The Descent of Man and Selection in Relation to Sex by Charles Darwin - HTML preview

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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 hummingbirds (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 breedingseason 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 lovesongs 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 blackcock, 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 firebacked 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 blackcock. 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 songbirds 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 armorous 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 Stubenvogel,' 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 Stubenvogel,' 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, 'Stubenvogel,' 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 halfgrown 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, beeeater, 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 i