An Historical Sketch Of The Progress Of Opinion On The Origin Of Species, Previously To The Publication Of The First Edition Of This Work

I will here give a brief sketch of the progress of opinion on the Origin of Species. Until recently the great majority of naturalists believed that species were immutable productions, and had been separately created. This view has been ably maintained by many authors. Some few naturalists, on the other hand, have believed that species undergo modification, and that the existing forms of life are the descendants by true generation of pre existing forms. Passing over allusions to the subject in the classical writers (Aristotle, in his "Physicae Auscultationes" (lib. 2, cap. 8, s. 2), after remarking that rain does not fall in order to make the corn grow, any more than it falls to spoil the farmer's corn when threshed out of doors, applies the same argument to organisation; and adds (as translated by Mr. Clair Grece, who first pointed out the passage to me), "So what hinders the different parts (of the body) from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity; and whatsoever things were not thus constituted, perished and still perish." We here see the principle of natural selection shadowed forth, but how little Aristotle fully comprehended the principle, is shown by his remarks on the formation of the teeth.), the first author who in modern times has treated it in a scientific spirit was Buffon. But as his opinions fluctuated greatly at different periods, and as he does not enter on the causes or means of the transformation of species, I need not here enter on details. Lamarck was the first man whose conclusions on the subject excited much attention. This justly celebrated naturalist first published his views in 1801; he much enlarged them in 1809 in his "Philosophie Zoologique", and subsequently, 1815, in the Introduction to his "Hist. Nat. des Animaux sans Vertebres". In these works he up holds the doctrine that all species, including man, are descended from other species. He first did the eminent service of arousing attention to the probability of all change in the organic, as well as in the inorganic world, being the result of law, and not of miraculous interposition. Lamarck seems to have been chiefly led to his conclusion on the gradual change of species, by the difficulty of distinguishing species and varieties, by the almost perfect gradation of forms in certain groups, and by the analogy of domestic productions. With respect to the means of modification, he attributed something to the direct action of the physical conditions of life, something to the crossing of already existing forms, and much to use and disuse, that is, to the effects of habit. To this latter agency he seems to attribute all the beautiful adaptations in nature; such as the long neck of the giraffe for browsing on the branches of trees. But he likewise believed in a law of progressive development, and as all the forms of life thus tend to progress, in order to account for the existence at the present day of simple productions, he maintains that such forms are now spontaneously generated. (I have taken the date of the first publication of Lamarck from Isidore Geoffroy Saint- Hilaire's (" Hist. Nat. Generale", tom. ii. page 405, 1859) excellent history of opinion on this subject. In this work a full account is given of Buffon's conclusions on the same subject. It is curious how largely my grandfather, Dr. Erasmus Darwin, anticipated the views and erroneous grounds of opinion of Lamarck in his "Zoonomia" (vol. i. pages 500-510), published in 1794. According to Isid. Geoffroy there is no doubt that Goethe was an extreme partisan of similar views, as shown in the introduction to a work written in 1794 and 1795, but not published till long afterward; he has pointedly remarked (" Goethe als Naturforscher", von Dr. Karl Meding, s. 34) that the future question for naturalists will be how, for instance, cattle got their horns and not for what they are used. It is rather a singular instance of the manner in which similar views arise at about the same time, that Goethe in Germany, Dr. Darwin in England, and Geoffroy Saint-Hilaire (as we shall immediately see) in France, came to the same conclusion on the origin of species, in the years 1794-5.) Geoffroy Saint-Hilaire, as is stated in his "Life", written by his son, suspected, as early as 1795, that what we call species are various degenerations of the same type. It was not until 1828 that he published his conviction that the same forms have not been perpetuated since the origin of all things. Geoffroy seems to have relied chiefly on the conditions of life, or the "monde ambiant" as the cause of change. He was cautious in drawing conclusions, and did not believe that existing species are now undergoing modification; and, as his son adds, "C'est donc un probleme a reserver entierement a l'avenir, suppose meme que l'avenir doive avoir prise sur lui." In 1813 Dr. W.C. Wells read before the Royal Society "An Account of a White Female, part of whose skin resembles that of a Negro"; but his paper was not published until his famous "Two Essays upon Dew and Single Vision" appeared in 1818. In this paper he distinctly recognises the principle of natural selection, and this is the first recognition which has been indicated; but he applies it only to the races of man, and to certain characters alone. After remarking that negroes and mulattoes enjoy an immunity from certain tropical diseases, he observes, firstly, that all animals tend to vary in some degree, and, secondly, that agriculturists improve their domesticated animals by selection; and then, he adds, but what is done in this latter case "by art, seems to be done with equal efficacy, though more slowly, by nature, in the formation of varieties of mankind, fitted for the country which they inhabit. Of the accidental varieties of man, which would occur among the first few and scattered inhabitants of the middle regions of Africa, some one would be better fitted than others to bear the diseases of the country. This race would consequently multiply, while the others would decrease; not only from their in ability to sustain the attacks of disease, but from their incapacity of contending with their more vigorous neighbours. The colour of this vigorous race I take for granted, from what has been already said, would be dark. But the same disposition to form varieties still existing, a darker and a darker race would in the course of time occur: and as the darkest would be the best fitted for the climate, this would at length become the most prevalent, if not the only race, in the particular country in which it had originated." He then extends these same views to the white inhabitants of colder climates. I am indebted to Mr. Rowley, of the United States, for having called my attention, through Mr. Brace, to the above passage of Dr. Wells' work. The Hon. and Rev. W. Herbert, afterward Dean of Manchester, in the fourth volume of the "Horticultural Transactions", 1822, and in his work on the "Amaryllidaceae" (1837, pages 19, 339), declares that "horticultural experiments have established, beyond the possibility of refutation, that botanical species are only a higher and more

On the origin of species

When on board H.M.S. Beagle, as naturalist, I was much struck with certain facts in the distribution of the organic beings inhabiting South America, and in the geological relations of the present to the past inhabitants of that continent. These facts, as will be seen in the latter chapters of this volume, seemed to throw some light on the origin of species— that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, it occurred to me, in 1837, that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it.

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As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be NATURALLY SELECTED. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.

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natural selection almost inevitably causes much extinction of the less improved forms of life, and leads to what I have called divergence of character.I can entertain no doubt, after the most deliberate study and dispassionate judgment of which I am capable, that the view which most naturalists until recently entertained, and which I formerly entertained— namely, that each species has been independently created— is erroneous. I am fully convinced that species are not immutable; but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species, in the same manner as the acknowledged varieties of any one species are the descendants of that species. Furthermore, I am convinced that natural selection has been the most important, but not the exclusive, means of modification.

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We see in definite variability in the endless slight peculiarities which distinguish the individuals of the same species, and which cannot be accounted for by inheritance from either parent or from some more remote ancestor. Even strongly-marked differences occasionally appear in the young of the same litter, and in seedlings from the same seed-capsule. At long intervals of time, out of millions of individuals reared in the same country and fed on nearly the same food, deviations of structure so strongly pronounced as to deserve to be called monstrosities arise; but monstrosities cannot be separated by any distinct line from slighter variation....

Important changes in the embryo or larva will probably entail changes in the mature animal...

Any variation which is not inherited is unimportant for us. But the number and diversity of inheritable deviations of structure, both those of slight and those of considerable physiological importance, are endless.

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.When any deviation of structure often appears, and we see it in the father and child, we cannot tell whether it may not be due to the same cause having acted on both; but when among individuals, apparently exposed to the same conditions, any very rare deviation, due to some extraordinary combination of circumstances, appears in the parent— say, once among several million individuals— and it reappears in the child, the mere doctrine of chances almost compels us to attribute its reappearance to inheritance.

Every one must have heard of cases of albinism, prickly skin, hairy bodies, etc., appearing in several members of the same family. If strange and rare deviations of structure are truly inherited, less strange and commoner deviations may be freely admitted to be inheritable. Perhaps the correct way of viewing the whole subject would be, to look at the inheritance of every character whatever as the rule, and non-inheritance as the anomaly. The laws governing inheritance are for the most part unknown; no one can say why the same peculiarity in different individuals of the same species, or in different species, is sometimes inherited and sometimes not so; why the child often reverts in certain characteristics to its grandfather or grandmother or more remote ancestor; why a peculiarity is often transmitted from one sex to both sexes, or to one sex alone, more commonly but not exclusively to the like sex.

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When we look to the hereditary varieties or races of our domestic animals and plants, and compare them with closely allied species, we generally perceive in each domestic race, as already remarked, less uniformity of character than in true species. Domestic races often have a somewhat monstrous character; by which I mean, that, although differing from each other and from other species of the same genus, in several trifling respects, they often differ in an extreme degree in some one part, both when compared one with another, and more especially when compared with the species under nature to which they are nearest allied.

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Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species.....The key is man's power of accumulative selection: nature gives successive variations; man adds them up in certain directions useful to him. In this sense he may be said to have made for himself useful breeds.

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Almost every part of every organic being is so beautifully related to its complex conditions of life that it seems as improbable that any part should have been suddenly produced perfect, as that a complex machine should have been invented by man in a perfect state....

If monstrous forms... ever do appear in a state of nature and are capable of reproduction (which is not always the case), as they occur rarely and singly, their preservation would depend on unusually favourable circumstances.

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A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season or occasional year, otherwise, on the principle of geometrical increase, its numbers would quickly become so inordinately great that no country could support the product. Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence, either one individual with another of the same species, or with the individuals of distinct species, or with the physical conditions of life.

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The amount of food for each species, of course, gives the extreme limit to which each can increase; but very frequently it is not the obtaining food, but the serving as prey to other animals, which determines the average number of a species.

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Climate plays an important part in determining the average numbers of a species, and periodical seasons of extreme cold or drought seem to be the most effective of all checks.

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The action of climate seems at first sight to be quite independent of the struggle for existence; but in so far as climate chiefly acts in reducing food, it brings on the most severe struggle between the individuals, whether of the same or of distinct species, which subsist on the same kind of food. Even when climate, for instance, extreme cold, acts directly, it will be the least vigorous individuals, or those which have got least food through the advancing winter, which will suffer the most. When we travel from south to north, or from a damp region to a dry, we invariably see some species gradually getting rarer and rarer, and finally disappearing; and the change of climate being conspicuous, we are tempted to attribute the whole effect to its direct action. But this is a false view; we forget that each species, even where it most abounds, is constantly suffering enormous destruction at some period of its life, from enemies or from competitors for the same place and food; and if these enemies or competitors be in the least degree favoured by any slight change of climate, they will increase in numbers; and as each area is already fully stocked with inhabitants, the other species must decrease.

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When a plant or animal is placed in a new country, among new competitors, the conditions of its life will generally be changed in an essential manner, although the climate may be exactly the same as in its former home. If its average numbers are to increase in its new home, we should have to modify it in a different way to what we should have had to do in its native country; for we should have to give it some advantage over a different set of competitors or enemies. It is good thus to try in imagination to give any one species an advantage over another. Probably in no single instance should we know what to do. This ought to convince us of our ignorance on the mutual relations of all organic beings; a conviction as necessary, as it is difficult to acquire. All that we can do is to keep steadily in mind that each organic being is striving to increase in a geometrical ratio; that each, at some period of its life, during some season of the year, during each generation, or at intervals, has to struggle for life and to suffer great destruction.When we reflect on this struggle we may console ourselves with the full belief that the war of nature is not incessant, that no fear is felt, that death is generally prompt, and that the vigorous, the healthy, and the happy survive and multipy.

This preservation of favourable individual differences and variations, and the destruction of those which are injurious, I have called Natural Selection, or the Survival of the Fittest.

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We shall best understand the probable course of natural selection by taking the case of a country undergoing some slight physical change, for instance, of climate. The proportional numbers of its inhabitants will almost immediately undergo a change, and some species will probably become extinct. We may conclude, from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together, that any change in the numerical proportions of the inhabitants, independently of the change of climate itself, would seriously affect the others. If the country were open on its borders, new forms would certainly immigrate, and this would likewise seriously disturb the relations of some of the former inhabitants. But in the case of an island, or of a country partly surrounded by barriers, into which new and better adapted forms could not freely enter, we should then have places in the economy of nature which would assuredly be better filled up if some of the original inhabitants were in some manner modified; for, had the area been open to immigration, these same places would have been seized on by intruders. In such cases, slight modifications, which in any way favoured the individuals of any species, by better adapting them to their altered conditions, would tend to be preserved; and natural selection would have free scope for the work of improvement.

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It may metaphorically be said that natural selection is daily and hourly scrutinising, throughout the world, the slightest variations; rejecting those that are bad, preserving and adding up all that are good; silently and insensibly working, WHENEVER AND WHEREVER OPPORTUNITY OFFERS, at the improvement of each organic being in relation to its organic and inorganic conditions of life. We see nothing of these slow changes in progress, until the hand of time has marked the long lapse of ages, and then so imperfect is our view into long-past geological ages that we see only that the forms of life are now different from what they formerly were.

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This leads me to say a few words on what I have called sexual selection. This form of selection depends, not on a struggle for existence in relation to other organic beings or to external conditions, but on a struggle between the individuals of one sex, generally the males, for the possession of the other sex. The result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection. Generally, the most vigorous males, those which are best fitted for their places in nature, will leave most progeny. But in many cases victory depends not so much on general vigour, but on having special weapons, confined to the male sex... Thus it is, as I believe, that when the males and females of any animal have the same general habits of life, but differ in structure, colour, or ornament, such differences have been mainly caused by sexual selection: that is, by individual males having had, in successive generations, some slight advantage over other males, in their weapons, means of defence, or charms; which they have transmitted to their male offspring alone.

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Isolation also is an important element in the modification of species through natural selection. In a confined or isolated area, if not very large, the organic and inorganic conditions of life will generally be almost uniform; so that natural selection will tend to modify all the varying individuals of the same species in the same manner. Intercrossing with the inhabitants of the surrounding districts, will also be thus prevented.

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Although isolation is of great importance in the production of new species, on the whole I am inclined to believe that largeness of area is still more important, especially for the production of species which shall prove capable of enduring for a long period, and of spreading widely. Throughout a great and open area, not only will there be a better chance of favourable variations, arising from the large number of individuals of the same species there supported, but the conditions of life are much more complex from the large number of already existing species; and if some of these many species become modified and improved, others will have to be improved in a corresponding degree, or they will be exterminated. Each new form, also, as soon as it has been much improved, will be able to spread over the open and continuous area, and will thus come into competition with many other forms.

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Slow though the process of selection may be, if feeble man can do much by artificial selection, I can see no limit to the amount of change, to the beauty and complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may have been effected in the long course of time through nature's power of selection, that is by the survival of the fittest.

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As new species in the course of time are formed through natural selection, others will become rarer and rarer, and finally extinct. The forms which stand in closest competition with those undergoing modification and improvement, will naturally suffer most.

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Natural selection acts exclusively by the preservation and accumulation of variations, which are beneficial under the organic and inorganic conditions to which each creature is exposed at all periods of life. The ultimate result is that each creature tends to become more and more improved in relation to its conditions. This improvement inevitably leads to the gradual advancement of the organisation of the greater number of living beings throughout the world. organisation. Among the vertebrata the degree of intellect and an approach in structure to man clearly come into play.

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part. If two species belonging to two distinct though allied genera, had both produced a large number of new and divergent forms, it is conceivable that these might approach each other so closely that they would have all to be classed under the same genus; and thus the descendants of two distinct genera would converge into one. But it would in most cases be extremely rash to attribute to convergence a close and general similarity of structure in the modified descendants of widely distinct forms.