By: Charles Darwin, 1859

AS THIS whole volume is one long argument, it may be convenient to the reader to have the leading facts and inferences briefly recapitulated.

That many and serious objections may be advanced against the theory of descent with modification through variation and natural selection, I do not deny. I have endeavoured to give to them their full force. Nothing at first can appear more difficult to believe than that the more complex organs and instincts have been perfected, not by means superior to, though analogous with, human reason, but by the accumulation of innumerable slight variations, each good for the individual possessor. Nevertheless, this difficulty, though appearing to our imagination insuperably great, cannot be considered real if we admit the following propositions, namely, that all parts of the organisation and instincts offer, at least, individual differences- that there is a struggle for existence leading to the preservation of profitable deviations of structure or instinct- and, lastly, that gradations in the state of perfection of each organ may have existed, each good of its kind. The truth of these propositions cannot, I think, be disputed.

It is, no doubt, extremely difficult even to conjecture by what gradations many structures have been perfected, more especially amongst broken and failing groups of organic beings, which have suffered much extinction, but we see so many strange gradations in nature, that we ought to be extremely cautious in saying that any organ or instinct, or any whole structure, could not have arrived at its present state by many graduated steps. There are, it must be admitted, cases of special difficulty opposed to the theory of natural selection; and one of the most curious of these is the existence in the same community of two or three defined castes of workers or sterile female ants; but I have attempted to show how these difficulties can be mastered.

With respect to the almost universal sterility of species when first crossed, which forms so remarkable a contrast with the almost universal fertility of varieties when crossed, I must refer the reader to the recapitulation of the facts given at the end of the ninth chapter, which seem to me conclusively to show that this sterility is no more a special endowment than is the incapacity of two distinct kinds of trees to be grafted together; but that it is incidental on differences confined to the reproductive systems of the intercrossed species. We see the truth of this conclusion in the vast difference in the results of crossing the same two species reciprocally,- that is, when one species is first used as the father and then as the mother. Analogy from the consideration of dimorphic and trimorphic plants clearly leads to the same conclusion, for when the forms are illegitimately united, they yield few or no seed, and their offspring are more or less sterile; and these forms belong to the same undoubted species, and differ from each other in no respect except in their reproductive organs and functions.

Although the fertility of varieties when intercrossed and of their mongrel offspring has been asserted by so many authors to be universal, this cannot be considered as quite correct after the facts given on the high authority of Gartner and Kolreuter. Most of the varieties which have been experimented on have been produced under domestication; and as domestication (I do not mean mere confinement) almost certainly tends to eliminate that sterility which, judging from analogy, would have affected the parent-species if intercrossed, we ought not to expect that domestication would likewise induce sterility in their modified descendants when crossed. This elimination of sterility apparently follows from the same cause which allows our domestic animals to breed freely under diversified circumstances; and this again apparently follows from their having been gradually accustomed to frequent changes in their conditions of life.

A double and parallel series of facts seems to throw much light on the sterility of species, when first crossed, and of their hybrid offspring. On the one side, there is good reason to believe that slight changes in the conditions of life give vigour and fertility to all organic beings. We know also that a cross between the distinct individuals of the same variety, and between distinct varieties, increases the number of their offspring, and certainly gives to them increased size and vigour. This is chiefly owing to the forms which are crossed having been exposed to somewhat different conditions of life; for I have ascertained by a laborious series of experiments that if all the individuals of the same variety be subjected during several generations to the same conditions, the good derived from crossing is often much diminished or wholly disappears. This is one side of the case. On the other side, we know that species which have long been exposed to nearly uniform conditions, when they are subjected under confinement to new and greatly changed conditions, either perish, or if they survive, are rendered sterile, though retaining perfect health. This does not occur, or only in a very slight degree, with our domesticated productions, which have long been exposed to fluctuating conditions. Hence when we find that hybrids produced by a cross between two distinct species are few in number, owing to their perishing soon after conception or at a very early age, or if surviving that they are rendered more or less sterile, it seems highly probable that this result is due to their having been in fact subjected to a great change in their conditions of life, from being compounded of two distinct organisations. He who will explain in a definite manner why, for instance, air elephant or a fox will not breed under confinement in its native country, whilst the domestic pig or dog will breed freely under the most diversified conditions, will at the same time be able to give a definite answer to the question why two distinct species, when crossed, as well as their hybrid offspring, are generally rendered more or less sterile, whilst two domesticated varieties when crossed and their mongrel offspring are perfectly fertile.

Turning to geographical distribution, the difficulties encountered on the theory of descent with modification are serious enough. All the individuals of the same species, and all the species of the same genus, or even higher group, are descended from common parents; and therefore, in however distant and isolated parts of the world they may now be found, they must in the course of successive generations have travelled from some one point to all the others. We are often wholly unable even to conjecture how this could have been effected. Yet, as we have reason to believe that some species have retained the same specific form for very long periods of time, immensely long as measured by years, too much stress ought not to be laid on the occasional wide diffusion of the same species; for during very long periods there will always have been a good chance for wide migration by many means. A broken or interrupted range may often be accounted for by the extinction of the species in the intermediate regions. It cannot be denied that we are as yet very ignorant as to the full extent of the various climatal and geographical changes which have affected the earth during modern periods; and such changes will often have facilitated migration. As an example, I have attempted to show how potent has been the influence of the Glacial period on the distribution of the same and of allied species throughout the world. We are as yet profoundly ignorant of the many occasional means of transport. With respect to distinct species of the same genus inhabiting distant and isolated regions, as the process of modification has necessarily been slow, all the means of migration will have been possible during a very long period; and consequently the difficulty of the wide diffusion of the species of the same genus is in some degree lessened.

As according to the theory of natural selection an interminable number of intermediate forms must have existed, linking together all the species in each group by gradations as fine as are our existing varieties, it may be asked: Why do we not see these linking forms all around us? Why are not all organic beings blended together in an inextricable chaos? With respect to existing forms, we should remember that we have no right to expect (excepting in rare cases) to discover directly connecting links between them, but only between each and some extinct and supplanted form. Even on a wide area, which has during a long period remained continuous, and of which the climatic and other conditions of life change insensibly in proceeding from a district occupied by one species into another district occupied by a closely allied species, we have no just right to expect often to find intermediate varieties in the intermediate zones. For we have reason to believe that only a few species of a genus ever undergo change; the other species becoming utterly extinct and leaving no modified progeny. Of the species which do change, only a few within the same country change at the same time; and all modifications are slowly effected. I have also shown that the intermediate varieties which probably at first existed in the intermediate zones, would be liable to be supplanted by the allied forms on either hand; for the latter, from existing in greater numbers, would generally be modified and improved at a quicker rate than the intermediate varieties, which existed in lesser numbers; so that the intermediate varieties would, in the long run, be supplanted and exterminated.

On this doctrine of the extermination of an infinitude of connecting links, between the living and extinct inhabitants of the world, and at each successive period between the extinct and still older species, why is not every geological formation charged with such links? Why does not every collection of fossil remains afford plain evidence of the gradation and mutation of the forms of life? Although geological research has undoubtedly revealed the former existence of many links, bringing numerous forms of life much closer together, it does not yield the infinitely many fine gradations between past and present species required on the theory; and this is the most obvious of the many objections which may be urged against it. Why, again, do whole groups of allied species appear, though this appearance is often false, to have come in suddenly on the successive geological stages? Although we now know that organic beings appeared on this globe, at a period incalculably remote, long before the lowest bed of the Cambrian system was deposited, why do we not find beneath this system great piles of strata stored with the remains of the progenitors of the Cambrian fossils? For on the theory, such strata must somewhere have been deposited at these ancient and utterly unknown epochs of the world's history.

I can answer these questions and objections only on the supposition that the geological record is far more imperfect than most geologists believe. The number of specimens in all our museums is absolutely as nothing compared with the countless generations of countless species which have certainly existed. The parent-form of any two or more species would not be in all its characters directly intermediate between its modified offspring, any more than the rock-pigeon is directly intermediate in crop and tail between its descendants, the pouter and fantail pigeons. We should not be able to recognise a species as the parent of another and modified species, if we were to examine the two ever so closely, unless we possessed most of the intermediate links; and owing to the imperfection of the geological record, we have no just right to expect to find so many links. If two or three, or even more linking forms were discovered, they would simply be ranked by many naturalists as so many new species, more especially if found in different geological sub-stages, let their differences be ever so slight. Numerous existing doubtful forms could be named which are probably varieties; but who will pretend that in future ages so many fossil links will be discovered, that naturalists will be able to decide whether or not these doubtful forms ought to be called varieties? Only a small portion of the world has been geologically explored. Only organic beings of certain classes can be preserved in a fossil condition, at least in any great number. Many species when once formed never undergo any further change but become extinct without leaving modified descendants; and the periods, during which species have undergone modification, though long as measured by years, have probably been short in comparison with the periods during which they retain the same form. It is the dominant and widely ranging species which vary most frequently and vary most, and varieties are often at first local- both causes rendering the discovery of intermediate links in any one formation less likely. Local varieties will not spread into other and distant regions until they are considerably modified and improved; and when they have spread, and are discovered in a geological formation, they appear as if suddenly created there, and will be simply classed as new species. Most formations have been intermittent in their accumulation; and their duration has probably been shorter than the average duration of specific forms. Successive formations are in most cases separated from each other by blank intervals of time of great length; for fossiliferous formations thick enough to resist future degradations can as a general rule be accumulated only where much sediment is deposited on the subsiding bed of the sea. During the alternate periods of elevation and of stationary level the record will generally be blank. During these latter periods there will probably be more variability in the forms of life; during periods of subsidence, more extinction.

With respect to the absence of strata rich in fossils beneath the Cambrian formation, I can recur only to the hypothesis given in the tenth chapter; namely, that though our continents and oceans have endured for an enormous period in nearly their present relative positions, we have no reason to assume that this has always been the case; consequently formations much older than any now known may lie buried beneath the great oceans. With respect to the lapse of time not having been sufficient since our planet was consolidated for the assumed amount of organic change, and this objection, as urged by Sir William Thompson, is probably one of the gravest as yet advanced, I can only say, firstly, that we do not know at what rate species change as measured by years, and secondly, that many philosophers are not as yet willing to admit that we know enough of the constitution of the universe and of the interior of our globe to speculate with safety on its past duration.

That the geological record is imperfect all will admit; but that it is imperfect to the degree required by our theory, few will be inclined to admit. If we look to long enough intervals of time, geology plainly declares that species have all changed; and they have changed in the manner required by the theory, for they have changed slowly and in a graduated manner. We clearly see this in the fossil remains from consecutive formations invariably being much more closely related to each other, than are the fossils from widely separated formations.

Such is the sum of the several chief objections and difficulties which may be justly urged against the theory; and I have now briefly recapitulated the answers and explanations which, as far as I can see, may be given. I have felt these difficulties far too heavily during many years to doubt their weight. But it deserves especial notice that the more important objections relate to questions on which we are confessedly ignorant; nor do we know how ignorant we are. We do not know all the possible transitional gradations between the simplest and the most perfect organs; it cannot be pretended that we know all the varied means of Distribution during the long lapse of years, or that we know how imperfect is the Geological Record. Serious as these several objections are, in my judgment they are by no means sufficient to overthrow the theory of descent with subsequent modification.

Now let us turn to the other side of the argument. Under domestication we see much variability, caused, or at least excited, by changed conditions of life; but often in so obscure a manner, that we are tempted to consider the variations as spontaneous. Variability is governed by many complex laws,- by correlated growth, compensation, the increased use and disuse of parts, and the definite action of the surrounding conditions. There is much difficulty in ascertaining how largely our domestic productions have been modified; but we may safely infer that the amount has been large, and that modifications can be inherited for long periods. As long as the conditions of life remain the same, we have reason to believe that a modification, which has already been inherited for many generations, may continue to be inherited for an almost infinite number of generations. On the other hand, we have evidence that variability when it has once come into play, does not cease under domestication for a very long period; nor do we know that it ever ceases, for new varieties are still occasionally produced by our oldest domesticated productions.

Variability is not actually caused by man; he only unintentionally exposes organic beings to new conditions of life, and then nature acts on the organisation and causes it to vary. But man can and does select the variations given to him by nature, and thus accumulates them in any desired manner. He thus adapts animals and plants for his own benefit or pleasure. He may do this methodically, or he may do it unconsciously by preserving the individuals most useful or pleasing to him without any intention of altering the breed. It is certain that he can largely influence the character of a breed by selecting, in each successive generation, individual differences so slight as to be inappreciable except by an educated eye. This unconscious process of selection has been the great agency in the formation of the most distinct and useful domestic breeds. That many breeds produced by man have to a large extent the character of natural species, is shown by the inextricable doubts whether many of them are varieties or aboriginally distinct species.

There is no reason why the principles which have acted so efficiently under domestication should not have acted under nature. In the survival of favoured individuals and races, during the constantly-recurrent Struggle for Existence, we see a powerful and ever-acting form of Selection. The struggle for existence inevitably follows from the high geometrical ratio of increase which is common, to all organic beings. This high rate of increase is proved by calculation,- by the rapid increase of many animals and plants during succession of peculiar seasons, and when naturalised in new countries. More individuals are born than can possibly survive. A grain in the balance may determine which individuals shall live and which shall die,- which variety or species shall increase in number, and which shall decrease, or finally become extinct. As the individuals of the same species come in all respects into the closest competition with each other, the struggle will generally be most severe between them; it will be almost equally severe between the varieties of the same species, and next in severity between the species of the same genus. On the other hand the struggle will often be severe between beings remote in the scale of nature. The slightest advantage in certain individuals, at any age or during any season, over those with which they come into competition, or better adaptation in however slight a degree to the surrounding physical conditions, will, in the long run, turn the balance.

With animals having separated sexes, there will be in most cases a struggle between the males for the possession of the females. The most vigorous males, or those which have most successfully struggled with their conditions of life, will generally leave most progeny. But success will often depend on the males having special weapons, or means of defence, or charms; add a slight advantage will lead to victory.

As geology plainly proclaims that each land has undergone great physical changes, we might have expected to find that organic beings have varied under nature, in the same way as they have varied under domestication. And if there has been any variability under nature, it would be an unaccountable fact if natural selection had not come into play. It has often been asserted, but the assertion is incapable of proof, that the amount of variation under nature is a strictly limited quantity. Man, though acting on external characters alone and often capriciously, can produce within a short period a great result by adding up mere individual differences in his domestic productions; and every one admits that species present individual differences. But, besides such differences, all naturalists admit that natural varieties exist, which are considered sufficiently distinct to be worthy of record in systematic works. No one has drawn any clear distinction between individual differences and slight varieties; or between more plainly marked varieties and sub-species, and species. On separate continents, and on different parts of the same continent when divided by barriers of any kind, and on outlying islands, what a multitude of forms exist, which some experienced naturalists rank as varieties, others as geographical races or sub-species, and others as distinct, though closely allied species!

If, then, animals and plants do vary, let it be ever so slightly or slowly, why should not variations or individual differences, which are in any way beneficial, be preserved and accumulated through natural selection, or the survival of the fittest? If man can by patience select variations useful to him, why, under changing and complex conditions of life, should not variations useful to nature's living products often arise, and be preserved or selected? What limit can be put to this power, acting during long ages and rigidly scrutinising the whole constitution, structure, and habits of each creature,- favouring the good and rejecting the bad? I can see no limit to this power, in slowly and beautifully adapting each form to the most complex relations of life. The theory of natural selection, even if we look no farther than this, seems to be in the highest degree probable. I have already recapitulated, as fairly as I could, the opposed difficulties and objections; now let us turn to the special facts and arguments in favour of the theory.

On the view that species are only strongly marked and permanent varieties, and that each species first existed as a variety, we can see why it is that no line of demarcation can be drawn between species, commonly supposed to have been produced by special acts of creation, and varieties which are acknowledged to have been produced by secondary laws. On this same view we can understand how it is that in a region where many species of a genus have been produced, and where they now flourish, these same species should present many varieties; for where the manufactory of species has been active, we might expect, as a general rule, to find it still in action; and this is the case if varieties be incipient species. Moreover, the species of the larger genera, which afford the greater number of varieties or incipient species, retain to a certain degree the character of varieties; for they differ from each other by a less amount of difference than do the species of smaller genera. The closely allied species also of the larger genera apparently have restricted ranges, and in their affinities they are clustered in little groups round other species- in both respects resembling varieties. These are strange relations on the view that each species was independently created, but are intelligible if each existed first as a variety.

As each species tends by its geometrical rate of reproduction to increase inordinately in number; and as the modified descendants of each species will be enabled to increase by as much as they become more diversified in habits and structure, so as to be able to seize on many and widely different places in the economy of nature, there will be a constant tendency in natural selection to preserve the most divergent offspring of any one species. Hence, during a long-continued course of modification, the slight differences, characteristic of varieties of the same species, tend to be augmented into the greater differences characteristic of the species of the same genus. New and improved varieties will inevitably supplant and exterminate the older, less improved, and intermediate varieties; and thus species are rendered to a large extent defined and distinct objects. Dominant species belonging to the larger groups within each class tend to give birth to new and dominant forms; so that each large group tends to become still larger, and at the same time more divergent in character. But as all groups cannot thus go on increasing in size, for the world would not hold them, the more dominant groups beat the less dominant. This tendency in the large groups to go on increasing in size and diverging in character, together with the inevitable contingency of much extinction, explains the arrangement of all the forms of life in groups subordinate to groups, all within a few great classes, which has prevailed throughout all time. This grand fact of the grouping of all organic beings under what is called the Natural System, is utterly inexplicable on the theory of creation.

As natural selection acts solely by accumulating slight, successive, favourable variations, it can produce no great or sudden modifications; it can act only by short and slow steps. Hence, the canon of "Natura non facit saltum," which every fresh addition to our knowledge tends to confirm, is on this theory intelligible. We can see why throughout nature the same general end is gained by an almost infinite diversity of means, for every peculiarity when once acquired is long inherited, and structures already modified in many different ways have to be adapted for the same general purpose. We can, in short, see why nature is prodigal in variety, though niggard in innovation. But why this should be a law of nature if each species has been independently created no man can explain.

Many other facts are, as it seems to me, explicable on this theory. How strange it is that a bird, under the form of a woodpecker, should prey on insects on the ground; that upland geese which rarely or never swim, should possess webbed feet; that a thrush-like bird should dive and feed on sub-aquatic insects; and that a petrel should have the habits and structure fitting it for the life of an awk! and so in endless other cases. But on the view of each species constantly trying to increase in number, with natural selection always ready to adapt the slowly varying descendants of each to any unoccupied or ill-occupied place in nature, these facts cease to be strange, or might even have been anticipated.

We can to a certain extent understand how it is that there is so much beauty throughout nature; for this may be largely attributed to the agency of selection. That beauty, according to our sense of it, is not universal, must be admitted by every one who will look at some venomous snakes, at some fishes, and at certain hideous bats with a distorted resemblance to the human face. Sexual selection has given the most brilliant colours, elegant patterns, and other ornaments to the males, and sometimes to both sexes of many birds, butterflies, and other animals. With birds it has often rendered the voice of the male musical to the female, as well as to our ears. Flowers and fruit have been rendered conspicuous by brilliant colours in contrast with the green foliage, in order that the flowers may be readily seen, visited and fertilised by insects, and the seeds disseminated by birds. How it comes that certain colours, sounds, and forms should give pleasure to man and the lower animals,- that is, that is, how the sense of beauty in its simplest form was first acquired,- we do not know any more than how certain odours and flavours were first rendered agreeable.

As natural selection acts by competition, it adapts and improves the inhabitants of each country only in relation to their co-inhabitants; so that we need feel no surprise at the species of any one country, although on the ordinary view supposed to have been created and specially adapted for that country, being beaten and supplanted by the naturalised productions from another land. Nor ought we to marvel if all the contrivances in nature be not, as far. as we can judge, absolutely perfect, as in the case even of the human eye; or if some of them be abhorrent to our ideas of fitness. We need not marvel at the sting of the bee, when used against an enemy, causing the bee's own death; at drones being produced in such great 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 the Ichneumonidae feeding within the living bodies of caterpillars; or 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 detected.

The complex and little known laws governing the production of varieties are the same, as far as we can judge, with the laws which have governed the production of distinct species. In both cases physical conditions seem to have produced some direct and definite effect, but how much we cannot say. Thus, when varieties enter any new station, they occasionally assume some of the characters proper to the species of that station. With both varieties and species, use and disuse seem to have produced a considerable effect; for it is impossible to resist this conclusion 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 tucu-tucu, 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. With varieties and species, correlated variation seems to have played an important part, so that when one part has been modified other parts have been necessarily modified. With both parties and.species, reversions to long-lost characters occasionally occur. How inexplicable on the theory of creation is the occasional appearance of stripes on the shoulders and legs of the several species of the horse-genus and of their hybrids! How simply is this fact explained if we believe that these species are all descended from a striped progenitor, in the same manner as the several domestic breeds of the pigeon are descended from the blue and barred rock-pigeon!

On the ordinary view of each species having been independently created, why should specific characters, or those by which the species of the same genus differ from each other, be more variable than 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 possess differently coloured flowers, than if all possessed 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; therefore these same characters would be more likely again to vary than the generic characters which have been inherited without change for an immense period. It is inexplicable on the theory of creation why a part developed in a very unusual manner in species alone of a genus, and therefore, as we may naturally infer, of great importance to that species, should be eminently liable to variation; but, on our 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 the 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 do corporeal structures 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 different animals of the same class with their several instincts. I have attempted to show how much light the principle of gradation throws on the admirable architectural powers of the hive-bee. Habit no doubt often 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 widely different conditions of life, yet follow nearly the same instincts; why the thrushes of tropical and temperate South America, for instance, line their nests 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 being not perfect and liable to mistakes, and at many instincts causing other animals to suffer.

If species be only well-marked and permanent varieties, 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 absorbed into each other by successive crosses, and in other such points,- as do the crossed offspring of acknowledged varieties. This similarity would be a strange fact, if species had been independently created and varieties had been produced through secondary laws.

If we admit that the geological record is imperfect to an extreme degree, then the facts, which the record does give, strongly 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 from the principle of natural selection; for old forms are supplanted by new and improved forms. Neither single species nor groups of species reappear when the chain of ordinary generation is once 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 beings can be classed with all recent beings, naturally follows from the living and the extinct being the offspring of common parents. As species have generally diverged in character during their long course of descent and modification, we can understand why it is that the more ancient forms, or early progenitors of each group, so often occupy a position in some degree intermediate between existing groups. Recent forms are generally looked upon as being, on the whole, higher in the scale of organisation than ancient forms; and they must be higher, insofar as the later and more improved forms have conquered the older and less improved forms in the struggle for life; they have also generally had their organs more specialised for different functions. This fact is perfectly compatible with numerous beings still retaining simple and but little improved structures, fitted for simple conditions of life; it is likewise compatible with some forms having retrograded in organisation, by having become at each stage of descent better fitted for new and degraded habits of life. Lastly, the wonderful 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 the same country the existing and the extinct will be closely 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 Distribution. We can see why there should be so striking a parallelism in the distribution of organic beings throughout 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 has 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 are the 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 mountains, and in the northern and southern temperate zones; and likewise the close alliance of some of the inhabitants of the sea in the northern and southern temperate latitudes, though separated by the whole intertropical ocean. Although two countries may present physical conditions as closely similar as the same species ever require, we need feel no surprise at their inhabitants being widely different, if they have been for a long period completely sundered from each other; for as the relation of organism to organism is the most important of all relations, and as the two countries will have received colonists at various periods and in different proportions, from some other country or from each other, the course of modification in the two areas will inevitably have been different.

On this view of migration, with subsequent modification, we see why oceanic islands are inhabited by only few species, but of these, why many are peculiar or endemic forms. We clearly see why species belonging to those groups of animals which cannot cross wide spaces of the ocean, as frogs and terrestrial mammals, do not inhabit oceanic islands; and why, on the other hand, new and peculiar species of bats, animals which can traverse the ocean, are found on islands far distant from any continent. Such cases as the presence of peculiar species of bats on oceanic islands and the absence of all other terrestrial mammals, are facts utterly inexplicable on the theory of independent acts of creation.

The existence of closely allied or representative species in any two areas, implies, on the theory of descent with modification, that the same parent-forms formerly inhabited both areas; and we almost invariably find that wherever many closely allied species inhabit two areas, some identical species are still common to both. Wherever many closely allied yet distinct species occur, doubtful forms and varieties belonging to the same groups 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 the striking relation of nearly all plants and animals of the Galapagos Archipelago, of Juan Fernandez, and of the other American islands, to the plants and animals of the neighbouring American mainland; and of those of the Cape de Verde Archipelago, and of the 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 can be arranged within a few great classes, in groups subordinate to groups, and with the extinct groups often falling in between the 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 forms 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 beings, are of hardly any importance in classification; why characters derived from rudimentary parts, though of no service to the beings, are often of high classificatory value; and why embryological characters are often the most valuable of all. The real affinities of all organic beings, in contradistinction to their adaptive resemblances, are due to inheritance or community of descent. The Natural System is a genealogical arrangement, with the acquired grades of difference, marked by the terms, varieties, species, genera, families, &c.; and we have to discover the lines of descent by the most permanent characters whatever they may be and of however slight vital importance.

The similar framework of bones in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse,- the same number of vertebrae forming the neck of the giraffe and of the elephant,- and innumerable 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 in the 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, to a large extent, intelligible on the view of the gradual modification of parts or organs, which were aboriginally alike in an early progenitor in each of these classes. 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 clearly see why the embryos of mammals, birds, reptiles, and fishes should be so closely similar, and 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 of a fish which has to breathe the air dissolved in water by the aid of well-developed branchiae.

Disuse, aided sometimes by natural selection, will often have reduced organs when rendered useless under changed habits or conditions of life; and we can understand on this view the meaning of rudimentary organs. But disuse and selection 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 on an organ during early life; hence the organ will not be 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 formerly reduced by disuse, owing to the tongue and palate, or lips, having become excellently fitted through natural selection to browse without their aid; whereas in the calf, the teeth have been left unaffected, 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 organism with all its separate parts having been specially created, how utterly inexplicable is it that organs bearing the plain stamp of inutility, such as the teeth in the embryonic calf or the shrivelled wings under the soldered wingcovers of many beetles, should so frequently occur. Nature may be said to have taken pains to reveal her scheme of modification, by means of rudimentary organs, of embryological and homologous structures, but we are too blind to understand her meaning.

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.

THE END

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