On the Origin of Species Part 17

In certain cases the successive steps of variation might supervene, from causes of which we are wholly ignorant, at a very early period of life, or each step might be inherited at an earlier period than that at which it first appeared. In either case (as with the short-faced tumbler) the young or embryo would closely resemble the mature parent-form. We have seen that this is the rule of development in certain whole groups of animals, as with cuttle-fish and spiders, and with a few members of the great cla.s.s of insects, as with Aphis. With respect to the final cause of the young in these cases not undergoing any metamorphosis, or closely resembling their parents from their earliest age, we can see that this would result from the two following contingencies; firstly, from the young, during a course of modification carried on for many generations, having to provide for their own wants at a very early stage of development, and secondly, from their following exactly the same habits of life with their parents; for in this case, it would be indispensable for the existence of the species, that the child should be modified at a very early age in the same manner with its parents, in accordance with their similar habits. Some further explanation, however, of the embryo not undergoing any metamorphosis is perhaps requisite. If, on the other hand, it profited the young to follow habits of life in any degree different from those of their parent, and consequently to be constructed in a slightly different manner, then, on the principle of inheritance at corresponding ages, the active young or larvae might easily be rendered by natural selection different to any conceivable extent from their parents. Such differences might, also, become correlated with successive stages of development; so that the larvae, in the first stage, might differ greatly from the larvae in the second stage, as we have seen to be the case with cirripedes. The adult might become fitted for sites or habits, in which organs of locomotion or of the senses, etc., would be useless; and in this case the final metamorphosis would be said to be retrograde.

As all the organic beings, extinct and recent, which have ever lived on this earth have to be cla.s.sed together, and as all have been connected by the finest gradations, the best, or indeed, if our collections were nearly perfect, the only possible arrangement, would be genealogical.

Descent being on my view the hidden bond of connexion which naturalists have been seeking under the term of the natural system. On this view we can understand how it is that, in the eyes of most naturalists, the structure of the embryo is even more important for cla.s.sification than that of the adult. For the embryo is the animal in its less modified state; and in so far it reveals the structure of its progenitor. In two groups of animal, however much they may at present differ from each other in structure and habits, if they pa.s.s through the same or similar embryonic stages, we may feel a.s.sured that they have both descended from the same or nearly similar parents, and are therefore in that degree closely related. Thus, community in embryonic structure reveals community of descent. It will reveal this community of descent, however much the structure of the adult may have been modified and obscured; we have seen, for instance, that cirripedes can at once be recognised by their larvae as belonging to the great cla.s.s of crustaceans. As the embryonic state of each species and group of species partially shows us the structure of their less modified ancient progenitors, we can clearly see why ancient and extinct forms of life should resemble the embryos of their descendants,--our existing species. Aga.s.siz believes this to be a law of nature; but I am bound to confess that I only hope to see the law hereafter proved true. It can be proved true in those cases alone in which the ancient state, now supposed to be represented in many embryos, has not been obliterated, either by the successive variations in a long course of modification having supervened at a very early age, or by the variations having been inherited at an earlier period than that at which they first appeared. It should also be borne in mind, that the supposed law of resemblance of ancient forms of life to the embryonic stages of recent forms, may be true, but yet, owing to the geological record not extending far enough back in time, may remain for a long period, or for ever, incapable of demonstration.

Thus, as it seems to me, the leading facts in embryology, which are second in importance to none in natural history, are explained on the principle of slight modifications not appearing, in the many descendants from some one ancient progenitor, at a very early period in the life of each, though perhaps caused at the earliest, and being inherited at a corresponding not early period. Embryology rises greatly in interest, when we thus look at the embryo as a picture, more or less obscured, of the common parent-form of each great cla.s.s of animals.

RUDIMENTARY, ATROPHIED, OR ABORTED ORGANS.

Organs or parts in this strange condition, bearing the stamp of inutility, are extremely common throughout nature. For instance, rudimentary mammae are very general in the males of mammals: I presume that the "b.a.s.t.a.r.d-wing" in birds may be safely considered as a digit in a rudimentary state: in very many snakes one lobe of the lungs is rudimentary; in other snakes there are rudiments of the pelvis and hind limbs. Some of the cases of rudimentary organs are extremely curious; for instance, the presence of teeth in foetal whales, which when grown up have not a tooth in their heads; and the presence of teeth, which never cut through the gums, in the upper jaws of our unborn calves. It has even been stated on good authority that rudiments of teeth can be detected in the beaks of certain embryonic birds. Nothing can be plainer than that wings are formed for flight, yet in how many insects do we see wings so reduced in size as to be utterly incapable of flight, and not rarely lying under wing-cases, firmly soldered together!

The meaning of rudimentary organs is often quite unmistakeable: for instance there are beetles of the same genus (and even of the same species) resembling each other most closely in all respects, one of which will have full-sized wings, and another mere rudiments of membrane; and here it is impossible to doubt, that the rudiments represent wings. Rudimentary organs sometimes retain their potentiality, and are merely not developed: this seems to be the case with the mammae of male mammals, for many instances are on record of these organs having become well developed in full-grown males, and having secreted milk. So again there are normally four developed and two rudimentary teats in the udders of the genus Bos, but in our domestic cows the two sometimes become developed and give milk. In individual plants of the same species the petals sometimes occur as mere rudiments, and sometimes in a well-developed state. In plants with separated s.e.xes, the male flowers often have a rudiment of a pistil; and Kolreuter found that by crossing such male plants with an hermaphrodite species, the rudiment of the pistil in the hybrid offspring was much increased in size; and this shows that the rudiment and the perfect pistil are essentially alike in nature.

An organ serving for two purposes, may become rudimentary or utterly aborted for one, even the more important purpose; and remain perfectly efficient for the other. Thus in plants, the office of the pistil is to allow the pollen-tubes to reach the ovules protected in the ovarium at its base. The pistil consists of a stigma supported on the style; but in some Compositae, the male florets, which of course cannot be fecundated, have a pistil, which is in a rudimentary state, for it is not crowned with a stigma; but the style remains well developed, and is clothed with hairs as in other compositae, for the purpose of brus.h.i.+ng the pollen out of the surrounding anthers. Again, an organ may become rudimentary for its proper purpose, and be used for a distinct object: in certain fish the swim-bladder seems to be rudimentary for its proper function of giving buoyancy, but has become converted into a nascent breathing organ or lung. Other similar instances could be given.

Rudimentary organs in the individuals of the same species are very liable to vary in degree of development and in other respects. Moreover, in closely allied species, the degree to which the same organ has been rendered rudimentary occasionally differs much. This latter fact is well exemplified in the state of the wings of the female moths in certain groups. Rudimentary organs may be utterly aborted; and this implies, that we find in an animal or plant no trace of an organ, which a.n.a.logy would lead us to expect to find, and which is occasionally found in monstrous individuals of the species. Thus in the snapdragon (antirrhinum) we generally do not find a rudiment of a fifth stamen; but this may sometimes be seen. In tracing the h.o.m.ologies of the same part in different members of a cla.s.s, nothing is more common, or more necessary, than the use and discovery of rudiments. This is well shown in the drawings given by Owen of the bones of the leg of the horse, ox, and rhinoceros.

It is an important fact that rudimentary organs, such as teeth in the upper jaws of whales and ruminants, can often be detected in the embryo, but afterwards wholly disappear. It is also, I believe, a universal rule, that a rudimentary part or organ is of greater size relatively to the adjoining parts in the embryo, than in the adult; so that the organ at this early age is less rudimentary, or even cannot be said to be in any degree rudimentary. Hence, also, a rudimentary organ in the adult, is often said to have retained its embryonic condition.

I have now given the leading facts with respect to rudimentary organs.

In reflecting on them, every one must be struck with astonishment: for the same reasoning power which tells us plainly that most parts and organs are exquisitely adapted for certain purposes, tells us with equal plainness that these rudimentary or atrophied organs, are imperfect and useless. In works on natural history rudimentary organs are generally said to have been created "for the sake of symmetry," or in order "to complete the scheme of nature;" but this seems to me no explanation, merely a restatement of the fact. Would it be thought sufficient to say that because planets revolve in elliptic courses round the sun, satellites follow the same course round the planets, for the sake of symmetry, and to complete the scheme of nature? An eminent physiologist accounts for the presence of rudimentary organs, by supposing that they serve to excrete matter in excess, or injurious to the system; but can we suppose that the minute papilla, which often represents the pistil in male flowers, and which is formed merely of cellular tissue, can thus act? Can we suppose that the formation of rudimentary teeth which are subsequently absorbed, can be of any service to the rapidly growing embryonic calf by the excretion of precious phosphate of lime? When a man's fingers have been amputated, imperfect nails sometimes appear on the stumps: I could as soon believe that these vestiges of nails have appeared, not from unknown laws of growth, but in order to excrete h.o.r.n.y matter, as that the rudimentary nails on the fin of the manatee were formed for this purpose.

On my view of descent with modification, the origin of rudimentary organs is simple. We have plenty of cases of rudimentary organs in our domestic productions,--as the stump of a tail in tailless breeds,--the vestige of an ear in earless breeds,--the reappearance of minute dangling horns in hornless breeds of cattle, more especially, according to Youatt, in young animals,--and the state of the whole flower in the cauliflower. We often see rudiments of various parts in monsters. But I doubt whether any of these cases throw light on the origin of rudimentary organs in a state of nature, further than by showing that rudiments can be produced; for I doubt whether species under nature ever undergo abrupt changes. I believe that disuse has been the main agency; that it has led in successive generations to the gradual reduction of various organs, until they have become rudimentary,--as in the case of the eyes of animals inhabiting dark caverns, and of the wings of birds inhabiting oceanic islands, which have seldom been forced to take flight, and have ultimately lost the power of flying. Again, an organ useful under certain conditions, might become injurious under others, as with the wings of beetles living on small and exposed islands; and in this case natural selection would continue slowly to reduce the organ, until it was rendered harmless and rudimentary.

Any change in function, which can be effected by insensibly small steps, is within the power of natural selection; so that an organ rendered, during changed habits of life, useless or injurious for one purpose, might easily be modified and used for another purpose. Or an organ might be retained for one alone of its former functions. An organ, when rendered useless, may well be variable, for its variations cannot be checked by natural selection. At whatever period of life disuse or selection reduces an organ, and this will generally be when the being has come to maturity and to its full powers of action, the principle of inheritance at corresponding ages will reproduce the organ in its reduced state at the same age, and consequently will seldom affect or reduce it in the embryo. Thus we can understand the greater relative size of rudimentary organs in the embryo, and their lesser relative size in the adult. But if each step of the process of reduction were to be inherited, not at the corresponding age, but at an extremely early period of life (as we have good reason to believe to be possible) the rudimentary part would tend to be wholly lost, and we should have a case of complete abortion. The principle, also, of economy, explained in a former chapter, by which the materials forming any part or structure, if not useful to the possessor, will be saved as far as is possible, will probably often come into play; and this will tend to cause the entire obliteration of a rudimentary organ.

As the presence of rudimentary organs is thus due to the tendency in every part of the organisation, which has long existed, to be inherited--we can understand, on the genealogical view of cla.s.sification, how it is that systematists have found rudimentary parts as useful as, or even sometimes more useful than, parts of high physiological importance. Rudimentary organs may be compared with the letters in a word, still retained in the spelling, but become useless in the p.r.o.nunciation, but which serve as a clue in seeking for its derivation. On the view of descent with modification, we may conclude that the existence of organs in a rudimentary, imperfect, and useless condition, or quite aborted, far from presenting a strange difficulty, as they a.s.suredly do on the ordinary doctrine of creation, might even have been antic.i.p.ated, and can be accounted for by the laws of inheritance.

SUMMARY.

In this chapter I have attempted to show, that the subordination of group to group in all organisms throughout all time; that the nature of the relations.h.i.+p, by which all living and extinct beings are united by complex, radiating, and circuitous lines of affinities into one grand system; the rules followed and the difficulties encountered by naturalists in their cla.s.sifications; the value set upon characters, if constant and prevalent, whether of high vital importance, or of the most trifling importance, or, as in rudimentary organs, of no importance; the wide opposition in value between a.n.a.logical or adaptive characters, and characters of true affinity; and other such rules;--all naturally follow on the view of the common parentage of those forms which are considered by naturalists as allied, together with their modification through natural selection, with its contingencies of extinction and divergence of character. In considering this view of cla.s.sification, it should be borne in mind that the element of descent has been universally used in ranking together the s.e.xes, ages, and acknowledged varieties of the same species, however different they may be in structure. If we extend the use of this element of descent,--the only certainly known cause of similarity in organic beings,--we shall understand what is meant by the natural system: it is genealogical in its attempted arrangement, with the grades of acquired difference marked by the terms varieties, species, genera, families, orders, and cla.s.ses.

On this same view of descent with modification, all the great facts in Morphology become intelligible,--whether we look to the same pattern displayed in the h.o.m.ologous organs, to whatever purpose applied, of the different species of a cla.s.s; or to the h.o.m.ologous parts constructed on the same pattern in each individual animal and plant.

On the principle of successive slight variations, not necessarily or generally supervening at a very early period of life, and being inherited at a corresponding period, we can understand the great leading facts in Embryology; namely, the resemblance in an individual embryo of the h.o.m.ologous parts, which when matured will become widely different from each other in structure and function; and the resemblance in different species of a cla.s.s of the h.o.m.ologous parts or organs, though fitted in the adult members for purposes as different as possible.

Larvae are active embryos, which have become specially modified in relation to their habits of life, through the principle of modifications being inherited at corresponding ages. On this same principle--and bearing in mind, that when organs are reduced in size, either from disuse or selection, it will generally be at that period of life when the being has to provide for its own wants, and bearing in mind how strong is the principle of inheritance--the occurrence of rudimentary organs and their final abortion, present to us no inexplicable difficulties; on the contrary, their presence might have been even antic.i.p.ated. The importance of embryological characters and of rudimentary organs in cla.s.sification is intelligible, on the view that an arrangement is only so far natural as it is genealogical.

Finally, the several cla.s.ses of facts which have been considered in this chapter, seem to me to proclaim so plainly, that the innumerable species, genera, and families of organic beings, with which this world is peopled, have all descended, each within its own cla.s.s or group, from common parents, and have all been modified in the course of descent, that I should without hesitation adopt this view, even if it were unsupported by other facts or arguments.

14. RECAPITULATION AND CONCLUSION.

Recapitulation of the difficulties on the theory of Natural Selection.

Recapitulation of the general and special circ.u.mstances 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 grave objections may be advanced against the theory of descent with modification through 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 should have been perfected, not by means superior to, though a.n.a.logous with, human reason, but by the acc.u.mulation 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 gradations in the perfection of any organ or instinct, which we may consider, either do now exist or could have existed, each good of its kind,--that all organs and instincts are, in ever so slight a degree, variable,--and, lastly, that there is a struggle for existence leading to the preservation of each profitable deviation of structure or instinct. 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; but we see so many strange gradations in nature, as is proclaimed by the canon, "Natura non facit saltum," that we ought to be extremely cautious in saying that any organ or instinct, or any whole being, could not have arrived at its present state by many graduated steps. There are, it must be admitted, cases of special difficulty on the theory of natural selection; and one of the most curious of these is the existence of two or three defined castes of workers or sterile females in the same community of ants; but I have attempted to show how this difficulty 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 eighth chapter, which seem to me conclusively to show that this sterility is no more a special endowment than is the incapacity of two trees to be grafted together, but that it is incidental on const.i.tutional differences in the reproductive systems of the intercrossed species. We see the truth of this conclusion in the vast difference in the result, when the same two species are crossed reciprocally; that is, when one species is first used as the father and then as the mother.

The fertility of varieties when intercrossed and of their mongrel offspring cannot be considered as universal; nor is their very general fertility surprising when we remember that it is not likely that either their const.i.tutions or their reproductive systems should have been profoundly modified. Moreover, most of the varieties which have been experimentised on have been produced under domestication; and as domestication apparently tends to eliminate sterility, we ought not to expect it also to produce sterility.

The sterility of hybrids is a very different case from that of first crosses, for their reproductive organs are more or less functionally impotent; whereas in first crosses the organs on both sides are in a perfect condition. As we continually see that organisms of all kinds are rendered in some degree sterile from their const.i.tutions having been disturbed by slightly different and new conditions of life, we need not feel surprise at hybrids being in some degree sterile, for their const.i.tutions can hardly fail to have been disturbed from being compounded of two distinct organisations. This parallelism is supported by another parallel, but directly opposite, cla.s.s of facts; namely, that the vigour and fertility of all organic beings are increased by slight changes in their conditions of life, and that the offspring of slightly modified forms or varieties acquire from being crossed increased vigour and fertility. So that, on the one hand, considerable changes in the conditions of life and crosses between greatly modified forms, lessen fertility; and on the other hand, lesser changes in the conditions of life and crosses between less modified forms, increase fertility.

Turning to geographical distribution, the difficulties encountered on the theory of descent with modification are grave enough. All the individuals of the same species, and all the species of the same genus, or even higher group, must have descended from common parents; and therefore, in however distant and isolated parts of the world they are now found, they must in the course of successive generations have pa.s.sed from some one part to 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, enormously 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 of time there will always be 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 of the full extent of the various climatal and geographical changes which have affected the earth during modern periods; and such changes will obviously have greatly facilitated migration. As an example, I have attempted to show how potent has been the influence of the Glacial period on the distribution both of the same and of representative 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 very 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 species of the same genus is in some degree lessened.

As on 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 present 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 climate and other conditions of life change insensibly in going 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 zone.

For we have reason to believe that only a few species are undergoing change at any one period; and all changes are slowly effected. I have also shown that the intermediate varieties which will at first probably exist in the intermediate zones, will be liable to be supplanted by the allied forms on either hand; and the latter, from existing in greater numbers, will generally be modified and improved at a quicker rate than the intermediate varieties, which exist in lesser numbers; so that the intermediate varieties will, 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? We meet with no such evidence, and this is the most obvious and forcible of the many objections which may be urged against my theory. Why, again, do whole groups of allied species appear, though certainly they often falsely appear, to have come in suddenly on the several geological stages? Why do we not find great piles of strata beneath the Silurian system, stored with the remains of the progenitors of the Silurian groups of fossils?

For certainly on my theory such strata must somewhere have been deposited at these ancient and utterly unknown epochs in the world's history.

I can answer these questions and grave objections only on the supposition that the geological record is far more imperfect than most geologists believe. It cannot be objected that there has not been time sufficient for any amount of organic change; for the lapse of time has been so great as to be utterly inappreciable by the human intellect. The number of specimens in all our museums is absolutely as nothing compared with the countless generations of countless species which certainly have existed. We should not be able to recognise a species as the parent of any one or more species if we were to examine them ever so closely, unless we likewise possessed many of the intermediate links between their past or parent and present states; and these many links we could hardly ever expect to discover, owing to the imperfection of the geological record. 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, on the common view, whether or not these doubtful forms are varieties? As long as most of the links between any two species are unknown, if any one link or intermediate variety be discovered, it will simply be cla.s.sed as another and distinct species. Only a small portion of the world has been geologically explored. Only organic beings of certain cla.s.ses can be preserved in a fossil condition, at least in any great number. Widely ranging species vary most, and varieties are often at first local,--both causes rendering the discovery of intermediate links less likely. Local varieties will not spread into other and distant regions until they are considerably modified and improved; and when they do spread, if discovered in a geological formation, they will appear as if suddenly created there, and will be simply cla.s.sed as new species. Most formations have been intermittent in their acc.u.mulation; and their duration, I am inclined to believe, has been shorter than the average duration of specific forms. Successive formations are separated from each other by enormous blank intervals of time; for fossiliferous formations, thick enough to resist future degradation, can be acc.u.mulated 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 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 fossiliferous formations beneath the lowest Silurian strata, I can only recur to the hypothesis given in the ninth chapter. That the geological record is imperfect all will admit; but that it is imperfect to the degree which I require, few will be inclined to admit. If we look to long enough intervals of time, geology plainly declares that all species have changed; and they have changed in the manner which my theory requires, 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 formations distant from each other in time.

Such is the sum of the several chief objections and difficulties which may justly be urged against my theory; and I have now briefly recapitulated the answers and explanations which can be given to them. 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 the Geological Record is. Grave as these several difficulties are, in my judgment they do not overthrow the theory of descent with modification.

Now let us turn to the other side of the argument. Under domestication we see much variability. This seems to be mainly due to the reproductive system being eminently susceptible to changes in the conditions of life; so that this system, when not rendered impotent, fails to reproduce offspring exactly like the parent-form. Variability is governed by many complex laws,--by correlation of growth, by use and disuse, and by the direct action of the physical conditions of life. There is much difficulty in ascertaining how much modification our domestic productions have undergone; 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 wholly cease; for new varieties are still occasionally produced by our most anciently domesticated productions.

Man does not actually produce variability; he only unintentionally exposes organic beings to new conditions of life, and then nature acts on the organisation, and causes variability. But man can and does select the variations given to him by nature, and thus acc.u.mulate 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 to him at the time, without any thought 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 quite inappreciable by an uneducated eye. This process of selection has been the great agency in the production of the most distinct and useful domestic breeds. That many of the breeds produced by man have to a large extent the character of natural species, is shown by the inextricable doubts whether very many of them are varieties or aboriginal species.

There is no obvious reason why the principles which have acted so efficiently under domestication should not have acted under nature.

In the preservation of favoured individuals and races, during the constantly-recurrent Struggle for Existence, we see the most powerful and ever-acting means 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 effects of a succession of peculiar seasons, and by the results of naturalisation, as explained in the third chapter. More individuals are born than can possibly survive. A grain in the balance will determine which individual 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 compet.i.tion 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. But the struggle will often be very severe between beings most remote in the scale of nature.

The slightest advantage in one being, at any age or during any season, over those with which it comes into compet.i.tion, or better adaptation in however slight a degree to the surrounding physical conditions, will turn the balance.

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

As geology plainly proclaims that each land has undergone great physical changes, we might have expected that organic beings would have varied under nature, in the same way as they generally have varied under the changed conditions of domestication. And if there be any variability under nature, it would be an unaccountable fact if natural selection had not come into play. It has often been a.s.serted, but the a.s.sertion is quite incapable of proof, that the amount of variation under nature is a strictly limited quant.i.ty. 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 there are at least individual differences in species under nature. But, besides such differences, all naturalists have admitted the existence of varieties, which they think sufficiently distinct to be worthy of record in systematic works. No one can draw any clear distinction between individual differences and slight varieties; or between more plainly marked varieties and sub-species, and species. Let it be observed how naturalists differ in the rank which they a.s.sign to the many representative forms in Europe and North America.

If then we have under nature variability and a powerful agent always ready to act and select, why should we doubt that variations in any way useful to beings, under their excessively complex relations of life, would be preserved, acc.u.mulated, and inherited? Why, if man can by patience select variations most useful to himself, should nature fail in selecting variations useful, under changing conditions of life, to her living products? What limit can be put to this power, acting during long ages and rigidly scrutinising the whole const.i.tution, 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 looked no further than this, seems to me to be in itself 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 each 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 they are cl.u.s.tered in little groups round other species--in which respects they resemble varieties. These are strange relations on the view of each species having been independently created, but are intelligible if all species first existed as varieties.

As each species tends by its geometrical ratio of reproduction to increase inordinately in number; and as the modified descendants of each species will be enabled to increase by so much the more as they become more diversified in habits and structure, so as to be enabled 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 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 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 succeed in 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 almost inevitable contingency of much extinction, explains the arrangement of all the forms of life, in groups subordinate to groups, all within a few great cla.s.ses, which we now see everywhere around us, and which has prevailed throughout all time. This grand fact of the grouping of all organic beings seems to me utterly inexplicable on the theory of creation.

As natural selection acts solely by acc.u.mulating slight, successive, favourable variations, it can produce no great or sudden modification; it can act only by very short and slow steps. Hence the canon of "Natura non facit saltum," which every fresh addition to our knowledge tends to make more strictly correct, is on this theory simply intelligible. We can plainly see why nature is prodigal in variety, though n.i.g.g.ard 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 woodp.e.c.k.e.r, should have been created to prey on insects on the ground; that upland geese, which never or rarely swim, should have been created with webbed feet; that a thrush should have been created to dive and feed on sub-aquatic insects; and that a petrel should have been created with habits and structure fitting it for the life of an auk or grebe! and so on 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 perhaps might even have been antic.i.p.ated.

As natural selection acts by compet.i.tion, it adapts the inhabitants of each country only in relation to the degree of perfection of their a.s.sociates; so that we need feel no surprise at the inhabitants of any one country, although on the ordinary view supposed to have been specially created and adapted for that country, being beaten and supplanted by the naturalised 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; and if some of them be abhorrent to our ideas of fitness. We need not marvel at the sting of the bee causing the bee's own death; at drones being produced in such vast numbers for one single act, and being then slaughtered by their sterile sisters; at the astonis.h.i.+ng 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 live bodies of caterpillars; and at other such cases. The wonder indeed is, on the theory of natural selection, that more cases of the want of absolute perfection have not been observed.

The complex and little known laws governing variation are the same, as far as we can see, with the laws which have governed the production of so-called specific forms. In both cases physical conditions seem to have produced but little direct effect; yet when varieties enter any zone, they occasionally a.s.sume some of the characters of the species proper to that zone. In both varieties and species, use and disuse seem to have produced some effect; for it is difficult 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 tucutucu, which is occasionally blind, and then at certain moles, which are habitually blind and have their eyes covered with skin; or when we look at the blind animals inhabiting the dark caves of America and Europe. In both varieties and species correlation of growth seems to have played a most important part, so that when one part has been modified other parts are necessarily modified. In both varieties and species reversions to long-lost characters occur. How inexplicable on the theory of creation is the occasional appearance of stripes on the shoulder and legs of the several species of the horse-genus and in their hybrids! How simply is this fact explained if we believe that these species have descended from a striped progenitor, in the same manner as the several domestic breeds of pigeon have descended from the blue and barred rock-pigeon!