"Recapitulation and Conclusion"
By Charles Darwin.

I have now recapitulated the facts and considerations which have thoroughly convinced me that species have been modified, during a long course of descent. This has been effected chiefly through the natural selection of numerous successive, slight, favourable variations; aided in an important manner by the inherited effects of the use and disuse of parts; and in an unimportant manner, that is in relation to adaptive structures, whether past or present, by the direct action of external conditions, and by variations which seem to us in our ignorance to arise spontaneously. It appears that I formerly underrated the frequency and value of these latter forms of variation, as leading to permanent modifications of structure independently of natural selection. But as my conclusions have lately been much misrepresented, and it has been stated that I attribute the modification of species exclusively to natural selection, I may be permitted to remark that in the first edition of this work, and subsequently, I placed in a most conspicuous position -- namely, at the close of the Introduction -- the following words: "I am convinced that natural selection has been the main but not the exclusive means of modification." This has been of no avail. Great is the power of steady misrepresentation; but the history of science shows that fortunately this power does not long endure.

It can hardly be supposed that a false theory would explain, in so satisfactory a manner as does the theory of natural selection, the several large classes of facts above specified. It has recently been objected that this is an unsafe method of arguing; but it is a method used in judging of the common events of life, and has often been used by the greatest natural philosophers. The undulatory theory of light has thus been arrived at; and the belief in the revolution of the earth on its own axis was until lately supported by hardly any direct evidence. It is no valid objection that science as yet throws no light on the far higher problem of the essence or origin of life. Who can explain what is the essence of the attraction of gravity? No one now objects to following out the results consequent on this unknown element of attraction; notwithstanding that Leibnitz formerly accused Newton of introducing "occult qualities and miracles into philosophy."

I see no good reason why the views given in this volume should shock the religious feelings of any one. It is satisfactory, as showing how transient such impressions are, to remember that the greatest discovery ever made by man, namely, the law of the attraction of gravity, was also attacked by Leibnitz, "as subversive of natural, and inferentially of revealed, religion." A celebrated author and divine has written to me that "he has gradually learnt to see that it is just as noble a conception of the Deity to believe that He created a few original forms capable of self-development into other and needful forms, as to believe that He required a fresh act of creation to supply the voids caused by the action of His laws."

Why, it may be asked, until recently did nearly all the most eminent living naturalists and geologists disbelieve in the mutability of species? It cannot be asserted that organic beings in a state of nature are subject to no variation; it cannot be proved that the amount of variation in the course of long ages is a limited quality; no clear distinction has been, or can be, drawn between species and well-marked varieties. It cannot be maintained that species when intercrossed are invariably sterile, and varieties invariably fertile; or that sterility is a special endowment and sign of creation. The belief that species were immutable productions was almost unavoidable as long as the history of the world was thought to be of short duration; and now that we have acquired some idea of the lapse of time, we are too apt to assume, without proof, that the geological record is so perfect that it would have afforded us plain evidence of the mutation of species, if they had undergone mutation.

But the chief cause of our natural unwillingness to admit that one species has given birth to clear and distinct species, is that we are always slow in admitting great changes of which we do not see the steps. The difficulty is the same as that felt by so many geologists, when Lyell first insisted that long lines of inland cliffs had been formed, and great valleys excavated, by the agencies which we see still at work. The mind cannot possibly grasp the full meaning of the term of even a million years; it cannot add up and perceive the full effects of many slight variations, accumulated during an almost infinite number of generations.

Although I am fully convinced of the truth of the views given in this volume under the form of an abstract, I by no means expect to convince experienced naturalists whose minds are stocked with a multitude of facts all viewed, during a long course of years, from a point of view directly opposite to mine. It is so easy to hide our ignorance under such expressions as the "plan of creation" or "unity of design," &c., and to think that we give an explanation when we only restate a fact. Any one whose disposition leads him to attach more weight to unexplained difficulties than to the explanation of a certain number of facts will certainly reject the theory. A few naturalists, endowed with much flexibility of mind, and who have already begun to doubt the immutability of species, may be influenced by this volume; but I look with confidence to the future, -- to young and rising naturalists, who will be able to view both sides of the question with impartiality. Whoever is led to believe that species are mutable will do good service by conscientiously expressing his conviction; for thus only can the load of prejudice by which this subject is overwhelmed be removed.

Several eminent naturalists have of late published their belief that a multitude of reputed species in each genus are not real species; but that other species are real, that is, have been independently created. This seems to me a strange conclusion to arrive at. They admit that a multitude of forms, which till lately they themselves thought were special creations, and which are still thus looked at by the majority of naturalists, and which consequently have all the external characteristic features of true species, -- they admit that these have been produced by variation, but they refuse to extend the same view to other and slightly different forms. Nevertheless they do not pretend that they can define, or even conjecture, which are the created forms of life, and which are those produced by secondary laws. They admit variation as a vera causa in one case, they arbitrarily reject it in another, without assigning any distinction in the two cases. The day will come when this will be given as a curious illustration of the blindness of preconceived opinion. These authors seem no more startled at a miraculous act of creation than at an ordinary birth. But do they really believe that at innumerable periods in the earth’s history certain elemental atoms have been commanded suddenly to flash into living tissues? Do they believe that at each supposed act of creation one individual or many were produced? Were all the infinitely numerous kinds of animals and plants created as eggs or seed, or as full grown? and in the case of mammals, were they created bearing the false marks of nourishment from the mother’s womb? Undoubtedly some of these same questions cannot be answered by those who believe in the appearance or creation of only a few forms of life, or of some one form alone. It has been maintained by several authors that it is as easy to believe in the creation of a million beings as of one; but Maupertuis’ philosophical axiom "of least action" leads the mind more willingly to admit the smaller number; and certainly we ought not to believe that innumerable beings within each great class have been created with plain, but deceptive, marks of descent from a single parent.

As a record of a former state of things, I have retained in the foregoing paragraphs, and elsewhere, several sentences which imply that naturalists believe in the separate creation of each species; and I have been much censured for having thus expressed myself. But undoubtedly this was the general belief when the first edition of the present work appeared. I formerly spoke to very many naturalists on the subject of evolution, and never once met with any sympathetic agreement. It is probable that some did then believe in evolution, but they were either silent, or expressed themselves so ambiguously that it was not easy to understand their meaning. Now things are wholly changed, and almost every naturalist admits the great principle of evolution. There are, however, some who still think that species have suddenly given birth, through quite unexplained means, to new and totally different forms: but, as I have attempted to show, weighty evidence can be opposed to the admission of great and abrupt modifications. Under a scientific point of view, and as leading to further investigation, but little advantage is gained by believing that new forms are suddenly developed in an inexplicable manner from old and widely different forms, over the old belief in the creation of species from the dust of the earth.

It may be asked how far I extend the doctrine of the modification of species. The question is difficult to answer, because the more distinct the forms are which we consider, by so much the arguments in favour of community of descent become fewer in number and less in force. But some arguments of the greatest weight extend very far. All the members of whole classes are connected together by a chain of affinities, and all can be classed on the same principle, in groups subordinate to groups. Fossil remains sometimes tend to fill up very wide intervals between existing orders.

Organs in a rudimentary condition plainly show that an early progenitor had the organ in a fully developed condition; and this in some cases implies an enormous amount of modification in the descendants. Throughout whole classes various structures are formed on the same pattern, and at a very early age the embryos closely resemble each other. Therefore I cannot doubt that the theory of descent with modification embraces all the members of the same great class or kingdom. I believe that animals are descended from at most only four or five progenitors, and plants from an equal or lesser number.

Analogy would lead me one step farther, namely, to the belief that all animals and plants are descended from some one prototype. But analogy may be a deceitful guide. Nevertheless all living things have much in common, in their chemical composition, their cellular structure, their laws of growth, and their liability to injurious influences. We see this even in so trifling a fact as that the same poison often similarly affects plants and animals; or that the poison secreted by the gallfly produces monstrous growths on the wild rose or oak-tree. With all organic beings excepting perhaps some of the very lowest, sexual production seems to be essentially similar. With all, as far as is at present known the germinal vesicle is the same; so that all organisms start from a common origin. If we look even to the two main divisions -- namely, to the animal and vegetable kingdoms -- certain low forms are so far intermediate in character that naturalists have disputed to which kingdom they should be referred. As Professor Asa Gray has remarked, "The spores and other reproductive bodies of many of the lower algae may claim to have first a characteristically animal, and then an unequivocally vegetable existence." Therefore, on the principle of natural selection with divergence of character, it does not seem incredible that, from such low and intermediate form, both animals and plants may have been developed; and, if we admit this, we must likewise admit that all the organic beings which have ever lived on this earth may be descended from some one primordial form. But this inference is chiefly grounded on analogy and it is immaterial whether or not it be accepted. No doubt it is possible, as Mr. G. H. Lewes has urged, that at the first commencement of life many different forms were evolved; but if so we may conclude that only a very few have left modified descendants. For, as I have recently remarked in regard to the members of each great kingdom, such as the Vertebrata, Articulata, &c., we have distinct evidence in their embryological homologous and rudimentary structures that within each kingdom all the members are descended from a single progenitor.

When the views advanced by me in this volume, and by Mr. Wallace, or when analogous views on the origin of species are generally admitted, we can dimly foresee that there will be a considerable revolution in natural history. Systematists will be able to pursue their labours as at present; but they will not be incessantly haunted by the shadowy doubt whether this or that form be a true species. This, I feel sure and I speak after experience, will be no slight relief. The endless disputes whether or not some fifty species of British brambles are good species will cease. Systematists will have only to decide (not that this will be easy) whether any form be sufficiently constant and distinct from other forms, to be capable of definition; and if definable, whether the differences be sufficiently important to deserve a specific name. This latter point will become a far more essential consideration than it is at present; for differences, however slight, between any two forms if not blended by intermediate gradations, are looked at by most naturalists as sufficient to raise both forms to the rank of species.

Hereafter we shall be compelled to acknowledge that the only distinction between species and well-marked varieties is, that the latter are known, or believed, to be connected at the present day by intermediate gradations, whereas species were formerly thus connected. Hence, without rejecting the consideration of the present existence of intermediate gradations between any two forms we shall be led to weigh more carefully and to value higher the actual amount of difference between them. It is quite possible that forms now generally acknowledged to be merely varieties may hereafter be thought worthy of specific names; and in this case scientific and common language will come into accordance. In short, we shall have to treat species in the same manner as those naturalists treat genera, who admit that genera are merely artificial combinations made for convenience. This may not be a cheering prospect; but we shall at least be free from the vain search for the undiscovered and undiscoverable essence of the term species.

The other and more general departments of natural history will rise greatly in interest. The terms used by naturalists, of affinity, relationship, community of type, paternity, morphology, adaptive characters, rudimentary and aborted organs, &c., will cease to be metaphorical, and will have a plain signification. When we no longer look at an organic being as a savage looks at a ship, as something wholly beyond his comprehension; when we regard every production of nature as one which has had a long history; when we contemplate every complex structure and instinct as the summing up of many contrivances, each useful to the possessor, in the same way as any great mechanical invention is the summing up of the labour, the experience, the reason, and even the blunders of numerous workmen; when we thus view each organic being, how far more interesting -- I speak from experience -- does the study of natural history become!

A grand and almost untrodden field of inquiry will be opened, on the causes and laws of variation, on correlation, on the effects of use and disuse, on the direct action of external conditions, and so forth. The study of domestic productions will rise immensely in value. A new variety raised by man will be a more important and interesting subject for study than one more species added to the infinitude of already recorded species. Our classifications will come to be, as far as they can be so made, genealogies; and will then truly give what may be called the plan of creation. The rules for classifying will no doubt become simpler when we have a definite object in view. We possess no pedigrees or armorial bearings; and we have to discover and trace the many diverging lines of descent in our natural genealogies, by characters of any kind which have long been inherited. Rudimentary organs will speak infallibly with respect to the nature of long-lost structures. Species and groups of species which are called aberrant, and which may fancifully be called living fossils, will aid us in forming a picture of the ancient forms of life. Embryology will often reveal to us the structure, in some degree obscured, of the prototype of each great class.

When we feel assured that all the individuals of the same species, and all the closely allied species of most genera, have within a not very remote period descended from one parent, and have migrated from some one birth-place; and when we better know the many means of migration, then, by the light which geology now throws, and will continue to throw, on former changes of climate and of the level of the land, we shall surely be enabled to trace in an admirable manner the former migrations of the inhabitants of the whole world. Even at present, by comparing the differences between the inhabitants of the sea on the opposite sides of a continent, and the nature of the various inhabitants on that continent, in relation to their apparent means of immigration, some light can be thrown on ancient geography.

The noble science of Geology loses glory from the extreme imperfection of the record. The crust of the earth with its imbedded remains must not be looked at as a well-filled museum, but as a poor collection made at hazard and at rare intervals. The accumulation of each great fossiliferous formation will be recognised as having depended on an unusual concurrence of favourable circumstances, and the blank intervals between the successive stages as having been of vast duration. But we shall be able to gauge with some security the duration of these intervals by a comparison of the preceding and succeeding organic forms. We must be cautious in attempting to correlate as strictly contemporaneous two formations, which do not include many identical species, by the general succession of the forms of life. As species are produced and exterminated by slowly acting and still existing causes, and not by miraculous acts of creation; and as the most important of all causes of organic change is one which is almost independent of altered and perhaps suddenly altered physical conditions, namely, the mutual relation of organism to organism, -- the improvement of one organism entailing the improvement or the extermination of others; it follows, that the amount of organic change in the fossils of consecutive formations probably serves as a fair measure of the relative though not actual lapse of time. A number of species, however, keeping in a body might remain for a long period unchanged, whilst within the same period several of these species by migrating into new countries and coming into competition with foreign associates, might become modified; so that we must not overrate the accuracy of organic change as a measure of time.

In the future I see open fields for far more important researches. Psychology will be securely based on the foundation already well laid by Mr. Herbert Spencer, that of the necessary acquirement of each mental power and capacity by gradation. Much light will be thrown on the origin of man and his history.

Authors of the highest eminence seem to be fully satisfied with the view that each species has been independently created. To my mind it accords better with what we know of the laws impressed on matter by the Creator, that the production and extinction of the past and present inhabitants of the world should have been due to secondary causes, like those determining the birth and death of the individual. When I view all beings not as special creations, but as the lineal descendants of some few beings which lived long before the first bed of the Cambrian system was deposited, they seem to me to become ennobled. Judging from the past, we may safely infer that not one living species will transmit its unaltered likeness to a distant futurity. And of the species now living very few will transmit progeny of any kind to a far distant futurity; for the manner in which all organic beings are grouped, shows that the greater number of species in each genus, and all the species in many genera, have left no descendants, but have become utterly extinct. We can so far take a prophetic glance into futurity as to foretell that it will be the common and widely-spread species, belonging to the larger and dominant groups within each class, which will ultimately prevail and procreate new and dominant species. As all the living forms of life are the lineal descendants of those which lived long before the Cambrian epoch, we may feel certain that the ordinary succession by generation has never once been broken, and that no cataclysm has desolated the whole world. Hence we may look with some confidence to secure future of great length. And as natural selection works solely by and for the good of each being, all corporeal and mental endowments will tend to progress towards perfection.

It is interesting to contemplate a tangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent upon each other in so complex a manner, have all been produced by laws acting around us. These laws, taken in the largest sense, being Growth with Reproduction; Inheritance which is almost implied by reproduction; Variability from the indirect and direct action of the conditions of life and from use and disuse: a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to Natural Selection, entailing Divergence of Character and the Extinction of less-improved forms. Thus, from the war of nature, from famine and death, the most exalted object which we are capable of conceiving, namely, the production of the higher animals, directly follows. There is grandeur in this view of life, with its several powers, having been originally breathed by the Creator into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being evolved.