The Variation of Animals and Plants under Domestication · Charles Darwin
CHAPTER XX. SELECTION BY MAN.
The Emperor Akbar Khan before the year l600 is said to have “wonderfully improved” his pigeons by crossing the breeds; and this necessarily implies careful selection. About the same period the Dutch attended with the greatest care to the breeding of these birds. Belon in 1555 says that good managers in France examined the colour of their goslings in order to get geese of a white colour and better kinds. Markham in 1631 tells the breeder “to elect the largest and goodliest conies,” and enters into minute details. Even with respect to seeds of plants for the flower-garden, Sir J. Hanmer writing about the year 1660[46] says, in “choosing seed, the best seed is the most weighty, and is had from the lustiest and most vigorous stems;” and he then gives rules about leaving only a few flowers on plants for seed; so that even such details were attended to in our flower-gardens two hundred years ago. In order to show that selection has been silently carried on in places where it would not have been expected, I may add that in the middle of the last century, in a remote part of North America, Mr. Cooper improved by careful selection all his vegetables, “so that they were greatly superior to those of any other person. When his radishes, for instance, are fit for use, he takes ten or twelve that he most approves, and plants them at least 100 yards from others that blossom at the same time. In the same manner he treats all his other plants, varying the circumstances according to their nature.”[47]
In the great work on China published in the last century by the Jesuits, and which is chiefly compiled from ancient Chinese encyclopaedias, it is said that with sheep “improving the breed consists in choosing with particular care the lambs which are destined for propagation, in nourishing them well, and in keeping the flocks separate.” The same principles were applied by the Chinese to various plants and fruit-trees.[48] An imperial edict recommends the choice of seed of remarkable size; and selection was practised even by imperial hands, for it is said that the Ya-mi, or imperial rice, was noticed at an ancient period in a field by the Emperor Khang-hi, was saved and cultivated in his garden, and has since become valuable from being the only kind which will grow north of the Great Wall.[49] Even with flowers, the tree paeony (_P. moutan_) has been cultivated, according to Chinese traditions, for 1400 years; between 200 and 300 varieties have been raised, which are cherished like tulips formerly were by the Dutch.[50]
Turning now to semi-civilised people and to savages: it occurred to me, from what I had seen of several parts of South America, where fences do not exist, and where the animals are of little value, that there would be absolutely no care in breeding or selecting them; and this to a large extent is true. Roulin,[51] however, describes in Columbia a naked race of cattle, which are not allowed to increase, on account of their delicate constitution. According to Azara[52] horses are often born in Paraguay with curly hair; but, as the natives do not like them, they are destroyed. On the other hand, Azara states that a hornless bull, born in 1770, was preserved and propagated its race. I was informed of the existence in Banda Oriental of a breed with reversed hair; and the extraordinary niata cattle first appeared and have since been kept distinct in La Plata. Hence certain conspicuous variations have been preserved, and others have been habitually destroyed, in these countries, which are so little favourable for careful selection. We have also seen that the inhabitants sometimes introduce fresh cattle on their estates to prevent the evil effects of close interbreeding. On the other hand, I have heard on reliable authority that the Gauchos of the Pampas never take any pains in selecting the best bulls or stallions for breeding; and this probably accounts for the cattle and horses being remarkably uniform in character throughout the immense range of the Argentine republic.
Looking to the Old World, in the Sahara Desert “The Touareg is as careful in the selection of his breeding Mahari (a fine race of the dromedary) as the Arab is in that of his horse. The pedigrees are handed down, and many a dromedary can boast a genealogy far longer than the descendants of the Darley Arabian.”[53] According to Pallas the Mongolians endeavour to breed the Yaks or horse-tailed buffaloes with white tails, for these are sold to the Chinese mandarins as fly-flappers; and Moorcroft, about seventy years after Pallas, found that white-tailed animals were still selected for breeding.[54] We have seen in the chapter on the Dog that savages in different parts of North America and in Guiana cross their dogs with wild Canidæ, as did the ancient Gauls, according to Pliny. This was done to give their dogs strength and vigour, in the same way as the keepers in large warrens now sometimes cross their ferrets (as I have been informed by Mr. Yarrell) with the wild polecat, “to give them more devil.” According to Varro, the wild ass was formerly caught and crossed with the tame animal to improve the breed, in the same manner as at the present day the natives of Java sometimes drive their cattle into the forests to cross with the wild Banteng (_Bos sondaicus_).[55] In Northern Siberia, among the Ostyaks, the dogs vary in markings in different districts, but in each place they are spotted black and white in a remarkably uniform manner;[56] and from this fact alone we may infer careful breeding, more especially as the dogs of one locality are famed throughout the country for their superiority. I have heard of certain tribes of Esquimaux who take pride in their teams of dogs being uniformly coloured. In Guiana, as Sir H. Schomburgk informs me,[57] the dogs of the Turuma Indians are highly valued and extensively bartered: the price of a good one is the same as that given for a wife: they are kept in a sort of cage, and the Indians “take great care when the female is in season to prevent her uniting with a dog of an inferior description.” The Indians told Sir Robert that, if a dog proved bad or useless, he was not killed, but was left to die from sheer neglect. Hardly any nation is more barbarous than the Fuegians, but I hear from Mr. Bridges, the Catechist to the Mission, that, “when these savages have a large, strong, and active bitch, they take care to put her to a fine dog, and even take care to feed her well, that her young may be strong and well favoured.”
In the interior of Africa, negroes, who have not associated with white men, show great anxiety to improve their animals; they “always choose the larger and stronger males for stock;” the Malakolo were much pleased at Livingstone’s promise to send them a bull, and some Bakalolo carried a live cock all the way from Loanda into the interior.[58] At Falaba Mr. Winwood Reade noticed an unusually fine horse, and the negro King informed him that “the owner was noted for his skill in breeding horses.” Further south on the same continent, Andersson states that he has known a Damara give two fine oxen for a dog which struck his fancy. The Damaras take great delight in having whole droves of cattle of the same colour, and they prize their oxen in proportion to the size of their horns. “The Namaquas have a perfect mania for a uniform team; and almost all the people of Southern Africa value their cattle next to their women, and take a pride in possessing animals that look high-bred. They rarely or never make use of a handsome animal as a beast of burden.”[59] The power of discrimination which these savages possess is wonderful, and they can recognise to which tribe any cattle belong. Mr. Andersson further informs me that the natives frequently match a particular bull with a particular cow.
The most curious case of selection by semi-civilised people, or indeed by any people, which I have found recorded, is that given by Garcilazo de la Vega, a descendant of the Incas, as having been practised in Peru before the country was subjugated by the Spaniards.[60] The Incas annually held great hunts, when all the wild animals were driven from an immense circuit to a central point. The beasts of prey were first destroyed as injurious. The wild Guanacos and Vicunas were sheared; the old males and females killed, and the others set at liberty. The various kinds of deer were examined; the old males and females were likewise killed, “but the young females, with a certain number of males, selected from the most beautiful and strong,” were given their freedom. Here, then, we have selection by man aiding natural selection. So that the Incas followed exactly the reverse system of that which our Scottish sportsman are accused of following, namely, of steadily killing the finest stags, thus causing the whole race to degenerate.[61] In regard to the domesticated llamas and alpacas, they were separated in the time of the Incas according to colour: and if by chance one in a flock was born of the wrong colour, it was eventually put into another flock.
In the genus Auchenia there are four forms,—the Guanaco and Vicuna, found wild and undoubtedly distinct species; the Llama and Alpaca, known only in a domesticated condition. These four animals appear so different, that most naturalists, especially those who have studied these animals in their native country, maintain that they are specifically distinct, notwithstanding that no one pretends to have seen a wild llama or alpaca. Mr. Ledger, however, who has closely studied these animals both in Peru and during their exportation to Australia, and who has made many experiments on their propagation, adduces arguments[62] which seem to me conclusive, that the llama is the domesticated descendant of the guanaco, and the alpaca of the vicuna. And now that we know that these animals were systematically bred and selected many centuries ago, there is nothing surprising in the great amount of change which they have undergone.
It appeared to me at one time probable that, though ancient and semi-civilised people might have attended to the improvement of their more useful animals in essential points, yet that they would have disregarded unimportant characters. But human nature is the same throughout the world: fashion everywhere reigns supreme, and man is apt to value whatever he may chance to possess. We have seen that in South America the niata cattle, which certainly are not made useful by their shortened faces and upturned nostrils, have been preserved. The Damaras of South Africa value their cattle for uniformity of colour and enormously long horns. And I will now show that there is hardly any peculiarity in our most useful animals which, from fashion, superstition, or some other motive, has not been valued, and consequently preserved. With respect to cattle, “an early record,” according to Youatt[63] “speaks of a hundred white cows with red ears being demanded as a compensation by the princes of North and South Wales. If the cattle were of a dark or black colour, 150 were to be presented.” So that colour was attended to in Wales before its subjugation by England.
In Central Africa, an ox that beats the ground with its tail is killed; and in South Africa some of the Damaras will not eat the flesh of a spotted ox. The Kaffirs value an animal with a musical voice; and “at a sale in British Kaffraria the low of a heifer excited so much admiration that a sharp competition sprung up for her possession, and she realised a considerable price.”[64] With respect to sheep, the Chinese prefer rams without horns; the Tartars prefer them with spirally wound horns, because the hornless are thought to lose courage.[65] Some of the Damaras will not eat the flesh of hornless sheep. In regard to horses, at the end of the fifteenth century animals of the colour described as liart pomme were most valued in France. The Arabs have a proverb, “Never buy a horse with four white feet, for he carries his shroud with him”;[66] the Arabs also, as we have seen, despise dun-coloured horses. So with dogs, Xenophon and others at an ancient period were prejudiced in favour of certain colours; and “white or slate-coloured hunting dogs were not esteemed.”[67] Turning to poultry, the old Roman gourmands thought that the liver of a white goose was the most savoury. In Paraguay black-skinned fowls are kept because they are thought to be more productive, and their flesh the most proper for invalids.[68] In Guiana, as I am informed by Sir R. Schomburgk, the aborigines will not eat the flesh or eggs of the fowl, but two races are kept distinct merely for ornament. In the Philippines, no less than nine sub-varieties of the game-cock are kept and named, so that they must be separately bred.
At the present time in Europe, the smallest peculiarities are carefully attended to in our most useful animals, either from fashion, or as a mark of purity of blood. Many examples could be given; two will suffice. “In the Western counties of England the prejudice against a white pig is nearly as strong as against a black one in Yorkshire.” In one of the Berkshire sub-breeds, it is said, “the white should be confined to four white feet, a white spot between the eyes, and a few white hairs behind each shoulder.” Mr. Saddler possessed “three hundred pigs, every one of which was marked in this manner.”[69] Marshall, towards the close of the last century, in speaking of a change in one of the Yorkshire breeds of cattle, says the horns have been considerably modified, as “a clean, small, sharp horn has been _fashionable_ for the last twenty years.”[70] In a part of Germany the cattle of the Race de Gfoehl are valued for many good qualities, but they must have horns of a particular curvature and tint, so much so that mechanical means are applied if they take a wrong direction; but the inhabitants “consider it of the highest importance that the nostrils of the bull should be flesh-coloured, and the eyelashes light; this is an indispensable condition. A calf with blue nostrils would not be purchased, or purchased at a very low price.”[71] Therefore let no man say that any point or character is too trifling to be methodically attended to and selected by breeders.
_Unconscious Selection._—By this term I mean, as already more than once explained, the preservation by man of the most valued, and the destruction of the least valued individuals, without any conscious intention on his part of altering the breed. It is difficult to offer direct proofs of the results which follow from this kind of selection; but the indirect evidence is abundant. In fact, except that in the one case man acts intentionally, and in the other unintentionally, there is little difference between methodical and unconscious selection. In both cases man preserves the animals which are most useful or pleasing to him, and destroys or neglects the others. But no doubt a far more rapid result follows from methodical than from unconscious selection. The “roguing” of plants by gardeners, and the destruction by law in Henry VIII.’s reign of all under-sized mares, are instances of a process the reverse of selection in the ordinary sense of the word, but leading to the same general result. The influence of the destruction of individuals having a particular character is well shown by the necessity of killing every lamb with a trace of black about it, in order to keep the flock white; or again, by the effects on the average height of the men of France of the destructive wars of Napoleon, by which many tall men were killed, the short ones being left to be the fathers of families. This at least is the conclusion of some of those who have closely studied the effects of the conscription; and it is certain that since Napoleon’s time the standard for the army has been lowered two or three times.
Unconscious selection blends with methodical, so that it is scarcely possible to separate them. When a fancier long ago first happened to notice a pigeon with an unusually short beak, or one with the tail-feathers unusually developed, although he bred from these birds with the distinct intention of propagating the variety, yet he could not have intended to make a short-faced tumbler or a fantail, and was far from knowing that he had made the first step towards this end. If he could have seen the final result, he would have been struck with astonishment, but, from what we know of the habits of fanciers, probably not with admiration. Our English carriers, barbs, and short-faced tumblers have been greatly modified in the same manner, as we may infer both from the historical evidence given in the chapters on the Pigeon, and from the comparison of birds brought from distant countries.
So it has been with dogs; our present fox-hounds differ from the old English hound; our greyhounds have become lighter: the Scotch deer-hound has been modified, and is now rare. Our bulldogs differ from those which were formerly used for baiting bulls. Our pointers and Newfoundlands do not closely resemble any native dog now found in the countries whence they were brought. These changes have been effected partly by crosses; but in every case the result has been governed by the strictest selection. Nevertheless, there is no reason to suppose that man intentionally and methodically made the breeds exactly what they now are. As our horses became fleeter, and the country more cultivated and smoother, fleeter fox-hounds were desired and produced, but probably without any one distinctly foreseeing what they would become. Our pointers and setters, the latter almost certainly descended from large spaniels, have been greatly modified in accordance with fashion and the desire for increased speed. Wolves have become extinct, and so has the wolf-dog; deer have become rarer, bulls are no longer baited, and the corresponding breeds of the dog have answered to the change. But we may feel almost sure that when, for instance, bulls were no longer baited, no man said to himself, I will now breed my dogs of smaller size, and thus create the present race. As circumstances changed, men unconsciously and slowly modified their course of selection.
With racehorses selection for swiftness has been followed methodically, and our horses now easily surpass their progenitors. The increased size and different appearance of the English racehorse led a good observer in India to ask, “Could any one in this year of 1856, looking at our racehorses, conceive that they were the result of the union of the Arab horse and the African mare?”[72] This change has, it is probable, been largely effected through unconscious selection, that is, by the general wish to breed as fine horses as possible in each generation, combined with training and high feeding, but without any intention to give to them their present appearance. According to Youatt,[73] the introduction in Oliver Cromwell’s time of three celebrated Eastern stallions speedily affected the English breed; “so that Lord Harleigh, one of the old school, complained that the great horse was fast disappearing.” This is an excellent proof how carefully selection must have been attended to; for without such care, all traces of so small an infusion of Eastern blood would soon have been absorbed and lost. Notwithstanding that the climate of England has never been esteemed particularly favourable to the horse, yet long-continued selection, both methodical and unconscious, together with that practised by the Arabs during a still longer and earlier period, has ended in giving us the best breed of horses in the world. Macaulay[74] remarks, “Two men whose authority on such subjects was held in great esteem, the Duke of Newcastle and Sir John Fenwick, pronounced that the meanest hack ever imported from Tangier would produce a finer progeny than could be expected from the best sire of our native breed. They would not readily have believed that a time would come when the princes and nobles of neighbouring lands would be as eager to obtain horses from England as ever the English had been to obtain horses from Barbary.”
The London dray-horse, which differs so much in appearance from any natural species, and which from its size has so astonished many Eastern princes, was probably formed by the heaviest and most powerful animals having been selected during many generations in Flanders and England, but without the least intention or expectation of creating a horse such as we now see. If we go back to an early period of history, we behold in the antique Greek statues, as Schaaffhausen has remarked,[75] a horse equally unlike a race or dray horse, and differing from any existing breed. The results of unconscious selection, in an early stage, are well shown in the difference between the flocks descended from the same stock, but separately reared by careful breeders. Youatt gives an excellent instance of this fact in the sheep belonging to Messrs. Buckley and Burgess, which “have been purely bred from the original stock of Mr. Bakewell for upwards of fifty years. There is not a suspicion existing in the mind of any one at all acquainted with the subject that the owner of either flock has deviated in any one instance from the pure blood of Mr. Bakewell’s flock; yet the difference between the sheep possessed by these two gentlemen is so great, that they have the appearance of being quite different varieties.”[76] I have seen several analogous and well marked cases with pigeons: for instance, I had a family of barbs descended from those long bred by Sir J. Sebright, and another family long bred by another fancier, and the two families plainly differed from each other.
Nathusius—and a more competent witness could not be cited—observes that, though the Shorthorns are remarkably uniform in appearance (except in colour), yet the individual character and wishes of each breeder become impressed on his cattle, so that different herds differ slightly from one another.[77] The Hereford cattle assumed their present well-marked character soon after the year 1769, through careful selection by Mr. Tomkins[78] and the breed has lately split into two strains—one strain having a white face, and differing slightly, it is said,[79] in some other points: but there is no reason to believe that this split, the origin of which is unknown, was intentionally made; it may with much more probability be attributed to different breeders having attended to different points. So again, the Berkshire breed of swine in the year 1810 had greatly changed from what it was in 1780; and since 1810 at least two distinct sub-breeds have arisen bearing the same name.[80] Keeping in mind how rapidly all animals increase, and that some must be annually slaughtered and some saved for breeding, then, if the same breeder during a long course of years deliberately settles which shall be saved and which shall be killed, it is almost inevitable that his individual turn of mind will influence the character of his stock, without his having had any intention to modify the breed.
Unconscious selection in the strictest sense of the word, that is, the saving of the more useful animals and the neglect or slaughter of the less useful, without any thought of the future, must have gone on occasionally from the remotest period and amongst the most barbarous nations. Savages often suffer from famines, and are sometimes expelled by war from their own homes. In such cases it can hardly be doubted that they would save their most useful animals. When the Fuegians are hard pressed by want, they kill their old women for food rather than their dogs; for, as we were assured, “old women no use—dogs catch otters.” The same sound sense would surely lead them to preserve their more useful dogs when still harder pressed by famine. Mr. Oldfield, who has seen so much of the aborigines of Australia, informs me that “they are all very glad to get a European kangaroo dog, and several instances have been known of the father killing his own infant that the mother might suckle the much-prized puppy.” Different kinds of dogs would be useful to the Australian for hunting opossums and kangaroos, and to the Fuegian for catching fish and otters; and the occasional preservation in the two countries of the most useful animals would ultimately lead to the formation of two widely distinct breeds.
Text from Project Gutenberg, public domain in the United States.