The Descent of Man, and Selection in Relation to Sex

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

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

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

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

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

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

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

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

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

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

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

VARIABILITY OF BIRDS, AND ESPECIALLY OF THEIR SECONDARY SEXUAL CHARACTERS

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

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

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

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

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

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