islands.

Howorth, H.H., extinction of savages.

Huber, P., on ants playing together; on memory in ants; on the intercommunication of ants; on the recognition of each other by ants after separation.

Huc, on Chinese opinions of the appearance of Europeans.

Huia, the, of New Zealand.

Human, man, classed alone in a kingdom.

Human sacrifices.

Humanity, unknown among some savages; deficiency of, among savages.

Humboldt, A. von, on the rationality of mules; on a parrot preserving the language of a lost tribe; on the cosmetic arts of savages; on the exaggeration of natural characters by man; on the red painting of American Indians.

Hume, D., on sympathetic feelings.

Humming-bird, racket-shaped feathers in the tail of a; display of plumage by the male.

Humming-birds, ornament their nests; polygamous; proportion of the sexes in; sexual differences in; pugnacity of male; modified primaries of male; coloration of the sexes of; display by; nidification of the; colours of female; young of.

Humour, sense of, in dogs.

Humphreys, H.N., on the habits of the stickleback.

Hunger, instinct of.

Huns, ancient, flattening of the nose by the.

Hunter, J., on the number of species of man; on secondary sexual characters; on the general behaviour of female animals during courtship; on the muscles of the larynx in song-birds; on strength of males; on the curled frontal hair of the bull; on the rejection of an ass by a female zebra.

Hunter, W.W., on the recent rapid increase of the Santali; on the Santali.

Huss, Dr. Max, on mammary glands.

Hussey, Mr., on a partridge distinguishing persons.

Hutchinson, Col., example of reasoning in a retriever.

Hutton, Captain, on the male wild goat falling on his horns.

Huxley, T.H., on the structural agreement of man with the apes; on the agreement of the brain in man with that of lower animals; on the adult age of the orang; on the embryonic development of man; on the origin of man; on variation in the skulls of the natives of Australia; on the abductor of the fifth metatarsal in apes; on the nature of the reasoning power; on the position of man; on the suborders of primates; on the Lemuridae; on the Dinosauria; on the amphibian affinities of the Ichthyosaurians; on variability of the skull in certain races of man; on the races of man; Supplement on the brain.

Hybrid birds, production of.

Hydrophobia, communicable between man and the lower animals.

Hydroporus, dimorphism of females of.

Hyelaphus porcinus.

Hygrogonus.

Hyla, singing species of.

Hylobates, absence of the thumb in; upright progression of some species of; maternal affection in a; direction of the hair on the arms of species of; females of, less hairy below than males.

Hylobates agilis, hair on the arms of; musical voice of the; superciliary ridge of; voice of.

Hylobates hoolock, sexual difference of colour in.

Hylobates lar, hair on the arms of; female less hairy.

Hylobates leuciscus, song of.

Hylobates syndactylus, laryngeal sac of.

Hylophila prasinana.

Hymenoptera, large size of the cerebral ganglia in; classification of; sexual differences in the wings of; aculeate, relative size of the sexes of.

Hymenopteron, parasitic, with a sedentary male.

Hyomoschus aquaticus.

Hyperythra, proportion of the sexes in.

Hypogymna dispar, sexual difference of colour in.

Hypopyra, coloration of.

Ibex, male, falling on his horns; beard of the.

Ibis, white, change of colour of naked skin in, during the breeding season; scarlet, young of the.

Ibis tantalus, age of mature plumage in; breeding in immature plumage.

Ibises, decomposed feathers in; white; and black.

Ichneumonidae, difference of the sexes in.

Ichthyopterygia.

Ichthyosaurians.

Idiots, microcephalous, their characters and habits; hairiness and animal nature of their actions; microcephalous, imitative faculties of.

Iguana tuberculata.

Iguanas.

Illegitimate and legitimate children, proportion of the sexes in.

Imagination, existence of, in animals.

Imitation, of man by monkeys; tendency to, in monkeys, microcephalous idiots and savages; influence of.

Immature plumage of birds.

Implacentata.

Implements, employed by monkeys; fashioning of, peculiar to man.

Impregnation, period of, influence of, upon sex.

Improvement, progressive, man alone supposed to be capable of.

Incisor teeth, knocked out or filed by some savages.

Increase, rate of; necessity of checks in.

Indecency, hatred of, a modern virtue.

India, difficulty of distinguishing the native races of; Cyprinidae of; colour of the beard in races of men of.

Indian, North American, honoured for scalping a man of another tribe.

Individuality, in animals.

Indolence of man, when free from a struggle for existence.

Indopicus carlotta, colours of the sexes of.

Infanticide, prevalence of; supposed cause of; prevalence and causes of.

Inferiority, supposed physical, of man.

Inflammation of the bowels, occurrence of, in Cebus Azarae.

Inheritance, of long and short sight; of effects of use of vocal and mental organs; of moral tendencies; laws of; sexual; sexually limited.

Inquisition, influence of the.

Insanity, hereditary.

insect, fossil, from the Devonian.

Insectivora, absence of secondary sexual characters in.

Insects, relative size of the cerebral ganglia in; male, appearance of, before the females; pursuit of female, by the males; period of development of sexual characters in; secondary sexual characters of; kept in cages; stridulation.

Insessores, vocal organs of.

Instep, depth of, in soldiers and sailors.

Instinct and intelligence.

Instinct, migratory, vanquishing the maternal.

Instinctive actions, the result of inheritance.

Instinctive impulses, difference of the force; and moral impulses, alliance of.

Instincts, complex origin of, through natural selection; possible origin of some; acquired, of domestic animals; variability of the force of; difference of force between the social and other; utilised for new purposes.

Instrumental music of birds.

Intellect, influence of, in natural selection in civilised society.

Intellectual faculties, their influence on natural selection in man; probably perfected through natural selection.

Intelligence, Mr. H. Spencer on the dawn of.

Intemperance, no reproach among savages; its destructiveness.

Intoxication in monkeys.

Iphias glaucippe.

Iris, sexual difference in the colour of the, in birds.

Ischio-pubic muscle.

Ithaginis cruentus, number of spurs in.

Iulus, tarsal suckers of the males of.

Jackals learning from dogs to bark.

Jack-snipe, coloration of the.

Jacquinot, on the number of species of man.

Jaeger, Dr., length of bones increased from carrying weights; on the difficulty of approaching herds of wild animals; male Silver-pheasant, rejected when his plumage was spoilt.

Jaguars, black.

Janson, E.W., on the proportions of the sexes in Tomicus villosus; on stridulant beetles.

Japan, encouragement of licentiousness in.

Japanese, general beardlessness of the; aversion of the, to whiskers.

Jardine, Sir W., on the Argus pheasant.

Jarrold, Dr., on modifications of the skull induced by unnatural position.

Jarves, Mr., on infanticide in the Sandwich Islands.

Javans, relative height of the sexes of; notions of female beauty.

Jaw, influence of the muscles of the, upon the physiognomy of the apes.

Jaws, smaller proportionately to the extremities; influence of food upon the size of; diminution of, in man; in man, reduced by correlation.

Jay, young of the; Canada, young of the.

Jays, new mates found by; distinguishing persons.

Jeffreys, J. Gwyn, on the form of the shell in the sexes of the Gasteropoda; on the influence of light upon the colours of shells.

Jelly-fish, bright colours of some.

Jenner, Dr., on the voice of the rook; on the finding of new mates by magpies; on retardation of the generative functions in birds.

Jenyns, L., on the desertion of their young by swallows; on male birds singing after the proper season.

Jerdon, Dr., on birds dreaming; on the pugnacity of the male bulbul; on the pugnacity of the male Ortygornis gularis; on the spurs of Galloperdix; on the habits of Lobivanellus; on the spoonbill; on the drumming of the Kalij-pheasant; on Indian bustards; on Otis bengalensis; on the ear-tufts of Sypheotides auritus; on the double moults of certain birds; on the moulting of the honeysuckers; on the moulting of bustards, plovers, and drongos; on the spring change of colour in some finches; on display in male birds; on the display of the under-tail coverts by the male bulbul; on the Indian honey-buzzard; on sexual differences in the colour of the eyes of hornbills; on the markings of the Tragopan pheasant; on the nidification of the Orioles; on the nidification of the hornbills; on the Sultan yellow-tit; on Palaeornis javanicus; on the immature plumage of birds; on representative species of birds; on the habits of Turnix; on the continued increase of beauty of the peacock; on coloration in the genus Palaeornis.

Jevons, W.S., on the migrations of man.

Jews, ancient use of flint tools by the; uniformity of, in various parts of the world; numerical proportion of male and female births among the; ancient, tattooing practised by.

Johnstone, Lieut., on the Indian elephant.

Jollofs, fine appearance of the.

Jones, Albert, proportion of sexes of Lepidoptera, reared by.

Juan Fernandez, humming-birds of.

Junonia, sexual differences of colouring in species of.

Jupiter, comparison with Assyrian effigies.

Kaffir skull, occurrence of the diastema in a.

Kaffirs, their cruelty to animals; lice of the; colour of the; engrossment of the handsomest women by the chiefs of the; marriage-customs of the.

Kalij-pheasant, drumming of the male; young of.

Kallima, resemblance of, to a withered leaf.

Kulmucks, general beardlessness of; aversion of, to hairs on the face; marriage-customs of the.

Kangaroo, great red, sexual difference in the colour of.

Kant, Imm., on duty; on self-restraint; on the number of species of man.

Katy-did, stridulation of the.

Keen, Dr., on the mental powers of snakes.

Keller, Dr., on the difficulty of fashioning stone implements.

Kent, W.S., elongation of dorsal fin of Callionymus lyra; courtship of Labrus mixtus; colours and courtship of Cantharus lineatus.

Kestrels, new mates found by.

Kidney, one, doing double work in disease.

King, W.R., on the vocal organs of Tetrao cupido; on the drumming of grouse; on the reindeer; on the attraction of male deer by the voice of the female.

King and Fitzroy, on the marriage-customs of the Fuegians.

King-crows, nidification of.

Kingfisher, racket-shaped feathers in the tail of a.

Kingfishers, colours and nidification of the; immature plumage of the; young of the.

King Lory, immature plumage of the.

Kingsley, C., on the sounds produced by the Umbrina.

Kirby and Spence, on sexual differences in the length of the snout in Curculionidae; on the courtship of insects; on the elytra of Dytiscus; on peculiarities in the legs of male insects; on the relative size of the sexes in insects; on the Fulgoridae; on the habits of the Termites; on difference of colour in the sexes of beetles; on the horns of the male lamellicorn beetles; on hornlike processes in male Curculionidae; on the pugnacity of the male stag-beetle.

Kite, killed by a game-cock.

Knot, retention of winter plumage by the.

Knox, R., on the semilunar fold; on the occurrence of the supra-condyloid foramen in the humerus of man; on the features of the young Memmon.

Koala, length of the caecum in.

Kobus ellipsiprymnus, proportion of the sexes in.

Kolreuter, on the sterility of hybrid plants.

Koodoo, development of the horns of the; markings of the.

Koppen, F.T., on the migratory locust.

Koraks, marriage customs of.

Kordofan, protuberances artificially produced by natives of.

Korte, on the proportion of sexes in locusts; Russian locusts.

Kovalevsky, A., on the affinity of the Ascidia to the Vertebrata.

Kovalevsky, W., on the pugnacity of the male capercailzie; on the pairing of the capercailzie.

Krause, on a convoluted body at the extremity of the tail in a Macacus and a cat.

Kupffer, Prof., on the affinity of the Ascidia to the Vertebrata.

Labidocera Darwinii, prehensile organs of the male.

Labrus, splendid colours of the species of.

Labrus mixtus, sexual differences in.

Labrus pavo.

Lacertilia, sexual differences of.

Lafresnaye, M. de, on birds of paradise.

Lamarck, on the origin of man.

Lamellibranchiata.

Lamellicorn beetles, horn-like processes from the head and thorax of; influence of sexual selection on.

Lamellicornia, stridulation of.

Lamont, Mr., on the tusks of the walrus; on the use of its tusks by the walrus; on the bladder-nose seal.

Lampornis porphyrurus, colours of the female.

Lampyridae, distasteful to mammals.

Lancelet.

Landois, H., gnats attracted by sound; on the production of sound by the Cicadae; on the stridulating organ of the crickets; on Decticus; on the stridulating organs of the Acridiidae; stridulating apparatus, in Orthoptera; on the stridulation of Necrophorus; on the stridulant organ of Cerambyx heros; on the stridulant organ of Geotrupes; on the stridulating organs in the Coleoptera; on the ticking of Anobium.

Landor, Dr., on remorse for not obeying tribal custom.

Language, an art; articulate, origin of; relation of the progress of, to the development of the brain; effects of inheritance in production of; complex structure of, among barbarous nations; natural selection in; gesture; primeval; of a lost tribe preserved by a parrot.

Languages, presence of rudiments in; classification of; variability of; crossing or blending of; complexity of, no test of perfection or proof of special creation; resemblance of, evidence of community of origin.

Languages and species, identity of evidence of their gradual development.

Lanius, characters of young.

Lanius rufus, anomalous young of.

Lankester, E.R., on comparative longevity; on the destructive effects of intemperance.

Lanugo of the human foetus.

Lapponian language, highly artificial.

Lark, proportion of the sexes in the; female, singing of the.

Larks, attracted by a mirror.

Lartet, E., comparison of cranial capacities of skulls of recent and tertiary mammals; on the size of the brain in mammals; on Dryopithecus; on pre-historic flutes.

Larus, seasonal change of plumage in.

Larva, luminous, of a Brazilian beetle.

Larynx, muscles of the, in songbirds.

Lasiocampa quercus, attraction of males by the female; sexual difference of colour in.

Latham, R.G., on the migrations of man.

Latooka, perforation of the lower lip by the women of.

Laurillard, on the abnormal division of the malar bone in man.

Lawrence, W., on the superiority of savages to Europeans in power of sight; on the colour of negro infants; on the fondness of savages for ornaments; on beardless races; on the beauty of the English aristocracy.

Layard, E.L., on the instance of rationality in a cobra; on the pugnacity of Gallus Stanleyi.

Laycock, Dr., on vital periodicity; theroid nature of idiots.

Leaves, autumn, tints useless.

Lecky, Mr., on the sense of duty; on suicide; on the practice of celibacy; his view of the crimes of savages; on the gradual rise of morality.

Leconte, J.L., on the stridulant organ in the Coprini and Dynastini.

Lee, H., on the numerical proportion of the sexes in the trout.

Leg, calf of the, artificially modified.

Legitimate and illegitimate children, proportion of the sexes in.

Legs, variation of the length of the, in man; proportions of, in soldiers and sailors; front, atrophied in some male butterflies; peculiarities of, in male insects.

Leguay, on the occurrence of the supra-condyloid foramen in the humerus of man.

Lek of the black-cock and capercailzie.

Lemoine, Albert, on the origin of language.

Lemur macaco, sexual difference of colour in.

Lemuridae, ears of the; variability of the muscles in the; position and derivation of the; their origin.

Lemurs, uterus in the.

Lenguas, disfigurement of the ears of the.

Leopards, black.

Lepidoptera, numerical proportions of the sexes in the; colouring of; ocellated spots of.

Lepidosiren.

Leptalides, mimicry of.

Leptorhynchus angustatus, pugnacity of male.

Leptura testacea, difference of colour in the sexes.

Leroy, on the wariness of young foxes in hunting-districts; on the desertion of their young by swallows.

Leslie, D., marriage customs of Kaffirs.

Lesse, valley of the.

Lesson, on the birds of paradise; on the sea-elephant.

Lessona, M., observations on Serranus.

Lethrus cephalotes, pugnacity of the males of.

Leuciscus phoxinus.

Leuckart, R., on the vesicula prostatica; on the influence of the age of parents on the sex of offspring.

Levator claviculae muscle.

Libellula depressa, colour of the male.

Libellulidae, relative size of the sexes of; difference in the sexes of.

Lice of domestic animals and man.

Licentiousness a check upon population; prevalence of, among savages.

Lichtenstein, on Chera progne.

Life, inheritance at corresponding periods of.

Light, effects on complexion; influence of, upon the colours of shells.

Lilford, Lord, the ruff attracted by bright objects.

Limosa lapponica.

Linaria.

Linaria montana.

Lindsay, Dr. W.L., diseases communicated from animals to man; madness in animals; the dog considers his master his God.

Linnaeus, views of, as to the position of man.

Linnet, numerical proportion of the sexes in the; crimson forehead and breast of the; courtship of the.

Lion, polygamous; mane of the, defensive; roaring of the.

Lions, stripes of young.

Lips, piercing of the, by savages.

Lithobius, prehensile appendages of the female.

Lithosia, coloration in.

Littorina littorea.

Livingstone, Dr., manner of sitting of gorilla; on the influence of dampness and dryness on the colour of the skin; on the liability of negroes to tropical fevers after residence in a cold climate; on the spur-winged goose; on weaverbirds; on an African night-jar; on the battle-scars of South African male mammals; on the removal of the upper incisors by the Batokas; on the perforation of the upper lip by the Makalolo; on the Banyai.

Livonia, numerical proportion of male and female births in.

Lizards, relative size of the sexes of; gular pouches of.

Lloyd, L., on the polygamy of the capercailzie and bustard; on the numerical proportion of the sexes in the capercailzie and blackcock; on the salmon; on the colours of the sea-scorpion; on the pugnacity of male grouse; on the capercailzie and blackcock; on the call of the capercailzie; on assemblages of grouse and snipes; on the pairing of a shield-drake with a common duck; on the battles of seals; on the elk.

Lobivanellus, wing-spurs in.

Local influences, effect of, upon stature.

Lockwood, Mr., on the development of Hippocampus.

Lockwood, Rev. S., musical mouse.

Locust, bright-coloured, rejected by lizards and birds.

Locust, migratory; selection by female.

Locustidae, stridulation of the; descent of the.

Locusts, proportion of sexes in; stridulation of.

Longicorn beetles, difference of the sexes of, in colour; stridulation of.

Lonsdale, Mr., on an example of personal attachment in Helix pomatia.

Lophobranchii, marsupial receptacles of the male.

Lophophorus, habits of.

Lophorina atra, sexual difference in coloration of.

Lophornis ornatus.

Lord, J.K., on Salmo lycaodon.

Lory, King; immature plumage of the.

Lory, King, constancy of.

Love-antics and dances of birds.

Lowne, B.T., on Musca vomitoria.

Loxia, characters of young of.

Lubbock, Sir J., on the antiquity of man; on the origin of man; on the mental capacity of savages; on the origin of implements; on the simplification of languages; on the absence of the idea of God among certain races of men; on the origin of the belief in spiritual agencies; on superstitions; on the sense of duty; on the practice of burying the old and sick among the Fijians; on the immorality of savages; on Mr. Wallace’s claim to the origination of the idea of natural selection; on the former barbarism of civilised nations; on improvements in the arts among savages; on resemblances of the mental characters in different races of men; on the arts practised by savages; on the power of counting in primeval man; on the prehensile organs of the male Labidocera Darwinii; on Chloeon; on Smynthurus luteus; finding of new mates by jays; on strife for women among the North American Indians; on music; on the ornamental practices of savages; on the estimation of the beard among the Anglo-Saxons; on artificial deformation of the skull; on “communal marriages;” on exogamy; on the Veddahs; on polyandry.

Lucanidae, variability of the mandibles in the male.

Lucanus, large size of males of.

Lucanus cervus, numerical proportion of sexes of; weapons of the male.

Lucanus elaphus, use of mandibles of; large jaws of male.

Lucas, Prosper, on pigeons; on sexual preference in horses and bulls.

Luminosity in insects.

Lunar periods.

Lund, Dr., on skulls found in Brazilian caves.

Lungs, enlargement of, in the Quichua and Aymara Indians; a modified swim-bladder; different capacity of, in races of man.

Luschka, Prof., on the termination of the coccyx.

Luxury, expectation of life uninfluenced by.

Lycaena, sexual differences of colour in species of.

Lycaenae, colours of.

Lyell, Sir C., on the antiquity of man; on the origin of man; on the parallelism of the development of species and languages; on the extinction of languages; on the Inquisition; on the fossil remains of vertebrata; on the fertility of mulattoes.

Lynx, Canadian throat-ruff of the.

Lyre-bird, assemblies of.

Macacus, ears of; convoluted body in the extremity of the tail of; variability of the tail in species of; whiskers of species of.

Macacus brunneus.

Macacus cynomolgus, superciliary ridge of; beard and whiskers of; becoming white with age.

Macacus ecaudatus.

Macacus lasiotus, facial spots of.

Macacus nemestrinus.

Macacus radiatus.

Macacus rhesus, sexual difference in the colour of.

Macalister, Prof., on variations of the palmaris accessorius muscle; on muscular abnormalities in man; on the greater variability of the muscles in men than in women.

Macaws, Mr. Buxton’s observations on.

McCann, J., on mental individuality.

McClelland, J., on the Indian Cyprinidae.

Macculloch, Col., on an Indian village without any female children.

Macculloch, Dr., on tertian ague in a dog.

Macgillivray, W., on the vocal organs of birds; on the Egyptian goose; on the habits of woodpeckers; on the habits of the snipe; on the whitethroat; on the moulting of the snipes; on the moulting of the Anatidae; on the finding of new mates by magpies; on the pairing of a blackbird and thrush; on pied ravens; on the guillemots; on the colours of the tits; on the immature plumage of birds.

Machetes, sexes and young of.

Machetes pugnax, supposed to be polygamous; numerical proportion of the sexes in; pugnacity of the male; double moult in.

McIntosh, Dr., colours of the Nemertians.

McKennan, marriage customs of Koraks.

Mackintosh, on the moral sense.

MacLachlan, R., on Apatania muliebris and Boreus hyemalis; on the anal appendages of male insects; on the pairing of dragon-flies; on dragon-flies; on dimorphism in Agrion; on the want of pugnacity in male dragon-flies; colour of ghost-moth in the Shetland Islands.

M’Lennan, Mr., on infanticide; on the origin of the belief in spiritual agencies; on the prevalence of licentiousness among savages; on the primitive barbarism of civilised nations; on traces of the custom of the forcible capture of wives; on polyandry.

Macnamara, Mr., susceptibility of Andaman islanders and Nepalese to change.

M’Neill, Mr., on the use of the antlers of deer; on the Scotch deerhound; on the long hairs on the throat of the stag; on the bellowing of stags.

Macropus, courtship of.

Macrorhinus proboscideus, structure of the nose of.

Magpie, power of speech of; vocal organs of the; nuptial assemblies of; new mates found by; stealing bright objects; young of the; coloration of the.

Maillard, M., on the proportion of the sexes in a species of Papilio from Bourbon.

Maine, Sir Henry, on the absorption of one tribe by another; a desire for improvement not general.

Major, Dr. C. Forsyth, on fossil Italian apes; skull of Bos etruscus; tusks of miocene pigs.

Makalolo, perforation of the upper lip by the.

Malar bone, abnormal division of, in man.

Malay Archipelago, marriage-customs of the savages of the.

Malays, line of separation between the Papuans and the; general beardlessness of the; staining of the teeth among; aversion of some, to hairs on the face.

Malays and Papuans, contrasted characters of.

Male animals, struggles of, for the possession of the females; eagerness of, in courtship; generally more modified than female; differ in the same way from females and young.

Male characters, developed in females; transfer of, to female birds.

Male, sedentary, of a hymenopterous parasite.

Malefactors.

Males, presence of rudimentary female organs in.

Males and females, comparative numbers of; comparative mortality of, while young.

Malherbe, on the woodpeckers.

Mallotus Peronii.

Mallotus villosus.

Malthus, T., on the rate of increase of population.

Maluridae, nidification of the.

Malurus, young of.

Mammae, rudimentary, in male mammals; supernumerary, in women; of male human subject.

Mammalia, Prof. Owen’s classification of; genealogy of the.

Mammals, recent and tertiary, comparison of cranial capacity of; nipples of; pursuit of female, by the males; secondary sexual characters of; weapons of; relative size of the sexes of; parallelism of, with birds in secondary sexual characters; voices of, used especially during the breeding season.

Man, variability of; erroneously regarded as more domesticated than other animals; migrations of; wide distribution of; causes of the nakedness of; supposed physical inferiority of; a member of the Catarrhine group; early progenitors of; transition from ape indefinite; numerical proportions of the sexes in; difference between the sexes; proportion of sexes amongst the illegitimate; different complexion of male and female negroes; secondary sexual characters of; primeval condition of.

Mandans, correlation of colour and texture of hair in the.

Mandible, left, enlarged in the male of Taphroderes distortus.

Mandibles, use of the, in Ammophila; large, of Corydalis cornutus; large, of male Lucanus elaphus.

Mandrill, number of caudal vertebrae in the; colours of the male.

Mantegazza, Prof., on last molar teeth of man; bright colours in male animals; on the ornaments of savages; on the beardlessness of the New Zealanders; on the exaggeration of natural characters by man.

Mantell, W., on the engrossment of pretty girls by the New Zealand chiefs.

Mantis, pugnacity of species of.

Maories, mortality of; infanticide and proportion of sexes; distaste for hairiness amongst men.

Marcus Aurelius, on the origin of the moral sense; on the influence of habitual thoughts.

Mareca penelope.

Marks, retained throughout groups of birds.

Marriage, restraints upon, among savages; influence of, upon morals; influence of, on mortality; development of.

Marriages, early; communal.

Marshall, Dr. W., protuberances on birds’ heads; on the moulting of birds; advantage to older birds of paradise.

Marshall, Col., interbreeding amongst Todas; infanticide and proportion of sexes with Todas; choice of husband amongst Todas.

Marshall, Mr., on the brain of a Bushwoman.

Marsupials, development of the nictitating membrane in; uterus of; possession of nipples by; their origin from Monotremata; abdominal sacs of; relative size of the sexes of; colours of.

Marsupium, rudimentary in male marsupials.

Martin, W.C.L., on alarm manifested by an orang at the sight of a turtle; on the hair in Hylobates; on a female American deer; on the voice of Hylobates agilis; on Semnopithecus nemaeus.

Martin, on the beards of the inhabitants of St. Kilda.

Martins deserting their young.

Martins, C., on death caused by inflammation of the vermiform appendage.

Mastoid processes in man and apes.

Maudsley, Dr., on the influence of the sense of smell in man; on idiots smelling their food; on Laura Bridgman; on the development of the vocal organs; moral sense failing in incipient madness; change of mental faculties at puberty in man.

Mayers, W.F., on the domestication of the goldfish in China.

Mayhew, E., on the affection between individuals of different sexes in the dog.

Maynard, C.J., on the sexes of Chrysemys picta.

Meckel, on correlated variation of the muscles of the arm and leg.

Medicines, effect produced by, the same in man and in monkeys.

Medusae, bright colours of some.

Megalithic structures, prevalence of.

Megapicus validus, sexual difference of colour in.

Megasoma, large size of males of.

Meigs, Dr. A., on variation in the skulls of the natives of America.

Meinecke, on the numerical proportion of the sexes in butterflies.

Melanesians, decrease of.

Meldola, Mr., colours and marriage flight of Colias and Pieris.

Meliphagidae, Australian, nidification of.

Melita, secondary sexual characters of.

Meloe, difference of colour in the sexes of a species of.

Memnon, young.

Memory, manifestations of, in animals.

Mental characters, difference of, in different races of men.

Mental faculties, diversity of, in the same race of men; inheritance of; variation of, in the same species; similarity of the, in different races of man; of birds.

Mental powers, difference of, in the two sexes in man.

Menura Alberti, song of.

Menura superba, long tails of both sexes of.

Merganser, trachea of the male.

Merganser serrator, male plumage of.

Mergus cucullatus, speculum of.

Mergus merganser, young of.

Metallura, splendid tail-feathers of.

Methoca ichneumonides, large male of.

Meves, M., on the drumming of the snipe.

Mexicans, civilisation of the, not foreign.

Meyer, on a convoluted body at the extremity of the tail in a Macacus and a cat.

Meyer, Dr. A., on the copulation of Phryganidae of distinct species.

Meyer, Prof. L., on development of helix of ear; men’s ears more variable than women’s; antennae serving as ears.

Migrations of man, effects of.

Migratory instinct of birds; vanquishing the maternal.

Mill, J.S., on the origin of the moral sense; on the “greatest happiness principle;” on the difference of the mental powers in the sexes of man.

Millipedes.

Milne-Edwards, H., on the use of enlarged chelae of the male Gelasimus.

Milvago leucurus, sexes and young of.

Mimicry.

Mimus polyglottus.

Mind, difference of, in man and the highest animals; similarity of the, in different races.

Minnow, proportion of the sexes in the.

Mirror, behaviour of monkeys before.

Mirrors, larks attracted by.

Mitchell, Dr., interbreeding in the Hebrides.

Mitford, selection of children in Sparta.

Mivart, St. George, on the reduction of organs; on the ears of the lemuroidea; on variability of the muscles in lemuroidea; on the caudal vertebrae of monkeys; on the classification of the primates; on the orang and on man; on differences in the lemuroidea; on the crest of the male newt.

Mobius, Prof., on reasoning powers in a pike.

Mocking-thrush, partial migration of; young of the.

Modifications, unserviceable.

Moggridge, J.T., on habits of spiders; on habits of ants.

Moles, numerical proportion of the sexes in; battles of male.

Mollienesia petenensis, sexual difference in.

Mollusca, beautiful colours and shapes of; absence of secondary sexual characters in the.

Molluscoida.

Monacanthus scopas and M. Peronii.

Monboddo, Lord, on music.

Mongolians, perfection of the senses in.

Monkey, protecting his keeper from a baboon; bonnet-; rhesus-, sexual difference in colour of the; moustache-, colours of the.

Monkeys, liability of, to the same diseases as man; male, recognition of women by; diversity of the mental faculties in; breaking hard fruits with stones; hands of the; basal caudal vertebrae of, imbedded in the body; revenge taken by; maternal affection in; variability of the faculty of attention in; American, manifestation of reason in; using stones and sticks; imitative faculties of; signal-cries of; mutual kindnesses of; sentinels posted by; human characters of; American, direction of the hair on the arms of some; gradation of species of; beards of; ornamental characters of; analogy of sexual differences of, with those of man; different degrees of difference in the sexes of; expression of emotions by; generally monogamous habits of; polygamous habits of some; naked surfaces of; courtship of.

Monogamy, not primitive.

Monogenists.

Mononychus pseudacori, stridulation of.

Monotremata, development of the nictitating membrane in; lactiferous glands of; connecting mammals with reptiles.

Monstrosities, analogous, in man and lower animals; caused by arrest of development; correlation of; transmission of.

Montagu, G., on the habits of the black and red grouse; on the pugnacity of the ruff; on the singing of birds; on the double moult of the male pintail.

Monteiro, Mr., on Bucorax abyssinicus.

Montes de Oca, M., on the pugnacity of male Humming-birds.

Monticola cyanea.

Monuments, as traces of extinct tribes.

Moose, battles of; horns of the, an incumbrance.

Moral and instinctive impulses, alliance of.

Moral faculties, their influence on natural selection in man.

Moral rules, distinction between the higher and lower.

Moral sense, so-called, derived from the social instincts; origin of the.

Moral tendencies, inheritance of.

Morality, supposed to be founded in selfishness; test of, the general welfare of the community; gradual rise of; influence of a high standard of.

Morgan, L.H., on the beaver; on the reasoning powers of the beaver; on the forcible capture of wives; on the castoreum of the beaver; marriage unknown in primeval times; on polyandry.

Morley, J., on the appreciation of praise and fear of blame.

Morris, F.O., on hawks feeding an orphan nestling.

Morse, Dr., colours of mollusca.

Morselli, E., division of the malar bone.

Mortality, comparative, of female and male.

Morton on the number of species of man.

Moschkau, Dr. A., on a speaking starling.

Moschus moschiferus, odoriferous organs of.

Motacillae, Indian, young of.

Moth, odoriferous.

Moths, absence of mouth in some males; apterous female; male, prehensile use of the tarsi by; male, attracted by females; sound produced by; coloration of; sexual differences of colour in.

Motmot, inheritance of mutilation of tail feathers; racket-shaped feathers in the tail of a.

Moult, double; double annual, in birds.

Moulting of birds.

Moults, partial.

Mouse, song of.

Moustache-monkey, colours of the.

Moustaches, in monkeys.

Mud-turtle, long claws of the male.

Mulattoes, persistent fertility of; immunity of, from yellow fever.

Mule, sterility and strong vitality of the.

Mules, rational.

Muller, Ferd., on the Mexicans and Peruvians.

Muller, Fritz, on astomatous males of Tanais; on the disappearance of spots and stripes in adult mammals; on the proportions of the sexes in some Crustacea; on secondary sexual characters in various Crustaceans; musical contest between male Cicadae; mode of holding wings in Castina; on birds shewing a preference for certain colours; on the sexual maturity of young amphipod Crustacea.

Muller, Hermann, emergence of bees, from pupa; pollen-gathering of bees; proportion of sexes in bees; courting of Eristalis; colour and sexual selection with bees.

Muller, J., on the nictitating membrane and semilunar fold.

Muller, Max, on the origin of language; language implies power of general conception; struggle for life among the words, etc., of languages.

Muller, S., on the banteng; on the colours of Semnopithecus chrysomelas.

Muntjac-deer, weapons of the.

Murie, J., on the reduction of organs; on the ears of the Lemuroidea; on variability of the muscles in the Lemuroidea; basal caudal vertebrae of Macacus brunneus imbedded in the body; on the manner of sitting in short-tailed apes; on differences in the Lemuroidea; on the throat-pouch of the male bustard; on the mane of Otaria jubata; on the sub-orbital pits of Ruminants; on the colours of the sexes in Otaria nigrescens.

Murray, A., on the Pediculi of different races of men.

Murray, T.A., on the fertility of Australian women with white men.

Mus coninga.

Mus minutus, sexual difference in the colour of.

Musca vomitoria.

Muscicapa grisola.

Muscicapa luctuosa.

Muscicapa ruticilla, breeding in immature plumage.

Muscle, ischio-pubic.

Muscles, rudimentary, occurrence of, in man; variability of the; effects of use and disuse upon; animal-like abnormalities of, in man; correlated variation of, in the arm and leg; variability of, in the hands and feet; of the jaws, influence of, on the physiognomy of the Apes; habitual spasms of, causing modifications of the facial bones, of the early progenitors of man; greater variability of the, in men than in women.

Musculus sternalis, Prof. Turner on the.

Music, of birds; discordant, love of savages for; reason of power of perception of notes in animals; power of distinguishing notes; its connection with primeval speech; different appreciation of, by different peoples; origin of; effects of.

Musical cadences, perception of, by animals; powers of man.

Musk-deer, canine teeth of male; male, odoriferous organs of the; winter change of the.

Musk-duck, Australian; large size of male; of Guiana, pugnacity of the male.

Musk-ox, horns of.

Musk-rat, protective resemblance of the, to a clod of earth.

Musophagae, colours and nidification of the; both sexes of, equally brilliant.

Mussels opened by monkeys.

Mustela, winter change of two species of.

Musters, Captain, on Rhea Darwinii; marriages amongst Patagonians.

Mutilations, healing of; inheritance of.

Mutilla europaea, stridulation of.

Mutillidae, absence of ocelli in female.

Mycetes caraya, polygamous; vocal organs of; beard of; sexual differences of colour in; voice of.

Mycetes seniculus, sexual differences of colour in.

Myriapoda.

Nageli, on the influence of natural selection on plants; on the gradation of species of plants.

Nails, coloured yellow or purple in part of Africa.

Narwhal, tusks of the.

Nasal cavities, large size of, in American aborigines.

Nascent organs.

Nathusius, H. von, on the improved breeds of pigs; male domesticated animals more variable than females; horns of castrated sheep; on the breeding of domestic animals.

Natural selection, its effects on the early progenitors of man; influence of, on man; limitation of the principle; influence of, on social animals; Mr. Wallace on the limitation of, by the influence of the mental faculties in man; influence of, in the progress of the United States; in relation to sex.

Natural and sexual selection contrasted.

Naulette, jaw from, large size of the canines in.

Neanderthal skull, capacity of the.

Neck, proportion of, in soldiers and sailors.

Necrophorus, stridulation of.

Nectarinia, young of.

Nectariniae, moulting of the; nidification of.

Negro, resemblance of a, to Europeans in mental characters.

Negro-women, their kindness to Mungo Park.

Negroes, Caucasian features in; character of; lice of; fertility of, when crossed with other races; blackness of; variability of; immunity of, from yellow fever; difference of, from Americans; disfigurements of the; colour of new-born children of; comparative beardlessness of; readily become musicians; appreciation of beauty of their women by; idea of beauty among; compression of the nose by some.

Nemertians, colours of.

Neolithic period.

Neomorpha, sexual difference of the beak in.

Nephila, size of male.

Nests, made by fishes; decoration of, by Humming-birds.

Neumeister, on a change of colour in pigeons after several moultings.

Neuration, difference of, in the two sexes of some butterflies and hymenoptera.

Neuroptera.

Neurothemis, dimorphism in.

New Zealand, expectation by the natives of, of their extinction; practice of tattooing in; aversion of natives of, to hairs on the face; pretty girls engrossed by the chiefs in.

Newton, A., on the throat-pouch of the male bustard; on the differences between the females of two species of Oxynotus; on the habits of the Phalarope, dotterel, and godwit.

Newts.

Nicholson, Dr., on the non-immunity of dark Europeans from yellow fever.

Nictitating membrane.

Nidification of fishes; relation of, to colour; of British birds.

Night-heron, cries of the.

Nightingale, arrival of the male before the female; object of the song of the.

Nightingales, new mates found by.

Nightjar, selection of a mate by the female; Australian, sexes of; coloration of the.

Nightjars, noise made by some male, with their wings; elongated feathers in.

Nilghau, sexual differences of colour in the.

Nilsson, Prof., on the resemblance of stone arrow-heads from various places; on the development of the horns of the reindeer.

Nipples, absence of, in Monotremata.

Nitsche, Dr., ear of foetal orang.

Nitzsch, C.L., on the down of birds.

Noctuae, brightly-coloured beneath.

Noctuidae, coloration of.

Nomadic habits, unfavourable to human progress.

Nordmann, A., on Tetrao urogalloides.

Norfolk Island, half-breeds on.

Norway, numerical proportion of male and female births in.

Nose, resemblance of, in man and the apes; piercing and ornamentation of the; very flat, not admired in negroes; flattening of the.

Nott and Gliddon, on the features of Rameses II.; on the features of Amunoph III.; on skulls from Brazilian caves; on the immunity of negroes and mulattoes from yellow fever; on the deformation of the skull among American tribes.

Novara, voyage of the, suicide in New Zealand.

Nudibranch Mollusca, bright colours of.

Numerals, Roman.

Nunemaya, natives of, bearded.

Nuthatch, of Japan, intelligence of; Indian.

Obedience, value of.

Observation, powers of, possessed by birds.

Occupations, sometimes a cause of diminished stature; effect of, upon the proportions of the body.

Ocelli, absence of, in female Mutilidae.

Ocelli of birds, formation and variability of the.

Ocelot, sexual differences in the colouring of the.

Ocyhaps lophotes.

Odonata.

Odonestis potatoria, sexual difference of colour in.

Odour, correlation of, with colour of skin; of moths; emitted by snakes in the breeding season; of mammals.

Oecanthus nivalis, difference of colour in the sexes of.

Oecanthus pellucidus.

Ogle, Dr. W., relation between colour and power of smell.

Oidemia.

Oliver, on sounds produced by Pimelia striata.

Omaloplia brunnea, stridulation of.

Onitis furcifer, processes of anterior femora of the male, and on the head and thorax of the female.

Onthophagus.

Onthophagus rangifer, sexual differences of; variations in the horns of the male.

Ophidia, sexual differences of.

Ophidium.

Opossum, wide range of, in America.

Optic nerve, atrophy of the, caused by destruction of the eye.

Orang-Outan, Bischoff on the agreement of the brain of the, with that of man; adult age of the; ears of the; vermiform appendage of; hands of the; absence of mastoid processes in the; platforms built by the; alarmed at the sight of a turtle; using a stick as a lever; using missiles; using the leaves of the Pandanus as a night covering; direction of the hair on the arms of the; its aberrant characters; supposed evolution of the; voice of the; monogamous habits of the; male, beard of the.

Oranges, treatment of, by monkeys.

Orange-tip butterfly.

Orchestia Darwinii, dimorphism of males of.

Orchestia Tucuratinga, limbs of.

Ordeal, trial by.

Oreas canna, colours of.

Oreas Derbianus, colours of.

Organs, prehensile; utilised for new purposes.

Organic scale, von Baer’s definition of progress in.

Orioles, nidification of.

Oriolus, species of, breeding in immature plumage.

Oriolus melanocephalus, coloration of the sexes in.

Ornaments, prevalence of similar; of male birds; fondness of savages for.

Ornamental characters, equal transmission of, to both sexes, in mammals; of monkeys.

Ornithoptera croesus.

Ornithorhynchus, reptilian tendency of; spur of the male.

Orocetes erythrogastra, young of.

Orrony, Grotto of.

Orsodacna atra, difference of colour in the sexes of.

Orsodacna ruficollis.

Orthoptera, metamorphosis of; stridulating apparatus of; colours of; rudimentary stridulating organs in female; stridulation of the, and Homoptera, discussed.

Ortygornis gularis, pugnacity of the male.

Oryctes, stridulation of; sexual differences in the stridulant organs of.

Oryx leucoryx, use of the horns of.

Osphranter rufus, sexual difference in the colour of.

Ostrich, African, sexes and incubation of the.

Ostriches, stripes of young.

Otaria jubata, mane of the male.

Otaria nigrescens, difference in the coloration of the sexes of.

Otis bengalensis, love-antics of the male.

Otis tarda, throat-pouch of the male; polygamous.

Ouzel, ring-, colours and nidification of the.

Ouzel, water-, singing in the autumn; colours and nidification of the.

Ovibos moschatus, horns of.

Ovipositor of insects.

Ovis cycloceros, mode of fighting of.

Ovule of man.

Owen, Prof., on the Corpora Wolffiana; on the great toe in man; on the nictitating membrane and semilunar fold; on the development of the posterior molars in different races of man; on the length of the caecum in the Koala; on the coccygeal vertebrae; on rudimentary structures belonging to the reproductive system; on abnormal conditions of the human uterus; on the number of digits in the Ichthyopterygia; on the canine teeth in man; on the walking of the chimpanzee and orang; on the mastoid processes in the higher apes; on the hairiness of elephants in elevated districts; on the caudal vertebrae of monkeys; classification of mammalia; on the hair in monkeys; on the piscine affinities of the Ichthyosaurians; on polygamy and monogamy among the antelopes; on the horns of Antilocapra Americana; on the musky odour of crocodiles during the breeding season; on the scent-glands of snakes; on the Dugong, Cachalot, and Ornithorhynchus; on the antlers of the red deer; on the dentition of the Camelidae; on the horns of the Irish elk; on the voice of the giraffe, porcupine, and stag; on the laryngeal sac of the gorilla and orang; on the odoriferous glands of mammals; on the effects of emasculation on the vocal organs of men; on the voice of Hylobates agilis; on American monogamous monkeys.

Chapter XXI: General a Summary and Conclusion (part 3)

Owls, white, new mates found by.

Oxynotus, difference of the females of two species of.

Pachydermata.

Pachytylus migratorius.

Paget, on the abnormal development of hairs in man; on the thickness of the skin on the soles of the feet of infants.

Pagurus, carrying the female.

Painting, pleasure of savages in.

Palaemon, chelae of a species of.

Palaeornis, sexual differences of colour in.

Palaeornis javanicus, colour of beak of.

Palaeornis rosa, young of.

Palamedea cornuta, spurs on the wings.

Paleolithic period.

Palestine, habits of the chaffinch in.

Pallas, on the perfection of the senses in the Mongolians; on the want of connexion between climate and the colour of the skin; on the polygamous habits of Antilope Saiga; on the lighter colour of horses and cattle in winter in Siberia; on the tusks of the musk-deer; on the odoriferous glands of mammals; on the odoriferous glands of the musk-deer; on winter changes of colour in mammals; on the ideal of female beauty in North China.

Palmaris accessorius, muscle variations of the.

Pampas, horses of the.

Pangenesis, hypothesis of.

Panniculus carnosus.

Pansch, on the brain of a foetal Cebus apella.

Papilio, proportion of the sexes in North American species of; sexual differences of colouring in species of; coloration of the wings in species of.

Papilio ascanius.

Papilio Sesostris and Childrenae, variability of.

Papilio Turnus.

Papilionidae, variability in the.

Papuans, line of separation between the, and the Malays; beards of the; teeth of.

Papuans and Malays, contrast in characters of.

Paradise, Birds of; supposed by Lesson to be polygamous; rattling of their quills by; racket-shaped feathers in; sexual differences in colour of; decomposed feathers in; display of plumage by the male; sexual differences in colour of.

Paradisea apoda, barbless feathers in the tail of; plumage of; and P. papuana; divergence of the females of; increase of beauty with age.

Paradisea papuana, plumage of.

Paraguay, Indians of, eradication of eyebrows and eyelashes by.

Parallelism of development of species and languages.

Parasites, on man and animals; as evidence of specific identity or distinctness; immunity from, correlated with colour.

Parental feeling in earwigs, starfishes, and spiders; affection, partly a result of natural selection.

Parents, age of, influence upon sex of offspring.

Parinae, sexual difference of colour in.

Park, Mungo, negro-women teaching their children to love the truth; his treatment by the negro-women; on negro opinions of the appearance of white men.

Parker, Mr., no bird or reptile in line of mammalian descent.

Parrakeet, young of; Australian, variation in the colour of the thighs of a male.

Parrot, racket-shaped feathers in the tail of a; instance of benevolence in a.

Parrots, change of colour in; imitative faculties of; living in triplets; affection of; colours and nidification of the; immature plumage of the; colours of; sexual differences of colour in; musical powers of.

Parthenogenesis in the Tenthredinae; in Cynipidae; in Crustacea.

Partridge, monogamous; proportion of the sexes in the; Indian; female.

Partridge-“dances.”

Partridges, living in triplets; spring coveys of male; distinguishing persons.

Parus coeruleus.

Passer, sexes and young of.

Passer brachydactylus.

Passer domesticus.

Passer montanus.

Patagonians, self-sacrifice by; marriages of.

Patterson, Mr., on the Agrionidae.

Patteson, Bishop, decrease of Melanesians.

Paulistas of Brazil.

Pavo cristatus.

Pavo muticus, possession of spurs by the female.

Pavo nigripennis.

Payaguas Indians, thin legs and thick arms of the.

Payan, Mr., on the proportion of the sexes in sheep.

Peacock, polygamous; sexual characters of; pugnacity of the; Javan, possessing spurs; rattling of the quills by; elongated tail-coverts of the; love of display of the; ocellated spots of the; inconvenience of long tail of the, to the female; continued increase of beauty of the.

Peacock-butterfly.

Peafowl, preference of females for a particular male; first advances made by the female.

Pediculi of domestic animals and man.

Pedigree of man.

Pedionomus torquatus, sexes of.

Peel, J., on horned sheep.

Peewit, wing-tubercles of the male.

Pelagic animals, transparency of.

Pelecanus erythrorhynchus, horny crest on the beak of the male, during the breeding season.

Pelecanus onocrotalus, spring plumage of.

Pelele, an African ornament.

Pelican, blind, fed by his companions; young, guided by old birds; pugnacity of the male.

Pelicans, fishing in concert.

Pelobius Hermanni, stridulation of.

Pelvis, alteration of, to suit the erect attitude of man; differences of the, in the sexes of man.

Penelope nigra, sound produced by the male.

Pennant, on the battles of seals; on the bladder-nose seal.

Penthe, antennal cushions of the male.

Perch, brightness of male, during breeding season.

Peregrine falcon, new mate found by.

Period of variability, relation of, to sexual selection.

Periodicity, vital, Dr. Laycock on.

Periods, lunar, followed by functions in man and animals.

Periods of life, inheritance at corresponding.

Perisoreus canadensis, young of.

Peritrichia, difference of colour in the sexes of a species of.

Periwinkle.

Pernis cristata.

Perrier, M., on sexual selection; on bees.

Perseverance, a characteristic of man.

Persians, said to be improved by intermixture with Georgians and Circassians.

Personnat, M., on Bombyx Yamamai.

Peruvians, civilisation of the, not foreign.

Petrels, colours of.

Petrocincla cyanea, young of.

Petrocossyphus.

Petronia.

Pfeiffer, Ida, on Javan ideas of beauty.

Phacochoerus aethiopicus, tusks and pads of.

Phalanger, Vulpine, black varieties of the.

Phalaropus fulicarius.

Phalaropus hyperboreus.

Phanaeus.

Phanaeus carnifex, variation of the horns of the male.

Phanaeus faunus, sexual differences of.

Phanaeus lancifer.

Phaseolarctus cinereus, taste for rum and tobacco.

Phasgonura viridissima, stridulation of.

Phasianus Soemmerringii.

Phasianus versicolor.

Phasianus Wallichii.

Pheasant, polygamous; and black grouse, hybrids of; production of hybrids with the common fowl; immature plumage of the.

Pheasant, Amherst, display of.

Pheasant, Argus, display of plumage by the male; ocellated spots of the; gradation of characters in the.

Pheasant, Blood- Pheasant, Cheer.

Pheasant, Eared, length of the tail in the; sexes alike in the.

Pheasant, Fire-backed, possessing spurs.

Pheasant, Golden, display of plumage by the male; age of mature plumage in the; sex of young, ascertained by pulling out head-feathers.

Pheasant, Kalij, drumming of the male.

Pheasant, Reeve’s, length of the tail in.

Pheasant, Silver, triumphant male, deposed on account of spoiled plumage; sexual coloration of the.

Pheasant, Soemmerring’s.

Pheasant, Tragopan, display of plumage by the male; marking of the sexes of the.

Pheasants, period of acquisition of male characters in the family of the; proportion of sexes in chicks of; length of the tail in.

Philters, worn by women.

Phoca groenlandica, sexual difference in the coloration of.

Phoenicura ruticilla.

Phosphorescence of insects.

Phryganidae, copulation of distinct species of.

Phryniscus nigricans.

Physical inferiority, supposed, of man.

Pickering, on the number of species of man.

Picton, J.A., on the soul of man.

Picus auratus.

Picus major.

Pieris.

Pigeon, female, deserting a weakened mate; carrier, late development of the wattle in; pouter, late development of crop in; domestic, breeds and sub-breeds of.

Pigeons, nestling, fed by the secretion of the crop of both parents;

changes of plumage in; transmission of sexual peculiarities in; Belgian, with black-streaked males; changing colour after several moultings; numerical proportion of the sexes in; cooing of; variations in plumage of; display of plumage by male; local memory of; antipathy of female, to certain males; pairing of; profligate male and female; wing-bars and tail-feathers of; supposititious breed of; pouter and carrier, peculiarities of, predominant in males; nidification of; Australian; immature plumage of the.

Pigs, origin of the improved breeds of; numerical proportion of the sexes in; stripes of young; tusks of miocene; sexual preference shewn by.

Pike, American, brilliant colours of the male, during the breeding season.

Pike, reasoning powers of; male, devoured by females.

Pike, L.O., on the psychical elements of religion.

Pimelia striata, sounds produced by the female.

Pinel, hairiness in idiots.

Pintail, drake, plumage of; pairing with a wild duck.

Pintail Duck, pairing with a widgeon.

Pipe-fish, filamentous; marsupial receptacles of the male.

Pipits, moulting of the.

Pipra, modified secondary wing-feathers of male.

Pipra deliciosa.

Pirates stridulus, stridulation of.

Pitcairn island, half-breeds on.

Pithecia leucocephala, sexual differences of colour in.

Pithecia Satanas, beard of; resemblance of, to a negro.

Pits, suborbital, of Ruminants.

Pittidae, nidification of.

Placentata.

Plagiostomous fishes.

Plain-wanderer, Australian.

Planariae, bright colours of some.

Plantain-eaters, colours and nidification of the; both sexes of, equally brilliant.

Plants, cultivated, more fertile than wild; Nageli, on natural selection in; male flowers of, mature before the female; phenomena of fertilisation in.

Platalea, change of plumage in.

Platyblemus.

Platycercus, young of.

Platyphyllum concavum.

Platyrrhine monkeys.

Platysma myoides.

Plecostomus, head-tentacles of the males of a species of.

Plecostomus barbatus, peculiar beard of the male.

Plectropterus gambensis, spurred wings of.

Ploceus.

Plovers, wing-spurs of; double moult in.

Plumage, changes of, inheritance of, by fowls; tendency to analogous variation in; display of, by male birds; changes of, in relation to season; immature, of birds; colour of, in relation to protection.

Plumes on the head in birds, difference of, in the sexes.

Pneumora, structure of.

Podica, sexual difference in the colour of the irides.

Poeppig, on the contact of civilised and savage races.

Poison, avoidance of, by animals.

Poisonous fruits and herbs avoided by animals.

Poisons, immunity from, correlated with colour.

Polish fowls, origin of the crest in.

Pollen and van Dam, on the colours of Lemur macaco.

Polyandry, in certain Cyprinidae; among the Elateridae.

Polydactylism in man.

Polygamy, influence of, upon sexual selection; superinduced by domestication; supposed increase of female births by. In the stickleback.

Polygenists.

Polynesia, prevalence of infanticide in.

Polynesians, wide geographical range of; difference of stature among the; crosses of; variability of; heterogeneity of the; aversion of, to hairs on the face.

Polyplectron, number of spurs in; display of plumage by the male; gradation of characters in; female of.

Polyplectron chinquis.

Polyplectron Hardwickii.

Polyplectron malaccense.

Polyplectron Napoleonis.

Polyzoa.

Pomotis.

Pontoporeia affinis.

Porcupine, mute, except in the rutting season.

Pores, excretory, numerical relation of, to the hairs in sheep.

Porpitae, bright colours of some.

Portax picta, dorsal crest and throat-tuft of; sexual differences of colour in.

Portunus puber, pugnacity of.

Potamochoerus pencillatus, tusks and facial knobs of the.

Pouchet, G., the relation of instinct to intelligence; on the instincts of ants; on the caves of Abou-Simbel; on the immunity of negroes from yellow fever; change of colour in fishes.

Pouter pigeon, late development of the large crop in.

Powell, Dr., on stridulation.

Power, Dr., on the different colours of the sexes in a species of Squilla.

Powys, Mr., on the habits of the chaffinch in Corfu.

Pre-eminence of man.

Preference for males by female birds; shewn by mammals, in pairing.

Prehensile organs.

Presbytis entellus, fighting of the male.

Preyer, Dr., on function of shell of ear; on supernumerary mammae in women.

Prichard, on the difference of stature among the Polynesians; on the connection between the breadth of the skull in the Mongolians and the perfection of their senses; on the capacity of British skulls of different ages; on the flattened heads of the Colombian savages; on Siamese notions of beauty; on the beardlessness of the Siamese; on the deformation of the head among American tribes and the natives of Arakhan.

Primary sexual organs.

Primates, sexual differences of colour in.

Primogeniture, evils of.

Prionidae, difference of the sexes in colour.

Proctotretus multimaculatus.

Proctotretus tenuis, sexual difference in the colour of.

Profligacy.

Progenitors, early, of man.

Progress, not the normal rule in human society; elements of.

Prong-horn antelope, horns of.

Proportions, difference of, in distinct races.

Protective colouring in butterflies; in lizards; in birds; in mammals.

Protective nature of the dull colouring of female Lepidoptera.

Protective resemblances in fishes.

Protozoa, absence of secondary sexual characters in.

Pruner-Bey, on the occurrence of the supra-condyloid foramen in the humerus of man; on the colour of negro infants.

Prussia, numerical proportion of male and female births in.

Psocus, proportions of the sexes in.

Ptarmigan, monogamous; summer and winter plumage of the; nuptial assemblages of; triple moult of the; protective coloration of.

Puff-birds, colours and nidification of the.

Pugnacity of fine-plumaged male birds.

Pumas, stripes of young.

Puppies learning from cats to clean their faces.

Pycnonotus haemorrhous, pugnacity of the male; display of under-tail coverts by the male.

Pyranga aestiva, male aiding in incubation; male characters in female of.

Pyrodes, difference of the sexes in colour.

Quadrumana, hands of; differences between man and the; sexual differences of colour in; ornamental characters of; analogy of sexual differences of, with those of man; fighting of males for the females; monogamous habits of; beards of the.

Quain, R., on the variation of the muscles in man.

Quatrefages, A. de, on the occurrence of a rudimentary tail in man; on variability; on the moral sense as a distinction between man and animals; civilised men stronger than savages; on the fertility of Australian women with white men; on the Paulistas of Brazil; on the evolution of the breeds of cattle; on the Jews; on the liability of negroes to tropical fevers after residence in a cold climate; on the difference between field-and house-slaves; on the influence of climate on colour; colours of annelids; on the Ainos; on the women of San Giuliano.

Quechua, see Quichua.

Querquedula acuta.

Quetelet, proportion of sexes in man; relative size in man and woman.

Quichua Indians; local variation of colour in the; no grey hair among the; hairlessness of the; long hair of the.

Quiscalus major, proportions of the sexes of, in Florida and Honduras.

Rabbit, white tail of the.

Rabbits, domestic, elongation of the skull in; modification of the skull in, by the lopping of the ear; danger-signals of; numerical proportion of the sexes in.

Races, distinctive characters of; or species of man; crossed, fertility or sterility of; of man, variability of the; of man, resemblance of, in mental characters; formation of; of man, extinction of; effects of the crossing of; of man, formation of the; of man, children of the; beardless, aversion of, to hairs on the face.

Raffles, Sir S., on the banteng.

Rafts, use of.

Rage, manifested by animals.

Raia batis, teeth of.

Raia clavata, female spined on the back; sexual difference in the teeth of.

Raia maculata, teeth of.

Rails, spur-winged.

Ram, mode of fighting of the; African, mane of an; fat-tailed.

Rameses II., features of.

Ramsay, Mr., on the Australian musk-duck; on the regent-bird; on the incubation of Menura superba.

Rana esculenta, vocal sacs of.

Rat, common, general dispersion of, a consequence of superior cunning; supplantation of the native in New Zealand, by the European rat; common, said to be polygamous; numerical proportion of the sexes in.

Rats, enticed by essential oils.

Rationality of birds.

Rattlesnakes, difference of the sexes in the; rattles as a call.

Raven, vocal organs of the; stealing bright objects; pied, of the Feroe Islands.

Rays, prehensile organs of male.

Razor-bill, young of the.

Reade, Winwood, suicide among savages in Africa; mulattoes not prolific; effect of castration of horned sheep; on the Guinea sheep; on the occurrence of a mane in an African ram; on singing of negroes; on the negroes’ appreciation of the beauty of their women; on the admiration of negroes for a black skin; on the idea of beauty among negroes; on the Jollofs; on the marriage-customs of the negroes.

Reason in animals.

Redstart, American, breeding in immature plumage.

Redstarts, new mates found by.

Reduvidae, stridulation of.

Reed-bunting, head-feathers of the male; attacked by a bullfinch.

Reefs, fishes frequenting.

Reeks, H., retention of horns by breeding deer; cow rejected by a bull; destruction of piebald rabbits by cats.

Regeneration, partial, of lost parts in man.

Regent bird.

Reindeer, horns of the; battles of; horns of the female; antlers of, with numerous points; winter change of the; sexual preferences shown by.

Relationship, terms of.

Religion, deficiency of among certain races; psychical elements of.

Remorse, deficiency of, among savages.

Rengger, on the diseases of Cebus Azarae; on the diversity of the mental faculties of monkeys; on the Payaguas Indians; on the inferiority of Europeans to savages in their senses; revenge taken by monkeys; on maternal affection in a Cebus; on the reasoning powers of American monkeys; on the use of stones by monkeys for cracking hard nuts; on the sounds uttered by Cebus Azarae; on the signal-cries of monkeys; on the polygamous habits of Mycetes caraya; on the voice of the howling monkeys; on the odour of Cervus campestris; on the beards of Mycetes caraya and Pithecia Satanas; on the colours of Felis mitis; on the colours of Cervus paludosus; on sexual differences of colour in Mycetes; on the colour of the infant Guaranys; on the early maturity of the female of Cebus Azarae; on the beards of the Guaranys; on the emotional notes employed by monkeys; on American polygamous monkeys.

Representative species, of birds.

Reproduction, unity of phenomena of, throughout the mammalia; period of, in birds.

Reproductive system, rudimentary structures in the; accessory parts of.

Reptiles.

Reptiles and birds, alliance of.

Resemblances, small, between man and the apes.

Retrievers, exercise of reasoning faculties by.

Revenge, manifested by animals.

Reversion, perhaps the cause of some bad dispositions.

Rhagium, difference of colour in the sexes of a species of.

Rhamphastos carinatus.

Rhea Darwinii.

Rhinoceros, nakedness of; horns of; horns of, used defensively; attacking white or grey horses.

Rhynchaea, sexes and young of.

Rhynchaea australis.

Rhynchaea bengalensis.

Rhynchaea capensis.

Rhythm, perception of, by animals.

Richard, M., on rudimentary muscles in man.

Richardson, Sir J., on the pairing of Tetrao umbellus; on Tetrao urophasianus; on the drumming of grouse; on the dances of Tetrao phasianellus; on assemblages of grouse; on the battles of male deer; on the reindeer; on the horns of the musk-ox; on antlers of the reindeer with numerous points; on the moose; on the Scotch deerhound.

Richter, Jean Paul, on imagination.

Riedel, on profligate female pigeons.

Riley, Mr., on mimicry in butterflies; bird’s disgust at taste of certain caterpillars.

Ring-ouzel, colours and nidification of the.

Ripa, Father, on the difficulty of distinguishing the races of the Chinese.

Rivalry, in singing, between male birds.

River-hog, African, tusks and knobs of the.

Rivers, analogy of, to islands.

Roach, brightness of the male during breeding-season.

Robbery, of strangers, considered honourable.

Robertson, Mr., remarks on the development of the horns in the roebuck and red deer.

Robin, pugnacity of the male; autumn song of the; female singing of the; attacking other birds with red in their plumage; young of the.

Robinet, on the difference of size of the male and female cocoons of the silk-moth.

Rodents, uterus in the; absence of secondary sexual characters in; sexual differences in the colours of.

Roe, winter changes of the.

Rohfs, Dr., Caucasian features in negro; fertility of mixed races in Sahara; colours of birds in Sahara; ideas of beauty amongst the Bornuans.

Rolle, F., on the origin of man; on a change in German families settled in Georgia.

Roller, harsh cry of.

Romans, ancient, gladiatorial exhibitions of the.

Rook, voice of the.

Rossler, Dr., on the resemblance of the lower surface of butterflies to the bark of trees.

Rostrum, sexual difference in the length of in some weevils.

Royer, Madlle., mammals giving suck.

Rudimentary organs, origin of.

Rudiments, presence of, in languages.

Rudolphi, on the want of connexion between climate and the colour of the skin.

Ruff, supposed to be polygamous; proportion of the sexes in the; pugnacity of the; double moult in; duration of dances of; attraction of the, to bright objects.

Ruminants, male, disappearance of canine teeth in; generally polygamous; suborbital pits of; sexual differences of colour in.

Rupicola crocea, display of plumage by the male.

Ruppell, on canine teeth in deer and antelopes.

Russia, numerical proportion of male and female births in.

Ruticilla.

Rutimeyer, Prof., on the physiognomy of the apes; on tusks of miocene boar; on the sexual differences of monkeys.

Rutlandshire, numerical proportion of male and female births in.

Sachs, Prof., on the behaviour of the male and female elements in fertilisation.

Sacrifices, human.

Sagittal crest, in male apes and Australians.

Sahara, fertility of mixed races in; birds of the; animal inhabitants of the.

Sailors, growth of, delayed by conditions of life; long-sighted.

Sailors and soldiers, difference in the proportions of.

St. John, Mr., on the attachment of mated birds.

St. Kilda, beards of the inhabitants of.

Salmo eriox, and Salmo umbla, colouring of the male, during the breeding season.

Salmo lycaodon.

Salmo salar.

Salmon, leaping out of fresh water; male, ready to breed before the female; proportion of the sexes in; male, pugnacity of the; male, characters of, during the breeding season; spawning of the; breeding of immature male.

Salvin, O., inheritance of mutilated feathers; on the Humming-birds; on the numerical proportion of the sexes in Humming-birds; on Chamaepetes and Penelope; on Selasphorus platycercus; Pipra deliciosa; on Chasmorhynchus.

Samoa Islands, beardlessness of the natives of.

Sandhoppers, claspers of male.

Sand-skipper.

Sandwich Islands, variation in the skulls of the natives of the; decrease of native population; population of; superiority of the nobles in the.

Sandwich Islanders, lice of.

San-Giuliano, women of.

Santali, recent rapid increase of the; Mr. Hunter on the.

Saphirina, characters of the males of.

Sarkidiornis melanonotus, characters of the young.

Sars, O., on Pontoporeia affinis.

Saturnia carpini, attraction of males by the female.

Saturnia Io, difference of coloration in the sexes of.

Saturniidae, coloration of the.

Savage, Dr., on the fighting of the male gorillas; on the habits of the gorilla.

Savage and Wyman on the polygamous habits of the gorilla.

Savages, uniformity of, exaggerated; long-sighted; rate of increase among, usually small; retention of the prehensile power of the feet by; imitative faculties of; causes of low morality of; tribes of, supplanting one another; improvements in the arts among; arts of; fondness of, for rough music; on long-enduring fashions among; attention paid by, to personal appearance; relation of the sexes among.

Saviotti, Dr., division of malar bone.

Saw-fly, pugnacity of a male.

Saw-flies, proportions of the sexes in.

Saxicola rubicola, young of.

Scalp, motion of the.

Scent-glands in snakes.

Schaaffhausen, Prof., on the development of the posterior molars in different races of man; on the jaw from La Naulette; on the correlation between muscularity and prominent supra-orbital ridges; on the mastoid processes of man; on modifications of the cranial bones; on human sacrifices; on the probable speedy extermination of the anthropomorphous apes; on the ancient inhabitants of Europe; on the effects of use and disuse of parts; on the superciliary ridge in man; on the absence of race-differences in the infant skull in man; on ugliness.

Schaum, H., on the elytra of Dytiscus and Hydroporus.

Scherzer and Schwarz, measurements of savages.

Schelver, on dragon-flies.

Schiodte, on the stridulation of Heterocerus.

Schlegel, F. von, on the complexity of the languages of uncivilised peoples.

Schlegel, Prof., on Tanysiptera.

Schleicher, Prof, on the origin of language.

Schomburgk, Sir R., on the pugnacity of the male musk-duck of Guiana; on the courtship of Rupicola crocea.

Schoolcraft, Mr., on the difficulty of fashioning stone implements.

Schopenhauer, on importance of courtship to mankind.

Schweinfurth, complexion of negroes.

Sciaena aquila.

Sclater, P.L., on modified secondary wing-feathers in the males of Pipra; on elongated feathers in nightjars; on the species of Chasmorhynchus; on the plumage of Pelecanus onocrotalus; on the plantain-eaters; on the sexes and young of Tadorna variegata; on the colours of Lemur macaco; on the stripes in asses.

Scolecida, absence of secondary sexual characters in.

Scolopax frenata, tail feathers of;

Scolopax gallinago, drumming of.

Scolopax javensis, tail-feathers of.

Scolopax major, assemblies of.

Scolopax Wilsonii, sound produced by.

Scolytus, stridulation of.

Scoter-duck, black, sexual difference in coloration of the; bright beak of male.

Scott, Dr., on idiots smelling their food.

Scott, J., on the colour of the beard in man.

Scrope, on the pugnacity of the male salmon; on the battles of stags.

Scudder, S.H., imitation of the stridulation of the Orthoptera; on the stridulation of the Acridiidae; on a Devonian insect; on stridulation.

Sculpture, expression of the ideal of beauty by.

Sea-anemones, bright colours of.

Sea-bear, polygamous.

Sea-elephant, male, structure of the nose of the; polygamous.

Sea-lion, polygamous.

Seal, bladder-nose.

Seals, their sentinels generally females; evidence furnished by, on classification; polygamous habits of; battles of male; canine teeth of male; sexual differences; pairing of; sexual peculiarities of; in the coloration of; appreciation of music by.

Sea-scorpion, sexual differences in.

Season, changes of colour in birds, in accordance with the; changes of plumage of birds in relation to.

Seasons, inheritance at corresponding.

Sebituani, African chief, trying to alter a fashion.

Sebright Bantam.

Secondary sexual characters; relations of polygamy to; transmitted through both sexes; gradation of, in birds.

Sedgwick, W., on hereditary tendency to produce twins.

Seemann, Dr., on the different appreciation of music by different peoples; on the effects of music.

Seidlitz, on horns of reindeer.

Selasphorus platycercus, acuminate first primary of the male.

Selby, P.J., on the habits of the black and red grouse.

Selection as applied to primeval man.

Selection, double.

Selection, injurious forms of, in civilised nations.

Selection of male by female birds.

Selection, methodical, of Prussian grenadiers.

Selection, sexual, explanation of; influence of, on the colouring of Lepidoptera.

Selection, sexual and natural, contrasted.

Self-command, habit of, inherited; estimation of.

Self-consciousness, in animals.

Self-preservation, instinct of.

Self-sacrifice, by savages; estimation of.

Semilunar fold.

Semnopithecus, long hair on the heads of species of.

Semnopithecus chrysomelas, sexual differences of colour in.

Semnopithecus comatus, ornamental hair on the head of.

Semnopithecus frontatus, beard etc., of.

Semnopithecus nasica, nose of.

Semnopithecus nemaeus, colouring of.

Semnopithecus rubicundus, ornamental hair on the head of.

Senses, inferiority of Europeans to savages in the.

Sentinels, among animals.

Serpents, instinctively dreaded by apes and monkeys.

Serranus, hermaphroditism in.

Setina, noise produced by.

Sex, inheritance limited by.

Sexes, relative proportions of, in man; proportions of, sometimes influenced by selection; probable relation of the, in primeval man.

Sexual and natural selection, contrasted.

Sexual characters, effects of the loss of; limitation of.

Sexual characters, secondary; relations of polygamy to; transmitted through both sexes; gradation of, in birds.

Sexual differences in man.

Sexual selection, explanation of; influence of, on the colouring of Lepidoptera; objections to; action of, in mankind.

Sexual selection in spiders.

Sexual selection, supplemental note on.

Sexual similarity.

Shaler, Prof., sizes of sexes in whales.

Shame.

Sharks, prehensile organs of male.

Sharpe, Dr., Europeans in the tropics.

Sharpe, R.B., on Tanysiptera sylvia; on Ceryle; on the young male of Dacelo Gaudi-chaudi.

Shaw, Mr., on the pugnacity of the male salmon.

Shaw, J., on the decorations of birds.

Sheep, danger-signals of; sexual differences in the horns of; horns of; domestic, sexual differences of, late developed; numerical proportion of the sexes in; inheritance of horns by one sex; effect of castration; mode of fighting of; arched foreheads of some.

Sheep, Merino, loss of horns in females of; horns of.

Shells, difference in form of, in male and female Gasteropoda; beautiful colours and shapes of.

Shield-drake, pairing with a common duck; New Zealand, sexes and young of.

Shooter, J., on the Kaffirs; on the marriage-customs of the Kaffirs.

Shrew-mice, odour of.

Shrike, Drongo.

Shrikes, characters of young.

Shuckard, W.E., on sexual differences in the wings of Hymenoptera.

Shyness of adorned male birds;

Siagonium, proportions of the sexes in; dimorphism in males of.

Siam, proportion of male and female births in.

Siamese, general beardlessness of the; notions of beauty of the; hairy family of.

Sidgwick, H., on morality in hypothetical bee community; our actions not entirely directed by pain and pleasure.

Siebold, C.T., von, on the proportion of sexes in the Apus; on the auditory apparatus of the stridulent Orthoptera.

Sight, inheritance of long and short.

Signal-cries of monkeys.

Silk-moth, proportion of the sexes in; Ailanthus, Prof. Canestrini, on the destruction of its larvae by wasps; difference of size of the male and female cocoons of the; pairing of the.

Simiadae, their origin and divisions.

Similarity, sexual.

Singing of the Cicadae and Fulgoridae; of tree-frogs; of birds, object of the.

Sirenia, nakedness of.

Sirex juvencus.

Siricidae, difference of the sexes in.

Siskin, pairing with a canary.

Sitana, throat-pouch of the males of.

Size, relative, of the sexes of insects.

Skin, dark colour of, a protection against heat.

Skin, movement of the; nakedness of, in man; colour of the.

Skin and hair, correlation of colour of.

Skull, variation of, in man; cubic contents of, no absolute test of intellect; Neanderthal, capacity of the; causes of modification of the; difference of, in form and capacity, in different races of men; variability of the shape of the; differences of, in the sexes in man; artificial modification of the shape of.

Skunk, odour emitted by the; white tail of, protective.

Slavery, prevalence of; of women.

Slaves, difference between field-and house-slaves.

Sloth, ornaments of male.

Smell, sense of, in man and animals.

Smith, Adam, on the basis of sympathy.

Smith, Sir A., on the recognition of women by male Cynocephali; on revenge by a baboon; on an instance of memory in a baboon; on the retention of their colour by the Dutch in South Africa; on the polygamy of the South African antelopes; on the polygamy of the lion; on the proportion of the sexes in Kobus ellipsiprymnus; on Bucephalus capensis; on South African lizards; on fighting gnus; on the horns of rhinoceroses; on the fighting of lions; on the colours of the Cape Eland; on the colours of the gnu; on Hottentot notions of beauty; disbelief in communistic marriages.

Smith, F., on the Cynipidae and Tenthredinidae; on the relative size of the sexes of Aculeate Hymenoptera; on the difference between the sexes of ants and bees; on the stridulation of Trox sabulosus; on the stridulation of Mononychus pseudacori.

Smynthurus luteus, courtship of.

Snakes, sexual differences of; mental powers of; male, ardency of.

“Snarling muscles.”

Snipe, drumming of the; coloration of the.

Snipe, painted, sexes and young of.

Snipe, solitary, assemblies of.

Snipes, arrival of male before the female; pugnacity of male; double moult in.

Snow-goose, whiteness of the.

Sociability, the sense of duty connected with; impulse to, in animals; manifestations of, in man; instinct of, in animals.

Social animals, affection of, for each other; defence of, by the males.

Sociality, probable, of primeval men; influence of, on the development of the intellectual faculties; origin of, in man.

Soldiers, American, measurements of.

Soldiers and sailors, difference in the proportions of.

Solenostoma, bright colours and marsupial sac of the females of.

Song, of male birds appreciated by their females; want of, in brilliant plumaged birds; of birds.

Sorex, odour of.

Sounds, admired alike by man and animals; produced by fishes; produced by male frogs and toads; instrumentally produced by birds.

Spain, decadence of.

Sparassus smaragdulus, difference of colour in the sexes of.

Sparrow, pugnacity of the male; acquisition of the Linnet’s song by a; coloration of the; immature plumage of the.

Sparrow, white-crowned, young of the.

Sparrows, house-and tree-.

Sparrows, new mates found by.

Sparrows, sexes and young of; learning to sing.

Spathura Underwoodi.

Spawning of fishes.

Spear, used before dispersion of man.

Species, causes of the advancement of; distinctive characters of; or races of man; sterility and fertility of, when crossed; supposed, of man; gradation of; difficulty of defining; representative, of birds; of birds, comparative differences between the sexes of distinct.

Spectrum femoratum, difference of colour in the sexes of.

Speech, connection between the brain and the faculty of; connection of intonation with music.

Spel, of the black-cock.

Spencer, Herbert, on the influence of food on the size of the jaws; on the dawn of intelligence; on the origin of the belief in spiritual agencies; on the origin of the moral sense; on music.

Spengel, disagrees with explanation of man’s hairlessness.

Sperm-whales, battles of male.

Sphingidae, coloration of the.

Sphinx, Humming-bird.

Sphinx, Mr. Bates on the caterpillar of a.

Sphinx moth, musky odour of.

Spiders, parental feeling in; male, more active than female; proportion of the sexes in; secondary sexual characters of; courtship of male; attracted by music; male, small size of.

Spilosoma menthastri, rejected by turkeys.

Spine, alteration of, to suit the erect attitude of man.

Spirits, fondness of monkeys for.

Spiritual agencies, belief in, almost universal.

Spiza cyanea and ciris.

Spoonbill, Chinese, change of plumage in.

Spots, retained throughout groups of birds; disappearance of, in adult mammals.

Sprengel, C.K., on the sexuality of plants.

Springboc, horns of the.

Sproat, Mr., on the extinction of savages in Vancouver Island; on the eradication of facial hair by the natives of Vancouver Island; on the eradication of the beard by the Indians of Vancouver Island.

Spurs, occurrence of, in female fowls; development of, in various species of Phasianidae; of Gallinaceous birds; development of, in female Gallinaceae.

Squilla, different colours of the sexes of a species of.

Squirrels, battles of male; African, sexual differences in the colouring of; black.

Stag, long hairs of the throat of; horns of the; battles of; horns of the, with numerous branches; bellowing of the; crest of the.

Stag-beetle, numerical proportion of sexes of; use of jaws; large size of male; weapons of the male.

Stainton, H.T., on the numerical proportion of the sexes in the smaller moths; habits of Elachista rufocinerea; on the coloration of moths; on the rejection of Spilosoma menthastri by turkeys; on the sexes of Agrotis exclamationis.

Staley, Bishop, mortality of infant Maories.

Stallion, mane of the.

Stallions, two, attacking a third; fighting; small canine teeth of.

Stansbury, Captain, observations on pelicans.

Staphylinidae, hornlike processes in male.

Starfishes, parental feeling in; bright colours of some.

Stark, Dr., on the death-rate in towns and rural districts; on the influence of marriage on mortality; on the higher mortality of males in Scotland.

Starling, American field-, pugnacity of male.

Starling, red-winged, selection of a mate by the female.

Starlings, three, frequenting the same nest; new mates found by.

Statues, Greek, Egyptian, Assyrian, etc., contrasted.

Stature, dependence of, upon local influences.

Staudinger, Dr., on breeding Lepidoptera; his list of Lepidoptera.

Staunton, Sir G., hatred of indecency a modern virtue.

Stealing of bright objects by birds.

Stebbing, T.R., on the nakedness of the human body.

Stemmatopus.

Stendhal, see Bombet.

Stenobothrus pratorum, stridulation.

Stephen, Mr. L., on the difference in the minds of men and animals; on general concepts in animals; distinction between material and formal morality.

Sterility, general, of sole daughters; when crossed, a distinctive character of species; under changed conditions.

Sterna, seasonal change of plumage in.

Stickleback, polygamous; male, courtship of the; male, brilliant colouring of, during the breeding season; nidification of the.

Sticks used as implements and weapons by monkeys.

Sting in bees.

Stokes, Captain, on the habits of the great bower-bird.

Stoliczka, Dr., on colours in snakes.

Stoliczka, on the pre-anal pores of lizards.

Stonechat, young of the.

Stone implements, difficulty of making; as traces of extinct tribes.

Stones, used by monkeys for breaking hard fruits and as missiles; piles of.

Stork, black, sexual differences in the bronchi of the; red beak of the.

Storks, sexual difference in the colour of the eyes of.

Strange, Mr., on the satin bowerbird.

Strepsiceros kudu, horns of; markings of.

Stretch, Mr., on the numerical proportion in the sexes of chickens.

Stridulation, by males of Theridion; of Hemiptera; of the Orthoptera and Homoptera discussed; of beetles.

Stripes, retained throughout groups of birds; disappearance of, in adult mammals.

Strix flammea.

Structure, existence of unserviceable modifications of.

Struggle for existence, in man.

Struthers, Dr., on the occurrence of the supra-condyloid foramen in the humerus of man.

Sturnella ludoviciana, pugnacity of the male.

Sturnus vulgaris.

Sub-species.

Suffering, in strangers, indifference of savages to.

Suicide, formerly not regarded as a crime; rarely practised among the lowest savages.

Suidae, stripes of the young.

Sulivan, Sir B.J., on speaking of parrots; on two stallions attacking a third.

Sumatra, compression of the nose by the Malays of.

Sumner, Archb., man alone capable of progressive improvement.

Sun-birds, nidification of.

Superciliary ridge in man.

Supernumerary digits, more frequent in men than in women; inheritance of; early development of.

Superstitions, prevalence of.

Superstitious customs.

Supra-condyloid foramen in the early progenitors of man.

Suspicion, prevalence of, among animals.

Swallow-tail butterfly.

Swallows deserting their young.

Swan, black, wild, trachea of the; white, young of; red beak of the; black-necked.

Swans, young.

Swaysland, Mr., on the arrival of migratory birds.

Swifts, migration of.

Swinhoe, R., on the common rat in Formosa and China; behaviour of lizards when caught; on the sounds produced by the male hoopoe; on Dicrurus macrocercus and the spoonbill; on the young of Ardeola; on the habits of Turnix; on the habits of Rhynchaea bengalensis; on Orioles breeding in immature plumage.

Sylvia atricapilla, young of.

Sylvia cinerea, aerial love-dance of the male.

Sympathy, among animals; its supposed basis.

Sympathies, gradual widening of.

Syngnathous fishes, abdominal pouch in male.

Sypheotides auritus, acuminated primaries of the male; ear-tufts of.

Tabanidae, habits of.

Tadorna variegata, sexes and young of.

Tadorna vulpanser.

Tahitians, compression of the nose by the.

Tail, rudimentary, occurrence of, in man; convoluted body in the extremity of the; absence of, in man and the higher apes; variability of, in species of Macacus and in baboons; presence of, in the early progenitors of man; length of, in pheasants; difference of length of the, in the two sexes of birds.

Tait, Lawson, on the effects of natural selection on civilised nations.

Tanager, scarlet, variation in the male.

Tanagra aestiva, age of mature plumage in.

Tanagra rubra, young of.

Tanais, absence of mouth in the males of some species of; relations of the sexes in; dimorphic males of a species of.

Tankerville, Earl, on the battles of wild bulls.

Tanysiptera, races of, determined from adult males.

Tanysiptera sylvia, long tail-feathers of.

Taphroderes distortus, enlarged left mandible of the male.

Tapirs, longitudinal stripes of young.

Tarsi, dilatation of front, in male beetles.

Tarsius.

Tasmania, half-castes killed by the natives of.

Tasmanians, extinction of.

Taste, in the Quadrumana.

Tattooing, universality of.

Taylor, G., on Quiscalus major.

Taylor, Rev. R., on tattooing in New Zealand.

Tea, fondness of monkeys for.

Teal, constancy of.

Tear-sacs, of Ruminants.

Teebay, Mr., on changes of plumage in spangled Hamburg fowls.

Teeth, rudimentary incisor, in Ruminants; posterior molar, in man; wisdom; diversity of; canine, in the early progenitors of man; canine, of male mammals; in man, reduced by correlation; staining of the; front, knocked out or filed by some savages.

Tegetmeier, Mr., on the transmission of colours in pigeons by one sex alone; numerical proportion of male and female births in dogs; on the abundance of male pigeons; on the wattles of game-cocks; on the courtship of fowls; on the loves of pigeons; on dyed pigeons; blue dragon pigeons.

Tembeta, S. American ornament.

Temper, in dogs and horses, inherited.

Tench, proportions of the sexes in the; brightness of male, during breeding season.

Tenebrionidae, stridulation of.

Tennent, Sir J.E., on the tusks of the Ceylon Elephant; on the frequent absence of beard in the natives of Ceylon; on the Chinese opinion of the aspect of the Cingalese.

Tennyson, A., on the control of thought.

Tenthredinidae, proportions of the sexes in; fighting habits of male; difference of the sexes in.

Tephrodornis, young of.

Terai, in India.

Termites, habits of.

Terns, white; and black.

Terns, seasonal change of plumage in.

Terror, common action of, upon the lower animals and man.

Testudo elegans.

Testudo nigra.

Tetrao cupido, battles of; sexual difference in the vocal organs of.

Tetrao phasianellus, dances of; duration of dances of.

Tetrao scoticus.

Tetrao tetrix, pugnacity of the male.

Tetrao umbellus, pairing of; battles of; drumming of the male.

Tetrao urogalloides, dances of.

Tetrao urogallus, pugnacity of the male.

Tetrao urophasianus, inflation of the oesophagus in the male.

Thamnobia, young of.

Thecla, sexual differences of colouring in species of.

Thecla rubi, protective colouring of.

Thecophora fovea.

Theognis, selection in mankind.

Theridion, stridulation of males of.

Theridion lineatum.

Thomisus citreus, and Thomisus floricolens, difference of colour in the sexes of.

Thompson, J.H., on the battles of sperm-whales.

Thompson, W., on the colouring of the male char during the breeding season; on the pugnacity of the males of Gallinula chloropus; on the finding of new mates by magpies; on the finding of new mates by Peregrine falcons.

Thorax, processes of, in male beetles.

Thorell, T., on the proportion of sexes in spiders.

Thornback, difference in the teeth of the two sexes of the.

Thoughts, control of.

Thrush, pairing with a blackbird; colours and nidification of the.

Thrushes, characters of young.

Thug, remorse of a.

Thumb, absence of, in Ateles and Hylobates.

Thury, M., on the numerical proportion of male and female births among the Jews.

Thylacinus, possession of the marsupial sac by the male.

Thysanura.

Tibia, dilated, of the male Crabro cribrarius.

Tibia and femur, proportions of, in the Aymara Indians.

Tierra del Fuego, marriage-customs of.

Tiger, colours and markings of the.

Tigers, depopulation of districts by, in India.

Tillus elongatus, difference of colour in the sexes of.

Timidity, variability of, in the same species.

Tinca vulgaris.

Tipula, pugnacity of male.

Tits, sexual difference of colour in.

Toads, male, treatment of ova by some; male, ready to breed before the female.

Todas, infanticide and proportion of sexes; practice polyandry; choice of husbands amongst.

Toe, great, condition of, in the human embryo.

Tomicus villosus, proportion of the sexes in.

Tomtit, blue, sexual difference of colour in the.

Tonga Islands, beardlessness of the natives of.

Tooke, Horne, on language.

Tools, flint; used by monkeys; use of.

Topknots in birds.

Tortoise, voice of the male.

Tortures, submitted to by American savages.

Totanus, double moult in.

Toucans, colours and nidification of the; beaks and ceres of the.

Towns, residence in, a cause of diminished stature.

Toynbee, J., on the external shell of the ear in man.

Trachea, convoluted and imbedded in the sternum, in some birds; structure of the, in Rhynchaea.

Trades, affecting the form of the skull.

Tragelaphus, sexual differences of colour in.

Tragelaphus scriptus, dorsal crest of; markings of.

Tragopan, swelling of the wattles of the male, during courtship; display of plumage by the male; marking of the sexes of the.

Tragops dispar, sexual difference in the colour of.

Training, effect of, on the mental difference between the sexes of man.

Transfer of male characters to female birds.

Transmission, equal, of ornamental characters, to both sexes in mammals.

Traps, avoidance of, by animals; use of.

Treachery, to comrades, avoidance of, by savages.

Tremex columbae.

Tribes, extinct; extinction of.

Trichius, difference of colour in the sexes of a species of.

Trigla.

Trigonocephalus, noise made by tail of.

Trimen, R., on the proportion of the sexes in South African butterflies; on the attraction of males by the female Lasiocampa quercus; on Pneumora; on difference of colour in the sexes of beetles; on moths brilliantly coloured beneath; on mimicry in butterflies; on Gynanisa Isis, and on the ocellated spots of Lepidoptera; on Cyllo Leda.

Tringa, sexes and young of.

Tringa cornuta.

Triphaena, coloration of the species of.

Tristram, H.B., on unhealthy districts in North Africa; on the habits of the chaffinch in Palestine; on the birds of the Sahara; on the animals inhabiting the Sahara.

Triton cristatus.

Triton palmipes.

Triton punctatus.

Troglodyte skulls, greater than those of modern Frenchmen.

Troglodytes vulgaris.

Trogons, colours and nidification of the.

Tropic-birds, white only when mature.

Tropics, freshwater fishes of the.

Trout, proportion of the sexes in; male, pugnacity of the.

Trox sabulosus, stridulation of.

Truth, not rare between members of the same tribe; more highly appreciated by certain tribes.

Tulloch, Major, on the immunity of the negro from certain fevers.

Tumbler, almond, change of plumage in the.

Turdus merula, young of.

Turdus migratorius.

Turdus musicus.

Turdus polyglottus, young of.

Turdus torquatus.

Turkey, wild, pugnacity of young male; wild, notes of the; swelling of the wattles of the male; variety of, with a top-knot; recognition of a dog by a; male, wild, acceptable to domesticated females; wild, first advances made by older females; wild, breast-tuft of bristles of the.

Turkey-cock, scraping of the wings of, upon the ground; wild, display of plumage by; fighting habits of.

Turner, Prof. W., on muscular fasciculi in man referable to the panniculus carnosus; on the occurrence of the supra-condyloid foramen in the human humerus; on muscles attached to the coccyx in man; on the filum terminale in man; on the variability of the muscles; on abnormal conditions of the human uterus; on the development of the mammary glands; on male fishes hatching ova in their mouths; on the external perpendicular fissure of the brain; on the bridging convolutions in the brain of a chimpanzee.

Turnix, sexes of some species of.

Turtle-dove, cooing of the.

Tuttle, H., on the number of species of man.

Tylor, E.B., on emotional cries, gestures, etc., of man; on the origin of the belief in spiritual agencies; remorse for violation of tribal usage in marrying; on the primitive barbarism of civilised nations; on the origin of counting; inventions of savages; on resemblances, of the mental characters in different races of man.

Type of structure, prevalence of.

Typhaeus, stridulating organs of; stridulation of.

Twins, tendency to produce, hereditary.

Twite, proportion of the sexes in.

Ugliness, said to consist in an approach to the lower animals.

Umbrella-bird.

Umbrina, sounds produced by.

United States, rate of increase in; influence of natural selection on the progress of; change undergone by Europeans in the.

Upupa epops, sounds produced by the male.

Uraniidae, coloration of the.

Uria troile, variety of (=U. lacrymans).

Urodela.

Urosticte Benjamini, sexual differences in.

Use and disuse of parts, effects of; influence of, on the races of man.

Uterus, reversion in the; more or less divided, in the human subject; double, in the early progenitors of man.

Vaccination, influence of.

Vancouver Island, Mr. Sproat on the savages of; natives of, eradication of facial hair by the.

Vanellus cristatus, wing tubercles of the male.

Vanessae, resemblance of lower surface of, to bark of trees.

Variability, causes of; in man, analogous to that in the lower animals; of the races of man; greater in men than in women; period of, relation of the, to sexual selection; of birds; of secondary sexual characters in man.

Variation, laws of; correlated; in man; analogous; analogous, in plumage of birds.

Variations, spontaneous.

Varieties, absence of, between two species, evidence of their distinctness.

Variety, an object in nature.

Variola, communicable between man and the lower animals.

Vaureal, human bones from.

Veddahs, monogamous habits of.

Veitch, Mr., on the aversion of Japanese ladies to whiskers.

Vengeance, instinct of.

Venus Erycina, priestesses of.

Vermes.

Vermiform appendage.

Verreaux, M., on the attraction of numerous males by the female of an Australian Bombyx.

Vertebrae, caudal, number of in macaques and baboons; of monkeys, partly imbedded in the body.

Vertebrata, common origin of the; most ancient progenitors of; origin of the voice in air-breathing.

Vesicula prostatica, the homologue of the uterus.

Vibrissae, represented by long hairs in the eyebrows.

Vidua.

Vidua axillaris.

Villerme, M., on the influence of plenty upon stature.

Vinson, Aug., courtship of male spider; on the male of Epeira nigra.

Viper, difference of the sexes in the.

Virey, on the number of species of man.

Virtues, originally social only; gradual appreciation of.

Viscera, variability of, in man.

Vlacovich, Prof., on the ischio-pubic muscle.

Vocal music of birds.

Vocal organs of man; of birds; of frogs; of the Insessores; difference of, in the sexes of birds; primarily used in relation to the propagation of the species.

Vogt, Karl, on the origin of species; on the origin of man; on the semilunar fold in man; on microcephalous idiots; on the imitative faculties of microcephalous idiots; on skulls from Brazilian caves; on the evolution of the races of man; on the formation of the skull in women; on the Ainos and negroes; on the increased cranial difference of the sexes in man with race development; on the obliquity of the eye in the Chinese and Japanese.

Voice in mammals; in monkeys and man; in man; origin of, in air-breathing vertebrates.

Von Baer, see Baer.

Vulpian, Prof., on the resemblance between the brains of man and the higher apes.

Vultures, selection of a mate by the female; colours of.

Waders, young of.

Wagner, R., on the occurrence of the diastema in a Kaffir skull; on the bronchi of the black stork.

Wagtail, Ray’s, arrival of the male before the female.

Wagtails, Indian, young of.

Waist, proportions of, in soldiers and sailors.

Waitz, Prof., on the number of species of man; on the liability of negroes to tropical fevers after residence in a cold climate; on the colour of Australian infants; on the beardlessness of negroes; on the fondness of mankind for ornaments; on negro ideas of female beauty; on Javan and Cochin Chinese ideas of beauty.

Waldeyer, M., on the hermaphroditism of the vertebrate embryo.

Wales, North, numerical proportion of male and female births in.

Walkenaer and Gervais, spider attracted by music; on the Myriapoda.

Walker, Alex., on the large size of the hands of labourers’ children.

Walker, F., on sexual differences in the diptera.

Wallace, Dr. A., on the prehensile use of the tarsi in male moths; on the rearing of the Ailanthus silkmoth; on breeding Lepidoptera; proportion of sexes of Bombyx cynthia, B. yamamai, and B. Pernyi reared by; on the development of Bombyx cynthia and B. yamamai; on the pairing of Bombyx cynthia.

Wallace, A.R., on the origin of man; on the power of imitation in man; on the use of missiles by the orang; on the varying appreciation of truth among different tribes; on the limits of natural selection in man; on the occurrence of remorse among savages; on the effects of natural selection on civilised nations; on the use of the convergence of the hair at the elbow in the orang; on the contrast in the characters of the Malays and Papuans; on the line of separation between the Papuans and Malays; on the birds of paradise; on the sexes of Ornithoptera Croesus; on protective resemblances; on the relative sizes of the sexes of insects; on Elaphomyia; on the pugnacity of the males of Leptorhynchus angustatus; on sounds produced by Euchirus longimanus; on the colours of Diadema; on Kallima; on the protective colouring of moths; on bright coloration as protective in butterflies; on variability in the Papilionidae; on male and female butterflies, inhabiting different stations; on the protective nature of the dull colouring of female butterflies; on mimicry in butterflies; on the bright colours of caterpillars; on brightly-coloured fishes frequenting reefs; on the coral snakes; on Paradisea apoda; on the display of plumage by male birds of paradise; on assemblies of birds of paradise; on the instability of the ocellated spots in Hipparchia Janira; on sexually limited inheritance; on the sexual coloration of birds; on the relation between the colours and nidification of birds; on the coloration of the Cotingidae; on the females of Paradisea apoda and papuana; on the incubation of the cassowary; on protective coloration in birds; on the Babirusa; on the markings of the tiger; on the beards of the Papuans; on the hair of the Papuans; on the distribution of hair on the human body.

Walrus, development of the nictitating membrane in the; tusks of the; use of the tusks by the.

Walsh, B.D., on the proportion of the sexes in Papilio Turnus; on the Cynipidae and Cecidomyidae; on the jaws of Ammophila; on Corydalis cornutus; on the prehensile organs of male insects; on the antennae of Penthe; on the caudal appendages of dragonflies; on Platyphyllum concavum; on the sexes of the Ephemeridae; on the difference of colour in the sexes of Spectrum femoratum; on sexes of dragon-flies; on the difference of the sexes in the Ichneumonidae; on the sexes of Orsodacna atra; on the variation of the horns of the male Phanaeas carnifex; on the coloration of the species of Anthocharis.

Wapiti, battles of; traces of horns in the female; attacking a man; crest of the male; sexual difference in the colour of the.

Warbler, hedge-; young of the.

Warblers, superb, nidification of.

Wariness, acquired by animals.

Warington, R., on the habits of the stickleback; on the brilliant colours of the male stickleback during the breeding season.

Wart-hog, tusks and pads of the.

Watchmakers, short-sighted.

Waterhen.

Waterhouse, C.O., on blind beetles; on difference of colour in the sexes of beetles.

Waterhouse, G.R., on the voice of Hylobates agilis.

Water-ouzel, autumn song of the.

Waterton, C., on the Bell-bird; on the pairing of a Canada goose with a Bernicle gander; on hares fighting.

Wattles, disadvantageous to male birds in fighting.

Weale, J., Mansel, on a South African caterpillar.

Wealth, influence of.

Weapons, used by man; employed by monkeys; offensive, of males; of mammals.

Weaver-bird.

Weaver-birds, rattling of the wings of; assemblies of.

Webb, Dr., on the wisdom teeth.

Wedderburn, Mr., assembly of black game.

Wedgwood, Hensleigh, on the origin of language.

Weevils, sexual difference in length of snout in some.

Weir, Harrison, on the numerical proportion of the sexes in pigs and rabbits; on the sexes of young pigeons; on the songs of birds; on pigeons; on the dislike of blue pigeons to other coloured varieties; on the desertion of their mates by female pigeons.

Weir, J. Jenner, on the nightingale and blackcap; on the relative sexual maturity of male birds; on female pigeons deserting a feeble mate; on three starlings frequenting the same nest; on the proportion of the sexes in Machetes pugnax and other birds; on the coloration of the Triphaenae; on the rejection of certain caterpillars by birds; on sexual differences of the beak in the goldfinch; on a piping bullfinch; on the object of the nightingale’s song; on song-birds; on the pugnacity of male fine-plumaged birds; on the courtship of birds; on the finding of new mates by Peregrine falcons and Kestrels; on the bullfinch and starling; on the cause of birds remaining unpaired; on starlings and parrots living in triplets; on recognition of colour by birds; on hybrid birds; on the selection of a greenfinch by a female canary; on a case of rivalry of female bullfinches; on the maturity of the golden pheasant.

Weisbach, Dr., measurement of men of different races; on the greater variability of men than of women; on the relative proportions of the body in the sexes of different races of man.

Weismann, Prof., colours of Lycaenae.

Welcker, M., on brachycephaly and dolichocephaly; on sexual differences in the skull in man.

Wells, Dr., on the immunity of coloured races from certain poisons.

Westring, on the stridulation of males of Theridion; on the stridulation of Reduvius personatus; on the stridulation of beetles; on the stridulation of Omaloplia brunnea; on the stridulating organs of the Coleoptera; on sounds produced by Cychrus.

Westropp, H.M., on reason in a bear; on the prevalence of certain forms of ornamentation.

Westwood, J.O., on the classification of the Hymenoptera; on the Culicidae and Tabanidae; on a Hymenopterous parasite with a sedentary male; on the proportions of the sexes in Lucanus cervus and Siagonium; on the absence of ocelli in female Mutillidae; on the jaws of Ammophila; on the copulation of insects of distinct species; on the male of Crabro cribrarius; on the pugnacity of the male Tipulae; on the stridulation of Pirates stridulus; on the Cicadae; on the stridulating organs of the cricket; on Ephippiger vitium; on Pneumora; on the pugnacity of the Mantides; on Platyblemnus; on difference in the sexes of the Agrionidae; on the pugnacity of the males of a species of Tenthredinae; on the pugnacity of the male stag-beetle; on Bledius taurus and Siagonium; on lamellicorn beetles; on the coloration of Lithosia.

Whale, Sperm-, battles of male.

Whales, nakedness of.

Whately, Arch., language not peculiar to man; on the primitive civilisation of man.

Whewell, Prof., on maternal affection.

Whiskers, in monkeys.

White, F.B., noise produced by Hylophila.

White, Gilbert, on the proportion of the sexes in the partridge; on the house-cricket; on the object of the song of birds; on the finding of new mates by white owls; on spring coveys of male partridges.

Whiteness, a sexual ornament in some birds; of mammals inhabiting snowy countries.

White-throat, aerial love-dance of the male.

Whitney, Prof., on the development of language; language not indispensable for thought.

Widgeon, pairing with a pintail duck.

Widow-bird, polygamous; breeding plumage of the male; female, rejecting the unadorned male.

Widows and widowers, mortality of.

Wilckens, Dr., on the modification of domestic animals in mountainous regions; on a numerical relation between the hairs and excretory pores in sheep.

Wilder, Dr. Burt, on the greater frequency of supernumerary digits in men than in women.

Williams, on the marriage-customs of the Fijians.

Wilson, Dr., on the conical heads of the natives of North-Western Africa; on the Fijians; on the persistence of the fashion of compressing the skull.

Wing-spurs.

Wings, differences of, in the two sexes of butterflies and Hymenoptera; play of, in the courtship of birds.

Winter, change of colour of mammals in.

Witchcraft.

Wives, traces of the forcible capture of.

Wolf, winter change of the.

Wolff, on the variability of the viscera in man.

Wollaston, T.V., on Eurygnathus; on musical Curculionidae; on the stridulation of Acalles.

Wolves, learning to bark from dogs; hunting in packs.

Wolves, black.

Wombat, black varieties of the.

Women, distinguished from men by male monkeys; preponderance of, in numbers; selection of, for beauty; effects of selection of, in accordance with different standards of beauty; practice of capturing; early betrothals and slavery of; freedom of selection by, in savage tribes.

Wonder, manifestations of, by animals.

Wonfor, Mr., on sexual peculiarities, in the wings of butterflies.

Wood, J., on muscular variations in man; on the greater variability of the muscles in men than in women.

Wood, T.W., on the colouring of the orange-tip butterfly; on the habits of the Saturniidae; quarrels of chamaeleons; on the habits of Menura Alberti; on Tetrao cupido; on the display of plumage by male pheasants; on the ocellated spots of the Argus pheasant; on fighting of Menura superba; on the habits of the female cassowary.

Woodcock, coloration of the.

Woodpecker, selection of a mate by the female.

Woodpeckers, tapping of; colours and nidification of the; characters of young.

Woolner, Mr., observations on the ear in man.

Wormald, Mr., on the coloration of Hypopyra.

Wounds, healing of.

Wren, young of the.

Wright, C.A., on the young of Orocetes and Petrocincla.

Wright, Chauncey, great brain-power requisite for language; on correlative acquisition; on the enlargement of the brain in man.

Wright, Mr., on the Scotch deer-hound; on sexual preference in dogs; on the rejection of a horse by a mare.

Wright, W. von, on the protective plumage of the Ptarmigan.

Writing.

Wyman, Prof., on the prolongation of the coccyx in the human embryo; on the condition of the great toe in the human embryo; on the occurrence of the supra-condyloid foramen in the humerus of man; on variation in the skulls of the natives of the Sandwich Islands; on the hatching of the eggs in the mouths and branchial cavities of male fishes.

Xenarchus, on the Cicadae.

Xenophon, selection in mankind advocated by.

Xenorhynchus, sexual difference in the colour of the eyes in.

Xiphophorus Hellerii, peculiar anal fin of the male.

Xylocopa, difference of the sexes in.

Yarrel, W., on the habits of the Cyprinidae; on Raia clavata; on the characters of the male salmon during the breeding season; on the characters of the rays; on the gemmeous dragonet; on colours of salmon; on the spawning of the salmon; on the incubation of the Lophobranchii; on rivalry in song-birds; on the trachea of the swan; on the moulting of the Anatidae; on the young of the waders.

Yellow fever, immunity of negroes and mulattoes from.

Youatt, Mr., on the development of the horns in cattle.

Yura-caras, their notions of beauty.

Zebra, rejection of an ass by a female; stripes of the.

Zebus, humps of.

Zigzags, prevalence of, as ornaments.

Zincke, Mr., on European emigration to America.

Zootoca vivipara, sexual difference in the colour of.

Zouteveen, Dr., polydactylism; proportion of sexes at Cape of Good Hope; spiders attracted by music; on sounds produced by fish.

Zygaenidae, coloration of the.

THE END.