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The Variation of Animals and Plants under Domestication
by
Charles Darwin
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The Variation of Animals and Plants under Domestication

The Variation of Animals and Plants under Domestication ............................................... 2

 

The Variation of Animals and Plants.................................................................................. 4

 

Introduction....................................................................................................................... 13

 

Chapter I: Domestic Dogs And Cats................................................................................. 22

 

Chapter II: Horses And Asses........................................................................................... 45

 

Chapter III: Pigs - Cattle - Sheep – Goats......................................................................... 56

 

Chapter IV: Domestic Rabbits .......................................................................................... 80

 

Chapter V: Domestic Pigeons........................................................................................... 98

 

Chapter VI: Pigeons – continued .................................................................................... 127

 

Chapter VII: Fowls ......................................................................................................... 156

 

Duck - Goose - Peacock - Turkey - Guinea-Fowl - Canary-Bird - Gold-Fish - River-Bees

 

- Silk-Moths .................................................................................................................... 188

 

THE TURKEY........................................................................................................ 198

 

THE GUINEA FOWL. ........................................................................................... 199

 

GOLD-FISH. .......................................................................................................... 200

 

HIVE-BEES............................................................................................................ 201

 

SILK-MOTHS. ....................................................................................................... 202

 

Chapter IX: Cultivated Plants: Cereal And Culinary Plants........................................... 209

 

Chapter X: Plants – continued ........................................................................................ 228

 

Chapter XI: On Bud-Variation........................................................................................ 256

 

Chapter XII: Inheritance ................................................................................................. 287

 

Chapter XIII: Inheritance – continued ............................................................................ 305 Chapter XIV: Inheritance – continued............................................................................ 328

 

Chapter XV: On Crossing............................................................................................... 344

 

Chapter XVI: Causes Which Interfere With The Free Crossing Of Varieties................ 354

 

Chapter XVII: On The Good Effects Of Crossing, And On The Evil Effects Of Close Interbreeding ................................................................................................................... 363

 

Chapter XVIII: On The Advantages And Disadvantages Of Changed Conditions Of Life ......................................................................................................................................... 385

 

Chapter XIX: Summary Of The Four Last Chapters, With Remarks On Hybridism..... 405

 

Chapter XX: Selection By Man ...................................................................................... 417

 

Chapter XXI: Selection, continued ................................................................................. 438

 

Chapter XXII: Causes Of Variability ............................................................................. 456

 

Chapter XXIII: Direct And Definite Action Of The External Conditions Of Life......... 470

 

Chapter XXIV: Laws Of Variation - Use And Disuse, Etc............................................ 485

 

Chapter XXV: Laws Of Variation, continued. - Correlated Variability......................... 503

 

Chapter XXVI: Laws Of Variation, continued – Summary ........................................... 517

 

Chapter XXVII: Provisional Hypothesis Of Pangenesis ................................................ 527 Chapter XXVIII: Concluding Remarks .......................................................................... 559

The Variation of Animals and Plants

FOREWORD
by
Harriet Ritvo

Charles Darwin wrote On the Origin of Species in a hurry. He had, it was true, been formulating his ideas and arguments for several decades—since his round-the-world Beagle voyage of 1831-1836. These ideas and arguments had been slow to take definitive shape; Darwin had nurtured and reworked them, amassing evidence for what he projected to be a weighty magnum opus. Although he had shared his developing evolutionary speculations with his closest professional colleagues, Darwin was reluctant to publish them on several grounds. He was aware that his theory of evolution by natural selection (or descent with modification) was complex, that it rested on vast but not incontrovertible evidence, and that the chain of his reasoning was not uniformly strong. Further, his conclusions challenged not only the scientific assumptions of many fellow specialists but also the theological convictions of a much wider circle of fellow citizens.

In 1859, Darwin did not feel quite ready to expose his cherished theory to the harsh light of public scrutiny. In the introduction to the Origin he confessed that although his work on evolution by natural selection was "nearly finished," he would need "two or three more years to complete it." The Origin was, he suggested, merely a stopgap, a schematic "abstract" of a much longer and more fully supported treatise yet to come. He had been moved to preview his labors in this way, he explained, because his health was "far from strong" and, perhaps more importantly, because Alfred Russel Wallace, a younger naturalist working in isolation in southeast Asia, had sent a paper to the Linnean Society of London in which he "arrived at almost exactly the same general conclusions that I have on the origin of species." If Darwin had not gone public with his theory at this point, he would have risked losing credit for the work of many years.

As its reception showed immediately and has continued to show, the Origin benefited from the succinctness imposed by circumstances. Darwin himself may have appreciated this point; at any rate, he never produced the massive treatise, although he repeatedly issued revised editions of the Origin. But he did not abandon his intention to buttress his initial schematic presentation with additional evidence. In the course of the next two decades he published several full-length elaborations of topics summarily discussed in the Origin: The Variation of Animals and Plants under Domestication; The Descent of Man, and Selection in Relation to Sex; and The Expression of the Emotions in Man and Animals. In addition to fleshing out the Origin, these subsequent studies bolstered its arguments and responded to questions raised by critical readers, especially pragmatic questions about the way that descent with modification actually operated.

In The Variation of Animals and Plants under Domestication, which appeared first in 1868 and in a revised edition in 1875, Darwin developed a theme to which he had accorded great rhetorical and evidentiary significance. He had begun the Origin with a description of artificial selection as practiced by farmers, stock breeders, and pet fanciers, thus using a reassuringly homely example—one recognizable by the general public as well as by members of the scientific community—to introduce the most innovative component of his evolutionary theory. In addition, domesticated animals and plants, because they were numerous and available for constant observation, provided a readily available body of evidence.

Reassuring as it was, the analogy between natural and artificial selection was far from perfect. The point of Darwin's analogy was to make the idea of natural selection seem plausible by characterizing it as a grander version of a well-known process while emphasizing its efficiency and shaping power. He noted, for example, that some of the prize birds bred by London pigeon fanciers diverged so strikingly in size, plumage, beak shape, flying technique, vocalizations. bone structure, and many other attributes, that if they had been presented to an ornithologist as wild specimens, they would unquestionably have been considered to represent distinct species, perhaps even distinct genera. Darwin argued that if the relatively brief and constrained selective efforts of human breeders had produced such impressive results, it was likely that the more protracted and thorough-going efforts of nature would work still more efficaciously.

But as Darwin acknowledged, there were some fairly obvious reasons why the two processes might diverge. The superior power of natural selection—"Man can act only on external and visible characters: nature . . . can act on . . . the whole machinery of life. Man selects only for his own good; Nature only for that of the being which she tends" (Origin, chap. 5)—might constitute a difference of kind rather than of degree, as might the much greater stretches of time available for natural selection. Further, although the mechanism of the two processes appeared superficially similar, their outcomes tended to be rather different. Natural selection produced a constantly increasing and diversifying variety of forms; it never reversed or exactly repeated itself.

Anyone familiar with artificial selection would have realized that, although new breeds were constantly being developed and although neither improved wheat nor improved cattle showed any tendency to revert to the condition of their aboriginal wild ancestors, the strains produced by human selection were neither as prolific nor as durable as those produced by nature. Indeed, the animals and plants celebrated as the noblest achievements of the breeder's art were especially liable to delicacy and infertility. Highly bred strains, long isolated from others of their species to preserve their genealogical purity, far from serving as a springboard for further variation, often had to be revivified with infusions of less-rarefied blood. Yet any relaxation of reproductive boundaries threatened subsidence into the common run of conspecifics.

Darwin firmly connected Variation to the Origin by devoting its introduction to an overview of his theory of evolution by natural selection. In particular, the two volumes of Variation, cumbersomely organized and packed with zoological and botanical detail, addressed some of the difficulties inherent in the attractive but paradoxical analogy between natural selection and artificial selection. For selection of any sort to operate, diversity already had to exist. With wild populations living under natural conditions, however, diversity was difficult to discern. It was widely believed that a heightened propensity to vary (at least in ways obvious to human observers) was one of the few general characteristics that differentiated domestic animals as a group from their wild relatives. This point was conventionally illustrated with reference to coat color and design. American bison, for example, were, on the whole, brown, and all Burchell's zebras shared similar black and white stripes. A single herd of either Bos tauras or Equus caballus (domestic cattle or horses), on the other hand, could display colors ranging from white through yellow, red, and brown to black, as well as a variety of spotted and blotched patterns.

In order to demonstrate that such populations spontaneously produced sufficient variation to support artificial selection, Darwin devoted most of the first volume of Variation to a species-by-species survey of domesticated plants and animals. He began with the dog, the breeds of which differed so greatly in size, shape, disposition, talents, and every other characteristic that Darwin attributed its exemplary plasticity to its derivation from several different species of wild canines. Domestic cats, on the other hand, differed relatively little from one another, at least, their variation tended to be individual, rather than consolidated into breeds. Darwin attributed this to the minimal influence exerted by cat owners over the mating behavior of their animals, so that, alone among fully domesticated animals, cats could not be said to have undergone a genuine process of artificial selection.

Farmyard ungulates, however, had all proved more susceptible to human manipulation, whether through the gradual enhancement of inherent tendencies, such as the relatively early maturation that distinguished shorthorn cattle, or through the preservation of spontaneously arising monstrosities, such as the short, broad foreheads and protruding lower jaws of the niata cattle of South America, the bulldogs of the bovine world. Among animals, fancy pigeons, with their short generations, devoted breeders, and lack of any pragmatic constraints on their extravagant deformations, provided Darwin with his most abundant material. He allotted less space to his survey of domesticated plants, although, with the exception of trees, they tended to he much shorter lived and more variable even than pigeons. For example, as Darwin pointed out, a single long-cultivated species— Brassica oleracea, the ordinary cabbage—had given rise to strains as distinctive as Brussels sprouts, cauliflower, broccoli, and kohl-rabi.

Darwin crammed in so much information of this sort that, in order to confine Variation to two volumes of manageable size, less crucial evidence was relegated to a smaller typeface. And so compendious was his survey of domesticates that he felt constrained to deny that it was intended to he an exhaustive catalog. After all, many such catalogs, devoted merely to the accumulation of species- or breed-specific data, existed already; Darwin cited them generously in his footnotes. The material included in Variation had been chosen to fulfill a more focused argumentative purpose. Darwin's theory of descent with modification required something further than the simple demonstration that abundant variation existed among domesticated animals and plants. The accumulated experience of naturalists and breeders offered no clear explanation of the causes of variation; indeed, no consensus existed on this issue. Variation under domestication was frequently attributed to accidental external influences, especially climate and food. But environmentally induced variation was not of much use to Darwin. Instead, he sought evidence not only that the tendency to vary was inherent in domesticated animals and plants but also that specific variations were inherited.

As a result, Darwin's wealth of detail in Variation disproportionately featured strong—as well as puzzling, problematic, or even questionable—versions of inheritance, in addition to the unsurprising, if still not completely understood, likelihood that children would resemble their parents. For example, he devoted an entire chapter to what he termed "atavism" or "reversion"—that is, the tendency for offspring to manifest traits apparently derived from their grandparents, collateral relations, or even remote ancestors, rather than from their mothers or their fathers. The existence of this tendency in the lineages of individuals, he argued, incontrovertibly demonstrated the fact of heritability; and in an extended or exaggerated version it also demonstrated evolutionary relations between species. Thus, many breeds of domesticated chickens revealed their ultimate ancestry by producing occasional sports with the red and orange plumage of the original Callus bankiva, or jungle fowl.

Like many other naturalists of his time, Darwin was receptive to the idea of telegony, also known as "the influence of the previous sire." He retailed the famous story of Lord Morton's mare, a chestnut of seven-eighths Arabian blood, whose first foal had been sired by a quagga (a now-extinct relative of the zebra) her owner was attempting to domesticate. It was not surprising that the young hybrid faintly echoed his father's stripes, but the fact that her next two foals, both sired by a black Arabian horse, also seemed to resemble the quagga in this regard, was more remarkable. Darwin pointed out that atavism offered one possible explanation of this phenomenon—infant horses and donkeys often showed evanescent striping, which might indicate the pattern of their ancient shared progenitor—but he was also drawn to the notion that the first male to impregnate a female left some permanent, heritable trace of himself behind.

He offered analogous examples from the vegetable kingdom, where the pollen of related varieties of apples, corn, or orchids, could not only produce hybrid offspring but occasionally also physically alter the reproductive tract of the female. Plants also, and more regularly, demonstrated a kind of variability that could arise independently of sexual reproduction, such as "bud variation," whereby what Darwin called a "monstrosity' might appear on a single branch or flower and then be transmitted, sexually or asexually, to future generations.

As he documented the profusion of variation among domesticated animals and plants, and the tendency of organisms to transmit these variations down the generations, Darwin did more than demonstrate that there was ample grist for the mill of natural selection. He also addressed the most serious weakness in the argument of the Origin. Despite the incompleteness of the fossil record, plenty of evidence suggested that evolution had taken place; indeed the idea of evolution had been current in one form or another for a century before 1859. Darwin's explanation of the way that natural selection should operate was also widely persuasive. The competitive metaphors with which he characterized it, especially the "struggle for life" prominently featured in the Origin's subtitle, fit well with Victorian understandings about how things worked in the human arenas of industry, commerce, and geopolitics. There was, however, a problem that troubled those inclined to sympathize with Darwin's reasoning as well as those inclined to reject it. The efficacy of natural selection, like that of artificial selection, depended on the inheritance of particular traits. But before the modern understanding of genetics became available, no satisfactory mechanism had been adduced to explain this phenomenon.

No consensus yet existed about the way that sexual reproduction worked, so there was also disagreement about which characteristics were inherited and which were the result of environment, and what could he contributed by the male as opposed to the female parent, let alone why offspring sometimes resembled a grandparent or some more distant relative rather than their parents. The special difficulty of accounting for the sudden emergence of monstrosities, or even less dramatically novel traits, led Darwin, in later editions of the Origin as well as in Variation, to become increasingly receptive to the notion that characteristics acquired by one generation might he inherited by the next.

In the penultimate chapter of Variation, Darwin attempted to strengthen the weak link in his chain of argument by proposing a mechanism for inheritance. He called his theory "pangenesis," and he claimed that it explained not only ordinary inheritance—the influence of parents on their children—but also reversion, telegony, the regeneration of amputated limbs in some kinds of animals, the inheritance of acquired characteristics, and the relationship between sexual and asexual modes of reproduction and inheritance. The operation of pangenesis depended on the posited existence of unobservable units that Darwin called "gemmules," tiny granules that were thrown off by individual cells and then circulated through the body.

They had, however, an affinity for each other, which led to their aggregation in the reproductive organs or in parthenogenetic buds. They could remain latent for years, until an organism reached a certain stage of development, or for generations, until they encountered other gemmules to which they bore some special relationship. In this way a long-dormant greatgrandparental gemmule might suddenly manifest itself in a child. Since gemmules could he altered by environmental influences, they could convert acquired characteristics into the stuff of heredity. And since they were vulnerable to error, they could occasionally make mistakes, causing organs, such as limbs or tails or even heads, to develop in inappropriate numbers or in the wrong places.

It has doubtless been fortunate for Darwin's reputation that his theory of pangenesis is not as well remembered as his theory of evolution by natural selection. As vague in detail as it was ambitious and comprehensive in scope, it was unpersuasive at the time and has since been proven completely wrong. But like Variation as a whole, which similarly illustrated the limitations of its author as well as his strengths, pangenesis does not therefore lack interest or significance. Despite recent excellent and well-appreciated studies of his entire life and extended oeuvre (Janet Browne, Charles Darwin: Voyaging [New York: Knopf, 1995] and Adrian Desmond and James Moore, Darwin [London: Michael Joseph, 1991], Darwin is known primarily as the author of the Origin, which is unrepresentative in its economy of structure, argument, and evidence, as well as on account of its historical notoriety. Its enforced streamlining has helped to preserve the Origin's accessibility, but its relative paucity of examples was particularly uncharacteristic of Darwin. Variation, with its accumulation of evidence about everything from the webbing between dogs' toes to the weight of gooseberries, was much more typical; in addition, it placed Darwin firmly—indeed, irretrievably—within his time, rather than in an achronological limbo reserved for intellectual heroes. As a graduate student from the People's Republic of China told me several years ago, after having participated in a seminar that read excerpts from Variation and The Expression of the Emotions, if the leaders of his government knew that Darwin had written such books, he would not be officially admired.

In science as in politics the victors tend to write the history books. As a result, the record of the past is edited, intentionally or unintentionally, so that it focuses mainly on the precursors of contemporary orthodoxy. Such a focus may accurately represent the genealogy of modem ideas, but it almost inevitably misrepresents the historical experience of their progenitors. Viewed without the benefit of hindsight, the marketplace of Victorian ideas seemed much more competitive than it does to us.

Even the powerful, persuasive, and ultimately triumphant theory of evolution by natural selection required not only defense, but repeated buttressing and revision. Variation showed Darwin hard at work on this rearguard action, using the materials he had at hand—for the most part, homely details about the domesticated animals and plants with which his audience was most familiar. His information was gleaned from the observations of fanciers, breeders, and amateur naturalists, as well as from the treatises of those on the cutting edge of zoology and botany. As hindsight narrows the historical spotlight, it imposes its own sense of hierarchy on the preoccupations of the past. But Darwin was interested in all of these topics, valued all of these sources, and belonged, to a greater or lesser extent, to all of these communities.

The author of Variation was a Victorian country gentleman, a lover of dogs and horses, a breeder of pigeons and peas. He was also, and equally, the author of On the Origin of Species.

PREFACE TO THE SECOND EDITION

During the seven years which have elapsed since the publication in 1868 of the first edition of this Work, I have continued to attend to the same subjects, as far as lay in my power; and I have thus accumulated a large body of additional facts, chiefly through the kindness of many correspondents. Of these facts I have been able here to use only those which seemed to me the more important. I have omitted some statements, and corrected some errors, the discovery of which I owe to my reviewers. Many additional references have been given. The eleventh chapter, and that on Pangenesis, are those which have been most altered, parts having been remodelled; but I will give a list of the more important alterations for the sake of those who may possess the first edition of this book.

TABLE OF PRINCIPAL ADDITIONS AND CORRECTIONS IN SECOND EDITION

First Second Edition Edition Vol. I Vol. I

Page Chapter 34 I

 

38 I 51 II

 

102 III 162 V

217 VI Dr. Burt Wilder's observations on the brains of different breeds of the Dog.

Degeneracy of Dogs imported into Guinea. Difference in the number of lumbar vertebræ in the races or species of the Horse.

 

Hairy appendages to the throats of Goats. Sexual differences in colour in the domestic Pigeon.

 

Movements like those of the Tumbler-pigeon, caused by injury to the brain.

 

290 VIII Additional facts with respect to the Blackshouldered Peacock.

296 VIII Ancient selection of Gold-fish in China.
314 IX Major Hallett's 'Pedigree Wheat.'

326 IX The common radish descended from Raphanus raphanistrum.

 

374 XI Several additional cases of bud-variation given.

396 XI An abstract of all the cases recently published of graft-hybrids in the potato, together with a general summary on graft-hybridisation.

399 XI An erroneous statement with respect to the pollen of the date-palm affecting the fruit of the Chamærops omitted.

400 XI New cases of the direct action of pollen on the mother-plant.

404 XI Additional and remarkable instances of the actions of the male parent on the future progeny of the female.

Vol.II

14 XII An erroneous statement corrected, with respect to the regrowth of supernumerary digits after amputation.

23 XII Additional facts with respect to the inherited effects of circumcision.

 

23 XII Dr. Brown-Séquard on the inherited effects of operations on the Guinea-pig.

24 XII Other cases of inherited mutilations.
Vol. II
43 XIII An additional case of reversion due to a cross.
72 XIV Inheritance as limited by sex.
105 XVI Two varieties of maize which cannot be crossed.

120 XVII Some additional facts on the advantages of crossbreeding in animals.

 

123 XVII Discussion on the effects of the close interbreeding in the case of man.

135 Additional cases of plants sterile with pollen from to XVII the same plant.
141

149 XVIII Mr. Sclater on the infertility of animals under confinement.

 

152 XVIII The Aperea a distinct species from the Guineapig.

 

230 XXI Prof. Jäger on hawks killing light-coloured pigeons.

 

273 XXIII Prof. Wisemann on the effects of isolation in the development of species.

 

281 XXIII The direct action of the conditions of life in causing variation.

 

317 XXIV Mr. Romanes on rudimentary parts.

324 Some additional cases of correlated variability. to XXV
328

339 XXVI On Geoffrey St. Hilaire's law of "soi pour soi."

357 The chapter on Pangenesis has been largely to XXVII altered and re-modelled; but the essential
404 principles remain the same.

Introduction

The object of this work is not to describe all the many races of animals which have been domesticated by man, and of the plants which have been cultivated by him; even if I possessed the requisite knowledge, so gigantic an undertaking would be here superfluous. It is my intention to give under the head of each species only such facts as I have been able to collect or observe, showing the amount and nature of the changes which animals and plants have undergone whilst under man's dominion, or which bear on the general principles of variation. In one case alone, namely in that of the domestic pigeon, I will describe fully all the chief races, their history, the amount and nature of their differences, and the probable steps by which they have been formed. I have selected this case, because, as we shall hereafter see, the materials are better than in any other; and one case fully described will in fact illustrate all others. But I shall also describe domesticated rabbits, fowls, and ducks, with considerable fulness.

The subjects discussed in this volume are so connected that it is not a little difficult to decide how they can be best arranged. I have determined in the first part to give, under the heads of the various animals and plants, a large body of facts, some of which may at first appear but little related to our subject, and to devote the latter part to general discussions. Whenever I have found it necessary to give numerous details, in support of any proposition or conclusion, small type has been used. The reader will, I think, find this plan a convenience, for, if he does not doubt the conclusion or care about the details, he can easily pass them over; yet I may be permitted to say that some of the discussions thus printed deserve attention, at least from the professed naturalist.

It may be useful to those who have read nothing about Natural Selection, if I here give a brief sketch of the whole subject and of its bearing on the origin of species.1 This is the more desirable, as it is impossible in the present work to avoid many allusions to questions which will be fully discussed in future volumes.

From a remote period, in all parts of the world, man has subjected many animals and plants to domestication or culture. Man has no power of altering the absolute conditions of life; he cannot change the climate of any country; he adds no new element to the soil; but he can remove an animal or plant from one climate or soil to another, and give it food on which it did not subsist in its natural state. It is an error to speak of man "tampering with nature" and causing variability. If a man drops a piece of iron into sulphuric acid, it cannot be said strictly that he makes the sulphate of iron, he only allows their elective affinities to come into play. If organic beings had not possessed an inherent tendency to vary, man could have done nothing.2 He unintentionally exposes his animals and plants to various conditions of life, and variability supervenes, which he cannot even prevent or check. Consider the simple case of a plant which has been cultivated during a long time in its native country, and which consequently has not been subjected to any change of climate. It has been protected to a certain extent from the competing roots of plants of other kinds; it has generally been grown in manured soil; but probably not richer than that of many an alluvial flat; and lastly, it has been exposed to changes in its conditions, being grown sometimes in one district and sometimes in another, in different soils. Under such circumstances, scarcely a plant can be named, though cultivated in the rudest manner, which has not given birth to several varieties. It can hardly be maintained that during the many changes which this earth has undergone, and during the natural migrations of plants from one land or island to another, tenanted by different species, that such plants will not often have been subjected to changes in their conditions analogous to those which almost inevitably cause cultivated plants to vary. No doubt man selects varying individuals, sows their seeds, and again selects their varying offspring. But the initial variation on which man works, and without which he can do nothing, is caused by slight changes in the conditions of life, which must often have occurred under nature. Man, therefore, may be said to have been trying an experiment on a gigantic scale; and it is an experiment which nature during the long lapse of time has incessantly tried. Hence it follows that the principles of domestication are important for us. The main result is that organic beings thus treated have varied largely, and the variations have been inherited. This has apparently been one chief cause of the belief long held by some few naturalists that species in a state of nature undergo change.

I shall in this volume treat, as fully as my materials permit, the whole subject of variation under domestication. We may thus hope to obtain some light, little though it be, on the causes of variability,—on the laws which govern it, such as the direct action of climate and food, the effects of use and disuse, and of correlation of growth,—and on the amount of change to which domesticated organisms are liable. We shall learn something of the laws of inheritance, of the effects of crossing different breeds, and on that sterility which often supervenes when organic beings are removed from their natural conditions of life, and likewise when they are too closely interbred. During this investigation we shall see that the principle of Selection is highly important. Although man does not cause variability and cannot even prevent it, he can select, preserve, and accumulate the variations given to him by the hand of nature almost in any way which he chooses; and thus he can certainly produce a great result. Selection may be followed either methodically and intentionally, or unconsciously and unintentionally. Man may select and preserve each successive variation, with the distinct intention of improving and altering a breed, in accordance with a preconceived idea; and by thus adding up variations, often so slight as to be imperceptible by an uneducated eye, he has effected wonderful changes and improvements. It can, also, be clearly shown that man, without any intention or thought of improving the breed, by preserving in each successive generation the individuals which he prizes most, and by destroying the worthless individuals, slowly, though surely, induces great changes. As the will of man thus comes into play, we can understand how it is that domesticated breeds show adaptation to his wants and pleasures. We can further understand how it is that domestic races of animals and cultivated races of plants often exhibit an abnormal character, as compared with natural species; for they have been modified not for their own benefit, but for that of man.

In another work I shall discuss, if time and health permit, the variability of organic beings in a state of nature; namely, the individual differences presented by animals and plants, and those slightly greater and generally inherited differences which are ranked by naturalists as varieties or geographical races. We shall see how difficult, or rather how impossible it often is, to distinguish between races and sub-species, as the less wellmarked forms have sometimes been denominated; and again between sub-species and true species. I shall further attempt to show that it is the common and widely ranging, or, as they may be called, the dominant species, which most frequently vary; and that it is the large and flourishing genera which include the greatest number of varying species. Varieties, as we shall see, may justly be called incipient species.

But it may be urged, granting that organic beings in a state of nature present some varieties,—that their organisation is in some slight degree plastic; granting that many animals and plants have varied greatly under domestication, and that man by his power of selection has gone on accumulating such variations until he has made strongly marked and firmly inherited races; granting all this, how, it may be asked, have species arisen in a state of nature? The differences between natural varieties are slight; whereas the differences are considerable between the species of the same genus, and great between the species of distinct genera. How do these lesser differences become augmented into the greater difference? How do varieties, or as I have called them incipient species, become converted into true and well-defined species? How has each new species been adapted to the surrounding physical conditions, and to the other forms of life on which it in any way depends? We see on every side of us innumerable adaptations and contrivances, which have justly excited the highest admiration of every observer. There is, for instance, a fly (Cecidomyia)3 which deposits its eggs within the stamens of a Scrophularia, and secretes a poison which produces a gall, on which the larva feeds; but there is another insect (Misocampus) which deposits its eggs within the body of the larva within the gall, and is thus nourished by its living prey; so that here a hymenopterous insect depends on a dipterous insect, and this depends on its power of producing a monstrous growth in a particular organ of a particular plant. So it is, in a more or less plainly marked manner, in thousands and tens of thousands of cases, with the lowest as well as with the highest productions of nature.

This problem of the conversion of varieties into species,—that is, the augmentation of the slight differences characteristic of varieties into the greater differences characteristic of species and genera, including the admirable adaptations of each being to its complex organic and inorganic conditions of life,—has been briefly treated in my 'Origin of Species.' It was there shown that all organic beings, without exception, tend to increase at so high a ratio, that no district, no station, not even the whole surface of the land or the whole ocean, would hold the progeny of a single pair after a certain number of generations. The inevitable result is an ever-recurrent Struggle for Existence. It has truly been said that all nature is at war; the strongest ultimately prevail, the weakest fail; and we well know that myriads of forms have disappeared from the face of the earth. If then organic beings in a state of nature vary even in a slight degree, owing to changes in the surrounding conditions, of which we have abundant geological evidence, or from any other cause; if, in the long course of ages, inheritable variations ever arise in any way advantageous to any being under its excessively complex and changing relations of life; and it would be a strange fact if beneficial variations did never arise, seeing how many have arisen which man has taken advantage of for his own profit or pleasure; if then these contingencies ever occur, and I do not see how the probability of their occurrence can be doubted, then the severe and often-recurrent struggle for existence will determine that those variations, however slight, which are favourable shall be preserved or selected, and those which are unfavourable shall be destroyed.

This preservation, during the battle for life, of varieties which possess any advantage in structure, constitution, or instinct, I have called Natural Selection; and Mr. Herbert Spencer has well expressed the same idea by the Survival of the Fittest. The term "natural selection" is in some respects a bad one, as it seems to imply conscious choice; but this will be disregarded after a little familiarity. No one objects to chemists speaking of "elective affinity;" and certainly an acid has no more choice in combining with a base, than the conditions of life have in determining whether or not a new form be selected or preserved. The term is so far a good one as it brings into connection the production of domestic races by man's power of selection, and the natural preservation of varieties and species in a state of nature. For brevity sake I sometimes speak of natural selection as an intelligent power;—in the same way as astronomers speak of the attraction of gravity as ruling the movements of the planets, or as agriculturists speak of man making domestic races by his power of selection. In the one case, as in the other, selection does nothing without variability, and this depends in some manner on the action of the surrounding circumstances on the organism. I have, also, often personified the word Nature; for I have found it difficult to avoid this ambiguity; but I mean by nature only the aggregate action and product of many natural laws,—and by laws only the ascertained sequence of events.

It has been shown from many facts that the largest amount of life can be supported on each area, by great diversification or divergence in the structure and constitution of its inhabitants. We have, also, seen that the continued production of new forms through natural selection, which implies that each new variety has some advantage over others, inevitably leads to the extermination of the older and less improved forms. These latter are almost necessarily intermediate in structure, as well as in descent, between the lastproduced forms and their original parent-species. Now, if we suppose a species to produce two or more varieties, and these in the course of time to produce other varieties, the principal of good being derived from diversification of structure will generally lead to the preservation of the most divergent varieties; thus the lesser differences characteristic of varieties come to be augmented into the greater differences characteristic of species, and, by the extermination of the older intermediate forms, new species end by being distinctly defined objects. Thus, also, we shall see how it is that organic beings can be classed by what is called a natural method in distinct groups—species under genera, and genera under families.

As all the inhabitants of each country may be said, owing to their high rate of reproduction, to be striving to increase in numbers; as each form comes into competition with many other forms in the struggle for life,—for destroy any one and its place will be seized by others; as every part of the organisation occasionally varies in some slight degree, and as natural selection acts exclusively by the preservation of variations which are advantageous under the excessively complex conditions to which each being is exposed, no limit exists to the number, singularity, and perfection of the contrivances and co-adaptations which may thus be produced. An animal or a plant may thus slowly become related in its structure and habits in the most intricate manner to many other animals and plants, and to the physical conditions of its home. Variations in the organisation will in some cases be aided by habit, or by the use and disuse of parts, and they will be governed by the direct action of the surrounding physical conditions and by correlation of growth.

On the principles here briefly sketched out, there is no innate or necessary tendency in each being to its own advancement in the scale of organisation. We are almost compelled to look at the specialisation or differentiation of parts or organs for different functions as the best or even sole standard of advancement; for by such division of labour each function of body and mind is better performed.

And as natural selection acts exclusively through the preservation of profitable modifications of structure, and as the conditions of life in each area generally become more and more complex from the increasing number of different forms which inhabit it and from most of these forms acquiring a more and more perfect structure, we may confidently believe, that, on the whole, organisation advances. Nevertheless a very simple form fitted for very simple conditions of life might remain for indefinite ages unaltered or unimproved; for what would it profit an infusorial animalcule, for instance, or an intestinal worm, to become highly organised?

Members of a high group might even become, and this apparently has often occurred, fitted for simpler conditions of life; and in this case natural selection would tend to simplify or degrade the organisation, for complicated mechanism for simple actions would be useless or even disadvantageous.

The arguments opposed to the theory of Natural Selection, have been discussed in my 'Origin of Species,' as far as the size of that work permitted, under the following heads: the difficulty in understanding how very simple organs have been converted by small and graduated steps into highly perfect and complex organs; the marvellous facts of Instinct; the whole question of Hybridity; and, lastly, the absence in our known geological formations of innumerable links connecting all allied species. Although some of these difficulties are of great weight, we shall see that many of them are explicable on the theory of natural selection, and are otherwise inexplicable.

In scientific investigations it is permitted to invent any hypothesis, and if it explains various large and independent classes of facts it rises to the rank of a well-grounded theory. The undulations of the ether and even its existence are hypothetical, yet every one now admits the undulatory theory of light. The principle of natural selection may be looked at as a mere hypothesis, but rendered in some degree probable by what we positively know of the variability of organic beings in a state of nature,—by what we positively know of the struggle for existence, and the consequent almost inevitable preservation of favourable variations,—and from the analogical formation of domestic races. Now this hypothesis may be tested,—and this seems to me the only fair and legitimate manner of considering the whole question,—by trying whether it explains several large and independent classes of facts; such as the geological succession of organic beings, their distribution in past and present times, and their mutual affinities and homologies. If the principle of natural selection does explain these and other large bodies of facts, it ought to be received. On the ordinary view of each species having been independently created, we gain no scientific explanation of any one of these facts. We can only say that it has so pleased the Creator to command that the past and present inhabitants of the world should appear in a certain order and in certain areas; that He has impressed on them the most extraordinary resemblances, and has classed them in groups subordinate to groups. But by such statements we gain no new knowledge; we do not connect together facts and laws; we explain nothing.

It was the consideration of such large groups of facts as these which first led me to take up the present subject. When I visited during the voyage of H.M.S. Beagle, the Galapagos Archipelago, situated in the Pacific Ocean about 500 miles from South America, I found myself surrounded by peculiar species of birds, reptiles, and plants, existing nowhere else in the world.

Yet they nearly all bore an American stamp. In the song of the mocking-thrush, in the harsh cry of the carrion-hawk, in the great candlestick-like opuntias, I clearly perceived the neighbourhood of America, though the islands were separated by so many miles of ocean from the mainland, and differed much in their geological constitution and climate. Still more surprising was the fact that most of the inhabitants of each separate island in this small archipelago were specifically different, though most closely related to each other.

The archipelago, with its innumerable craters and bare streams of lava, appeared to be of recent origin; and thus I fancied myself brought near to the very act of creation. I often asked myself how these many peculiar animals and plants had been produced: the simplest answer seemed to be that the inhabitants of the several islands had descended from each other, undergoing modification in the course of their descent; and that all the inhabitants of the archipelago were descended from those of the nearest land, namely America, whence colonists would naturally have been derived. But it long remained to me an inexplicable problem how the necessary degree of modification could have been effected, and it would have thus remained for ever, had I not studied domestic productions, and thus acquired a just idea of the power of Selection. As soon as I had fully realised this idea, I saw, on reading Malthus on Population, that Natural Selection was the inevitable result of the rapid increase of all organic beings; for I was prepared to appreciate the struggle for existence by having long studied the habits of animals.

Before visiting the Galapagos I had collected many animals whilst travelling from north to south on both sides of America, and everywhere, under conditions of life as different as it is possible to conceive, American forms were met with—species replacing species of the same peculiar genera. Thus it was when the Cordilleras were ascended, or the thick tropical forests penetrated, or the fresh waters of America searched. Subsequently I visited other countries, which in all their conditions of life were incomparably more like parts of South America, than the different parts of that continent are to each other; yet in these countries, as in Australia or Southern Africa, the traveller cannot fail to be struck with the entire difference of their productions. Again the reflection was forced on me that community of descent from the early inhabitants of South America would alone explain the wide prevalence of American types throughout that immense area.

To exhume with one's own hands the bones of extinct and gigantic quadrupeds brings the whole question of the succession of species vividly before one's mind; and I found in South America great pieces of tesselated armour exactly like, but on a magnificent scale, that covering the pigmy armadillo; I had found great teeth like those of the living sloth, and bones like those of the cavy. An analogous succession of allied forms had been previously observed in Australia.

Here then we see the prevalence, as if by descent, in time as in space, of the same types in the same areas; and in neither the case does the similarity of the conditions by any means seem sufficient to account for the similarity of the forms of life. It is notorious that the fossil remains of closely consecutive formations are closely allied in structure, and we can at once understand the fact if they are closely allied by descent. The succession of the many distinct species of the same genus throughout the long series of geological formations seems to have been unbroken or continuous.

New species come in gradually one by one. Ancient and extinct forms of life are often intermediate in character, like the words of a dead language with respect to its several offshoots or living tongues. All these facts seemed to me to point to descent with modification as the means of production of new species.

The innumerable past and present inhabitants of the world are connected together by the most singular and complex affinities, and can be classed in groups under groups, in the same manner as varieties can be classed under species and sub-varieties under varieties, but with much higher grades of difference. These complex affinities and the rules for classification, receive a rational explanation on the theory of descent, combined with the principle of natural selection, which entails divergence of character and the extinction of intermediate forms.

How inexplicable is the similar pattern of the hand of a man, the foot of a dog, the wing of a bat, the flipper of a seal, on the doctrine of independent acts of creation! how simply explained on the principle of the natural selection of successive slight variations in the diverging descendants from a single progenitor! So it is with certain parts or organs in the same individual animal or plant, for instance, the jaws and legs of a crab, or the petals, stamens, and pistils of a flower.

During the many changes to which in the course of time organic beings have been subjected, certain organs or parts have occasionally become at first of little use and ultimately superfluous; and the retention of such parts in a rudimentary and useless condition is intelligible on the theory of descent. It can be shown that modifications of structure are generally inherited by the offspring at the same age at which each successive variation appeared in the parents; it can further be shown that variations do not commonly supervene at a very early period of embryonic growth, and on these two principles we can understand that most wonderful fact in the whole circuit of natural history, namely, the close similarity of the embryos within the same great class—for instance, those of mammals, birds, reptiles, and fish.

It is the consideration and explanation of such facts as these which has convinced me that the theory of descent with modification by means of natural selection is in the main true. These facts have as yet received no explanation on the theory of independent Creation; they cannot be grouped together under one point of view, but each has to be considered as an ultimate fact. As the first origin of life on this earth, as well as the continued life of each individual, is at present quite beyond the scope of science, I do not wish to lay much stress on the greater simplicity of the view of a few forms or of only one form having been originally created, instead of innumerable miraculous creations having been necessary at innumerable periods; though this more simple view accords well with Maupertuis's philosophical axiom of "least action."

In considering how far the theory of natural selection may be extended, —that is, in determining from how many progenitors the inhabitants of the world have descended,— we may conclude that at least all the members of the same class have descended from a single ancestor. A number of organic beings are included in the same class, because they present, independently of their habits of life, the same fundamental type of structure, and because they graduate into each other.

Moreover, members of the same class can in most cases be shown to be closely alike at an early embryonic age. These facts can be explained on the belief of their descent from a common form; therefore it may be safely admitted that all the members of the same class are descended from one progenitor. But as the members of quite distinct classes have something in common in structure and much in common in constitution, analogy would lead us one step further, and to infer as probable that all living creatures are descended from a single prototype.

I hope that the reader will pause before coming to any final and hostile conclusion on the theory of natural selection. The reader may consult my 'Origin of Species' for a general sketch of the whole subject; but in that work he has to take many statements on trust. In considering the theory of natural selection, he will assuredly meet with weighty difficulties, but these difficulties relate chiefly to subjects—such as the degree of perfection of the geological record, the means of distribution, the possibility of transitions in organs, etc.—on which we are confessedly ignorant; nor do we know how ignorant we are. If we are much more ignorant than is generally supposed, most of these difficulties wholly disappear. Let the reader reflect on the difficulty of looking at whole classes of facts from a new point of view.

Let him observe how slowly, but surely, the noble views of Lyell on the gradual changes now in progress on the earth's surface have been accepted as sufficient to account for all that we see in its past history. The present action of natural selection may seem more or less probable; but I believe in the truth of the theory, because it collects, under one point of view, and gives a rational explanation of, many apparently independent classes of facts.4

REFERENCES

1. To any one who has attentively read my 'Origin of Species' this Introduction will be superfluous. As I stated in that work that I should soon publish the facts on which the conclusions given in it were founded, I here beg permission to remark that the great delay in publishing this first work has been caused by continued ill-health.

2. M. Pouchet has recently ('Plurality of Races,' Eng. Translat., 1864, p. 83, etc.) insisted that variation under domestication throws no light on the natural modification of species. I cannot perceive the force of his arguments, or, to speak more accurately, of his assertions to this effect.

3. Léon Dufour in 'Annales des Science. Nat.' (3rd series, Zoolog.), tom. v. p. 6.
4. In treating the several subjects included in the present and my other works I have continually been led to ask for information from many zoologists, botanists, geologists, breeders of animals, and horticulturists, and I have invariably received from them the most generous assistance. Without such aid I could have effected little. I have repeatedly applied for information and specimens to foreigners, and to British merchants and officers of the Government residing in distant lands, and, with the rarest exceptions, I have received prompt, open-handed, and valuable assistance. I cannot express too strongly my obligations to the many persons who have assisted me, and who, I am convinced, would be equally willing to assist others in any scientific investigation.