Read Ebook: The Descent of Man and Selection in Relation to Sex Vol. II (1st Edition) by Darwin Charles

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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 it is more deeply embedded in the adult male than in the female or young male. In the male Merganser the enlarged portion of the trachea is furnished with an additional pair of muscles. But the meaning of these differences between the sexes of many Anatidae is not at all 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. In both sexes of one of the cranes 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. So that highly important structures have in these cases been modified according to sex.

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 the sounds. The drumming, or bleating, or neighing, or thundering noise, as expressed by different observers, which is made by the common snipe 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 in a curved line, with outspread tail and quivering pinions, with surprising velocity to the earth. 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 , having a stiff sabre-shaped shaft, with the oblique barbs of unusual length, the outer webs being strongly bound together.

The diversity of the sounds, both vocal and instrumental, made by the males of many species 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 same conclusion with respect 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. This is somewhat more difficult in the case of the modified feathers, by which the drumming, whistling, or roaring noises are produced. But we have seen 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 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, 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."

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 , which, as we shall hereafter see, are highly 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 the habits of some Satin Bower-birds, which he kept in his 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, and 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 assemblages, 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.

The males are often ornamented with elongated feathers or plumes springing 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 of the Argus pheasant. The body of this latter bird 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. The wing-feathers are not elongated nearly so often as the tail-feathers; for their elongation would impede the act of flight. Yet the beautifully ocellated secondary wing-feathers of the male Argus pheasant are nearly three feet in length; and in a small African nightjar 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 nightjars, the shafts of the elongated wing-feathers are naked, except at the extremity, where there is a disc. Again, in another genus of nightjars, the tail-feathers are even still more prodigiously developed; so that we see the same kind of ornament gained by the males of closely-allied birds, through the development of widely different feathers.

Male humming-birds 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 the 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 belonging to the same species in other characters; and these have been seized on by man and augmented to an extreme point--as 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, &c., it is due to sexual selection,--that is 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 of S. America, the note of which can be distinguished at the distance of nearly three miles, and astonishes every one who first hears it. The male is pure white, whilst the female is dusky-green; and the former colour with terrestrial species of moderate size and inoffensive habits is very rare. 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 most 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 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 the head and neck alone of the male are white, the rest of the body being chesnut-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.

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 by very young birds of their down; for the down in most cases arises from the summits of the first true feathers.

With respect to the birds which annually undergo a double moult, there are, firstly, some kinds, for instance snipes, swallow-plovers , 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, which seems, when there is no change of colour, the most probable cause of a double moult. Secondly, there are birds, for instance certain species of Totanus and other grallatores, the sexes of which resemble each other, but have a slightly different summer and winter plumage. The difference, however, in colour in these cases is so slight 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, sometimes, as with certain bustards, a great change of colour. 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 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, 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, &c., for they acquire their beautiful plumes only during the breeding-season. Moreover, such plumes, top-knots, &c., though possessed by both sexes, are occasionally a little more highly 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 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.

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 moulting twice in the year has not been acquired in order that the male should assume during the breeding-season an ornamental character; 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.

The gold pheasant during his courtship not only expands and raises his splendid frill, but turns it, 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. Mr. Bartlett has observed a male Polyplectron 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 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.

The case of the Argus pheasant is still more striking. The immensely developed secondary wing-feathers, which are confined to the male, are ornamented with a row of from twenty to twenty-three ocelli, each above an inch in diameter. The feathers are also elegantly marked with oblique dark stripes and rows of spots, like those on the skin of a tiger and leopard combined. The ocelli are so beautifully shaded that, as the Duke of Argyll remarks, they stand out like a ball lying loosely within a socket. But when I looked at the specimen in the British Museum, which is mounted with the wings expanded and trailing downwards, I was greatly disappointed, for the ocelli appeared flat or even concave. Mr. Gould, however, soon made the case clear to me, for he had made a drawing of a male whilst he was displaying himself. At such times the long secondary feathers in both wings are vertically erected and expanded; and these, together with the enormously elongated tail-feathers, make a grand semicircular upright fan. Now as soon as the wing-feathers are held in this position, and the light shines on them from above, the full effect of the shading comes out, and each ocellus at once resembles the ornament called a ball and socket. These feathers have been shewn 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 primary wing-feathers, which in most gallinaceous birds are uniformly coloured, are in the Argus pheasant not less wonderful objects than the secondary wing-feathers. 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 chesnut, 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 when the long secondary feathers are erected, though in a widely different manner; for they are expanded in front like two little fans or shields, one on each side of the breast near the ground.

The case of the male Argus pheasant is eminently interesting, because it affords good evidence that the most refined beauty may serve as a charm for the female, and for no other purpose. We must conclude that this is the case, as the primary wing-feathers are never displayed, and the ball-and-socket ornaments are not exhibited in full perfection, except when the male assumes the attitude of courtship. The Argus pheasant does not possess brilliant colours, so that his success in courtship appears to have depended 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, though perhaps she admires the general effect rather than each separate detail. 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 beautiful 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 ; 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 identically 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 very 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, selected from a large body of valuable notes, sent to me by Mr. Jenner Weir, who has long kept birds of many kinds, including all the British Fringillidae and Emberizidae. 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 act thus 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. No other British finch, as Mr. Weir informs me, turns during his courtship from side to side in this manner; not even the closely-allied male siskin, for he would not thus add to his beauty.

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 they have been acquired in some cases at the expense of greatly impeded powers of flight or of running. The African nightjar , which during the pairing-season has one of its primary wing-feathers developed into a streamer of extreme length, is thus 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 are 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 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 a more easy 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 it probably is, as Mr. Gould has remarked, 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.

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 comb 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." 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, when distended during the act of 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 male must be of the highest importance to him; and we further see that beauty in some cases is even more important than success in battle.

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, in the power of singing, or in producing what I have called instrumental music, it is almost invariably the male which excels 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, this is always shortly 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 all 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 equally excite and attract the female? or does she exert a choice, and prefer certain males? This question can be answered in the affirmative by much direct and indirect evidence. It is much 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.

The common magpie , 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. Whilst they were so numerous, they had the habit very early in the spring of assembling 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 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 never met with any account of regular assemblages of black game in Scotland, yet these assemblages are so well known in Germany and Scandinavia that they have special names.

Sir J. Lubbock's gamekeeper has repeatedly shot, but how many times he could not say, one of a pair of jays , and has never failed shortly afterwards to find the survivor rematched. The Rev. W. D. Fox, Mr. F. Bond, and others, have shot one of a pair of carrion-crows , but the nest was soon again tenanted by a pair. These birds are rather common; but the peregrine falcon is rare, yet Mr. Thompson states that in Ireland "if either an old male or female be killed in the breeding-season , 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 to occur with the peregrine falcons at Beachy Head. The same observer informs me that three kestrels, all males , were killed one after the other whilst attending the same nest; two of these were in mature plumage, and the third in the plumage of the previous year. Even with the rare golden eagle , Mr. Birkbeck was assured by a trustworthy gamekeeper in Scotland, that if one is killed, another is soon found. So with the white owl , it has been observed that "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 was always soon provided with 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 , the writer remarks that it was by no means common in the neighbourhood, and he expresses much surprise how the sitting female could so soon give effectual notice that she was a widow. 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 far from loud. I will give only one other fact, on the authority of this same observer; one of a pair of starlings 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.

These facts are certainly remarkable. How is it that so many birds are ready immediately to replace a lost mate? Magpies, jays, carrion-crows, partridges, and some other birds, are never seen during the spring by themselves, and these offer at first sight the most perplexing case. 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. 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 either one of a pair, would leave the other bird 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. Such contingencies as these probably explain most of the foregoing cases. 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 inasmuch as the act of courtship appears to be with many birds a prolonged and tedious affair, 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.

Birds sometimes exhibit benevolent feelings; they will feed the deserted young even of distinct species, but this perhaps ought to be considered as a mistaken instinct. They will also 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."

Birds possess acute powers of observation. Every mated bird, of course, recognises its fellow. Audubon states that with the mocking-thrushes of the United States a certain number remain all the year round in Louisiana, whilst the 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." Mr. Hewitt has carefully described the habits of some ducks , 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 so well Mr. Hewitt's own dogs and cats 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.

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 , which had acquired its black head, 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. Mr. Weir was also obliged to turn out a robin, as it fiercely attacked all birds 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 into his aviary, fly towards the species which resemble them most in colour, and settle by their sides.

As male birds display with so much care their fine plumage and other ornaments in the presence of 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 we cannot feel sure that it is not from jealousy at 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, attracts the Ruff strongly 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 outside 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 , where the sexes congregate and play strange antics, are differently constructed, but what most concerns us is, that they are decorated in a different manner by the several species. 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 rearranged, 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 shewing a decided taste for the beautiful." Well may Mr. Gould say "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.

The Rev. W. D. Fox informs me that he possessed at the same time a pair of Chinese geese , 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 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.

With the members of the same species the fact of the female preferring to pair with one male rather than with another is not so likely to excite attention, as when this occurs between distinct species. Such 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 referred to; 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 everyone 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 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 nightjars 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 of the United States, parties of eight or 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 , 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." 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 they have long been kept. Mr. Tegetmeier is convinced that a game-cock, though disfigured by being dubbed 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.

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. 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 be matched, after repeated trials, with a black male, but immediately paired with a dun. Generally colour alone 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 Corbi?, whose experience extended over forty-five years, state: "Quand une femelle ?prouve de l'antipathie pour un m?le avec lequel on veut l'accoupler, malgr? tous les feux de l'amour, malgr? l'alpiste et le ch?nevis dont on la nourrit pour augmenter son ardeur, malgr? un emprisonnement de six mois et m?me d'un an, elle refuse constamment ses caresses; les avances empress?es, les agaceries, les tournoiemens, les tendres roucoulemens, rien ne peut lui plaire ni l'?mouvoir; gonfl?e, boudeuse, blottie dans un coin de sa prison, elle n'en sort que pour boire et manger, ou pour repousser avec une esp?ce de rage des caresses devenues trop pressantes." 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, 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.

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 peacock. 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." This rival was a japanned or black-winged peacock, which to our eyes is 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 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. Here is another striking case; Dr. Jaeger, director of the Zoological Gardens of Vienna, states that a male silver pheasant, who had been triumphant over the 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.

Not only does the female exert a choice, but in some few cases she courts the male, or even fights 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. We have seen that a tame wild-duck seduced after a long courtship an unwilling Pintail drake. 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, Mr. Weir replaced 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 exceptions to this rule, as we shall hereafter see, apparently occur in some few groups. With domesticated birds, I have heard of only one case in which the males shew any preference for particular 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." From some inexplicable cause he shews the most determined aversion to certain hens, which no care on the part of the breeder can overcome. Some hens, as Mr. Hewitt informs me, 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 , "it has been remarked," says M. Ekstr?m, "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.

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 placing ourselves in imagination in the same position. If an inhabitant of another planet were to behold a number of young rustics at a fair, courting and quarrelling over a pretty girl, like birds at one of their places of assemblage, he would be able to infer that she had the power of choice only by observing the eagerness of the wooers to please her, and to display their finery. 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 most highly 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 have been thought, would have altogether depended 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 the analogy of our own minds; and the mental powers of birds, if reason be excluded, do not fundamentally differ 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.

Mr. Gould is well known rarely to admit the existence of varieties, for he esteems very slight differences as specific; now he states 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." In the United States some few of the males of the Scarlet Tanager have "a beautiful transverse band of glowing red on the smaller wing-coverts;" but this variation seems to be somewhat rare, so that its preservation through sexual selection would follow only under unusually favourable circumstances. In Bengal the Honey buzzard 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."

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 Br?nnich to conclude that it was specifically distinct; but this is now known to be an error.

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. 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 exclusively in most cases, as I hope to shew in the following chapter, on the form of inheritance which prevails in the groups in question.

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 Cr?ve-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 Golden 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 modified in the same manner. We are thus enabled without much trouble to rear breeds with their plumage marked and coloured 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 see feathers becoming 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.

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