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
