Several writers, however, not content with the indirect effects of isolation here indicated, maintain that it is in itself a cause of modification, and ultimately of the origination of new species. This was the keynote of Mr. Vernon Wollaston's essay on "Variation of Species," published in 1856, and it is adopted by the Rev. J.G. Gulick in his paper on "Diversity of Evolution under one Set of External Conditions" (Journ. Linn. Soc. Zool., vol. xi. p. 496). The idea seems to be that there is an inherent tendency to variation in certain divergent lines, and that when one portion of a species is isolated, even though under identical conditions, that tendency sets up a divergence which carries that portion farther and farther away from the original species. This view is held to be supported by the case of the land shells of the Sandwich Islands, which certainly present some very remarkable phenomena. In this comparatively small area there are about 300 species of land shells, almost all of which belong to one family (or sub-family), the Achatinellidae, found nowhere else in the world. The interesting point is the extreme restriction of the species and varieties. The average range of each species is only five or six miles, while some are restricted to but one or two square miles, and only a very few range over a whole island. The forest region that extends over one of the mountain-ranges of the island of Oahu, is about forty miles in length and five or six miles in breadth; and this small territory furnishes about 175 species, represented by 700 or 800 varieties. Mr. Gulick states, that the vegetation of the different valleys on the same side of this range is much the same, yet each has a molluscan fauna differing in some degree from that of any other. "We frequently find a genus represented in several successive valleys by allied species, sometimes feeding on the same, sometimes on different plants. In every such case the valleys that are nearest to each other furnish the most nearly allied forms; and a full set of the varieties of each species presents a minute gradation of forms between the more divergent types found in the more widely separated localities." He urges, that these constant differences cannot be attributed to natural selection, because they occur in different valleys on the same side of the mountain, where food, climate, and enemies are the same; and also, because there is no greater difference in passing from the rainy to the dry side of the mountains than in passing from one valley to another on the same side an equal distance apart. In a very lengthy paper, presented to the Linnean Society last year, on "Divergent Evolution through Cumulative Segregation," Mr. Gulick endeavours to work out his views into a complete theory, the main point of which may perhaps be indicated by the following passage: "No two portions of a species possess exactly the same average character, and the initial differences are for ever reacting on the environment and on each other in such a way as to ensure increasing divergence in each successive generation as long as the individuals of the two groups are kept from intercrossing."[49]

It need hardly be said that the views of Mr. Darwin and myself are inconsistent with the notion that, if the environment were absolutely similar for the two isolated portions of the species, any such necessary and constant divergence would take place. It is an error to assume that what seem to us identical conditions are really identical to such small and delicate organisms as these land molluscs, of whose needs and difficulties at each successive stage of their existence, from the freshly-laid egg up to the adult animal, we are so profoundly ignorant. The exact proportions of the various species of plants, the numbers of each kind of insect or of bird, the peculiarities of more or less exposure to sunshine or to wind at certain critical epochs, and other slight differences which to us are absolutely immaterial and unrecognisable, may be of the highest significance to these humble creatures, and be quite sufficient to require some slight adjustments of size, form, or colour, which natural selection will bring about. All we know of the facts of variation leads us to believe that, without this action of natural selection, there would be produced over the whole area a series of inconstant varieties mingled together, not a distinct segregation of forms each confined to its own limited area.

Mr. Darwin has shown that, in the distribution and modification of species, the biological is of more importance than the physical environment, the struggle with other organisms being often more severe than that with the forces of nature. This is particularly evident in the case of plants, many of which, when protected from competition, thrive in a soil, climate, and atmosphere widely different from those of their native habitat. Thus, many alpine plants only found near perpetual snow thrive well in our gardens at the level of the sea; as do the tritomas from the sultry plains of South Africa, the yuccas from the arid hills of Texas and Mexico, and the fuchsias from the damp and dreary shores of the Straits of Magellan. It has been well said that plants do not live where they like, but where they can; and the same remark will apply to the animal world. Horses and cattle run wild and thrive both in North and South America; rabbits, once confined to the south of Europe, have established themselves in our own country and in Australia; while the domestic fowl, a native of tropical India, thrives well in every part of the temperate zone.

If, then, we admit that when one portion of a species is separated from the rest, there will necessarily be a slight difference in the average characters of the two portions, it does not follow that this difference has much if any effect upon the characteristics that are developed by a long period of isolation. In the first place, the difference itself will necessarily be very slight unless there is an exceptional amount of variability in the species; and in the next place, if the average characters of the species are the expression of its exact adaptation to its whole environment, then, given a precisely similar environment, and the isolated portion will inevitably be brought back to the same average of characters. But, as a matter of fact, it is impossible that the environment of the isolated portion can be exactly like that of the bulk of the species. It cannot be so physically, since no two separated areas can be absolutely alike in climate and soil; and even if these are the same, the geographical features, size, contour, and relation to winds, seas, and rivers, would certainly differ. Biologically, the differences are sure to be considerable. The isolated portion of a species will almost always be in a much smaller area than that occupied by the species as a whole, hence it is at once in a different position as regards its own kind. The proportions of all the other species of animals and plants are also sure to differ in the two areas, and some species will almost always be absent in the smaller which are present in the larger country. These differences will act and react on the isolated portion of the species. The struggle for existence will differ in its severity and in its incidence from that which affects the bulk of the species. The absence of some one insect or other creature inimical to the young animal or plant may cause a vast difference in its conditions of existence, and may necessitate a modification of its external or internal characters in quite a different direction from that which happened to be present in the average of the individuals which were first isolated.

On the whole, then, we conclude that, while isolation is an important factor in effecting some modification of species, it is so, not on account of any effect produced, or influence exerted by isolation per se, but because it is always and necessarily accompanied by a change of environment, both physical and biological. Natural selection will then begin to act in adapting the isolated portion to its new conditions, and will do this the more quickly and the more effectually because of the isolation. We have, however, seen reason to believe that geographical or local isolation is by no means essential to the differentiation of species, because the same result is brought about by the incipient species acquiring different habits or frequenting a different station; and also by the fact that different varieties of the same species are known to prefer to pair with their like, and thus to bring about a physiological isolation of the most effective kind. This part of the subject will be again referred to when the very difficult problems presented by hybridity are discussed.[50]

Cases in which Isolation is Ineffective.

One objection to the views of those who, like Mr. Gulick, believe isolation itself to be a cause of modification of species deserves attention, namely, the entire absence of change where, if this were a vera causa, we should expect to find it. In Ireland we have an excellent test case, for we know that it has been separated from Britain since the end of the glacial epoch, certainly many thousand years. Yet hardly one of its mammals, reptiles, or land molluscs has undergone the slightest change, even although there is certainly a distinct difference in the environment both inorganic and organic. That changes have not occurred through natural selection, is perhaps due to the less severe struggle for existence owing to the smaller number of competing species; but, if isolation itself were an efficient cause, acting continuously and cumulatively, it is incredible that a decided change should not have been produced in thousands of years. That no such change has occurred in this, and many other cases of isolation, seems to prove that it is not in itself a cause of modification.

There yet remain a number of difficulties and objections relating to the question of hybridity, which are so important as to require a separate chapter for their adequate discussion.

FOOTNOTES:

[41] See Origin of Species, pp. 176-198.