The Origin of Species by Means of Natural Selection

In the case of a large tree covered with innumerable flowers, it may be objected that pollen could seldom be carried from tree to tree, and at most only from flower to flower on the same tree; and flowers on the same tree can be considered as distinct individuals only in a limited sense. I believe this objection to be valid, but that nature has largely provided against it by giving to trees a strong tendency to bear flowers with separated sexes. When the sexes are separated, although the male and female flowers may be produced on the same tree, pollen must be regularly carried from flower to flower; and this will give a better chance of pollen being occasionally carried from tree to tree. That trees belonging to all orders have their sexes more often separated than other plants, I find to be the case in this country; and at my request Dr. Hooker tabulated the trees of New Zealand, and Dr. Asa Gray those of the United States, and the result was as I anticipated. On the other hand, Dr. Hooker informs me that the rule does not hold good in Australia: but if most of the Australian trees are dichogamous, the same result would follow as if they bore flowers with separated sexes. I have made these few remarks on trees simply to call attention to the subject.

Turning for a brief space to animals: various terrestrial species are hermaphrodites, such as the land-mollusca and earth-worms; but these all pair. As yet I have not found a single terrestrial animal which can fertilise itself. This remarkable fact, which offers so strong a contrast with terrestrial plants, is intelligible on the view of an occasional cross being indispensable; for owing to the nature of the fertilising element there are no means, analogous to the action of insects and of the wind with plants, by which an occasional cross could be effected with terrestrial animals without the concurrence of two individuals. Of aquatic animals, there are many self-fertilising hermaphrodites; but here the currents of water offer an obvious means for an occasional cross. As in the case of flowers, I have as yet failed, after consultation with one of the highest authorities, namely, Professor Huxley, to discover a single hermaphrodite animal with the organs of reproduction so perfectly enclosed that access from without, and the occasional influence of a distinct individual, can be shown to be physically impossible. Cirripedes long appeared to me to present, under this point of view, a case of great difficulty; but I have been enabled, by a fortunate chance, to prove that two individuals, though both are self-fertilising hermaphrodites, do sometimes cross.

It must have struck most naturalists as a strange anomaly that, both with animals and plants, some species of the same family and even of the same genus, though agreeing closely with each other in their whole organisation, are hermaphrodites, and some unisexual. But if, in fact, all hermaphrodites do occasionally intercross, the difference between them and unisexual species is, as far as function is concerned, very small.

From these several considerations and from the many special facts which I have collected, but which I am unable here to give, it appears that with animals and plants an occasional intercross between distinct individuals is a very general, if not universal, law of nature.

CIRCUMSTANCES FAVOURABLE FOR THE PRODUCTION OF NEW FORMS THROUGH NATURAL SELECTION.

This is an extremely intricate subject. A great amount of variability, under which term individual differences are always included, will evidently be favourable. A large number of individuals, by giving a better chance within any given period for the appearance of profitable variations, will compensate for a lesser amount of variability in each individual, and is, I believe, a highly important element of success. Though nature grants long periods of time for the work of natural selection, she does not grant an indefinite period; for as all organic beings are striving to seize on each place in the economy of nature, if any one species does not become modified and improved in a corresponding degree with its competitors it will be exterminated. Unless favourable variations be inherited by some at least of the offspring, nothing can be effected by natural selection. The tendency to reversion may often check or prevent the work; but as this tendency has not prevented man from forming by selection numerous domestic races, why should it prevail against natural selection?

In the case of methodical selection, a breeder selects for some definite object, and if the individuals be allowed freely to intercross, his work will completely fail. But when many men, without intending to alter the breed, have a nearly common standard of perfection, and all try to procure and breed from the best animals, improvement surely but slowly follows from this unconscious process of selection, notwithstanding that there is no separation of selected individuals. Thus it will be under nature; for within a confined area, with some place in the natural polity not perfectly occupied, all the individuals varying in the right direction, though in different degrees, will tend to be preserved. But if the area be large, its several districts will almost certainly present different conditions of life; and then, if the same species undergoes modification in different districts, the newly formed varieties will intercross on the confines of each. But we shall see in the sixth chapter that intermediate varieties, inhabiting intermediate districts, will in the long run generally be supplanted by one of the adjoining varieties. Intercrossing will chiefly affect those animals which unite for each birth and wander much, and which do not breed at a very quick rate. Hence with animals of this nature, for instance birds, varieties will generally be confined to separated countries; and this I find to be the case. With hermaphrodite organisms which cross only occasionally, and likewise with animals which unite for each birth, but which wander little and can increase at a rapid rate, a new and improved variety might be quickly formed on any one spot, and might there maintain itself in a body and afterward spread, so that the individuals of the new variety would chiefly cross together. On this principle nurserymen always prefer saving seed from a large body of plants, as the chance of intercrossing is thus lessened.

Even with animals which unite for each birth, and which do not propagate rapidly, we must not assume that free intercrossing would always eliminate the effects of natural selection; for I can bring forward a considerable body of facts showing that within the same area two varieties of the same animal may long remain distinct, from haunting different stations, from breeding at slightly different seasons, or from the individuals of each variety preferring to pair together.

Intercrossing plays a very important part in nature by keeping the individuals of the same species, or of the same variety, true and uniform in character. It will obviously thus act far more efficiently with those animals which unite for each birth; but, as already stated, we have reason to believe that occasional intercrosses take place with all animals and plants. Even if these take place only at long intervals of time, the young thus produced will gain so much in vigour and fertility over the offspring from long-continued self-fertilisation, that they will have a better chance of surviving and propagating their kind; and thus in the long run the influence of crosses, even at rare intervals, will be great. With respect to organic beings extremely low in the scale, which do not propagate sexually, nor conjugate, and which cannot possibly intercross, uniformity of character can be retained by them under the same conditions of life, only through the principle of inheritance, and through natural selection which will destroy any individuals departing from the proper type. If the conditions of life change and the form undergoes modification, uniformity of character can be given to the modified offspring, solely by natural selection preserving similar favourable variations.

Isolation also is an important element in the modification of species through natural selection. In a confined or isolated area, if not very large, the organic and inorganic conditions of life will generally be almost uniform; so that natural selection will tend to modify all the varying individuals of the same species in the same manner. Intercrossing with the inhabitants of the surrounding districts, will also be thus prevented. Moritz Wagner has lately published an interesting essay on this subject, and has shown that the service rendered by isolation in preventing crosses between newly-formed varieties is probably greater even than I supposed. But from reasons already assigned I can by no means agree with this naturalist, that migration and isolation are necessary elements for the formation of new species. The importance of isolation is likewise great in preventing, after any physical change in the conditions, such as of climate, elevation of the land, etc., the immigration of better adapted organisms; and thus new places in the natural economy of the district will be left open to be filled up by the modification of the old inhabitants. Lastly, isolation will give time for a new variety to be improved at a slow rate; and this may sometimes be of much importance. If, however, an isolated area be very small, either from being surrounded by barriers, or from having very peculiar physical conditions, the total number of the inhabitants will be small; and this will retard the production of new species through natural selection, by decreasing the chances of favourable variations arising.

The mere lapse of time by itself does nothing, either for or against natural selection. I state this because it has been erroneously asserted that the element of time has been assumed by me to play an all-important part in modifying species, as if all the forms of life were necessarily undergoing change through some innate law. Lapse of time is only so far important, and its importance in this respect is great, that it gives a better chance of beneficial variations arising and of their being selected, accumulated, and fixed. It likewise tends to increase the direct action of the physical conditions of life, in relation to the constitution of each organism.

If we turn to nature to test the truth of these remarks, and look at any small isolated area, such as an oceanic island, although the number of the species inhabiting it is small, as we shall see in our chapter on Geographical Distribution; yet of these species a very large proportion are endemic, – that is, have been produced there and nowhere else in the world. Hence an oceanic island at first sight seems to have been highly favourable for the production of new species. But we may thus deceive ourselves, for to ascertain whether a small isolated area, or a large open area like a continent, has been most favourable for the production of new organic forms, we ought to make the comparison within equal times; and this we are incapable of doing.

Although isolation is of great importance in the production of new species, on the whole I am inclined to believe that largeness of area is still more important, especially for the production of species which shall prove capable of enduring for a long period, and of spreading widely. Throughout a great and open area, not only will there be a better chance of favourable variations, arising from the large number of individuals of the same species there supported, but the conditions of life are much more complex from the large number of already existing species; and if some of these many species become modified and improved, others will have to be improved in a corresponding degree, or they will be exterminated. Each new form, also, as soon as it has been much improved, will be able to spread over the open and continuous area, and will thus come into competition with many other forms. Moreover, great areas, though now continuous, will often, owing to former oscillations of level, have existed in a broken condition, so that the good effects of isolation will generally, to a certain extent, have concurred. Finally, I conclude that, although small isolated areas have been in some respects highly favourable for the production of new species, yet that the course of modification will generally have been more rapid on large areas; and what is more important, that the new forms produced on large areas, which already have been victorious over many competitors, will be those that will spread most widely, and will give rise to the greatest number of new varieties and species. They will thus play a more important part in the changing history of the organic world.

In accordance with this view, we can, perhaps, understand some facts which will be again alluded to in our chapter on Geographical Distribution; for instance, the fact of the productions of the smaller continent of Australia now yielding before those of the larger Europaeo-Asiatic area. Thus, also, it is that continental productions have everywhere become so largely naturalised on islands. On a small island, the race for life will have been less severe, and there will have been less modification and less extermination. Hence, we can understand how it is that the flora of Madeira, according to Oswald Heer, resembles to a certain extent the extinct tertiary flora of Europe. All fresh water basins, taken together, make a small area compared with that of the sea or of the land. Consequently, the competition between fresh water productions will have been less severe than elsewhere; new forms will have been more slowly produced, and old forms more slowly exterminated. And it is in fresh water basins that we find seven genera of Ganoid fishes, remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders at present widely separated in the natural scale. These anomalous forms may be called living fossils; they have endured to the present day, from having inhabited a confined area, and from having been exposed to less varied, and therefore less severe, competition.

To sum up, as far as the extreme intricacy of the subject permits, the circumstances favourable and unfavourable for the production of new species through natural selection. I conclude that for terrestrial productions a large continental area, which has undergone many oscillations of level, will have been the most favourable for the production of many new forms of life, fitted to endure for a long time and to spread widely. While the area existed as a continent the inhabitants will have been numerous in individuals and kinds, and will have been subjected to severe competition. When converted by subsidence into large separate islands there will still have existed many individuals of the same species on each island: intercrossing on the confines of the range of each new species will have been checked: after physical changes of any kind immigration will have been prevented, so that new places in the polity of each island will have had to be filled up by the modification of the old inhabitants; and time will have been allowed for the varieties in each to become well modified and perfected. When, by renewed elevation, the islands were reconverted into a continental area, there will again have been very severe competition; the most favoured or improved varieties will have been enabled to spread; there will have been much extinction of the less improved forms, and the relative proportional numbers of the various inhabitants of the reunited continent will again have been changed; and again there will have been a fair field for natural selection to improve still further the inhabitants, and thus to produce new species.

That natural selection generally act with extreme slowness I fully admit. It can act only when there are places in the natural polity of a district which can be better occupied by the modification of some of its existing inhabitants. The occurrence of such places will often depend on physical changes, which generally take place very slowly, and on the immigration of better adapted forms being prevented. As some few of the old inhabitants become modified the mutual relations of others will often be disturbed; and this will create new places, ready to be filled up by better adapted forms; but all this will take place very slowly. Although all the individuals of the same species differ in some slight degree from each other, it would often be long before differences of the right nature in various parts of the organisation might occur. The result would often be greatly retarded by free intercrossing. Many will exclaim that these several causes are amply sufficient to neutralise the power of natural selection. I do not believe so. But I do believe that natural selection will generally act very slowly, only at long intervals of time, and only on a few of the inhabitants of the same region. I further believe that these slow, intermittent results accord well with what geology tells us of the rate and manner at which the inhabitants of the world have changed.

Slow though the process of selection may be, if feeble man can do much by artificial selection, I can see no limit to the amount of change, to the beauty and complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may have been effected in the long course of time through nature's power of selection, that is by the survival of the fittest.

EXTINCTION CAUSED BY NATURAL SELECTION.

This subject will be more fully discussed in our chapter on Geology; but it must here be alluded to from being intimately connected with natural selection. Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure. Owing to the high geometrical rate of increase of all organic beings, each area is already fully stocked with inhabitants, and it follows from this, that as the favoured forms increase in number, so, generally, will the less favoured decrease and become rare. Rarity, as geology tells us, is the precursor to extinction. We can see that any form which is represented by few individuals will run a good chance of utter extinction, during great fluctuations in the nature or the seasons, or from a temporary increase in the number of its enemies. But we may go further than this; for as new forms are produced, unless we admit that specific forms can go on indefinitely increasing in number, many old forms must become extinct. That the number of specific forms has not indefinitely increased, geology plainly tells us; and we shall presently attempt to show why it is that the number of species throughout the world has not become immeasurably great.

We have seen that the species which are most numerous in individuals have the best chance of producing favourable variations within any given period. We have evidence of this, in the facts stated in the second chapter, showing that it is the common and diffused or dominant species which offer the greatest number of recorded varieties. Hence, rare species will be less quickly modified or improved within any given period; they will consequently be beaten in the race for life by the modified and improved descendants of the commoner species.

From these several considerations I think it inevitably follows, that as new species in the course of time are formed through natural selection, others will become rarer and rarer, and finally extinct. The forms which stand in closest competition with those undergoing modification and improvement, will naturally suffer most. And we have seen in the chapter on the Struggle for Existence that it is the most closely-allied forms, – varieties of the same species, and species of the same genus or related genera, – which, from having nearly the same structure, constitution and habits, generally come into the severest competition with each other. Consequently, each new variety or species, during the progress of its formation, will generally press hardest on its nearest kindred, and tend to exterminate them. We see the same process of extermination among our domesticated productions, through the selection of improved forms by man. Many curious instances could be given showing how quickly new breeds of cattle, sheep and other animals, and varieties of flowers, take the place of older and inferior kinds. In Yorkshire, it is historically known that the ancient black cattle were displaced by the long-horns, and that these "were swept away by the short-horns" (I quote the words of an agricultural writer) "as if by some murderous pestilence."

DIVERGENCE OF CHARACTER.

The principle, which I have designated by this term, is of high importance, and explains, as I believe, several important facts. In the first place, varieties, even strongly-marked ones, though having somewhat of the character of species – as is shown by the hopeless doubts in many cases how to rank them – yet certainly differ far less from each other than do good and distinct species. Nevertheless according to my view, varieties are species in the process of formation, or are, as I have called them, incipient species. How, then, does the lesser difference between varieties become augmented into the greater difference between species? That this does habitually happen, we must infer from most of the innumerable species throughout nature presenting well-marked differences; whereas varieties, the supposed prototypes and parents of future well-marked species, present slight and ill-defined differences. Mere chance, as we may call it, might cause one variety to differ in some character from its parents, and the offspring of this variety again to differ from its parent in the very same character and in a greater degree; but this alone would never account for so habitual and large a degree of difference as that between the species of the same genus.

As has always been my practice, I have sought light on this head from our domestic productions. We shall here find something analogous. It will be admitted that the production of races so different as short-horn and Hereford cattle, race and cart horses, the several breeds of pigeons, etc., could never have been effected by the mere chance accumulation of similar variations during many successive generations. In practice, a fancier is, for instance, struck by a pigeon having a slightly shorter beak; another fancier is struck by a pigeon having a rather longer beak; and on the acknowledged principle that "fanciers do not and will not admire a medium standard, but like extremes," they both go on (as has actually occurred with the sub-breeds of the tumbler-pigeon) choosing and breeding from birds with longer and longer beaks, or with shorter and shorter beaks. Again, we may suppose that at an early period of history, the men of one nation or district required swifter horses, while those of another required stronger and bulkier horses. The early differences would be very slight; but, in the course of time, from the continued selection of swifter horses in the one case, and of stronger ones in the other, the differences would become greater, and would be noted as forming two sub-breeds. Ultimately after the lapse of centuries, these sub-breeds would become converted into two well-established and distinct breeds. As the differences became greater, the inferior animals with intermediate characters, being neither very swift nor very strong, would not have been used for breeding, and will thus have tended to disappear. Here, then, we see in man's productions the action of what may be called the principle of divergence, causing differences, at first barely appreciable, steadily to increase, and the breeds to diverge in character, both from each other and from their common parent.

But how, it may be asked, can any analogous principle apply in nature? I believe it can and does apply most efficiently (though it was a long time before I saw how), from the simple circumstance that the more diversified the descendants from any one species become in structure, constitution, and habits, by so much will they be better enabled to seize on many and widely diversified places in the polity of nature, and so be enabled to increase in numbers.

We can clearly discern this in the case of animals with simple habits. Take the case of a carnivorous quadruped, of which the number that can be supported in any country has long ago arrived at its full average. If its natural power of increase be allowed to act, it can succeed in increasing (the country not undergoing any change in conditions) only by its varying descendants seizing on places at present occupied by other animals: some of them, for instance, being enabled to feed on new kinds of prey, either dead or alive; some inhabiting new stations, climbing trees, frequenting water, and some perhaps becoming less carnivorous. The more diversified in habits and structure the descendants of our carnivorous animals become, the more places they will be enabled to occupy. What applies to one animal will apply throughout all time to all animals – that is, if they vary – for otherwise natural selection can effect nothing. So it will be with plants. It has been experimentally proved, that if a plot of ground be sown with one species of grass, and a similar plot be sown with several distinct genera of grasses, a greater number of plants and a greater weight of dry herbage can be raised in the latter than in the former case. The same has been found to hold good when one variety and several mixed varieties of wheat have been sown on equal spaces of ground. Hence, if any one species of grass were to go on varying, and the varieties were continually selected which differed from each other in the same manner, though in a very slight degree, as do the distinct species and genera of grasses, a greater number of individual plants of this species, including its modified descendants, would succeed in living on the same piece of ground. And we know that each species and each variety of grass is annually sowing almost countless seeds; and is thus striving, as it may be said, to the utmost to increase in number. Consequently, in the course of many thousand generations, the most distinct varieties of any one species of grass would have the best chance of succeeding and of increasing in numbers, and thus of supplanting the less distinct varieties; and varieties, when rendered very distinct from each other, take the rank of species.

The truth of the principle that the greatest amount of life can be supported by great diversification of structure, is seen under many natural circumstances. In an extremely small area, especially if freely open to immigration, and where the contest between individual and individual must be very severe, we always find great diversity in its inhabitants. For instance, I found that a piece of turf, three feet by four in size, which had been exposed for many years to exactly the same conditions, supported twenty species of plants, and these belonged to eighteen genera and to eight orders, which shows how much these plants differed from each other. So it is with the plants and insects on small and uniform islets: also in small ponds of fresh water. Farmers find that they can raise more food by a rotation of plants belonging to the most different orders: nature follows what may be called a simultaneous rotation. Most of the animals and plants which live close round any small piece of ground, could live on it (supposing its nature not to be in any way peculiar), and may be said to be striving to the utmost to live there; but, it is seen, that where they come into the closest competition, the advantages of diversification of structure, with the accompanying differences of habit and constitution, determine that the inhabitants, which thus jostle each other most closely, shall, as a general rule, belong to what we call different genera and orders.

The same principle is seen in the naturalisation of plants through man's agency in foreign lands. It might have been expected that the plants which would succeed in becoming naturalised in any land would generally have been closely allied to the indigenes; for these are commonly looked at as specially created and adapted for their own country. It might also, perhaps, have been expected that naturalised plants would have belonged to a few groups more especially adapted to certain stations in their new homes. But the case is very different; and Alph. de Candolle has well remarked, in his great and admirable work, that floras gain by naturalisation, proportionally with the number of the native genera and species, far more in new genera than in new species. To give a single instance: in the last edition of Dr. Asa Gray's "Manual of the Flora of the Northern United States," 260 naturalised plants are enumerated, and these belong to 162 genera. We thus see that these naturalised plants are of a highly diversified nature. They differ, moreover, to a large extent, from the indigenes, for out of the 162 naturalised genera, no less than 100 genera are not there indigenous, and thus a large proportional addition is made to the genera now living in the United States.

By considering the nature of the plants or animals which have in any country struggled successfully with the indigenes, and have there become naturalised, we may gain some crude idea in what manner some of the natives would have had to be modified in order to gain an advantage over their compatriots; and we may at least infer that diversification of structure, amounting to new generic differences, would be profitable to them.