These curious and inexplicable correlations of colour with constitutional peculiarities, both in animals and plants, render it probable that the correlation of colour with infertility, which has been detected in several cases in plants, may also extend to animals in a state of nature; and if so, the fact is of the highest importance as throwing light on the origin of the infertility of many allied species. This will be better understood after considering the facts which will be now described.
The Isolation of Varieties by Selective Association.
In the last chapter I have shown that the importance of geographical isolation for the formation of new species by natural selection has been greatly exaggerated, because the very change of conditions, which is the initial power in starting such new forms, leads also to a local or stational segregation of the forms acted upon. But there is also a very powerful cause of isolation in the mental nature—the likes and dislikes—of animals; and to this is probably due the fact of the comparative rarity of hybrids in a state of nature. The differently coloured herds of cattle in the Falkland Islands, each of which keeps separate, have been already mentioned; and it may be added, that the mouse-coloured variety seem to have already developed a physiological peculiarity in breeding a month earlier than the others. Similar facts occur, however, among our domestic animals and are well known to breeders. Professor Low, one of the greatest authorities on our domesticated animals, says: "The female of the dog, when not under restraint, makes selection of her mate, the mastiff selecting the mastiff, the terrier the terrier, and so on." And again: "The Merino sheep and Heath sheep of Scotland, if two flocks are mixed together, each will breed with its own variety." Mr. Darwin has collected many facts illustrating this point. One of the chief pigeon-fanciers in England informed him that, if free to choose, each breed would prefer pairing with its own kind. Among the wild horses in Paraguay those of the same colour and size associate together; while in Circassia there are three races of horses which have received special names, and which, when living a free life, almost always refuse to mingle and cross, and will even attack one another. On one of the Faroe Islands, not more than half a mile in diameter, the half-wild native black sheep do not readily mix with imported white sheep. In the Forest of Dean, and in the New Forest, the dark and pale coloured herds of fallow deer have never been known to mingle; and even the curious Ancon sheep of quite modern origin have been observed to keep together, separating themselves from the rest of the flock when put into enclosures with other sheep. The same rule applies to birds, for Darwin was informed by the Rev. W.D. Fox that his flocks of white and Chinese geese kept distinct.[61]
This constant preference of animals for their like, even in the case of slightly different varieties of the same species, is evidently a fact of great importance in considering the origin of species by natural selection, since it shows us that, so soon as a slight differentiation of form or colour has been effected, isolation will at once arise by the selective association of the animals themselves; and thus the great stumbling-block of "the swamping effects of intercrossing," which has been so prominently brought forward by many naturalists, will be completely obviated.
If now we combine with this fact the correlation of colour with important constitutional peculiarities, and, in some cases, with infertility; and consider, further, the curious parallelism that has been shown to exist between the effects of changed conditions and the intercrossing of varieties in producing either an increase or a decrease of fertility, we shall have obtained, at all events, a starting-point for the production of that infertility which is so characteristic a feature of distinct species when intercrossed. All we need, now, is some means of increasing or accumulating this initial tendency; and to a discussion of this problem we will therefore address ourselves.
The Influence of Natural Selection upon Sterility and Fertility.
It will occur to many persons that, as the infertility or sterility of incipient species would be useful to them when occupying the same or adjacent areas, by neutralising the effects of intercrossing, this infertility might have been increased by the action of natural selection; and this will be thought the more probable if we admit, as we have seen reason to do, that variations in fertility occur, perhaps as frequently as other variations. Mr. Darwin tells us that, at one time, this appeared to him probable, but he found the problem to be one of extreme complexity; and he was also influenced against the view by many considerations which seemed to render such an origin of the sterility or infertility of species when intercrossed very improbable. The fact that species which occupy distinct areas, and which nowhere come in contact with each other, are often sterile when crossed, is one of the difficulties; but this may perhaps be overcome by the consideration that, though now isolated, they may, and often must, have been in contact at their origination. More important is the objection that natural selection could not possibly have produced the difference that often occurs between reciprocal crosses, one of these being sometimes fertile, while the other is sterile. The extremely different amounts of infertility or sterility between different species of the same genus, the infertility often bearing no proportion to the difference between the species crossed, is also an important objection. But none of these objections would have much weight if it could be clearly shown that natural selection is able to increase the infertility variations of incipient species, as it is certainly able to increase and develop all useful variations of form, structure, instincts, or habits. Ample causes of infertility have been shown to exist, in the nature of the organism and the laws of correlation; the agency of natural selection is only needed to accumulate the effects produced by these causes, and to render their final results more uniform and more in accordance with the facts that exist.
About twenty years ago I had much correspondence and discussion with Mr. Darwin on this question. I then believed that I was able to demonstrate the action of natural selection in accumulating infertility; but I could not convince him, owing to the extreme complexity of the process under the conditions which he thought most probable. I have recently returned to the question; and, with the fuller knowledge of the facts of variation we now possess, I think it may be shown that natural selection is, in some probable cases at all events, able to accumulate variations in infertility between incipient species.
The simplest case to consider, will be that in which two forms or varieties of a species, occupying an extensive area, are in process of adaptation to somewhat different modes of life within the same area. If these two forms freely intercross with each other, and produce mongrel offspring which are quite fertile inter se, then the further differentiation of the forms into two distinct species will be retarded, or perhaps entirely prevented; for the offspring of the crossed unions will be, perhaps, more vigorous on account of the cross, although less perfectly adapted to the conditions of existence than either of the pure breeds; and this would certainly establish a powerful antagonistic influence to the further differentiation of the two forms.
Now, let us suppose that a partial sterility of the hybrids between the two forms arises, in correlation with the different modes of life and the slight external or internal peculiarities that exist between them, both of which we have seen to be real causes of infertility. The result will be that, even if the hybrids between the two forms are still freely produced, these hybrids will not themselves increase so rapidly as the two pure forms; and as these latter are, by the terms of the problem, better suited to their conditions of life than are the hybrids between them, they will not only increase more rapidly, but will also tend to supplant the hybrids altogether whenever the struggle for existence becomes exceptionally severe. Thus, the more complete the sterility of the hybrids the more rapidly will they die out and leave the two parent forms pure. Hence it will follow that, if there is greater infertility between the two forms in one part of the area than the other, these forms will be kept more pure wherever this greater infertility prevails, will therefore have an advantage at each recurring period of severe struggle for existence, and will thus ultimately supplant the less infertile or completely fertile forms that may exist in other portions of the area. It thus appears that, in such a case as here supposed, natural selection would preserve those portions of the two breeds which were most infertile with each other, or whose hybrid offspring were most infertile; and would, therefore, if variations in fertility continued to arise, tend to increase that infertility. It must particularly be noted that this effect would result, not by the preservation of the infertile variations on account of their infertility, but by the inferiority of the hybrid offspring, both as being fewer in numbers, less able to continue their race, and less adapted to the conditions of existence than either of the pure forms. It is this inferiority of the hybrid offspring that is the essential point; and as the number of these hybrids will be permanently less where the infertility is greatest, therefore those portions of the two forms in which infertility is greatest will have the advantage, and will ultimately survive in the struggle for existence.