On the Origin of Species by Charles Darwin - HTML preview

Recapitulation And Conclusion]

Recapitulation of the objections to the theory of Natural Selection -- Recapitulation of the general and special circumstances in its favour -- Causes of the general belief in the immutability of species -- How far the theory of Natural Selection may be extended -Effects of its adoption on the study of Natural History -- Concluding remarks.

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 among 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 labourious 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, an 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, while 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 climatical 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 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 substages, 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 retained 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 degradation 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 justly be 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 a 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; and 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 subspecies 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 further 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 a 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, would 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 auk! 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 easily 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, 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 the 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 ichneumonidae feeding within the living 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 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 loggerheaded 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 varieties 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 the g