The Variation of Animals and Plants under Domestication/XXVIII

by Charles Darwin

CHAPTER XXVIII.

CONCLUDING REMARKS.

DOMESTICATION—NATURE AND CAUSES OF VARIABILITY—SELECTION—DIVERGENCE AND DISTINCTNESS OF CHARACTER—EXTINCTION OF RACES—CIRCUMSTANCES FAVOURABLE TO SELECTION BY MAN—ANTIQUITY OF CERTAIN RACES—THE QUESTION WHETHER EACH PARTICULAR VARIATION HAS BEEN SPECIALLY PREORDAINED.

As summaries have been added to nearly all the chapters, and as, in the chapter on pangenesis, various subjects, such as the forms of reproduction, inheritance, reversion, the causes and laws of variability, &c., have been recently discussed, I will here only make a few general remarks on the more important conclusions which may be deduced from the multifarious details given throughout this work.

Savages in all parts of the world easily succeed in taming wild animals; and those inhabiting any country or island, when first invaded by man, would probably have been still more easily tamed. Complete subjugation generally depends on an animal being social in its habits, and on receiving man as the chief of the herd or family. Domestication implies almost complete fertility under new and changed conditions of life, and this is far from being invariably the case. An animal would not have been worth the labour of domestication, at least during early times, unless of service to man. From these circumstances the number of domesticated animals has never been large. With respect to plants, I have shown in the ninth chapter how their varied uses were probably first discovered, and the early steps in their cultivation. Man could not have known, when he first domesticated an animal or plant, whether it would flourish and multiply when transported to other countries, therefore he could not have been thus influenced in his choice. We see that the close adaptation of the reindeer and camel to extremely cold and hot countries has not prevented their domestication. Still less could man have foreseen whether his animals and plants would vary in succeeding generations and thus give birth to new races; and the small capacity of variability in the goose and ass has not prevented their domestication from the remotest epoch.

With extremely few exceptions, all animals and plants which have been long domesticated, have varied greatly. It matters not under what climate, or for what purpose, they are kept, whether as food for man or beast, for draught or hunting, for clothing or mere pleasure,—under all these circumstances domesticated animals and plants have varied to a much greater extent than the forms which in a state of nature are ranked as one species. Why certain animals and plants have varied more under domestication than others we do not know, any more than why some are rendered more sterile than others under changed conditions of life. But we frequently judge of the amount of variation by the production of numerous and diversified races, and we can clearly see why in many cases this has not occurred, namely, because slight successive variations have not been steadily accumulated; and such variations will never be accumulated when an animal or plant is not closely observed, or much valued, or kept in large numbers.

The fluctuating, and, as far as we can judge, never-ending variability of our domesticated productions,—the plasticity of their whole organisation,—is one of the most important facts which we learn from the numerous details given in the earlier chapters of this work. Yet domesticated animals and plants can hardly have been exposed to greater changes in their conditions than have many natural species during the incessant geological, geographical, and climatal changes of the whole world. The former will, however, commonly have been exposed to more sudden changes and to less continuously uniform conditions. As man has domesticated so many animals and plants belonging to widely different classes, and as he certainly did not with prophetic instinct choose those species which would vary most, we may infer that all natural species, if subjected to analogous conditions, would, on an average, vary to the same degree. Few men at the present day will maintain that animals and plants were created with a tendency to vary, which long remained dormant, in order that fanciers in after ages might rear, for instance, curious breeds of the fowl, pigeon, or canary-bird.

From several causes it is difficult to judge of the amount of modification which our domestic productions have undergone. In some cases the primitive parent-stock has become extinct, or cannot be recognised with certainty owing to its supposed descendants having been so much modified. In other cases two or more closely allied forms, after being domesticated, have crossed; and then it is difficult to estimate how much of the change ought to be attributed to variation. But the degree to which our domestic breeds have been modified by the crossing of distinct natural forms has probably been exaggerated by some authors. A few individuals of one form would seldom permanently affect another form existing in much greater numbers; for, without careful selection, the stain of the foreign blood would soon be obliterated, and during early and barbarous times, when our animals were first domesticated, such care would seldom have been taken.

There is good reason to believe that several of the breeds of the dog, ox, pig, and of some other animals, are respectively descended from distinct wild prototypes; nevertheless the belief in the multiple origin of our domesticated animals has been extended by some few naturalists and by many breeders to an unauthorised extent. Breeders refuse to look at the whole subject under a single point of view; I have heard one, who maintained that our fowls were the descendants of at least half-a-dozen aboriginal species, protest that he was in no way concerned with the origin of pigeons, ducks, rabbits, horses, or any other animal. They overlook the improbability of many species having been domesticated at an early and barbarous period. They do not consider the improbability of species having existed in a state of nature which, if like our present domestic breeds, would have been highly abnormal in comparison with all their congeners. They maintain that certain species, which formerly existed, have become extinct or unknown, although the world is now so much better known. The assumption of so much recent extinction is no difficulty in their eyes; for they do not judge of its probability by the facility or difficulty of the extinction of other closely allied wild forms. Lastly, they often ignore the whole subject of geographical distribution as completely as if its laws were the result of chance.

Although from the reasons just assigned it is often difficult to judge accurately of the amount of change which our domesticated productions have undergone, yet this can be ascertained in the cases in which we know that all the breeds are descended from a single species, as with the pigeon, duck, rabbit, and almost certainly with the fowl; and by the aid of analogy this is to a certain extent possible in the case of animals descended from several wild stocks. It is impossible to read the details given in the earlier chapters, and in many published works, or to visit our various exhibitions, without being deeply impressed with the extreme variability of our domesticated animals and cultivated plants. I have in many instances purposely given details on new and strange peculiarities which have arisen. No part of the organisation escapes the tendency to vary. The variations generally affect parts of small vital or physiological importance, but so it is with the differences which exist between closely allied species. In these unimportant characters there is often a greater difference between the breeds of the same species than between the natural species of the same genus, as Isidore Geoffroy has shown to be the case with size, and as is often the case with the colour, texture, form, &c., of the hair, feathers, horns, and other dermal appendages.

It has often been asserted that important parts never vary under domestication, but this is a complete error. Look at the skull of the pig in any one of the highly improved breeds, with the occipital condyles and other parts greatly modified; or look at that of the niata ox. Or again, in the several breeds of the rabbit, observe the elongated skull, with the differently shaped occipital foramen, atlas, and other cervical vertebræ. The whole shape of the brain, together with the skull, has been modified in Polish fowls; in other breeds of the fowl the number of the vertebræ and the forms of the cervical vertebræ have been changed. In certain pigeons the shape of the lower jaw, the relative length of the tongue, the size of the nostrils and eyelids, the number and shape of the ribs, the form and size of the œsophagus, have all varied. In certain quadrupeds the length of the intestines has been much increased or diminished. With plants we see wonderful differences in the stones of various fruits. In the Cucurbitaceæ several highly important characters have varied, such as the sessile position of the stigmas on the ovarium, the position of the carpels within the ovarium, and its projection out of the receptacle. But it would be useless to run through the many facts given in the earlier chapters.

It is notorious how greatly the mental disposition, tastes, habits, consensual movements, loquacity or silence, and the tone of voice have varied and been inherited with our domesticated animals. The dog offers the most striking instance of changed mental attributes, and these differences cannot be accounted for by descent from distinct wild types. New mental characters have certainly often been acquired, and natural ones lost, under domestication.

New characters may appear and disappear at any stage of growth, and be inherited at a corresponding period. We see this in the difference between the eggs of various breeds of the fowl, and in the down on chickens; and still more plainly in the differences between the caterpillars and cocoons of various breeds of the silk-moth. These facts, simple as they appear, throw light on the characters which distinguish the larval and adult states of natural species, and on the whole great subject of embryology. New characters are liable to become attached exclusively to that sex in which they first appeared, or they may be developed in a much higher degree in the one than the other sex; or again, after having become attached to one sex, they may be partially transferred to the opposite sex. These facts, and more especially the circumstance that new characters seem to be particularly liable, from some unknown cause, to become attached to the male sex, have an important bearing on the acquirement by animals in a state of nature of secondary sexual characters.

It has sometimes been said that our domestic productions do not differ in constitutional peculiarities, but this cannot be maintained. In our improved cattle, pigs, &c., the period of maturity, including that of the second dentition, has been much hastened. The period of gestation varies much, but has been modified in a fixed manner in only one or two cases. In our poultry and pigeons the acquirement of down and of the first plumage by the young, and of the secondary sexual characters by the males, differ. The number of moults through which the larvæ of silk-moths pass, varies. The tendency to fatten, to yield much milk, to produce many young or eggs at a birth or during life, differs in different breeds. We find different degrees of adaptation to climate, and different tendencies to certain diseases, to the attacks of parasites, and to the action of certain vegetable poisons. With plants, adaptation to certain soils, as with some kinds of plums, the power of resisting frost, the period of flowering and fruiting, the duration of life, the period of shedding the leaves and of retaining them throughout the winter, the proportion and nature of certain chemical compounds in the tissues or seeds, all vary.

There is, however, one important constitutional difference between domestic races and species; I refer to the sterility which almost invariably follows, in a greater or less degree, when species are crossed, and to the perfect fertility of the most distinct domestic races, with the exception of a very few plants, when similarly crossed. It certainly appears a remarkable fact that many closely allied species which in appearance differ extremely little should yield when united only a few, more or less sterile offspring, or none at all; whilst domestic races which differ conspicuously from each other, are when united remarkably fertile, and yield perfectly fertile offspring. But this fact is not in reality so inexplicable as it at first appears. In the first place, it was clearly shown in the nineteenth chapter that the sterility of crossed species does not closely depend on differences in their external structure or general constitution, but results exclusively from differences in the reproductive system, analogous with those which cause the lessened fertility of the illegitimate unions and illegitimate offspring of dimorphic and trimorphic plants. In the second place, the Pallasian doctrine, that species after having been long domesticated lose their natural tendency to sterility when crossed, has been shown to be highly probable; we can scarcely avoid this conclusion when we reflect on the parentage and present fertility of the several breeds of the dog, of Indian and European cattle, sheep, and pigs. Hence it would be unreasonable to expect that races formed under domestication should acquire sterility when crossed, whilst at the same time we admit that domestication eliminates the normal sterility of crossed species. Why with closely allied species their reproductive systems should almost invariably have been modified in so peculiar a manner as to be mutually incapable of acting on each other—though in unequal degrees in the two sexes, as shown by the difference in fertility between reciprocal crosses in the same species—we do not know, but may with much probability infer the cause to be as follows. Most natural species have been habituated to nearly uniform conditions of life for an incomparably longer period of time than have domestic races; and we positively know that changed conditions exert an especial and powerful influence on the reproductive system. Hence this difference in habituation may well account for the different action of the reproductive organs when domestic races and when species are crossed. It is a nearly analogous fact, that most domestic races may be suddenly transported from one climate to another, or be placed under widely different conditions, and yet retain their fertility unimpaired; whilst a multitude of species subjected to lesser changes are rendered incapable of breeding.

With the exception of fertility, domestic varieties resemble species when crossed in transmitting their characters in the same unequal manner to their offspring, in being subject to the prepotency of one form over the other, and in their liability to reversion. By repeated crosses a variety or a species may be made completely to absorb another. Varieties, as we shall see when we treat of their antiquity, sometimes inherit their new characters almost, or even quite, as firmly as species. With both, the conditions leading to variability and the laws governing its nature appear to be the same. Domestic varieties can be classed in groups under groups, like species under genera, and these under families and orders; and the classification may be either artificial,—that is, founded on any arbitrary character,—or natural. With varieties a natural classification is certainly founded, and with species is apparently founded, on community of descent, together with the amount of modification which the forms have undergone. The characters by which domestic varieties differ from each other are more variable than those distinguishing species, though hardly more so than with certain protean species; but this greater degree of variability is not surprising, as varieties have generally been exposed within recent times to fluctuating conditions of life, are much more liable to have been crossed, and are still in many cases undergoing, or have recently undergone, modification by man's methodical or unconscious selection.

Domestic varieties as a general rule certainly differ from each other in less important parts of their organisation than do species; and when important differences occur, they are seldom firmly fixed; but this fact is intelligible if we consider man's method of selection. In the living animal or plant he cannot observe internal modifications in the more important organs; nor does he regard them as long as they are compatible with health and life. What does the breeder care about any slight change in the molar teeth of his pigs, or for an additional molar tooth in the dog; or for any change in the intestinal canal or other internal organ? The breeder cares for the flesh of his cattle being well marbled with fat, and for an accumulation of fat within the abdomen of his sheep, and this he has effected. What would the floriculturist care for any change in the structure of the ovarium or of the ovules? As important internal organs are certainly liable to numerous slight variations, and as these would probably be inherited, for many strange monstrosities are transmitted, man could undoubtedly effect a certain amount of change in these organs. When he has produced any modification in an important part, it has generally been unintentionally in correlation with some other conspicuous part, as when he has given ridges and protuberances to the skulls of fowls, by attending to the form of the comb, and in the case of the Polish fowl to the plume of feathers on the head. By attending to the external form of the pouter-pigeon, he has enormously increased the size of the œsophagus, and has added to the number of the ribs, and given them greater breadth. With the carrier-pigeon, by increasing, through steady selection, the wattles on the upper mandible, he has greatly modified the form of the lower mandible; and so in many other cases. Natural species, on the other hand, have been modified exclusively for their own good, to fit them for infinitely diversified conditions of life, to avoid enemies of all kinds, and to struggle against a host of competitors. Hence, under such complex conditions, it would often happen that modifications of the most varied kinds, in important as well as in unimportant parts, would be advantageous or even necessary; and they would slowly but surely be acquired through the survival of the fittest. Various indirect modifications would likewise arise through the law of correlated variation.

Domestic breeds often have an abnormal or semi-monstrous character, as the Italian greyhound, bulldog, Blenheim spaniel, and bloodhound amongst dogs,—some breeds of cattle and pigs, several breeds of the fowl, and the chief breeds of the pigeon. The differences between such abnormal breeds occur in parts which in closely-allied natural species differ but slightly or not at all. This may be accounted for by man's often selecting, especially at first, conspicuous and semi-monstrous deviations of structure. We should, however, be cautious in deciding what deviations ought to be called monstrous: there can hardly be a doubt that, if the brush of horse-like hair on the breast of the turkey-cock had first appeared on the domesticated bird, it would have been considered a monstrosity; the great plume of feathers on the head of the Polish cock has been thus designated, though plumes are common with many kinds of birds; we might call the wattle or corrugated skin round the base of the beak of the English carrier-pigeon a monstrosity, but we do not thus speak of the globular fleshy excrescence at the base of the beak of the male Carpophaga oceanica.

Some authors have drawn a wide distinction between artificial and natural breeds; although in extreme cases the distinction is plain, in many other cases an arbitrary line has to be drawn. The difference depends chiefly on the kind of selection which has been applied. Artificial breeds are those which have been intentionally improved by man; they frequently have an unnatural appearance, and are especially liable to loss of excellence through reversion and continued variability. The so-called natural breeds, on the other hand, are those which are now found in semi-civilised countries, and which formerly inhabited separate districts in nearly all the European kingdoms. They have been rarely acted on by man's intentional selection; more frequently, it is probable, by unconscious selection, and partly by natural selection, for animals kept in semi-civilised countries have to provide largely for their own wants. Such natural breeds will also, it may be presumed, have been directly acted on to some extent by the differences, though slight, in the surrounding physical conditions.

It is a much more important distinction that some breeds have been from their first origin modified in so slow and insensible a manner, that if we could see their early progenitors we should hardly be able to say when or how the breed first arose; whilst other breeds have originated from a strongly-marked or semi-monstrous deviation of structure, which, however, may subsequently have been augmented by selection. From what we know of the history of the racehorse, greyhound, gamecock, &c., and from their general appearance, we may feel nearly confident that they were formed by a slow process of improvement: and with the carrier-pigeon, as well as with some other pigeons, we know that this has been the case. On the other hand, it is certain that the ancon and mauchamp breeds of sheep, and almost certain that the niata cattle, turnspit and pug-dogs, jumper and frizzled fowls, short-faced tumbler pigeons, hook-billed ducks, &c., and with plants a multitude of varieties, suddenly appeared in nearly the same state as we now see them. The frequency of these cases is likely to lead to the false belief that natural species have often originated in the same abrupt manner. But we have no evidence of the appearance, or at least of the continued procreation, under nature, of abrupt modifications of structure; and various general reasons could be assigned against such a belief: for instance, without separation a single monstrous variation would almost certainly be soon obliterated by crossing.

On the other hand, we have abundant evidence of the constant occurrence under nature of slight individual differences of the most diversified kinds; and thus we are led to conclude that species have generally originated by the natural selection, not of abrupt modifications, but of extremely slight differences. This process may be strictly compared with the slow and gradual improvement of the racehorse, greyhound, and gamecock. As every detail of structure in each species is closely adapted to its general habits of life, it will rarely happen that one part alone will be modified; but the co-adapted modifications, as formerly shown, need not be absolutely simultaneous. Many variations, however, are from the first connected by the law of correlation. Hence it follows that even closely-allied species rarely or never differ from each other by some one character alone; and this same remark applies to a certain extent to domestic races; for these, if they differ much, generally differ in many respects.

Some naturalists boldly insist^[1] that species are absolutely distinct productions, never passing by intermediate links into each other; whilst they maintain that domestic varieties can always be connected either with each other or with their parent-forms. But if we could always find the links between the several breeds of the dog, horse, cattle, sheep, pigs, &c., the incessant doubts whether they are descended from one or several species would not have arisen. The greyhound genus, if such a term may be used, cannot be closely connected with any other breed, unless, perhaps, we go back to the ancient Egyptian monuments. Our English bulldog also forms a very distinct breed. In all these cases crossed breeds must of course be excluded, for the most distinct natural species can thus be connected. By what links can the Cochin fowl be closely united with others? By searching for breeds still preserved in distant lands, and by going back to historical records, tumbler-pigeons, carriers, and barbs can be closely connected with the parent rock-pigeon; but we cannot thus connect the turbit or the pouter. The degree of distinctness between the various domestic breeds depends on the amount of modification which they have undergone, and especially on the neglect and final extinction of the linking, intermediate, and less valued forms.

It has often been argued that no light is thrown, from the admitted changes of domestic races, on the changes which natural species are believed to undergo, as the former are said to be mere temporary productions, always reverting, as soon as they become feral, to their pristine form. This argument has been well combated by Mr. Wallace;^[2] and full details were given in the thirteenth chapter, showing that the tendency to reversion in feral animals and plants has been greatly exaggerated, though no doubt to a certain extent it exists. It would be opposed to all the principles inculcated in this work, if domestic animals, when exposed to new conditions and compelled to struggle for their own wants against a host of foreign competitors, were not in the course of time in some manner modified. It should also be remembered that many characters lie latent in all organic beings ready to be evolved under fitting conditions; and in breeds modified within recent times the tendency to reversion is particularly strong. But the antiquity of various breeds clearly proves that they remain nearly constant as long as their conditions of life remain the same.

It has been boldly maintained by some authors that the amount of variation to which our domestic productions are liable is strictly limited; but this is an assertion resting on little evidence. Whether or not the amount in any particular direction is fixed, the tendency to general variability seems unlimited. Cattle, sheep, and pigs have been domesticated and have varied from the remotest period, as shown by the researches of Rütimeyer and others, yet these animals have, within quite recent times, been improved in an unparalleled degree; and this implies continued variability of structure. Wheat, as we know from the remains found in the Swiss lake-habitations, is one of the most anciently cultivated plants, yet at the present day new and better varieties occasionally arise. It may be that an ox will never be produced of larger size or finer proportions than our present animals, or a race-horse fleeter than Eclipse, or a gooseberry larger than the London variety; but he would be a bold man who would assert that the extreme limit in these respects has been finally attained. With flowers and fruit it has repeatedly been asserted that perfection has been reached, but the standard has soon been excelled. A breed of pigeons may never be produced with a beak shorter than that of the present short-faced tumbler, or with one longer than that of the English carrier, for these birds have weak constitutions and are bad breeders; but the shortness and length of the beak are the points which have been steadily improved during at least the last 150 years; and some of the best judges deny that the goal has yet been reached. We may, also, reasonably suspect, from what we see in natural species of the variability of extremely modified parts, that any structure, after remaining constant during a long series of generations, would, under new and changed conditions of life, recommence its course of variability, and might again be acted on by selection. Nevertheless, as Mr. Wallace^[3] has recently remarked with much force and truth, there must be both with natural and domestic productions a limit to change in certain directions; for instance, there must be a limit to the fleetness of any terrestrial animal, as this will be determined by the friction to be overcome, the weight to be carried, and the power of contraction in the muscular fibres. The English racehorse may have reached this limit; but it already surpasses in fleetness its own wild progenitor, and all other equine species.

It is not surprising, seeing the great difference between many domestic breeds, that some few naturalists have concluded that all are descended from distinct aboriginal stocks, more especially as the principle of selection has been ignored, and the high antiquity of man, as a breeder of animals, has only recently become known. Most naturalists, however, freely admit that various extremely dissimilar breeds are descended from a single stock, although they do not know much about the art of breeding, cannot show the connecting links, nor say where and when the breeds arose. Yet these same naturalists will declare, with an air of philosophical caution, that they can never admit that one natural species has given birth to another until they behold all the transitional steps. But fanciers have used exactly the same language with respect to domestic breeds; thus an author of an excellent treatise says he will never allow that carrier and fantail pigeons are the descendants of the wild rock-pigeon, until the transitions have "actually been observed, and can be repeated whenever man chooses to set about the task." No doubt it is difficult to realise that slight changes added up during long centuries can produce such results; but he who wishes to understand the origin of domestic breeds or natural species must overcome this difficulty.

The causes inducing and the laws governing variability have been so lately discussed, that I need here only enumerate the leading points. As domesticated organisms are much more liable to slight deviations of structure and to monstrosities, than species living under their natural conditions, and as widely-ranging species vary more than those which inhabit restricted areas, we may infer that variability mainly depends on changed conditions of life. We must not overlook the effects of the unequal combination of the characters derived from both parents, nor reversion to former progenitors. Changed conditions have an especial tendency to render the reproductive organs more or less impotent, as shown in the chapter devoted to this subject; and these organs consequently often fail to transmit faithfully the parental characters. Changed conditions also act directly and definitely on the organisation, so that all or nearly all the individuals of the same species thus exposed become modified in the same manner; but why this or that part is especially affected we can seldom or never say. In most cases, however, of the direct action of changed conditions, independently of the indirect variability caused by the reproductive organs being affected, indefinite modifications are the result; in nearly the same manner as exposure to cold or the absorption of the same poison affects different individuals in various ways. We have reason to suspect that an habitual excess of highly nutritious food, or an excess relatively to the wear and tear of the organisation from exercise, is a powerful exciting cause of variability. When we see the symmetrical and complex outgrowths, caused by a minute atom of the poison of a gall-insect, we may believe that slight changes in the chemical nature of the sap or blood would lead to extraordinary modifications of structure.

The increased use of a muscle with its various attached parts, and the increased activity of a gland or other organ, lead to their increased development. Disuse has a contrary effect. With domesticated productions organs sometimes become rudimentary through abortion; but we have no reason to suppose that this has ever followed from mere disuse. With natural species, on the contrary, many organs appear to have been rendered rudimentary through disuse, aided by the principle of the economy of growth, and by the hypothetical principle discussed in the last chapter, namely, the final destruction of the germs or gemmules of such useless parts. This difference may be partly accounted for by disuse having acted on domestic forms for an insufficient length of time, and partly from their exemption from any severe struggle for existence, entailing rigid economy in the development of each part, to which all species under nature are subjected. Nevertheless the law of compensation or balancement apparently affects, to a certain extent, our domesticated productions.

We must not exaggerate the importance of the definite action of changed conditions in modifying all the individuals of the same species in the same manner, or of use and disuse. As every part of the organisation is highly variable, and as variations are so easily selected, both consciously and unconsciously, it is very difficult to distinguish between the effects of the selection of indefinite variations, and the direct action of the conditions of life. For instance, it is possible that the feet of our water-dogs, and of the American dogs which have to travel much over the snow, may have become partially webbed from the stimulus of widely extending their toes; but it is far more probable that the webbing, like the membrane between the toes of certain pigeons, spontaneously appeared and was afterwards increased by the best swimmers and the best snow-travellers being preserved during many generations. A fancier who wished to decrease the size of his bantams or tumbler-pigeons would never think of starving them, but would select the smallest individuals which spontaneously appeared. Quadrupeds are sometimes born destitute of hair, and hairless breeds have been formed, but there is no reason to believe that this is caused by a hot climate. Within the tropics heat often causes sheep to lose their fleeces, and on the other hand wet and cold act as a direct stimulus to the growth of hair; it is, however, possible that these changes may merely be an exaggeration of the regular yearly change of coat; and who will pretend to decide how far this yearly change, or the thick fur of arctic animals, or as I may add their white colour, is due to the direct action of a severe climate, and how far to the preservation of the best protected individuals during a long succession of generations?

Of all the laws governing variability, that of correlation is the most important. In many cases of slight deviations of structure as well as of grave monstrosities, we cannot even conjecture what is the nature of the bond of connexion. But between homologous parts—between the fore and hind limbs—between the hair, hoofs, horns, and teeth—we can see that parts which are closely similar during their early development, and which are exposed to similar conditions, would be liable to be modified in the same manner. Homologous parts, from having the same nature, are apt to blend together and, when many exist, to vary in number.

Although every variation is either directly or indirectly caused by some change in the surrounding conditions, we must never forget that the nature of the organisation which is acted on essentially governs the result. Distinct organisms, when placed under similar conditions, vary in different manners, whilst closely-allied organisms under dissimilar conditions often vary in nearly the same manner. We see this in the same modification frequently reappearing at long intervals of time in the same variety, and likewise in the several striking cases given of analogous or parallel varieties. Although some of these latter cases are simply due to reversion, others cannot thus be accounted for.

From the indirect action of changed conditions on the organisation, through the impaired state of the reproductive organs—from the direct action of such conditions (and this will cause the individuals of the same species either to vary in the same manner, or differently in accordance with slight differences in their constitution)—from the effects of the increased or decreased use of parts,—and from correlation,—the variability of our domesticated productions is complicated in an extreme degree. The whole organisation becomes slightly plastic. Although each modification must have its proper exciting cause, and though each is subjected to law, yet we can so rarely trace the precise relation between cause and effect, that we are tempted to speak of variations as if they spontaneously arose. We may even call them accidental, but this must be only in the sense in which we say that a fragment of rock dropped from a height owes its shape to accident.

 

It may be worth while briefly to consider the results of the exposure to unnatural conditions of a large number of animals of the same species, allowed to cross freely, with no selection of any kind; and afterwards to consider the results when selection is brought into play. Let us suppose that 500 wild rock-pigeons were confined in their native land in an aviary, and fed in the same manner as pigeons usually are; and that they were not allowed to increase in number. As pigeons propagate so rapidly, I suppose that a thousand or fifteen hundred birds would have to be annually killed by mere chance. After several generations had been thus reared, we may feel sure that some of the young birds would vary, and the variations would tend to be inherited; for at the present day slight deviations of structure often occur, but, as most breeds are already well established, these modifications are rejected as blemishes. It would be tedious even to enumerate the multitude of points which still go on varying or have recently varied. Many variations would occur in correlation, as the length of the wing and tail feathers—the number of the primary wing-feathers, as well as the number and breadth of the ribs, in correlation with the size and form of the body—the number of the scutellæ, with the size of the feet—the length of the tongue, with the length of the beak—the size of the nostrils and eyelids and the form of lower jaw in correlation with the development of wattle—the nakedness of the young with the future colour of the plumage—the size of the feet and beak, and other such points. Lastly, as our birds are supposed to be confined in an aviary, they would use their wings and legs but little, and certain parts of the skeleton, such as the sternum and scapulæ and the feet, would in consequence become slightly reduced in size.

As in our assumed case many birds have to be indiscriminately killed every year, the chances are against any new variety surviving long enough to breed. And as the variations which arise are of an extremely diversified nature, the chances are very great against two birds pairing which have varied in the same manner; nevertheless, a varying bird even when not thus paired would occasionally transmit its character to its young; and these would not only be exposed to the same conditions which first caused the variation in question to appear, but would in addition inherit from their one modified parent a tendency again to vary in the same manner. So that, if the conditions decidedly tended to induce some particular variation, all the birds might in the course of time become similarly modified. But a far commoner result would be, that one bird would vary in one way and another bird in another way; one would be born with a little longer beak, and another with a shorter beak; one would gain some black feathers, another some white or red feathers. And as these birds would be continually intercrossing, the final result would be a body of individuals differing from each other slightly in many ways, yet far more than did the original rock-pigeons. But there would not be the least tendency to the formation of distinct breeds.

If two separate lots of pigeons were to be treated in the manner just described, one in England and the other in a tropical country, the two lots being supplied with different food, would they, after many generations had passed, differ? When we reflect on the cases given in the twenty-third chapter, and on such facts as the difference in former times between the breeds of cattle, sheep, &c., in almost every district of Europe, we are strongly inclined to admit that the two lots would be differently modified through the influence of climate and food. But the evidence on the definite action of changed conditions is in most cases insufficient; and, with respect to pigeons, I have had the opportunity of examining a large collection of domesticated birds, sent to me by Sir W. Elliot from India, and they varied in a remarkably similar manner with our European birds.

If two distinct breeds were to be confined together in equal numbers, there is reason to suspect that they would to a certain extent prefer pairing with their own kind; but they would likewise intercross. From the greater vigour and fertility of the crossed offspring, the whole body would by this means become interblended sooner than would otherwise have occurred. From certain breeds being prepotent over others, it does not follow that the interblended progeny would be strictly intermediate in character. I have, also, proved that the act of crossing in itself gives a strong tendency to reversion, so that the crossed offspring would tend to revert to the state of the aboriginal rock-pigeon. In the course of time they would probably be not much more heterogeneous in character than in our first case, when birds of the same breed were confined together.

I have just said that the crossed offspring would gain in vigour and fertility. From the facts given in the seventeenth chapter there can be no doubt of this; and there can be little doubt, though the evidence on this head is not so easily acquired, that long-continued close interbreeding leads to evil results. With hermaphrodites of all kinds, if the sexual elements of the same individual habitually acted on each other, the closest possible interbreeding would be perpetual. Therefore we should bear in mind that with all hermaphrodite animals, as far as I can learn, their structure permits and frequently necessitates a cross with a distinct individual. With hermaphrodite plants we incessantly meet with elaborate and perfect contrivances for this same end. It is no exaggeration to assert that, if the use of the talons and tusks of a carnivorous animal, or the use of the viscid threads of a spider's web, or of the plumes and hooks on a seed may be safely inferred from their structure, we may with equal safety infer that many flowers are constructed for the express purpose of ensuring a cross with a distinct plant. From these various considerations, the conclusion arrived at in the chapter just referred to—namely, that great good of some kind is derived from the sexual concourse of distinct individuals—must be admitted.

To return to our illustration: we have hitherto assumed that the birds were kept down to the same number by indiscriminate slaughter; but if the least choice be permitted in their preservation and slaughter, the whole result will be changed. Should the owner observe any slight variation in one of his birds, and wish to obtain a breed thus characterised, he would succeed in a surprisingly short time by carefully selecting and pairing the young. As any part which has once varied generally goes on varying in the same direction, it is easy, by continually preserving the most strongly marked individuals, to increase the amount of difference up to a high, predetermined standard of excellence. This is methodical selection.

If the owner of the aviary, without any thought of making a new breed, simply admired, for instance, short-beaked more than long-beaked birds, he would, when he had to reduce the number, generally kill the latter; and there can be no doubt that he would thus in the course of time sensibly modify his stock. It is improbable, if two men were to keep pigeons and act in this manner, that they would prefer exactly the same characters; they would, as we know, often prefer directly opposite characters, and the two lots would ultimately come to differ. This has actually occurred with strains or families of cattle, sheep, and pigeons, which have been long kept and carefully attended to by different breeders without any wish on their part to form new and distinct sub-breeds. This unconscious kind of selection will more especially come into action with animals which are highly serviceable to man; for every one tries to get the best dog, horse, cow, or sheep, and these animals will transmit more or less surely their good qualities to their offspring. Hardly any one is so careless as to breed from his worst animals. Even savages, when compelled from extreme want to kill some of their animals, would destroy the worst and preserve the best. With animals kept for use and not for mere amusement, different fashions prevail in different districts, leading to the preservation, and consequently to the transmission, of all sorts of trifling peculiarities of character. The same process will have been pursued with our fruit-trees and vegetables, for the best will always have been the most largely cultivated, and will occasionally have yielded seedlings better than their parents.

The different strains, just alluded to, which have been raised by different breeders without any wish for such a result, and the unintentional modification of foreign breeds in their new homes, both afford excellent evidence of the power of unconscious selection. This form of selection has probably led to far more important results than methodical selection, and is likewise more important under a theoretical point of view from closely resembling natural selection. For during this process the best or most valued individuals are not separated and prevented crossing with others of the same breed, but are simply preferred and preserved; but this inevitably leads during a long succession of generations to their increase in number and to their gradual improvement; so that finally they prevail to the exclusion of the old parent-form.

With our domesticated animals natural selection checks the production of races with any injurious deviation of structure. In the case of animals kept by savages and semi-civilised people, which have to provide largely for their own wants under different circumstances, natural selection will probably play a more important part. Hence such animals often closely resemble natural species.

As there is no limit to man's desire to possess animals and plants more and more useful in any respect, and as the fancier always wishes, from fashion running into extremes, to produce each character more and more strongly pronounced, there is a constant tendency in every breed, through the prolonged action of methodical and unconscious selection, to become more and more different from its parent-stock; and when several breeds have been produced and are valued for different qualities, to differ more and more from each other. This leads to Divergence of Character. As improved sub-varieties and races are slowly formed, the older and less improved breeds are neglected and decrease in number. When few individuals of any breed exist within the same locality, close interbreeding, by lessening their vigour and fertility, aids in their final extinction. Thus the intermediate links are lost, and breeds which have already diverged gain Distinctness of Character.

In the chapters on the Pigeon, it was proved by historical details and by the existence of connecting sub-varieties in distant lands that several breeds have steadily diverged in character, and that many old and intermediate sub-breeds have become extinct. Other cases could be adduced of the extinction of domestic breeds, as of the Irish wolf-dog, the old English hound, and of two breeds in France, one of which was formerly highly valued.^[4] Mr. Pickering remarks^[5] that "the sheep figured on the most ancient Egyptian monuments is unknown at the present day; and at least one variety of the bullock, formerly known in Egypt, has in like manner become extinct." So it has been with some animals, and with several plants cultivated by the ancient inhabitants of Europe during the neolithic period. In Peru, Von Tschudi^[6] found in certain tombs, apparently prior to the dynasty of the Incas, two kinds of maize not now known in the country. With our flowers and culinary vegetables, the production of new varieties and their extinction has incessantly recurred. At the present time improved breeds sometimes displace at an extraordinarily rapid rate older breeds; as has recently occurred throughout England with pigs. The Long-horn cattle in their native home were "suddenly swept away as if by some murderous pestilence," by the introduction of Short-horns.^[7]

What grand results have followed from the long-continued action of methodical and unconscious selection, checked and regulated to a certain extent by natural selection, is seen on every side of us. Compare the many animals and plants which are displayed at our exhibitions with their parent-forms when these are known, or consult old historical records with respect to their former state. Almost all our domesticated animals have given rise to numerous and distinct races, excepting those which cannot be easily subjected to selection—such as cats, the cochineal insect, and the hive-bee,—and excepting those animals which are not much valued. In accordance with what we know of the process of selection, the formation of our many races has been slow and gradual. The man who first observed and preserved a pigeon with its œsophagus a little enlarged, its beak a little longer, or its tail a little more expanded than usual, never dreamed that he had made the first step in the creation of the pouter, carrier, and fantail-pigeon. Man can create not only anomalous breeds, but others with their whole structure admirably co-ordinated for certain purposes, such as the race-horse and dray-horse, or the greyhound. It is by no means necessary that each small change of structure throughout the body, leading towards excellence, should simultaneously arise and be selected. Although man seldom attends to differences in organs which are important under a physiological point of view, yet he has so profoundly modified some breeds, that assuredly, if found wild, they would be ranked under distinct genera.

The best proof of what selection has effected is perhaps afforded by the fact that whatever part or quality in any animal, and more especially in any plant, is most valued by man, that part or quality differs most in the several races. This result is well seen by comparing the amount of difference between the fruits produced by the varieties of the same fruit-tree, between the flowers of the varieties in our flower-garden, between the seeds, roots, or leaves of our culinary and agricultural plants, in comparison with the other and not valued parts of the same plants. Striking evidence of a different kind is afforded by the fact ascertained by Oswald Heer,^[8] namely, that the seeds of a large number of plants,—wheat, barley, oats, peas, beans, lentils, poppies,—cultivated for their seed by the ancient Lake-inhabitants of Switzerland, were all smaller than the seeds of our existing varieties. Rütimeyer has shown that the sheep and cattle which were kept by the earlier Lake-inhabitants were likewise smaller than our present breeds. In the middens of Denmark, the earliest dog of which the remains have been found was the weakest; this was succeeded during the Bronze age by a stronger kind, and this again during the Iron age by one still stronger. The sheep of Denmark during the Bronze period had extraordinarily slender limbs, and the horse was smaller than our present animal.^[9] No doubt in these cases the new and larger breeds were generally introduced from foreign lands by the immigration of new hordes of men. But it is not probable that each larger breed, which in the course of time supplanted a previous and smaller breed, was the descendant of a distinct and larger species; it is far more probable that the domestic races of our various animals were gradually improved in different parts of the great Europæo-Asiatic continent, and thence spread to other countries. This fact of the gradual increase in size of our domestic animals is all the more striking as certain wild or half-wild animals, such as red-deer, aurochs, park-cattle, and boars,^[10] have within nearly the same period decreased in size.

The conditions favourable to selection by man are,—the closest attention being paid to every character,—long-continued perseverance,—facility in matching or separating animals,—and especially a large number being kept, so that the inferior individuals may be freely rejected or destroyed, and the better ones preserved. When many are kept there will also be a greater chance of the occurrence of well-marked deviations of structure. Length of time is all-important; for as each character, in order to become strongly pronounced, has to be augmented by the selection of successive variations of the same nature, this can only be effected during a long series of generations. Length of time will, also, allow any new feature to become fixed by the continued rejection of those individuals which revert or vary, and the preservation of those which inherit the new character. Hence, although some few animals have varied rapidly in certain respects under new conditions of life, as dogs in India and sheep in the West Indies, yet all the animals and plants which have produced strongly marked races were domesticated at an extremely remote epoch, often before the dawn of history. As a consequence of this, no record has been preserved of the origin of our chief domestic breeds. Even at the present day new strains or sub-breeds are formed so slowly that their first appearance passes unnoticed. A man attends to some particular character, or merely matches his animals with unusual care, and after a time a slight difference is perceived by his neighbours;—the difference goes on being augmented by unconscious and methodical selection, until at last a new sub-breed is formed, receives a local name, and spreads; but, by this time, its history is almost forgotten. When the new breed has spread widely, it gives rise to new strains and sub-breeds, and the best of these succeed and spread, supplanting other and older breeds; and so always onwards in the march of improvement.

When a well-marked breed has once been established, if not supplanted by still improving sub-breeds, and if not exposed to greatly changed conditions of life, inducing further variability or reversion to long-lost characters, it may apparently last for an enormous period. We may infer that this is the case from the high antiquity of certain races; but some caution is necessary on this head, for the same variation may appear independently after long intervals of time, or in distant places. We may safely assume that this has occurred with the turnspit-dog which is figured on the ancient Egyptian monuments, with the solid-hoofed swine^[11] mentioned by Aristotle, with five-toed fowls described by Columella, and certainly with the nectarine. The dogs represented on the Egyptian monuments, about 2000 B.C., show us that some of the chief breeds then existed, but it is extremely doubtful whether any are identically the same with our present breeds. A great mastiff sculptured on an Assyrian tomb, 640 B.C., is said to be the same with the dog still imported into the same region from Thibet. The true greyhound existed during the Roman classical period. Coming down to a later period, we have seen that, though most of the chief breeds of the pigeon existed between two and three centuries ago, they have not all retained to the present day exactly the same character; but this has occurred in certain cases in which improvement was not desired, for instance in the case of the Spot or the Indian ground-tumbler.

De Candolle^[12] has fully discussed the antiquity of various races of plants; he states that the black-seeded poppy was known in the time of Homer, the white-seeded sesamum by the ancient Egyptians, and almonds with sweet and bitter kernels by the Hebrews; but it does not seem improbable that some of these varieties may have been lost and reappeared. One variety of barley and apparently one of wheat, both of which were cultivated at an immensely remote period by the Lake-inhabitants of Switzerland, still exist. It is said^[13] that "specimens of a small variety of gourd which is still common in the market of Lima were exhumed from an ancient cemetery in Peru." De Candolle remarks that, in the books and drawings of the sixteenth century, the principal races of the cabbage, turnip, and gourd can be recognised; this might have been expected at so late a period, but whether any of these plants are absolutely identical with our present sub-varieties is not certain. It is, however, said that the Brussels sprout, a variety which in some places is liable to degeneration, has remained genuine for more than four centuries in the district where it is believed to have originated.^[14]

 

In accordance with the views maintained by me in this work and elsewhere, not only the various domestic races, but the most distinct genera and orders within the same great class,—for instance, whales, mice, birds, and fishes—are all the descendants of one common progenitor, and we must admit that the whole vast amount of difference between these forms of life has primarily arisen from simple variability. To consider the subject under this point of view is enough to strike one dumb with amazement. But our amazement ought to be lessened when we reflect that beings, almost infinite in number, during an almost infinite lapse of time, have often had their whole organisation rendered in some degree plastic, and that each slight modification of structure which was in any way beneficial under excessively complex conditions of life, will have been preserved, whilst each which was in any way injurious will have been rigorously destroyed. And the long-continued accumulation of beneficial variations will infallibly lead to structures as diversified, as beautifully adapted for various purposes, and as excellently co-ordinated, as we see in the animals and plants all around us. Hence I have spoken of selection as the paramount power, whether applied by man to the formation of domestic breeds, or by nature to the production of species. I may recur to the metaphor given in a former chapter: if an architect were to rear a noble and commodious edifice, without the use of cut stone, by selecting from the fragments at the base of a precipice wedge-formed stones for his arches, elongated stones for his lintels, and flat stones for his roof, we should admire his skill and regard him as the paramount power. Now, the fragments of stone, though indispensable to the architect, bear to the edifice built by him the same relation which the fluctuating variations of each organic being bear to the varied and admirable structures ultimately acquired by its modified descendants.

Some authors have declared that natural selection explains nothing, unless the precise cause of each slight individual difference be made clear. Now, if it were explained to a savage utterly ignorant of the art of building, how the edifice had been raised stone upon stone, and why wedge-formed fragments were used for the arches, flat stones for the roof, &c.; and if the use of each part and of the whole building were pointed out, it would be unreasonable if he declared that nothing had been made clear to him, because the precise cause of the shape of each fragment could not be given. But this is a nearly parallel case with the objection that selection explains nothing, because we know not the cause of each individual difference in the structure of each being.

The shape of the fragments of stone at the base of our precipice may be called accidental, but this is not strictly correct; for the shape of each depends on a long sequence of events, all obeying natural laws; on the nature of the rock, on the lines of deposition or cleavage, on the form of the mountain which depends on its upheaval and subsequent denudation, and lastly on the storm or earthquake which threw down the fragments. But in regard to the use to which the fragments may be put, their shape may be strictly said to be accidental. And here we are led to face a great difficulty, in alluding to which I am aware that I am travelling beyond my proper province. An omniscient Creator must have foreseen every consequence which results from the laws imposed by Him. But can it be reasonably maintained that the Creator intentionally ordered, if we use the words in any ordinary sense, that certain fragments of rock should assume certain shapes so that the builder might erect his edifice? If the various laws which have determined the shape of each fragment were not predetermined for the builder's sake, can it with any greater probability be maintained that He specially ordained for the sake of the breeder each of the innumerable variations in our domestic animals and plants;—many of these variations being of no service to man, and not beneficial, far more often injurious, to the creatures themselves? Did He ordain that the crop and tail-feathers of the pigeon should vary in order that the fancier might make his grotesque pouter and fantail breeds? Did He cause the frame and mental qualities of the dog to vary in order that a breed might be formed of indomitable ferocity, with jaws fitted to pin down the bull for man's brutal sport? But if we give up the principle in one case,—if we do not admit that the variations of the primeval dog were intentionally guided in order that the greyhound, for instance, that perfect image of symmetry and vigour, might be formed,—no shadow of reason can be assigned for the belief that variations, alike in nature and the result of the same general laws, which have been the groundwork through natural selection of the formation of the most perfectly adapted animals in the world, man included, were intentionally and specially guided. However much we may wish it, we can hardly follow Professor Asa Gray in his belief "that variation has been led along certain beneficial lines," like a stream "along definite and useful lines of irrigation." If we assume that each particular variation was from the beginning of all time preordained, the plasticity of organisation, which leads to many injurious deviations of structure, as well as that redundant power of reproduction which inevitably leads to a struggle for existence, and, as a consequence, to the natural selection or survival of the fittest, must appear to us superfluous laws of nature. On the other hand, an omnipotent and omniscient Creator ordains everything and foresees everything. Thus we are brought face to face with a difficulty as insoluble as is that of free will and predestination.

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1. Godron, 'De l'Espèce,' 1859, tom. ii. p. 44, &c.
2. Journal Proc. Linn. Soc., 1858, vol. iii. p. 60.
3. 'The Quarterly Journal of Science,' Oct. 1867, p. 486.
4. M. Rufz de Lavison, in 'Bull. Soc. Imp. d'Acclimat.,' Dec. 1862, p. 1009.
5. 'Races of Man,' 1850, p. 315.
6. 'Travels in Peru,' Eng. translat., p. 177.
7. Youatt on Cattle, 1834, p 200: on Pigs; see 'Gard. Chronicle,' 1854, p. 410.
8. 'Die Pflanzen der Pfahlbauten,' 1865.
9. Morlot, 'Soc. Vaud. des Scien. Nat,' Mars 1860, p. 298.
10. Rütimeyer, 'Die Fauna der Pfahlbauten,' 1861, s. 30.
11. Godron, 'De l'Espèce,' tom. i., 1859, p. 368.
12. 'Géographie Botan.,' 1855, p. 989.
13. Pickering, 'Races of Man,' 1850, p. 318.
14. 'Journal of a Horticultural Tour,' by a Deputation of the Caledonian Hist. Soc., 1823, p. 293.