But even if there are no well-marked physical barriers, the members of a species on a continent or large island tend to fall into local groups, between which, unless the animal be of a widely ranging habit, there will be little intercrossing. Hence local varieties are apt to occur, and varieties show the first beginnings of that divergence which, if carried further and more deeply ingrained, results in the differentiation of species. Geographically, therefore, we may have either complete isolation or local segregation, and in both cases the possibility of divergence.
Another mode of segregation arises also out of geographical conditions. If variations of habits occur (and structure is closely correlated with habit) such that certain individuals take to the mountains, others to the plains or valleys; or that certain individuals take to the forests, others to the open country; the probabilities are that the forest forms will interbreed frequently with each other, but seldom with those in the open, and so with the other varieties. The conditions of forest life or mountain life being thus similar throughout a large area, and life being through elimination slowly but surely adapted to its environment, there might thus arise two distinct varieties scattered throughout the length and breadth of the area, the one inhabiting the mountains, the other the forests. In illustration of this mode of segregation, we may take the case of two species of rats which have recently been found by Mr. C. M. Woodford on one of the Solomon Islands. These two quite distinct species are regarded by Mr. Oldfield Thomas as slightly modified descendants of one parent species, the modifications resulting from the fact that of this original species some individuals have adopted a terrestrial, others an arboreal life, and their respective descendants have been modified accordingly. Thus Mus rex lives in trees, has broad foot-pads, and a long rasp-like, probably semi-prehensile, tail; while Mus imperator lives on the ground, has smaller pads, and a short, smooth tail. The segregation of these two species has probably been effected by the difference of their mode of life, and each has been adapted to its special environment through the elimination of those individuals which were not in harmony with the condition of their life. It is probable that this mode of segregation has been an important one. And it is clear that in many cases competition would be a co-operating factor in this process, weaker organisms being forced into otherwise uncongenial habitats through the stress of competitive elimination, the weaker forms not perishing, but being eliminated from more favoured areas.
Protective coloration may also be a means of segregation. A species of insects having no protective resemblance might vary in two directions—in the direction of green tints, assimilating their hue to that of vegetation; and in the direction of sandy or dull earthy colours, assimilating them to the colour of the soil. In the one variety elimination would weed out all but the green forms, and these would be left to intercross. In the other variety, green forms would be eliminated, dull-brown forms being left to interbreed. Stragglers from one group into the other would stand a chance of elimination before interbreeding was effected.[AO]
In the case of birds whose freedom of flight gives them a wide range, sometimes almost a world-wide range, it would seem at first sight that their facilities for interbreeding and intercrossing are so great that divergence is well-nigh impossible. And yet the examples of divergence I cited from Mr. Wallace were taken from birds, and it is well known that divergence is particularly well shown in this class. But when the habits of birds are studied attentively, it is found that, wide as is their range, their breeding area is often markedly restricted. The sanderling and knot range freely during the winter throughout the Northern hemisphere; but their breeding area is restricted to the north polar region. The interbreeding within this area keeps the species one and homogeneous, notwithstanding its wide range, and, at the same time, prevents intercrossing with allied species with different breeding-grounds.
Another most important mode of segregation among animals arises out of habitual or instinctive preferences. Where varieties are formed there is a tendency for like to breed with like. In the Falkland Islands the differently coloured herds of cattle, all descended from the same stock, keep separate, and interbreed with each other, but not with individuals outside their own colour-caste. If two flocks of merino sheep and heath sheep be mixed together, they do not interbreed. In the Forest of Dean and in the New Forest, the dark and pale coloured herds of fallow deer have never been known to intermingle.[AP] Here we have a case of selective segregation through preferential mating, and may find therein the basis of sexual selection in its higher ranges as advocated by Darwin.
The question of sexual selection will, however, be briefly considered in the chapter on "Organic Evolution." At present what we have to notice is that, through preferential mating, segregation is effected. The forms that interbreed have a distinguishing colour. From this it is but a step to the possession, not merely of a distinguishing colour, but of distinguishing colour-markings. Hence, through preferential mating, may arise those special markings which so frequently distinguish allied species. They not only enable us to recognize species as distinct, but enable the species which possess them to recognize the members of their own kind. Mr. Wallace calls these diacritical marks recognition-marks, and gives many illustrative examples.[AQ] They are especially noticeable in gregarious animals and in birds which congregate in flocks or which migrate together. Mr. Wallace considers that they "have in all probability been acquired in the process of differentiation for the purpose of checking the intercrossing of allied forms;" for "one of the first needs of a new species would be to keep separate from its nearest allies, and this could be more readily done by some easily seen external mark of difference." This language seems, however, to savour of teleology (that pitfall of the evolutionist). The cart is placed before the horse. The recognition-marks were, I believe, not produced to prevent intercrossing, but intercrossing has been prevented because of preferential mating between individuals possessing special recognition-marks. To miss this point is to miss an important segregation-factor. Undoubtedly, other tendencies co-operate in maintaining the standard of the recognition-marks. Stragglers who failed in the matter of recognition would get separated from their fellows, and stand a greater chance of elimination by enemies; young who failed in this respect would be in like condemnation. Still, I cannot doubt that the foundations of recognition-marks were laid in preferential mating, and that in this we have an important factor in segregation.
We may here note, in passing, as also arising out of preference, how the selection of flowers by insects may lead to segregation; for insects seem often to have habitual or instinctive colour-preferences. Flowers of similar colour would be thus cross-fertilized, but would not intercross with those of different colour, whence colour-varieties might arise. It is important to note that in these cases there is a psychological factor in evolution.
We have so far assumed that intercrossing of parents and interblending of their characters in the offspring always go together. This, we must now notice, is not always the fact. If a blue-eyed Saxon marry a dark-eyed Italian, the children will have blue eyes or dark eyes, not eyes of an intermediate tint. The characters do not interblend. The ancon, or otter-sheep, a breed with a long body and short, bandy legs, appeared in Massachusetts as a chance sport in a single lamb. The offspring of this ram were either ancons or ordinary sheep. The ancon characters did not blend. Hence for a time a definite breed was maintained. We may call this mode of isolation isolation by exclusive inheritance.
A further mode of isolation or segregation, for which Mr. Romanes[AR] claims a foremost, indeed, the foremost, place, is physiological isolation as due to differential fertility. One among the many variations to which organisms are subject is a variation in fertility, which may reach the climax of absolute sterility. But it is clear that a sterile variation carries with it its own death-warrant, since the sterile individual leaves no descendants to inherit its peculiarity. Relative infertility, too, unless it chances to be correlated with some unusual excellence, would be no advantage, would be transmitted to few descendants, and would tend to be extinguished. The same is not true, however, of differential fertility. "It is by no means rare," said Darwin,[AS] "to find certain males and females which will not breed together, though both are known to be perfectly fertile with other males and females." Mr. Romanes assumes, as a starting-point, the converse of this, namely, that certain males and females will breed together, though they are infertile with all other members of the species.
Suppose, then, a variety to arise which is perfectly fertile within the limits of the varietal form, but imperfectly fertile or infertile with the parent species. Such a variety would have to run the risks of those ill effects which, as Darwin showed,[AT] are attendant upon close interbreeding. But Mr. Wallace points out[AU] that these ill effects may not be so marked under nature as they are under domestication. Suppose, then, that it escapes these ill effects. In this case, Mr. Romanes urges, it would neither be swamped by intercrossing nor die out on account of sterility. But although it could not be swamped by intercrossing, still, if it arose sporadically, here a case, there a case, and so on, the chances would be enormously against the perpetuation of the variety, unless some co-operating mode of segregation aided in bringing together the varying individuals. If, for example, there were a segregation of these variants in a particular habitat—all the variants meeting in some definite locality for breeding purposes; or if there were a further segregation through mutual preferences; or if, again, there were a further segregation in time the variety might obtain a firm footing. But without these co-operating factors it is clear that if one male and one female in a hundred individuals varied in this particular way, the chances would be at least forty-nine to one against their happening to mate together.