On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life

species constantly trying to mcrease in number, with natural selection always ready to adapt the slowly vary- ing descendants of each to any unoccupied or ill-occu- pied place in nature, these facts cease to be strange, or perhaps might even have been anticipated. As natural selection acts by competition, it adapts the inhabitants of each country only in relation to the degree of perfection of their associates; so that we need feel no surprise at the inhabitants of any one country, although on the ordinary view supposed to have been specially created and adapted for that country, being beaten and supplanted by the naturalised produc- tions from another land. Nor ought we to marvel if all the contrivances in nature be not, as far as we can judge, absolutely perfect ; and if some of them be ab- horrent to our ideas of fitness. We need not marvel at the sting of the bee causing the bee’s own death; at drones being produced in such vast numbers for one single act, and being then slaughtered by their sterile sisters; at the astonishing waste of pollen by our fir- trees; at the instinctive hatred of the queen bee for her own fertile daughters ; at ichneumonide feeding within the live bodies of caterpillars; and at other such cases. The wonder indeed is, on the theory of natural selection, that more cases of the want of absolute perfection have not been observed.

The complex and little known laws governing varia- tion are the same, as far as we can see, with the laws which have governed the production of so-called specific forms. In both cases physical conditions seem to have produced but little direct effect; yet when varieties enter any zone, they occasionally assume some of the characters of the species proper to that zone. In both varieties and species, use and disuse seem to have pro- duced some effect; for it is difficult to resist this con-

clusion when we look, for instance, at the logger-headed duck, which has wings incapable of flight, in nearly the same condition as in the domestic duck; or when we look at the burrowing tucutucu, which is occasionally blind, and then at certain moles, which are habitually blind and have their eyes covered with skin; or when we look at the blind animals inhabiting the dark caves of America and Europe. In both varieties and species correlation of growth seems to have played a most im- portant part, so that when one part has been modified other parts are necessarily modified. In both varieties and species reversions to long-lost characters occur. How inexplicable on the theory of creation is the occa- sional appearance of stripes on the shoulder and legs of the several species of the horse-genus and in their hybrids! How simply is this fact explained if we believe that these species have descended from a striped progenitor, in the same manner as the several domestic breeds of pigeon have descended from the blue and barred rock-pigeon !

On the ordinary view of each species having been independently created, why should the specific charac- ters, or those by which the species of the same genus differ from each other, be more variable than the generic characters in which they all agree ? Why, for instance, should the colour of a flower be more likely to vary in any one species of a genus, if the other species, supposed to have been created independently, have differently co- loured flowers, than if all the species of the genus have the same coloured flowers? If species are only well-marked varieties, of which the characters have become in a high degree permanent, we can understand this fact; for they have already varied since they branched off from a common progenitor in certain characters, by which they have come to be specifically distinct from each other ;

and therefore these same characters would be more likely still to be variable than the generic characters which have been inherited without change for an enor- mous period. It is inexplicable on the theory of crea- tion why a part developed in a very unusual manner in any one species of a genus, and therefore, as we may naturally infer, of great importance to the species, should be eminently liable to variation; but, on my view, this part has undergone, since the several species branched off from a common progenitor, an unusual amount of variability and modification, and therefore we might expect this part generally to be still variable. But a part may be developed in the most unusual manner, like the wing of a bat, and yet not be more variable than any other structure, if the part be common to many subordinate forms, that is, if it has been inherited for a very long period; for in this case it will have been rendered constant by long-continued natural selection. Glancing at instincts, marvellous as some are, they offer no greater difficulty than does corporeal structure on the theory of the natural selection of successive, slight, but profitable modifications. We can thus understand why nature moves by graduated steps in endowing dif- ferent animals of the same class with their several in- stincts. J have attempted to show how much light the principle of gradation throws on the admirable archi- tectural powers of the hive-bee. Habit no doubt some- times comes into play in modifying instincts; but it certainly is not indispensable, as we see, in the case of neuter insects, which leave no progeny to inherit the effects of long-continued habit.

On the view of all the species of the same genus having descended from a common parent, and having inherited much in common, we can understand how it is that allied species, when placed under considerably different conditions of life, yet should follow nearly the same instincts; why the thrush of South America, for instance, lines her nest with mud like our British species. On the view of instincts having been slowly acquired through natural selection we need not marvel at some instincts beg apparently not perfect and liable to mistakes, and at many instincts causing other animals to suffer.

If species be only well-marked and permanent varie- ties, we can at once see why their crossed offspring should follow the same complex laws in their degrees and kinds of resemblance to their parents,—in being ab- sorbed into each other by successive crosses, and in other such points,—as do the crossed offspring of acknow- ledged varieties. On the other hand, these would be strange facts if species have been independently created, and varieties have been produced by secondary laws.

If we admit that the geological record is imperfect in an extreme degree, then such facts as the record gives, support the theory of descent with modification. New species have come on the stage slowly and at successive intervals; and the amount of change, after equal intervals of time, is widely different in different groups. The extinction of species and of whole groups of species, which has played so conspicuous a part in the history of the organic world, almost inevitably follows on the principle of natural selection ; for old forms will be supplanted by new and improved forms. Neither single species nor groups of species reappear when the chain of ordinary generation has once been broken. The gradual diffusion of dominant forms, with the slow modification of their descendants, causes the forms of life, after long intervals of time, to appear as if they had changed simultaneously throughout the world. ‘The fact of the fossil remains of each formation being in some degree intermediate in character between the

fossils in the formations above and below, is simply explained by their intermediate position in the chain of descent. The grand fact that all extinct organic beings belong to the same system with recent beings, falling either into the same or into intermediate groups, follows from the living and the extinct beimg the offspring of common parents. As the groups which have de- scended from an ancient progenitor have generally diverged in character, the progenitor with its early de- scendants will often be intermediate in character in comparison with its later descendants ; and thus we can see why the more ancient a fossil is, the oftener it stands in some degree intermediate between existing and allied groups. Recent forms are generally looked at as being, in some vague sense, higher than ancient and extinct forms; and they are in so far higher as the later and more improved forms have conquered the older and less improved organic beings in the struggle for life. Lastly, the law of the long endurance of allied forms on the same continent,—of marsupials in Australia, of edentata in America, and other such cases,—is intelligible, for within a confined country, the recent and the extinct will naturally be allied by descent.

Looking to geographical distribution, if we admit that there has been during the long course of ages much migration from one part of the world to another, owing to former climatal and geographical changes and to the many occasional and unknown means of dispersal, then we can understand, on the theory of descent with modification, most of the great leading facts in Distribu- tion. We can see why there should be so striking a parallelism in the distribution of organic beings through- out space, and in their geological succession throughout time ; for in both cases the beings have been connected by the bond of ordinary generation, and the means of

modification have been the same. We see the full meaning of the wonderful fact, which must have struck every traveller, namely, that on the same continent, under the most diverse conditions, under heat and cold, on mountain and lowland, on deserts and marshes, most of the inhabitants within each great class are plainly related; for they will generally be descendants of the same progenitors and early colonists. On this same principle of former migration, combined in most cases with modification, we can understand, by the aid of the Glacial period, the identity of some few plants, and the close alliance of many others, on the most distant moun- tains, under the most different climates; and likewise the close alliance of some of the inhabitants of the sea in the northern and southern temperate zones, though separated by the whole intertropical ocean. Although two areas may present the same physical conditions of life, we need feel no surprise at their inhabitants being widely different, if they have been for a long period completely separated from each other; for as the rela- tion of organism to organism is the most important of all relations, and as the two areas will have received colonists from some third source or from each other, at various periods and in different proportions, the course of modification in the two areas will inevitably be different.

On this view of migration, with subsequent modifica- tion, we can see why oceanic islands should be inhabited by few species, but of these, that many should be peculiar. We can clearly see why those animals which cannot cross wide spaces of ocean, as frogs and terrestrial mammals, should not inhabit oceanic islands ; and why, on the other hand, new and peculiar species of bats, which can traverse the ocean, should so often be found on islands far distant from any continent. Such facts

as the presence of peculiar species of bats, and the ab- sence of all other mammals, on oceanic islands, are utterly mexplicable on the theory of independent acts of creation. The existence of closely allied or representative spe- cles in any two areas, implies, on the theory of descent with modification, that the same parents formerly in- habited both areas; and we almost invariably find that wherever many closely allied species inhabit two areas, some identical species common to both still exist. Wherever many closely allied yet distinct species occur, many doubtful forms and varieties of the same species likewise occur. It is a rule of high generality that the inhabitants of each area are related to the inhabitants of the nearest source whence immigrants might have been derived. We see this in nearly all the plants and animals of the Galapagos archipelago, of Juan Fernandez, and of the other American islands being related in the most striking manner to the plants and animals of the neighbouring American mainland; and those of the Cape de Verde archipelago and other African islands to the African mainland. It must be admitted that these facts receive no explanation on the theory of creation.

The fact, as we have seen, that all past and present organic beings constitute one grand natural system, with group subordinate to group, and with extinct groups often falling in between recent groups, is intelligible on the theory of natural selection with its contingencies of extinction and divergence of character. On these same principles we see how it is, that the mutual affinities of the species and genera within each class are so complex and circuitous. We see why certain characters are far more serviceable than others for classification ; — why adaptive characters, though of paramount importance to the being, are of hardly any

importance in classification; why characters derived from rudimentary parts, though of no service to the being, are often of high classificatory value; and why embryological characters are the most valuable of all. The real affinities of all organic beings are due to inhe- ritance or community of descent. The natural system is a genealogical arrangement, in which we have to discover the lines of descent by the most permanent characters, however slight their vital importance may be.

The framework of bones being the same in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse,—the same number of vertebre forming the neck of the giraffe and of the elephant,—and innu- merable other such facts, at once explain themselves on the theory of descent with slow and slight successive modifications. The similarity of pattern in the wing and leg of a bat, though used for such different purpose,—in the jaws and legs of a crab,—in the petals, stamens, and pistils of a flower, is likewise intelligible on the view of the gradual modification of parts or organs, which were alike in the early progenitor of each class. On the principle of successive variations not always supervening at an early age, and being inherited at a corresponding not early period of life, we can clearly see why the em- bryos of mammals, birds, reptiles, and fishes should be so closely alike, and should be so unlike the adult forms. We may cease marvelling at the embryo of an air- breathing mammal or bird having branchial slits and arteries running in loops, like those in a fish which has to breathe the air dissolved in water, by the aid of well- developed branchiz.

Disuse, aided sometimes by natural selection, will often tend to reduce an organ, when it has become useless by changed habits or under changed conditions of life; and we can clearly understand on this view the meaning of rudimentary organs. But disuse and selec- tion will generally act on each creature, when it has come to maturity and has to play its full part in the struggle for existence, and will thus have little power of acting on an organ during early life; hence the organ will not be much reduced or rendered rudimentary at this early age. The calf, for instance, has inherited teeth, which never cut through the gums of the upper jaw, from an early progenitor having well-developed teeth ; and we may believe, that the teeth in the mature animal were reduced, during successive gene- rations, by disuse or by the tongue and palate having been fitted by natural selection to browse without their aid; whereas in the calf, the teeth have been left un- touched by selection or disuse, and on the principle of inheritance at corresponding ages have been inherited from a remote period to the present day. On the view of each organic being and each separate organ having been specially created, how utterly inexplicable it is that parts, like the teeth in the embryonic calf or like the shrivelled wings under the soldered wing-covers of some beetles, should thus so frequently bear the plam stamp of inutility! Nature may be said to have taken pains to reveal, by rudimentary organs and by homologous structures, her scheme of modification, which it seems that we wilfully will not understand.

I have now recapitulated the chief facts and consider- ations which have thoroughly convinced me that species have changed, and are still slowly changing by the pre- servation and accumulation of successive slight favour- able variations. Why, it may be asked, have all the most eminent living naturalists and geologists rejected this view of 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 quantity; 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 pro- ductions 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 other and distinct species, is that we are always slow in admitting any great change of which we do not see the interme- diate 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 slow action of the coast-waves. The mind cannot possibly grasp the full meaning of the term of a hundred million years; it cannot add up and perceive the full effects of many slight variations, accumulated during an almost infinite number of ge- nerations.

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,” “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 explana- tion of a certain number of facts will certainly reject my theory. A few naturalists, endowed with much flexibility of mind, and who have already begun to doubt on 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. Who- ever is led to believe that species are mutable will do good service by conscientiously expressing his convic- tion; for only thus 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 natu- ralists, and which consequently have every external characteristic feature of true species,—they admit that these have been produced by variation, but they refuse to extend the same view to other and very 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 cer- tain 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? Although naturalists very properly demand a full explanation of every difficulty from those who believe in the mutability of species, on their own side they ignore the whole subject of the first appearance of species in what they consider reverent silence.

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 may consider, by so much the arguments fall away in force. But some arguments of the greatest weight extend very far. All the members of whole classes can be connected together by chains of affinities, and all can be classified on the same principle, in groups sub- ordinate to groups. Jossil 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 state; and this in some instances necessarily implies an enormous amount of modification in the descendants. Throughout whole classes various structures are formed on the same pattern, and at an embryonic age the species closely resemble each other. Therefore I can- not doubt that the theory of descent with modification

embraces all the members of the same class. I believe that animals have descended from at most only four or five progenitors, and plants from an equal or lesser number. Analogy would lead me one step further, namely, to the belief that all animals and plants have descended from some one prototype. But analogy may be a de- ceitful guide. Nevertheless all living things have much in common, in their chemical composition, their germinal vesicles, their cellular structure, and their laws of growth and reproduction. We see this even in so trifling a cir- cumstance as that the same poison often similarly affects plants and animals; or that the poison secreted by the gall-fly produces monstrous growths on the wild rose or oak-tree. Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed.

When the views entertained in this volume on the origin of species, or when analogous views are generally admitted, we can dimly foresee that there will be a con- siderable 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 in essence a species. This I feel sure, and I speak after experience, will be no slight relief. The endiess disputes whether or not some fifty species of British brambles are true 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 con-

sideration than it is at present; for differences, how- ever slight, between any two forms, if not blended by intermediate gradations, are looked at by most natural- ists as sufficient to raise both forms to the rank of species. Hereafter we shall be compelled to acknow- ledge 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 inter- mediate gradations, whereas species were formerly thus connected. Hence, without quite rejecting the con- sideration of the present existence of intermediate gra- dations 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 varie- ties may hereafter be thought worthy of specific names, as with the primrose and cowslip; and in this case scientific and common language will come into aecord- ance. 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 freed 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 at something wholly beyond his comprehension; when we regard every production of nature as one which has had a history ; when we contemplate every complex structure

and instinct as the summing up of many contrivances, each useful to the possessor, nearly in the same way as when we look at any great mechanical invention as 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, will 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 corre- lation of growth, 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 far 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 dis- cover 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. Hmbryology will reveal to us the structure, in some degree obscured, of the prototypes of each great class.

When we can 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 de- scended from one parent, and have migrated from some one birthplace; 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 of the inhabitants of the sea on the opposite sides of a conti- nent, and the nature of the various inhabitants of that continent in relation to their apparent means of immigra- tion, 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 embedded 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 haying depended on an unusual concurrence of circum- stances, 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 include few identical species, by the general succession of their forms of life. As species are produced and exterminated by slowly act- ing and still existing causes, and not by miraculous acts of creation and by catastrophes; 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 being 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 lapse of actual time. A number of species, however, keeping in a body might remain for a long period unchanged, whilst within this 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. During early periods of the earth’s history, when the forms of life were probably fewer and simpler, the rate of change was probably slower; and at the first dawn of life, when very few forms of the simplest structure existed, the rate of change may have been slow in an extreme degree. The whole history of the world, as at present known, although of a length quite incompre- hensible by us, will hereafter be recognised as a mere fragment of time, compared with the ages which have elapsed since the first creature, the progenitor of innu- merable extinct and living descendants, was created.