Introduction

When on board H.M.S. Beagle, as naturalist, I was much struck with certain facts in the distribution of the organic beings inhabiting South America, and in the geological relations of the present to the past inhabitants of that continent. These facts, as will be seen in the latter chapters of this volume, seemed to throw some light on the origin of species--that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, it occurred to me, in 1837, that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it. After five years' work I allowed myself to speculate on the subject, and drew up some short notes; these I enlarged in 1844 into a sketch of the conclusions, which then seemed to me probable: from that period to the present day I have steadily pursued the same object. I hope that I may be excused for entering on these personal details, as I give them to show that I have not been hasty in coming to a decision.

My work is now (1859) nearly finished; but as it will take me many more years to complete it, and as my health is far from strong, I have been urged to publish this abstract. I have more especially been induced to do this, as Mr. Wallace, who is now studying the natural history of the Malay Archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. In 1858 he sent me a memoir on this subject, with a request that I would forward it to Sir Charles Lyell, who sent it to the Linnean Society, and it is published in the third volume of the Journal of that Society. Sir C. Lyell and Dr. Hooker, who both knew of my work--the latter having read my sketch of 1844--honoured me by thinking it advisable to publish, with Mr. Wallace's excellent memoir, some brief extracts from my manuscripts.

This abstract, which I now publish, must necessarily be imperfect. I cannot here give references and authorities for my several statements; and I must trust to the reader reposing some confidence in my accuracy. No doubt errors may have crept in, though I hope I have always been cautious in trusting to good authorities alone. I can here give only the general conclusions at which I have arrived, with a few facts in illustration, but which, I hope, in most cases will suffice. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts, with references, on which my conclusions have been grounded; and I hope in a future work to do this. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced, often apparently leading to conclusions directly opposite to those at which I have arrived. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question; and this is here impossible.

I much regret that want of space prevents my having the satisfaction of acknowledging the generous assistance which I have received from very many naturalists, some of them personally unknown to me. I cannot, however, let this opportunity pass without expressing my deep obligations to Dr. Hooker, who, for the last fifteen years, has aided me in every possible way by his large stores of knowledge and his excellent judgment.

In considering the origin of species, it is quite conceivable that a naturalist, reflecting on the mutual affinities of organic beings, on their embryological relations, their geographical distribution, geological succession, and other such facts, might come to the conclusion that species had not been independently created, but had descended, like varieties, from other species. Nevertheless, such a conclusion, even if well founded, would be unsatisfactory, until it could be shown how the innumerable species, inhabiting this world have been modified, so as to acquire that perfection of structure and coadaptation which justly excites our admiration. Naturalists continually refer to external conditions, such as climate, food, etc., as the only possible cause of variation. In one limited sense, as we shall hereafter see, this may be true; but it is preposterous to attribute to mere external conditions, the structure, for instance, of the woodpecker, with its feet, tail, beak, and tongue, so admirably adapted to catch insects under the bark of trees. In the case of the mistletoe, which draws its nourishment from certain trees, which has seeds that must be transported by certain birds, and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its relations to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the plant itself.

It is, therefore, of the highest importance to gain a clear insight into the means of modification and coadaptation. At the commencement of my observations it seemed to me probable that a careful study of domesticated animals and of cultivated plants would offer the best chance of making out this obscure problem. Nor have I been disappointed; in this and in all other perplexing cases I have invariably found that our knowledge, imperfect though it be, of variation under domestication, afforded the best and safest clue. I may venture to express my conviction of the high value of such studies, although they have been very commonly neglected by naturalists.

From these considerations, I shall devote the first chapter of this abstract to variation under domestication. We shall thus see that a large amount of hereditary modification is at least possible; and, what is equally or more important, we shall see how great is the power of man in accumulating by his selection successive slight variations. I will then pass on to the variability of species in a state of nature; but I shall, unfortunately, be compelled to treat this subject far too briefly, as it can be treated properly only by giving long catalogues of facts. We shall, however, be enabled to discuss what circumstances are most favourable to variation. In the next chapter the struggle for existence among all organic beings throughout the world, which inevitably follows from the high geometrical ratio of their increase, will be considered. This is the doctrine of Malthus, applied to the whole animal and vegetable kingdoms. As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be NATURALLY SELECTED. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form...

Chapter I.

Variation Under Domestication

Causes of Variability

When we compare the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us is, that they generally differ more from each other than do the individuals of any one species or variety in a state of nature. And if we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, we are driven to conclude that this great variability is due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent species had been exposed under nature. There is, also, some probability in the view propounded by Andrew Knight, that this variability may be partly connected with excess of food. It seems clear that organic beings must be exposed during several generations to new conditions to cause any great amount of variation; and that, when the organisation has once begun to vary, it generally continues varying for many generations. No case is on record of a variable organism ceasing to vary under cultivation. Our oldest cultivated plants, such as wheat, still yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification...

Effects of Habit and the use and disuse of Parts

Changed habits produce an inherited effect as in the period of the flowering of plants when transported from one climate to another. With animals the increased use or disuse of parts has had a more marked influence; thus I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more, in proportion to the whole skeleton, than do the same bones in the wild duck; and this change may be safely attributed to the domestic duck flying much less, and walking more, than its wild parents. The great and inherited development of the udders in cows and goats in countries where they are habitually milked, in comparison with these organs in other countries, is probably another instance of the effects of use. Not one of our domestic animals can be named which has not in some country drooping ears; and the view which has been suggested that the drooping is due to disuse of the muscles of the ear, from the animals being seldom much alarmed, seems probable...

Correlated Variation

When we look to the hereditary varieties or races of our domestic animals and plants, and compare them with closely allied species, we generally perceive in each domestic race, as already remarked, less uniformity of character than in true species. Domestic races often have a somewhat monstrous character; by which I mean, that, although differing from each other and from other species of the same genus, in several trifling respects, they often differ in an extreme degree in some one part, both when compared one with another, and more especially when compared with the species under nature to which they are nearest allied. With these exceptions (and with that of the perfect fertility of varieties when crossed—a subject hereafter to be discussed), domestic races of the same species differ from each other in the same manner as do the closely allied species of the same genus in a state of nature, but the differences in most cases are less in degree. This must be admitted as true, for the domestic races of many animals and plants have been ranked by some competent judges as the descendants of aboriginally distinct species, and by other competent judges as mere varieties. If any well marked distinction existed between a domestic race and a species, this source of doubt would not so perpetually recur. It has often been stated that domestic races do not differ from each other in characters of generic value. It can be shown that this statement is not correct; but naturalists differ much in determining what characters are of generic value; all such valuations being at present empirical. When it is explained how genera originate under nature, it will be seen that we have no right to expect often to find a generic amount of difference in our domesticated races...

Chapter II.

Variation Under Nature

Variability

Before applying the principles arrived at in the last chapter to organic beings in a state of nature, we must briefly discuss whether these latter are subject to any variation. To treat this subject properly, a long catalogue of dry facts ought to be given; but these I shall reserve for a future work. Nor shall I here discuss the various definitions which have been given of the term species. No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species. Generally the term includes the unknown element of a distinct act of creation. The term "variety" is almost equally difficult to define; but here community of descent is almost universally implied, though it can rarely be proved. We have also what are called monstrosities; but they graduate into varieties. By a monstrosity I presume is meant some considerable deviation of structure, generally injurious, or not useful to the species. Some authors use the term "variation" in a technical sense, as implying a modification directly due to the physical conditions of life; and "variations" in this sense are supposed not to be inherited; but who can say that the dwarfed condition of shells in the brackish waters of the Baltic, or dwarfed plants on Alpine summits, or the thicker fur of an animal from far northwards, would not in some cases be inherited for at least a few generations? And in this case I presume that the form would be called a variety.

It may be doubted whether sudden and considerable deviations of structure, such as we occasionally see in our domestic productions, more especially with plants, are ever permanently propagated in a state of nature. Almost every part of every organic being is so beautifully related to its complex conditions of life that it seems as improbable that any part should have been suddenly produced perfect, as that a complex machine should have been invented by man in a perfect state. Under domestication monstrosities sometimes occur which resemble normal structures in widely different animals. Thus pigs have occasionally been born with a sort of proboscis, and if any wild species of the same genus had naturally possessed a proboscis, it might have been argued that this had appeared as a monstrosity; but I have as yet failed to find, after diligent search, cases of monstrosities resembling normal structures in nearly allied forms, and these alone bear on the question. If monstrous forms of this kind ever do appear in a state of nature and are capable of reproduction (which is not always the case), as they occur rarely and singly, their preservation would depend on unusually favourable circumstances. They would, also, during the first and succeeding generations cross with the ordinary form, and thus their abnormal character would almost inevitably be lost. But I shall have to return in a future chapter to the preservation and perpetuation of single or occasional variations...

Chapter III.

Struggle for Existence

Its bearing on natural selection

Before entering on the subject of this chapter I must make a few preliminary remarks to show how the struggle for existence bears on natural selection. It has been seen in the last chapter that among organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life and of one organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.

Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow from the struggle for life. Owing to this struggle, variations, however slight and from whatever cause proceeding, if they be in any degree profitable to the individuals of a species, in their infinitely complex relations to other organic beings and to their physical conditions of life, will tend to the preservation of such individuals, and will generally be inherited by the offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term natural selection, in order to mark its relation to man's power of selection. But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man's feeble efforts, as the works of Nature are to those of Art.

We will now discuss in a little more detail the struggle for existence. In my future work this subject will be treated, as it well deserves, at greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult--at least I found it so--than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that, though food may be now superabundant, it is not so at all seasons of each recurring year...

Chapter IV.

Natural Selection; or the Survival of the Fittest

Natural Selection

How will the struggle for existence, briefly discussed in the last chapter, act in regard to variation? Can the principle of selection, which we have seen is so potent in the hands of man, apply under nature? I think we shall see that it can act most efficiently. Let the endless number of slight variations and individual differences occurring in our domestic productions, and, in a lesser degree, in those under nature, be borne in mind; as well as the strength of the hereditary tendency. Under domestication, it may truly be said that the whole organisation becomes in some degree plastic. But the variability, which we almost universally meet with in our domestic productions is not directly produced, as Hooker and Asa Gray have well remarked, by man; he can neither originate varieties nor prevent their occurrence; he can only preserve and accumulate such as do occur. Unintentionally he exposes organic beings to new and changing conditions of life, and variability ensues; but similar changes of conditions might and do occur under nature. Let it also be borne in mind how infinitely complex and close-fitting are the mutual relations of all organic beings to each other and to their physical conditions of life; and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life. Can it then be thought improbable, seeing that variations useful to man have undoubtedly occurred, that other variations useful in some way to each being in the great and complex battle of life, should occur in the course of many successive generations? If such do occur, can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage, however slight, over others, would have the best chance of surviving and procreating their kind? On the other hand, we may feel sure that any variation in the least degree injurious would be rigidly destroyed. This preservation of favourable individual differences and variations, and the destruction of those which are injurious, I have called Natural Selection, or the Survival of the Fittest. Variations neither useful nor injurious would not be affected by natural selection, and would be left either a fluctuating element, as perhaps we see in certain polymorphic species, or would ultimately become fixed, owing to the nature of the organism and the nature of the conditions.

Several writers have misapprehended or objected to the term Natural Selection. Some have even imagined that natural selection induces variability, whereas it implies only the preservation of such variations as arise and are beneficial to the being under its conditions of life. No one objects to agriculturists speaking of the potent effects of man's selection; and in this case the individual differences given by nature, which man for some object selects, must of necessity first occur. Others have objected that the term selection implies conscious choice in the animals which become modified; and it has even been urged that, as plants have no volition, natural selection is not applicable to them! In the literal sense of the word, no doubt, natural selection is a false term; but who ever objected to chemists speaking of the elective affinities of the various elements?--and yet an acid cannot strictly be said to elect the base with which it in preference combines. It has been said that I speak of natural selection as an active power or Deity; but who objects to an author speaking of the attraction of gravity as ruling the movements of the planets? Every one knows what is meant and is implied by such metaphorical expressions; and they are almost necessary for brevity. So again it is difficult to avoid personifying the word Nature; but I mean by nature, only the aggregate action and product of many natural laws, and by laws the sequence of events as ascertained by us. With a little familiarity such superficial objections will be forgotten.

We shall best understand the probable course of natural selection by taking the case of a country undergoing some slight physical change, for instance, of climate. The proportional numbers of its inhabitants will almost immediately undergo a change, and some species will probably become extinct. We may conclude, from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together, that any change in the numerical proportions of the inhabitants, independently of the change of climate itself, would seriously affect the others. If the country were open on its borders, new forms would certainly immigrate, and this would likewise seriously disturb the relations of some of the former inhabitants. Let it be remembered how powerful the influence of a single introduced tree or mammal has been shown to be. But in the case of an island, or of a country partly surrounded by barriers, into which new and better adapted forms could not freely enter, we should then have places in the economy of nature which would assuredly be better filled up if some of the original inhabitants were in some manner modified; for, had the area been open to immigration, these same places would have been seized on by intruders. In such cases, slight modifications, which in any way favoured the individuals of any species, by better adapting them to their altered conditions, would tend to be preserved; and natural selection would have free scope for the work of improvement.

We have good reason to believe, as shown in the first chapter, that changes in the conditions of life give a tendency to increased variability; and in the foregoing cases the conditions the changed, and this would manifestly be favourable to natural selection, by affording a better chance of the occurrence of profitable variations. Unless such occur, natural selection can do nothing. Under the term of "variations," it must never be forgotten that mere individual differences are included. As man can produce a great result with his domestic animals and plants by adding up in any given direction individual differences, so could natural selection, but far more easily from having incomparably longer time for action. Nor do I believe that any great physical change, as of climate, or any unusual degree of isolation, to check immigration, is necessary in order that new and unoccupied places should be left for natural selection to fill up by improving some of the varying inhabitants. For as all the inhabitants of each country are struggling together with nicely balanced forces, extremely slight modifications in the structure or habits of one species would often give it an advantage over others; and still further modifications of the same kind would often still further increase the advantage, as long as the species continued under the same conditions of life and profited by similar means of subsistence and defence. No country can be named in which all the native inhabitants are now so perfectly adapted to each other and to the physical conditions under which they live, that none of them could be still better adapted or improved; for in all countries, the natives have been so far conquered by naturalised productions that they have allowed some foreigners to take firm possession of the land. And as foreigners have thus in every country beaten some of the natives, we may safely conclude that the natives might have been modified with advantage, so as to have better resisted the intruders.

Chapter V.

Laws of Variation

Effects of changed conditions

I have hitherto sometimes spoken as if the variations—so common and multiform with organic beings under domestication, and in a lesser degree with those under nature—were due to chance. This, of course is a wholly incorrect expression, but it serves to acknowledge plainly our ignorance of the cause of each particular variation. Some authors believe it to be as much the function of the reproductive system to produce individual differences, or slight deviations of structure, as to make the child like its parents. But the fact of variations and monstrosities occurring much more frequently under domestication than under nature, and the greater variability of species having wide ranges than of those with restricted ranges, lead to the conclusion that variability is generally related to the conditions of life to which each species has been exposed during several successive generations. In the first chapter I attempted to show that changed conditions act in two ways, directly on the whole organisation or on certain parts alone, and indirectly through the reproductive system. In all cases there are two factors, the nature of the organism, which is much the most important of the two, and the nature of the conditions. The direct action of changed conditions leads to definite or indefinite results. In the latter case the organisation seems to become plastic, and we have much fluctuating variability. In the former case the nature of the organism is such that it yields readily, when subjected to certain conditions, and all, or nearly all, the individuals become modified in the same way.

It is very difficult to decide how far changed conditions, such as of climate, food, etc., have acted in a definite manner. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence. But we may safely conclude that the innumerable complex co-adaptations of structure, which we see throughout nature between various organic beings, cannot be attributed simply to such action. In the following cases the conditions seem to have produced some slight definite effect: E. Forbes asserts that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species from further north or from a greater depth; but this certainly does not always hold good. Mr. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere, than when living near the coast or on islands; and Wollaston is convinced that residence near the sea affects the colours of insects. Moquin-Tandon gives a list of plants which, when growing near the sea-shore, have their leaves in some degree fleshy, though not elsewhere fleshy. These slightly varying organisms are interesting in as far as they present characters analogous to those possessed by the species which are confined to similar conditions.

When a variation is of the slightest use to any being, we cannot tell how much to attribute to the accumulative action of natural selection, and how much to the definite action of the conditions of life. Thus, it is well known to furriers that animals of the same species have thicker and better fur the further north they live; but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations, and how much to the action of the severe climate? For it would appear that climate has some direct action on the hair of our domestic quadrupeds.

Instances could be given of similar varieties being produced from the same species under external conditions of life as different as can well be conceived; and, on the other hand, of dissimilar varieties being produced under apparently the same external conditions. Again, innumerable instances are known to every naturalist, of species keeping true, or not varying at all, although living under the most opposite climates. Such considerations as these incline me to lay less weight on the direct action of the surrounding conditions, than on a tendency to vary, due to causes of which we are quite ignorant...

Chapter VI.

Difficulties Of The Theory

Difficulties of the theory of descent with modification

Long before the reader has arrived at this part of my work, a crowd of difficulties will have occurred to him. Some of them are so serious that to this day I can hardly reflect on them without being in some degree staggered; but, to the best of my judgment, the greater number are only apparent, and those that are real are not, I think, fatal to the theory.

These difficulties and objections may be classed under the following heads: First, why, if species have descended from other species by fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion, instead of the species being, as we see them, well defined?

Secondly, is it possible that an animal having, for instance, the structure and habits of a bat, could have been formed by the modification of some other animal with widely different habits and structure? Can we believe that natural selection could produce, on the one hand, an organ of trifling importance, such as the tail of a giraffe, which serves as a fly-flapper, and, on the other hand, an organ so wonderful as the eye?

Thirdly, can instincts be acquired and modified through natural selection? What shall we say to the instinct which leads the bee to make cells, and which has practically anticipated the discoveries of profound mathematicians?

Fourthly, how can we account for species, when crossed, being sterile and producing sterile offspring, whereas, when varieties are crossed, their fertility is unimpaired?

The two first heads will be here discussed; some miscellaneous objections in the following chapter; Instinct and Hybridism in the two succeeding chapters.

On the Absence of Rarity of Transitional Varieties

As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to take the place of, and finally to exterminate, its own less improved parent-form and other less-favoured forms with which it comes into competition. Thus extinction and natural selection go hand in hand. Hence, if we look at each species as descended from some unknown form, both the parent and all the transitional varieties will generally have been exterminated by the very process of the formation and perfection of the new form.

But, as by this theory innumerable transitional forms must have existed, why do we not find them embedded in countless numbers in the crust of the earth? It will be more convenient to discuss this question in the chapter on the imperfection of the geological record; and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed. The crust of the earth is a vast museum; but the natural collections have been imperfectly made, and only at long intervals of time.

But it may be urged that when several closely allied species inhabit the same territory, we surely ought to find at the present time many transitional forms. Let us take a simple case: in travelling from north to south over a continent, we generally meet at successive intervals with closely allied or representative species, evidently filling nearly the same place in the natural economy of the land. These representative species often meet and interlock; and as the one becomes rarer and rarer, the other becomes more and more frequent, till the one replaces the other. But if we compare these species where they intermingle, they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis inhabited by each. By my theory these allied species are descended from a common parent; and during the process of modification, each has become adapted to the conditions of life of its own region, and has supplanted and exterminated its original parent-form and all the transitional varieties between its past and present states. Hence we ought not to expect at the present time to meet with numerous transitional varieties in each region, though they must have existed there, and may be embedded there in a fossil condition. But in the intermediate region, having intermediate conditions of life, why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. But I think it can be in large part explained.

In the first place we should be extremely cautious in inferring, because an area is now continuous, that it has been continuous during a long period. Geology would lead us to believe that most continents have been broken up into islands even during the later tertiary periods; and in such islands distinct species might have been separately formed without the possibility of intermediate varieties existing in the intermediate zones. By changes in the form of the land and of climate, marine areas now continuous must often have existed within recent times in a far less continuous and uniform condition than at present. But I will pass over this way of escaping from the difficulty; for I believe that many perfectly defined species have been formed on strictly continuous areas; though I do not doubt that the formerly broken condition of areas now continuous, has played an important part in the formation of new species, more especially with freely-crossing and wandering animals...

Chapter VII.

Miscellaneous Objections To The Theory Of Natural Selection

Longevity

I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views, as some of the previous discussions may thus be made clearer; but it would be useless to discuss all of them, as many have been made by writers who have not taken the trouble to understand the subject. Thus a distinguished German naturalist has asserted that the weakest part of my theory is, that I consider all organic beings as imperfect: what I have really said is, that all are not as perfect as they might have been in relation to their conditions; and this is shown to be the case by so many native forms in many quarters of the world having yielded their places to intruding foreigners. Nor can organic beings, even if they were at any one time perfectly adapted to their conditions of life, have remained so, when their conditions changed, unless they themselves likewise changed; and no one will dispute that the physical conditions of each country, as well as the number and kinds of its inhabitants, have undergone many mutations.

A critic has lately insisted, with some parade of mathematical accuracy, that longevity is a great advantage to all species, so that he who believes in natural selection "must arrange his genealogical tree" in such a manner that all the descendants have longer lives than their progenitors! Cannot our critics conceive that a biennial plant or one of the lower animals might range into a cold climate and perish there every winter; and yet, owing to advantages gained through natural selection, survive from year to year by means of its seeds or ova? Mr. E. Ray Lankester has recently discussed this subject, and he concludes, as far as its extreme complexity allows him to form a judgment, that longevity is generally related to the standard of each species in the scale of organisation, as well as to the amount of expenditure in reproduction and in general activity. And these conditions have, it is probable, been largely determined through natural selection.

It has been argued that, as none of the animals and plants of Egypt, of which we know anything, have changed during the last three or four thousand years, so probably have none in any part of the world. But, as Mr. G.H. Lewes has remarked, this line of argument proves too much, for the ancient domestic races figured on the Egyptian monuments, or embalmed, are closely similar or even identical with those now living; yet all naturalists admit that such races have been produced through the modification of their original types. The many animals which have remained unchanged since the commencement of the glacial period, would have been an incomparably stronger case, for these have been exposed to great changes of climate and have migrated over great distances; whereas, in Egypt, during the last several thousand years, the conditions of life, as far as we know, have remained absolutely uniform. The fact of little or no modification having been effected since the glacial period, would have been of some avail against those who believe in an innate and necessary law of development, but is powerless against the doctrine of natural selection or the survival of the fittest, which implies that when variations or individual differences of a beneficial nature happen to arise, these will be preserved; but this will be effected only under certain favourable circumstances.

The celebrated palaeontologist, Bronn, at the close of his German translation of this work, asks how, on the principle of natural selection, can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions, they might live together; and if we lay on one side polymorphic species, in which the variability seems to be of a peculiar nature, and all mere temporary variations, such as size, albinism, etc., the more permanent varieties are generally found, as far as I can discover, inhabiting distinct stations, such as high land or low land, dry or moist districts. Moreover, in the case of animals which wander much about and cross freely, their varieties seem to be generally confined to distinct regions...

Chapter VIII.

Instinct

Instincts comparable with habits, but different in their origin

Many instincts are so wonderful that their development will probably appear to the reader a difficulty sufficient to overthrow my whole theory. I may here premise, that I have nothing to do with the origin of the mental powers, any more than I have with that of life itself. We are concerned only with the diversities of instinct and of the other mental faculties in animals of the same class.

I will not attempt any definition of instinct. It would be easy to show that several distinct mental actions are commonly embraced by this term; but every one understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds' nests. An action, which we ourselves require experience to enable us to perform, when performed by an animal, more especially by a very young one, without experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive. But I could show that none of these characters are universal. A little dose of judgment or reason, as Pierre Huber expresses it, often comes into play, even with animals low in the scale of nature.

Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. This comparison gives, I think, an accurate notion of the frame of mind under which an instinctive action is performed, but not necessarily of its origin. How unconsciously many habitual actions are performed, indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason. Habits easily become associated with other habits, with certain periods of time and states of the body. When once acquired, they often remain constant throughout life. Several other points of resemblance between instincts and habits could be pointed out. As in repeating a well-known song, so in instincts, one action follows another by a sort of rhythm; if a person be interrupted in a song, or in repeating anything by rote, he is generally forced to go back to recover the habitual train of thought: so P. Huber found it was with a caterpillar, which makes a very complicated hammock; for if he took a caterpillar which had completed its hammock up to, say, the sixth stage of construction, and put it into a hammock completed up only to the third stage, the caterpillar simply re-performed the fourth, fifth, and sixth stages of construction. If, however, a caterpillar were taken out of a hammock made up, for instance, to the third stage, and were put into one finished up to the sixth stage, so that much of its work was already done for it, far from deriving any benefit from this, it was much embarrassed, and, in order to complete its hammock, seemed forced to start from the third stage, where it had left off, and thus tried to complete the already finished work.

If we suppose any habitual action to become inherited--and it can be shown that this does sometimes happen--then the resemblance between what originally was a habit and an instinct becomes so close as not to be distinguished. If Mozart, instead of playing the pianoforte at three years old with wonderfully little practice, had played a tune with no practice at all, be might truly be said to have done so instinctively. But it would be a serious error to suppose that the greater number of instincts have been acquired by habit in one generation, and then transmitted by inheritance to succeeding generations. It can be clearly shown that the most wonderful instincts with which we are acquainted, namely, those of the hive-bee and of many ants, could not possibly have been acquired by habit.

It will be universally admitted that instincts are as important as corporeal structures for the welfare of each species, under its present conditions of life. Under changed conditions of life, it is at least possible that slight modifications of instinct might be profitable to a species; and if it can be shown that instincts do vary ever so little, then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that was profitable. It is thus, as I believe, that all the most complex and wonderful instincts have originated. As modifications of corporeal structure arise from, and are increased by, use or habit, and are diminished or lost by disuse, so I do not doubt it has been with instincts. But I believe that the effects of habit are in many cases of subordinate importance to the effects of the natural selection of what may be called spontaneous variations of instincts;--that is of variations produced by the same unknown causes which produce slight deviations of bodily structure.

No complex instinct can possibly be produced through natural selection, except by the slow and gradual accumulation of numerous, slight, yet profitable, variations. Hence, as in the case of corporeal structures, we ought to find in nature, not the actual transitional gradations by which each complex instinct has been acquired--for these could be found only in the lineal ancestors of each species--but we ought to find in the collateral lines of descent some evidence of such gradations; or we ought at least to be able to show that gradations of some kind are possible; and this we certainly can do. I have been surprised to find, making allowance for the instincts of animals having been but little observed, except in Europe and North America, and for no instinct being known among extinct species, how very generally gradations, leading to the most complex instincts, can be discovered. Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life, or at different seasons of the year, or when placed under different circumstances, etc.; in which case either the one or the other instinct might be preserved by natural selection. And such instances of diversity of instinct in the same species can be shown to occur in nature.

Chapter IX.

Hybridism

Distinction between the sterility of first crosses and of hybrids

The view commonly entertained by naturalists is that species, when intercrossed, have been specially endowed with sterility, in order to prevent their confusion. This view certainly seems at first highly probable, for species living together could hardly have been kept distinct had they been capable of freely crossing. The subject is in many ways important for us, more especially as the sterility of species when first crossed, and that of their hybrid offspring, cannot have been acquired, as I shall show, by the preservation of successive profitable degrees of sterility. It is an incidental result of differences in the reproductive systems of the parent-species.

In treating this subject, two classes of facts, to a large extent fundamentally different, have generally been confounded; namely, the sterility of species when first crossed, and the sterility of the hybrids produced from them.

Pure species have of course their organs of reproduction in a perfect condition, yet when intercrossed they produce either few or no offspring. Hybrids, on the other hand, have their reproductive organs functionally impotent, as may be clearly seen in the state of the male element in both plants and animals; though the formative organs themselves are perfect in structure, as far as the microscope reveals. In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed, or are imperfectly developed. This distinction is important, when the cause of the sterility, which is common to the two cases, has to be considered. The distinction probably has been slurred over, owing to the sterility in both cases being looked on as a special endowment, beyond the province of our reasoning powers.

The fertility of varieties, that is of the forms known or believed to be descended from common parents, when crossed, and likewise the fertility of their mongrel offspring, is, with reference to my theory, of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species.

Degrees of Sterility

First, for the sterility of species when crossed and of their hybrid offspring. It is impossible to study the several memoirs and works of those two conscientious and admirable observers, Kolreuter and Gartner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility. Kolreuter makes the rule universal; but then he cuts the knot, for in ten cases in which he found two forms, considered by most authors as distinct species, quite fertile together, he unhesitatingly ranks them as varieties. Gartner, also, makes the rule equally universal; and he disputes the entire fertility of Kolreuter's ten cases. But in these and in many other cases, Gartner is obliged carefully to count the seeds, in order to show that there is any degree of sterility. He always compares the maximum number of seeds produced by two species when first crossed, and the maximum produced by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature. But causes of serious error here intervene: a plant, to be hybridised, must be castrated, and, what is often more important, must be secluded in order to prevent pollen being brought to it by insects from other plants. Nearly all the plants experimented on by Gartner were potted, and were kept in a chamber in his house. That these processes are often injurious to the fertility of a plant cannot be doubted; for Gartner gives in his table about a score of cases of plants which he castrated, and artificially fertilised with their own pollen, and (excluding all cases such as the Leguminosae, in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. Moreover, as Gartner repeatedly crossed some forms, such as the common red and blue pimpernels (Anagallis arvensis and coerulea), which the best botanists rank as varieties, and found them absolutely sterile, we may doubt whether many species are really so sterile, when intercrossed, as he believed.

It is certain, on the one hand, that the sterility of various species when crossed is so different in degree and graduates away so insensibly, and, on the other hand, that the fertility of pure species is so easily affected by various circumstances, that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. I think no better evidence of this can be required than that the two most experienced observers who have ever lived, namely Kolreuter and Gartner, arrived at diametrically opposite conclusions in regard to some of the very same forms. It is also most instructive to compare--but I have not space here to enter on details--the evidence advanced by our best botanists on the question whether certain doubtful forms should be ranked as species or varieties, with the evidence from fertility adduced by different hybridisers, or by the same observer from experiments made during different years. It can thus be shown that neither sterility nor fertility affords any certain distinction between species and varieties. The evidence from this source graduates away, and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences...

Chapter X.

On The Imperfection Of The Geological Record

On the absence of intermediate varieties at the present day

In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. Most of them have now been discussed. One, namely, the distinctness of specific forms and their not being blended together by innumerable transitional links, is a very obvious difficulty. I assigned reasons why such links do not commonly occur at the present day under the circumstances apparently most favourable for their presence, namely, on an extensive and continuous area with graduated physical conditions. I endeavoured to show, that the life of each species depends in a more important manner on the presence of other already defined organic forms, than on climate, and, therefore, that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. I endeavoured, also, to show that intermediate varieties, from existing in lesser numbers than the forms which they connect, will generally be beaten out and exterminated during the course of further modification and improvement. The main cause, however, of innumerable intermediate links not now occurring everywhere throughout nature depends, on the very process of natural selection, through which new varieties continually take the places of and supplant their parent-forms. But just in proportion as this process of extermination has acted on an enormous scale, so must the number of intermediate varieties, which have formerly existed, be truly enormous. Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this, perhaps, is the most obvious and serious objection which can be urged against my theory. The explanation lies, as I believe, in the extreme imperfection of the geological record.

In the first place, it should always be borne in mind what sort of intermediate forms must, on the theory, have formerly existed. I have found it difficult, when looking at any two species, to avoid picturing to myself forms DIRECTLY intermediate between them. But this is a wholly false view; we should always look for forms intermediate between each species and a common but unknown progenitor; and the progenitor will generally have differed in some respects from all its modified descendants. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon; if we possessed all the intermediate varieties which have ever existed, we should have an extremely close series between both and the rock-pigeon; but we should have no varieties directly intermediate between the fantail and pouter; none, for instance, combining a tail somewhat expanded with a crop somewhat enlarged, the characteristic features of these two breeds. These two breeds, moreover, have become so much modified, that, if we had no historical or indirect evidence regarding their origin, it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon, C. livia, whether they had descended from this species or from some other allied species, such as C. oenas.

So with natural species, if we look to forms very distinct, for instance to the horse and tapir, we have no reason to suppose that links directly intermediate between them ever existed, but between each and an unknown common parent. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse; but in some points of structure may have differed considerably from both, even perhaps more than they differ from each other. Hence, in all such cases, we should be unable to recognise the parent-form of any two or more species, even if we closely compared the structure of the parent with that of its modified descendants, unless at the same time we had a nearly perfect chain of the intermediate links.

It is just possible, by the theory, that one of two living forms might have descended from the other; for instance, a horse from a tapir; and in this case DIRECT intermediate links will have existed between them. But such a case would imply that one form had remained for a very long period unaltered, whilst its descendants had undergone a vast amount of change; and the principle of competition between organism and organism, between child and parent, will render this a very rare event; for in all cases the new and improved forms of life tend to supplant the old and unimproved forms.

By the theory of natural selection all living species have been connected with the parent-species of each genus, by differences not greater than we see between the natural and domestic varieties of the same species at the present day; and these parent-species, now generally extinct, have in their turn been similarly connected with more ancient forms; and so on backwards, always converging to the common ancestor of each great class. So that the number of intermediate and transitional links, between all living and extinct species, must have been inconceivably great. But assuredly, if this theory be true, such have lived upon the earth...

Chapter XI.

On The Geological Succession Of Organic Beings

On the slow and successive appearance of new species

Let us now see whether the several facts and laws relating to the geological succession of organic beings accord best with the common view of the immutability of species, or with that of their slow and gradual modification, through variation and natural selection.

New species have appeared very slowly, one after another, both on the land and in the waters. Lyell has shown that it is hardly possible to resist the evidence on this head in the case of the several tertiary stages; and every year tends to fill up the blanks between the stages, and to make the proportion between the lost and existing forms more gradual. In some of the most recent beds, though undoubtedly of high antiquity if measured by years, only one or two species are extinct, and only one or two are new, having appeared there for the first time, either locally, or, as far as we know, on the face of the earth. The secondary formations are more broken; but, as Bronn has remarked, neither the appearance nor disappearance of the many species embedded in each formation has been simultaneous.

Species belonging to different genera and classes have not changed at the same rate, or in the same degree. In the older tertiary beds a few living shells may still be found in the midst of a multitude of extinct forms. Falconer has given a striking instance of a similar fact, for an existing crocodile is associated with many lost mammals and reptiles in the sub-Himalayan deposits. The Silurian Lingula differs but little from the living species of this genus; whereas most of the other Silurian Molluscs and all the Crustaceans have changed greatly. The productions of the land seem to have changed at a quicker rate than those of the sea, of which a striking instance has been observed in Switzerland. There is some reason to believe that organisms high in the scale, change more quickly than those that are low: though there are exceptions to this rule. The amount of organic change, as Pictet has remarked, is not the same in each successive so-called formation. Yet if we compare any but the most closely related formations, all the species will be found to have undergone some change. When a species has once disappeared from the face of the earth, we have no reason to believe that the same identical form ever reappears. The strongest apparent exception to this latter rule is that of the so-called "colonies" of M. Barrande, which intrude for a period in the midst of an older formation, and then allow the pre-existing fauna to reappear; but Lyell's explanation, namely, that it is a case of temporary migration from a distinct geographical province, seems satisfactory.

These several facts accord well with our theory, which includes no fixed law of development, causing all the inhabitants of an area to change abruptly, or simultaneously, or to an equal degree. The process of modification must be slow, and will generally affect only a few species at the same time; for the variability of each species is independent of that of all others. Whether such variations or individual differences as may arise will be accumulated through natural selection in a greater or less degree, thus causing a greater or less amount of permanent modification, will depend on many complex contingencies--on the variations being of a beneficial nature, on the freedom of intercrossing, on the slowly changing physical conditions of the country, on the immigration of new colonists, and on the nature of the other inhabitants with which the varying species come into competition. Hence it is by no means surprising that one species should retain the same identical form much longer than others; or, if changing, should change in a less degree. We find similar relations between the existing inhabitants of distinct countries; for instance, the land-shells and coleopterous insects of Madeira have come to differ considerably from their nearest allies on the continent of Europe, whereas the marine shells and birds have remained unaltered. We can perhaps understand the apparently quicker rate of change in terrestrial and in more highly organised productions compared with marine and lower productions, by the more complex relations of the higher beings to their organic and inorganic conditions of life, as explained in a former chapter. When many of the inhabitants of any area have become modified and improved, we can understand, on the principle of competition, and from the all-important relations of organism to organism in the struggle for life, that any form which did not become in some degree modified and improved, would be liable to extermination. Hence, we see why all the species in the same region do at last, if we look to long enough intervals of time, become modified; for otherwise they would become extinct.

In members of the same class the average amount of change, during long and equal periods of time, may, perhaps, be nearly the same; but as the accumulation of enduring formations, rich in fossils, depends on great masses of sediment being deposited on subsiding areas, our formations have been almost necessarily accumulated at wide and irregularly intermittent intervals of time; consequently the amount of organic change exhibited by the fossils embedded in consecutive formations is not equal. Each formation, on this view, does not mark a new and complete act of creation, but only an occasional scene, taken almost at hazard, in an ever slowly changing drama.

We can clearly understand why a species when once lost should never reappear, even if the very same conditions of life, organic and inorganic, should recur. For though the offspring of one species might be adapted (and no doubt this has occurred in innumerable instances) to fill the place of another species in the economy of nature, and thus supplant it; yet the two forms--the old and the new--would not be identically the same; for both would almost certainly inherit different characters from their distinct progenitors; and organisms already differing would vary in a different manner. For instance, it is possible, if all our fantail-pigeons were destroyed, that fanciers might make a new breed hardly distinguishable from the present breed; but if the parent rock-pigeon were likewise destroyed, and under nature we have every reason to believe that parent forms are generally supplanted and exterminated by their improved offspring, it is incredible that a fantail, identical with the existing breed, could be raised from any other species of pigeon, or even from any other well established race of the domestic pigeon, for the successive variations would almost certainly be in some degree different, and the newly-formed variety would probably inherit from its progenitor some characteristic differences...

Chapter XII.

Geographical Distribution

Present distribution cannot be accounted for by differences in physical conditions

In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the dissimilarity of the inhabitants of various regions can be wholly accounted for by climatal and other physical conditions. Of late, almost every author who has studied the subject has come to this conclusion. The case of America alone would almost suffice to prove its truth; for if we exclude the arctic and northern temperate parts, all authors agree that one of the most fundamental divisions in geographical distribution is that between the New and Old Worlds; yet if we travel over the vast American continent, from the central parts of the United States to its extreme southern point, we meet with the most diversified conditions; humid districts, arid deserts, lofty mountains, grassy plains, forests, marshes, lakes and great rivers, under almost every temperature. There is hardly a climate or condition in the Old World which cannot be paralleled in the New--at least so closely as the same species generally require. No doubt small areas can be pointed out in the Old World hotter than any in the New World; but these are not inhabited by a fauna different from that of the surrounding districts; for it is rare to find a group of organisms confined to a small area, of which the conditions are peculiar in only a slight degree. Notwithstanding this general parallelism in the conditions of Old and New Worlds, how widely different are their living productions!

In the southern hemisphere, if we compare large tracts of land in Australia, South Africa, and western South America, between latitudes 25 and 35 degrees, we shall find parts extremely similar in all their conditions, yet it would not be possible to point out three faunas and floras more utterly dissimilar. Or, again, we may compare the productions of South America south of latitude 35 degrees with those north of 25 degrees, which consequently are separated by a space of ten degrees of latitude, and are exposed to considerably different conditions; yet they are incomparably more closely related to each other than they are to the productions of Australia or Africa under nearly the same climate. Analogous facts could be given with respect to the inhabitants of the sea.

A second great fact which strikes us in our general review is, that barriers of any kind, or obstacles to free migration, are related in a close and important manner to the differences between the productions of various regions. We see this in the great difference in nearly all the terrestrial productions of the New and Old Worlds, excepting in the northern parts, where the land almost joins, and where, under a slightly different climate, there might have been free migration for the northern temperate forms, as there now is for the strictly arctic productions. We see the same fact in the great difference between the inhabitants of Australia, Africa, and South America under the same latitude; for these countries are almost as much isolated from each other as is possible. On each continent, also, we see the same fact; for on the opposite sides of lofty and continuous mountain-ranges, and of great deserts and even of large rivers, we find different productions; though as mountain chains, deserts, etc., are not as impassable, or likely to have endured so long, as the oceans separating continents, the differences are very inferior in degree to those characteristic of distinct continents.

Turning to the sea, we find the same law. The marine inhabitants of the eastern and western shores of South America are very distinct, with extremely few shells, crustacea, or echinodermata in common; but Dr. Gunther has recently shown that about thirty per cent of the fishes are the same on the opposite sides of the isthmus of Panama; and this fact has led naturalists to believe that the isthmus was formerly open. Westward of the shores of America, a wide space of open ocean extends, with not an island as a halting-place for emigrants; here we have a barrier of another kind, and as soon as this is passed we meet in the eastern islands of the Pacific with another and totally distinct fauna. So that three marine faunas range northward and southward in parallel lines not far from each other, under corresponding climate; but from being separated from each other by impassable barriers, either of land or open sea, they are almost wholly distinct. On the other hand, proceeding still further westward from the eastern islands of the tropical parts of the Pacific, we encounter no impassable barriers, and we have innumerable islands as halting-places, or continuous coasts, until, after travelling over a hemisphere, we come to the shores of Africa; and over this vast space we meet with no well-defined and distinct marine faunas. Although so few marine animals are common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands, yet many fishes range from the Pacific into the Indian Ocean, and many shells are common to the eastern islands of the Pacific and the eastern shores of Africa on almost exactly opposite meridians of longitude.

A third great fact, partly included in the foregoing statement, is the affinity of the productions of the same continent or of the same sea, though the species themselves are distinct at different points and stations. It is a law of the widest generality, and every continent offers innumerable instances. Nevertheless, the naturalist, in travelling, for instance, from north to south, never fails to be struck by the manner in which successive groups of beings, specifically distinct, though nearly related, replace each other. He hears from closely allied, yet distinct kinds of birds, notes nearly similar, and sees their nests similarly constructed, but not quite alike, with eggs coloured in nearly the same manner. The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich), and northward the plains of La Plata by another species of the same genus; and not by a true ostrich or emu, like those inhabiting Africa and Australia under the same latitude. On these same plains of La Plata we see the agouti and bizcacha, animals having nearly the same habits as our hares and rabbits, and belonging to the same order of Rodents, but they plainly display an American type of structure. We ascend the lofty peaks of the Cordillera, and we find an alpine species of bizcacha; we look to the waters, and we do not find the beaver or muskrat, but the coypu and capybara, rodents of the South American type. Innumerable other instances could be given. If we look to the islands off the American shore, however much they may differ in geological structure, the inhabitants are essentially American, though they may be all peculiar species. We may look back to past ages, as shown in the last chapter, and we find American types then prevailing on the American continent and in the American seas. We see in these facts some deep organic bond, throughout space and time, over the same areas of land and water, independently of physical conditions. The naturalist must be dull who is not led to inquire what this bond is...

Chapter XIII.

Geographical Distribution - Continued

Present distribution cannot be accounted for by differences in physical conditions

The bond is simply inheritance, that cause which alone, as far as we positively know, produces organisms quite like each other, or, as we see in the case of varieties, nearly alike. The dissimilarity of the inhabitants of different regions may be attributed to modification through variation and natural selection, and probably in a subordinate degree to the definite influence of different physical conditions. The degrees of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been more or less effectually prevented, at periods more or less remote--on the nature and number of the former immigrants--and on the action of the inhabitants on each other in leading to the preservation of different modifications; the relation of organism to organism in the struggle for life being, as I have already often remarked, the most important of all relations. Thus the high importance of barriers comes into play by checking migration; as does time for the slow process of modification through natural selection. Widely-ranging species, abounding in individuals, which have already triumphed over many competitors in their own widely-extended homes, will have the best chance of seizing on new places, when they spread out into new countries. In their new homes they will be exposed to new conditions, and will frequently undergo further modification and improvement; and thus they will become still further victorious, and will produce groups of modified descendants. On this principle of inheritance with modification we can understand how it is that sections of genera, whole genera, and even families, are confined to the same areas, as is so commonly and notoriously the case...

Chapter XIV.

Mutual Affinities Of Organic Beings

Classification, groups subordinate to groups

From the most remote period in the history of the world organic beings have been found to resemble each other in descending degrees, so that they can be classed in groups under groups. This classification is not arbitrary like the grouping of the stars in constellations. The existence of groups would have been of simple significance, if one group had been exclusively fitted to inhabit the land, and another the water; one to feed on flesh, another on vegetable matter, and so on; but the case is widely different, for it is notorious how commonly members of even the same subgroup have different habits. In the second and fourth chapters, on Variation and on Natural Selection, I have attempted to show that within each country it is the widely ranging, the much diffused and common, that is the dominant species, belonging to the larger genera in each class, which vary most. The varieties, or incipient species, thus produced, ultimately become converted into new and distinct species; and these, on the principle of inheritance, tend to produce other new and dominant species. Consequently the groups which are now large, and which generally include many dominant species, tend to go on increasing in size. I further attempted to show that from the varying descendants of each species trying to occupy as many and as different places as possible in the economy of nature, they constantly tend to diverge in character. This latter conclusion is supported by observing the great diversity of forms, which, in any small area, come into the closest competition, and by certain facts in naturalisation.

I attempted also to show that there is a steady tendency in the forms which are increasing in number and diverging in character, to supplant and exterminate the preceding, less divergent and less improved forms. I request the reader to turn to the diagram illustrating the action, as formerly explained, of these several principles; and he will see that the inevitable result is, that the modified descendants proceeding from one progenitor become broken up into groups subordinate to groups. In the diagram each letter on the uppermost line may represent a genus including several species; and the whole of the genera along this upper line form together one class, for all are descended from one ancient parent, and, consequently, have inherited something in common. But the three genera on the left hand have, on this same principle, much in common, and form a subfamily, distinct from that containing the next two genera on the right hand, which diverged from a common parent at the fifth stage of descent. These five genera have also much in common, though less than when grouped in subfamilies; and they form a family distinct from that containing the three genera still further to the right hand, which diverged at an earlier period. And all these genera, descended from (A), form an order distinct from the genera descended from (I). So that we here have many species descended from a single progenitor grouped into genera; and the genera into subfamilies, families and orders, all under one great class. The grand fact of the natural subordination of organic beings in groups under groups, which, from its familiarity, does not always sufficiently strike us, is in my judgment thus explained. No doubt organic beings, like all other objects, can be classed in many ways, either artificially by single characters, or more naturally by a number of characters. We know, for instance, that minerals and the elemental substances can be thus arranged. In this case there is of course no relation to genealogical succession, and no cause can at present be assigned for their falling into groups. But with organic beings the case is different, and the view above given accords with their natural arrangement in group under group; and no other explanation has ever been attempted.

Naturalists, as we have seen, try to arrange the species, genera and families in each class, on what is called the Natural System. But what is meant by this system? Some authors look at it merely as a scheme for arranging together those living objects which are most alike, and for separating those which are most unlike; or as an artificial method of enunciating, as briefly as possible, general propositions--that is, by one sentence to give the characters common, for instance, to all mammals, by another those common to all carnivora, by another those common to the dog-genus, and then, by adding a single sentence, a full description is given of each kind of dog. The ingenuity and utility of this system are indisputable. But many naturalists think that something more is meant by the Natural System; they believe that it reveals the plan of the Creator; but unless it be specified whether order in time or space, or both, or what else is meant by the plan of the Creator, it seems to me that nothing is thus added to our knowledge. Expressions such as that famous one by Linnaeus, which we often meet with in a more or less concealed form, namely, that the characters do not make the genus, but that the genus gives the characters, seem to imply that some deeper bond is included in our classifications than mere resemblance. I believe that this is the case, and that community of descent--the one known cause of close similarity in organic beings--is the bond, which, though observed by various degrees of modification, is partially revealed to us by our classifications.

Let us now consider the rules followed in classification, and the difficulties which are encountered on the view that classification either gives some unknown plan of creation, or is simply a scheme for enunciating general propositions and of placing together the forms most like each other. It might have been thought (and was in ancient times thought) that those parts of the structure which determined the habits of life, and the general place of each being in the economy of nature, would be of very high importance in classification. Nothing can be more false. No one regards the external similarity of a mouse to a shrew, of a dugong to a whale, of a whale to a fish, as of any importance. These resemblances, though so intimately connected with the whole life of the being, are ranked as merely "adaptive or analogical characters;" but to the consideration of these resemblances we shall recur. It may even be given as a general rule, that the less any part of the organisation is concerned with special habits, the more important it becomes for classification. As an instance: Owen, in speaking of the dugong, says, "The generative organs, being those which are most remotely related to the habits and food of an animal, I have always regarded as affording very clear indications of its true affinities. We are least likely in the modifications of these organs to mistake a merely adaptive for an essential character." With plants how remarkable it is that the organs of vegetation, on which their nutrition and life depend, are of little signification; whereas the organs of reproduction, with their product the seed and embryo, are of paramount importance! So again, in formerly discussing certain morphological characters which are not functionally important, we have seen that they are often of the highest service in classification. This depends on their constancy throughout many allied groups; and their constancy chiefly depends on any slight deviations not having been preserved and accumulated by natural selection, which acts only on serviceable characters...

Chapter XV.

Recapitulation And Conclusion

Recapitulation of the objections to the theory of Natural Selection

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...