RECAPITULATION AND CONCLUSION
I will now recapitulate the course of this work, more fully with respect to the former parts, and briefly [[as to]] the latter. In the first chapter we have seen that most, if not all, organic beings, when taken by man out of their natural condition, and bred during several generations, vary; that is variation is partly due to the direct effect of the new external influences, and partly to the indirect effect on the reproductive system rendering the organization of the offspring in some degree plastic. Of the variations thus produced, man when uncivilised naturally preserves the life, and therefore unintentionally breeds from those individuals most useful to him in his different states: when even semi-civilised, he intentionally separates and breeds from such individuals. Every part of the structure seems occasionally to vary in a very slight degree, and the extent to which all kinds of peculiarities in mind and body, when congenital and when slowly acquired either from external influences, from exercise, or from disuse [[are inherited]], is truly wonderful. When several breeds are once formed, then crossing is the most fertile source of new breeds. Variation must be ruled, of course, by the health of the new race, by the tendency to return to the ancestral forms, and by unknown laws determining the proportional increase and symmetry of the body. The amount of variation, which has been effected under domestication, is quite unknown in the majority of domestic beings.
 Compare however Darwin's later view:--"The possibility of making distinct races by crossing has been greatly exaggerated," Origin, Ed. i. p. 20, vi. p. 23. The author's change of opinion was no doubt partly due to his experience in breeding pigeons.
In the second chapter it was shown that wild organisms undoubtedly vary in some slight degree: and that the kind of variation, though much less in degree, is similar to that of domestic organisms. It is highly probable that every organic being, if subjected during several generations to new and varying conditions, would vary. It is certain that organisms, living in an isolated country which is undergoing geological changes, must in the course of time be so subjected to new conditions; moreover an organism, when by chance transported into a new station, for instance into an island, will often be exposed to new conditions, and be surrounded by a new series of organic beings. If there were no power at work selecting every slight variation, which opened new sources of subsistence to a being thus situated, the effects of crossing, the chance of death and the constant tendency to reversion to the old parent-form, would prevent the production of new races. If there were any selective agency at work, it seems impossible to assign any limit to the complexity and beauty of the adaptive structures, which might thus be produced: for certainly the limit of possible variation of organic beings, either in a wild or domestic state, is not known.
 In the Origin, Ed. i. p. 469, vi. p. 644, Darwin makes a strong statement to this effect.
It was then shown, from the geometrically increasing tendency of each species to multiply (as evidenced from what we know of mankind and of other animals when favoured by circumstances), and from the means of subsistence of each species on an average remaining constant, that during some part of the life of each, or during every few generations, there must be a severe struggle for existence; and that less than a grain in the balance will determine which individuals shall live and which perish. In a country, therefore, undergoing changes, and cut off from the free immigration of species better adapted to the new station and conditions, it cannot be doubted that there is a most powerful means of selection, tending to preserve even the slightest variation, which aided the subsistence or defence of those organic beings, during any part of their whole existence, whose organization had been rendered plastic. Moreover, in animals in which the sexes are distinct, there is a sexual struggle, by which the most vigorous, and consequently the best adapted, will oftener procreate their kind.
 "A grain in the balance will determine which individual shall live and which shall die," Origin, Ed. i. p. 467, vi. p. 642. A similar statement occurs in the 1842 Essay, p. 8, note 3.[Note 59]
A new race thus formed by natural selection would be undistinguishable from a species. For comparing, on the one hand, the several species of a genus, and on the other hand several domestic races from a common stock, we cannot discriminate them by the amount of external difference, but only, first, by domestic races not remaining so constant or being so "true" as species are; and secondly by races always producing fertile offspring when crossed. And it was then shown that a race naturally selected--from the variation being slower--from the selection steadily leading towards the same ends, and from every new slight change in structure being adapted (as is implied by its selection) to the new conditions and being fully exercised, and lastly from the freedom from occasional crosses with other species, would almost necessarily be "truer" than a race selected by ignorant or capricious and short-lived man. With respect to the sterility of species when crossed, it was shown not to be a universal character, and when present to vary in degree: sterility also was shown probably to depend less on external than on constitutional differences. And it was shown that when individual animals and plants are placed under new conditions, they become, without losing their healths, as sterile, in the same manner and to the same degree, as hybrids; and it is therefore conceivable that the cross-bred offspring between two species, having different constitutions, might have its constitution affected in the same peculiar manner as when an individual animal or plant is placed under new conditions. Man in selecting domestic races has little wish and still less power to adapt the whole frame to new conditions; in nature, however, where each species survives by a struggle against other species and external nature, the result must be very different.
 Thus according to the author what is now known as orthogenesis is due to selection.
Races descending from the same stock were then compared with species of the same genus, and they were found to present some striking analogies. The offspring also of races when crossed, that is mongrels, were compared with the cross-bred offspring of species, that is hybrids, and they were found to resemble each other in all their characters, with the one exception of sterility, and even this, when present, often becomes after some generations variable in degree. The chapter was summed up, and it was shown that no ascertained limit to the amount of variation is known; or could be predicted with due time and changes of condition granted. It was then admitted that although the production of new races, undistinguishable from true species, is probable, we must look to the relations in the past and present geographical distribution of the infinitely numerous beings, by which we are surrounded--to their affinities and to their structure--for any direct evidence.
In the third chapter the inheritable variations in the mental phenomena of domestic and of wild organic beings were considered. It was shown that we are not concerned in this work with the first origin of the leading mental qualities; but that tastes, passions, dispositions, consensual movements, and habits all became, either congenitally or during mature life, modified and were inherited. Several of these modified habits were found to correspond in every essential character with true instincts, and they were found to follow the same laws. Instincts and dispositions &c. are fully as important to the preservation and increase of a species as its corporeal structure; and therefore the natural means of selection would act on and modify them equally with corporeal structures. This being granted, as well as the proposition that mental phenomena are variable, and that the modifications are inheritable, the possibility of the several most complicated instincts being slowly acquired was considered, and it was shown from the very imperfect series in the instincts of the animals now existing, that we are not justified in prima facie rejecting a theory of the common descent of allied organisms from the difficulty of imagining the transitional stages in the various now most complicated and wonderful instincts. We were thus led on to consider the same question with respect both to highly complicated organs, and to the aggregate of several such organs, that is individual organic beings; and it was shown, by the same method of taking the existing most imperfect series, that we ought not at once to reject the theory, because we cannot trace the transitional stages in such organs, or conjecture the transitional habits of such individual species.
In the Second Part the direct evidence of allied forms having descended from the same stock was discussed. It was shown that this theory requires a long series of intermediate forms between the species and groups in the same classes--forms not directly intermediate between existing species, but intermediate with a common parent. It was admitted that if even all the preserved fossils and existing species were collected, such a series would be far from being formed; but it was shown that we have not good evidence that the oldest known deposits are contemporaneous with the first appearance of living beings; or that the several subsequent formations are nearly consecutive; or that any one formation preserves a nearly perfect fauna of even the hard marine organisms, which lived in that quarter of the world. Consequently, we have no reason to suppose that more than a small fraction of the organisms which have lived at any one period have ever been preserved; and hence that we ought not to expect to discover the fossilised sub-varieties between any two species. On the other hand, the evidence, though extremely imperfect, drawn from fossil remains, as far as it does go, is in favour of such a series of organisms having existed as that required. This want of evidence of the past existence of almost infinitely numerous intermediate forms, is, I conceive, much the weightiest difficulty on the theory of common descent; but I must think that this is due to ignorance necessarily resulting from the imperfection of all geological records.
 Part II begins with Ch. IV. See the Introduction, where the absence of division into two parts (in the Origin) is discussed.
 In the recapitulation in the last chapter of the Origin, Ed. i. p. 475, vi. p. 651, the author does not insist on this point as the weightiest difficulty, though he does so in Ed. i. p. 299. It is possible that he had come to think less of the difficulty in question: this was certainly the case when he wrote the 6th edition, see p. 438.
In the fifth chapter it was shown that new species gradually appear, and that the old ones gradually disappear, from the earth; and this strictly accords with our theory. The extinction of species seems to be preceded by their rarity; and if this be so, no one ought to feel more surprise at a species being exterminated than at its being rare. Every species which is not increasing in number must have its geometrical tendency to increase checked by some agency seldom accurately perceived by us. Each slight increase in the power of this unseen checking agency would cause a corresponding decrease in the average numbers of that species, and the species would become rarer: we feel not the least surprise at one species of a genus being rare and another abundant; why then should we be surprised at its extinction, when we have good reason to believe that this very rarity is its regular precursor and cause.
 [[The following words:]] The fauna changes singly [[were inserted by the author, apparently to replace a doubtful erasure]].
In the sixth chapter the leading facts in the geographical distribution of organic beings were considered--namely, the dissimilarity in areas widely and effectually separated, of the organic beings being exposed to very similar conditions (as for instance, within the tropical forests of Africa and America, or on the volcanic islands adjoining them). Also the striking similarity and general relations of the inhabitants of the same great continents, conjoined with a lesser degree of dissimilarity in the inhabitants living on opposite sides of the barriers intersecting it--whether or not these opposite sides are exposed to similar conditions. Also the dissimilarity, though in a still lesser degree, in the inhabitants of different islands in the same archipelago, together with their similarity taken as a whole with the inhabitants of the nearest continent, whatever its character may be. Again, the peculiar relations of Alpine floras; the absence of mammifers on the smaller isolated islands; and the comparative fewness of the plants and other organisms on islands with diversified stations; the connection between the possibility of occasional transportal from one country to another, with an affinity, though not identity, of the organic beings inhabiting them. And lastly, the clear and striking relations between the living and the extinct in the same great divisions of the world; which relation, if we look very far backward, seems to die away. These facts, if we bear in mind the geological changes in progress, all simply follow from the proposition of allied organic beings having lineally descended from common parent-stocks. On the theory of independent creations they must remain, though evidently connected together, inexplicable and disconnected.
In the seventh chapter, the relationship or grouping of extinct and recent species; the appearance and disappearance of groups; the ill-defined objects of the natural classification, not depending on the similarity of organs physiologically important, not being influenced by adaptive or analogical characters, though these often govern the whole economy of the individual, but depending on any character which varies least, and especially on the forms through which the embryo passes, and, as was afterwards shown, on the presence of rudimentary and useless organs. The alliance between the nearest species in distinct groups being general and not especial; the close similarity in the rules and objects in classifying domestic races and true species. All these facts were shown to follow on the natural system being a genealogical system.
In the eighth chapter, the unity of structure throughout large groups, in species adapted to the most different lives, and the wonderful metamorphosis (used metaphorically by naturalists) of one part or organ into another, were shown to follow simply on new species being produced by the selection and inheritance of successive small changes of structure. The unity of type is wonderfully manifested by the similarity of structure, during the embryonic period, in the species of entire classes. To explain this it was shown that the different races of our domestic animals differ less, during their young state, than when full grown; and consequently, if species are produced like races, the same fact, on a greater scale, might have been expected to hold good with them. This remarkable law of nature was attempted to be explained through establishing, by sundry facts, that slight variations originally appear during all periods of life, and that when inherited they tend to appear at the corresponding period of life; according to these principles, in several species descended from the same parent-stock, their embryos would almost necessarily much more closely resemble each other than they would in their adult state. The importance of these embryonic resemblances, in making out a natural or genealogical classification, thus becomes at once obvious. The occasional greater simplicity of structure in the mature animal than in the embryo; the gradation in complexity of the species in the great classes; the adaptation of the larvę of animals to independent powers of existence; the immense difference in certain animals in their larval and mature states, were all shown on the above principles to present no difficulty.
In the [[ninth]] chapter, the frequent and almost general presence of organs and parts, called by naturalists abortive or rudimentary, which, though formed with exquisite care, are generally absolutely useless [[was considered]]. [[These structures,]] though sometimes applied to uses not normal,--which cannot be considered as mere representative parts, for they are sometimes capable of performing their proper function,--which are always best developed, and sometimes only developed, during a very early period of life,--and which are of admitted high importance in classification,--were shown to be simply explicable on our theory of common descent.
Why do we wish to reject the theory of common descent?
Thus have many general facts, or laws, been included under one explanation; and the difficulties encountered are those which would naturally result from our acknowledged ignorance. And why should we not admit this theory of descent? Can it be shown that organic beings in a natural state are all absolutely invariable? Can it be said that the limit of variation or the number of varieties capable of being formed under domestication are known? Can any distinct line be drawn between a race and a species? To these three questions we may certainly answer in the negative. As long as species were thought to be divided and defined by an impassable barrier of sterility, whilst we were ignorant of geology, and imagined that the world was of short duration, and the number of its past inhabitants few, we were justified in assuming individual creations, or in saying with Whewell that the beginnings of all things are hidden from man. Why then do we feel so strong an inclination to reject this theory--especially when the actual case of any two species, or even of any two races, is adduced--and one is asked, have these two originally descended from the same parent womb? I believe it is because we are always slow in admitting any great change of which we do not see the intermediate steps. The mind cannot grasp the full meaning of the term of a million or hundred million years, and cannot consequently add up and perceive the full effects of small successive variations accumulated during almost infinitely many generations. The difficulty is the same with that which, with most geologists, it has taken long years to remove, as when Lyell propounded that great valleys were hollowed out [and long lines of inland cliffs had been formed] by the slow action of the waves of the sea. A man may long view a grand precipice without actually believing, though he may not deny it, that thousands of feet in thickness of solid rock once extended over many square miles where the open sea now rolls; without fully believing that the same sea which he sees beating the rock at his feet has been the sole removing power.
 This question forms the subject of what is practically a section of the final chapter of the Origin (Ed. i. p. 480, vi. p. 657).
 Origin, Ed. i. p. 481, vi. p. 659.
Shall we then allow that the three distinct species of rhinoceros which separately inhabit Java and Sumatra and the neighbouring mainland of Malacca were created, male and female, out of the inorganic materials of these countries? Without any adequate cause, as far as our reason serves, shall we say that they were merely, from living near each other, created very like each other, so as to form a section of the genus dissimilar from the African section, some of the species of which section inhabit very similar and some very dissimilar stations? Shall we say that without any apparent cause they were created on the same generic type with the ancient woolly rhinoceros of Siberia and of the other species which formerly inhabited the same main division of the world: that they were created, less and less closely related, but still with interbranching affinities, with all the other living and extinct mammalia? That without any apparent adequate cause their short necks should contain the same number of vertebrę with the giraffe; that their thick legs should be built on the same plan with those of the antelope, of the mouse, of the hand of the monkey, of the wing of the bat, and of the fin of the porpoise. That in each of these species the second bone of their leg should show clear traces of two bones having been soldered and united into one; that the complicated bones of their head should become intelligible on the supposition of their having been formed of three expanded vertebrę; that in the jaws of each when dissected young there should exist small teeth which never come to the surface. That in possessing these useless abortive teeth, and in other characters, these three rhinoceroses in their embryonic state should much more closely resemble other mammalia than they do when mature. And lastly, that in a still earlier period of life, their arteries should run and branch as in a fish, to carry the blood to gills which do not exist. Now these three species of rhinoceros closely resemble each other; more closely than many generally acknowledged races of our domestic animals; these three species if domesticated would almost certainly vary, and races adapted to different ends might be selected out of such variations. In this state they would probably breed together, and their offspring would possibly be quite, and probably in some degree, fertile; and in either case, by continued crossing, one of these specific forms might be absorbed and lost in another. I repeat, shall we then say that a pair, or a gravid female, of each of these three species of rhinoceros, were separately created with deceptive appearances of true relationship, with the stamp of inutility on some parts, and of conversion in other parts, out of the inorganic elements of Java, Sumatra and Malacca? or have they descended, like our domestic races, from the same parent-stock? For my own part I could no more admit the former proposition than I could admit that the planets move in their courses, and that a stone falls to the ground, not through the intervention of the secondary and appointed law of gravity, but from the direct volition of the Creator.
 The discussion on the three species of Rhinoceros which also occurs in the Essay of 1842, p. 48, was omitted in Ch. XIV of the Origin, Ed. i.
Before concluding it will be well to show, although this has incidentally appeared, how far the theory of common descent can legitimately be extended. If we once admit that two true species of the same genus can have descended from the same parent, it will not be possible to deny that two species of two genera may also have descended from a common stock. For in some families the genera approach almost as closely as species of the same genus; and in some orders, for instance in the monocotyledonous plants, the families run closely into each other. We do not hesitate to assign a common origin to dogs or cabbages, because they are divided into groups analogous to the groups in nature. Many naturalists indeed admit that all groups are artificial; and that they depend entirely on the extinction of intermediate species. Some naturalists, however, affirm that though driven from considering sterility as the characteristic of species, that an entire incapacity to propagate together is the best evidence of the existence of natural genera. Even if we put on one side the undoubted fact that some species of the same genus will not breed together, we cannot possibly admit the above rule, seeing that the grouse and pheasant (considered by some good ornithologists as forming two families), the bull-finch and canary-bird have bred together.
 This corresponds to a paragraph in the Origin, Ed. i. p. 483, vi. p. 662, where it is assumed that animals have descended "from at most only four or five progenitors, and plants from an equal or lesser number." In the Origin, however, the author goes on, Ed. i. p. 484, vi. p. 663: "Analogy would lead me one step further, namely, to the belief that all animals and plants have descended from some one prototype."
No doubt the more remote two species are from each other, the weaker the arguments become in favour of their common descent. In species of two distinct families the analogy, from the variation of domestic organisms and from the manner of their intermarrying, fails; and the arguments from their geographical distribution quite or almost quite fails. But if we once admit the general principles of this work, as far as a clear unity of type can be made out in groups of species, adapted to play diversified parts in the economy of nature, whether shown in the structure of the embryonic or mature being, and especially if shown by a community of abortive parts, we are legitimately led to admit their community of descent. Naturalists dispute how widely this unity of type extends: most, however, admit that the vertebrata are built on one type; the articulata on another; the mollusca on a third; and the radiata on probably more than one. Plants also appear to fall under three or four great types. On this theory, therefore, all the organisms yet discovered are descendants of probably less than ten parent-forms.
My reasons have now been assigned for believing that specific forms are not immutable creations. The terms used by naturalists of affinity, unity of type, adaptive characters, the metamorphosis and abortion of organs, cease to be metaphorical expressions and become intelligible facts. We no longer look at an organic being as a savage does at a ship or other great work of art, as at a thing wholly beyond his comprehension, but as a production that has a history which we may search into. How interesting do all instincts become when we speculate on their origin as hereditary habits, or as slight congenital modifications of former instincts perpetuated by the individuals so characterised having been preserved. When we look at every complex instinct and mechanism as the summing up of a long history of contrivances, each most useful to its possessor, nearly in the same way as when we look at a great mechanical invention as the summing up of the labour, the experience, the reason, and even the blunders of numerous workmen. How interesting does the geographical distribution of all organic beings, past and present, become as throwing light on the ancient geography of the world. Geology loses glory from the imperfection of its archives, but it gains in the immensity of its subject. There is much grandeur in looking at every existing organic being either as the lineal successor of some form now buried under thousands of feet of solid rock, or as being the co-descendant of that buried form of some more ancient and utterly lost inhabitant of this world. It accords with what we know of the laws impressed by the Creator on matter that the production and extinction of forms should, like the birth and death of individuals, be the result of secondary means. It is derogatory that the Creator of countless Universes should have made by individual acts of His will the myriads of creeping parasites and worms, which since the earliest dawn of life have swarmed over the land and in the depths of the ocean. We cease to be astonished that a group of animals should have been formed to lay their eggs in the bowels and flesh of other sensitive beings; that some animals should live by and even delight in cruelty; that animals should be led away by false instincts; that annually there should be an incalculable waste of the pollen, eggs and immature beings; for we see in all this the inevitable consequences of one great law, of the multiplication of organic beings not created immutable. From death, famine, and the struggle for existence, we see that the most exalted end which we are capable of conceiving, namely, the creation of the higher animals, has directly proceeded. Doubtless, our first impression is to disbelieve that any secondary law could produce infinitely numerous organic beings, each characterised by the most exquisite workmanship and widely extended adaptations: it at first accords better with our faculties to suppose that each required the fiat of a Creator. There is a [simple] grandeur in this view of life with its several powers of growth, reproduction and of sensation, having been originally breathed into matter under a few forms, perhaps into only one, and that whilst this planet has gone cycling onwards according to the fixed laws of gravity and whilst land and water have gone on replacing each other--that from so simple an origin, through the selection of infinitesimal varieties, endless forms most beautiful and most wonderful have been evolved.
 This sentence corresponds, not to the final section of the Origin, Ed. i. p. 484, vi. p. 664, but rather to the opening words of the section already referred to (Origin, Ed. i. p. 480, vi. p. 657).
 This simile occurs in the Essay of 1842, p. 50, and in the Origin, Ed. i. p. 485, vi. p. 665, i.e. in the final section of Ch. XIV (vi. Ch. XV). In the MS. there is some erasure in pencil of which I have taken no notice.
 An almost identical sentence occurs in the Origin, Ed. i. p. 487, vi. p. 667. The fine prophecy (in the Origin, Ed. i. p. 486, vi. p. 666) on "the almost untrodden field of inquiry" is wanting in the present Essay.
 See the last paragraph on p. 488 of the Origin, Ed. i., vi. p. 668.
 A passage corresponding to this occurs in the sketch of 1842, p. 51, but not in the last chapter of the Origin.
 This sentence occurs in an almost identical form in the Origin, Ed. i. p. 490, vi. p. 669. It will be noted that man is not named though clearly referred to. Elsewhere (Origin, Ed. i. p. 488) the author is bolder and writes "Light will be thrown on the origin of man and his history." In Ed. vi. p. 668, he writes "Much light &c."
 For the history of this sentence (with which the Origin of Species closes) see the Essay of 1842, p. 52, note 2[Note 184]: also the concluding pages of the Introduction.
 These four words are added in pencil between the lines.
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