Thus we can understand the identity of many plants at points so immensely remote as the mountains of the United States and those of Europe. We can thus also understand the fact that the Alpine plants of each mountain-range are more especially related to the arctic forms living due north or nearly due north of them: for the first migration when the cold came on, and the re-migration on the returning warmth, would generally have been due south and north. The Alpine plants, for example, of Scotland, as remarked by Mr. H.C. Watson, and those of the Pyrenees, as remarked by Ramond, are more especially allied to the plants of northern Scandinavia; those of the United States to Labrador; those of the mountains of Siberia to the arctic regions of that country. These views, grounded as they are on the perfectly well-ascertained occurrence of a former Glacial period, seem to me to explain in so satisfactory a manner the present distribution of the Alpine and Arctic productions of Europe and America, that when in other regions we find the same species on distant mountain-summits, we may almost conclude, without other evidence, that a colder climate formerly permitted their migration across the intervening lowlands, now become too warm for their existence.
As the arctic forms moved first southward and afterwards backward to the north, in unison with the changing climate, they will not have been exposed during their long migrations to any great diversity of temperature; and as they all migrated in a body together, their mutual relations will not have been much disturbed. Hence, in accordance with the principles inculcated in this volume, these forms will not have been liable to much modification. But with the Alpine productions, left isolated from the moment of the returning warmth, first at the bases and ultimately on the summits of the mountains, the case will have been somewhat different; for it is not likely that all the same arctic species will have been left on mountain ranges far distant from each other, and have survived there ever since; they will also, in all probability, have become mingled with ancient Alpine species, which must have existed on the mountains before the commencement of the Glacial epoch, and which during the coldest period will have been temporarily driven down to the plains; they will, also, have been subsequently exposed to somewhat different climatical influences. Their mutual relations will thus have been in some degree disturbed; consequently they will have been liable to modification; and they have been modified; for if we compare the present Alpine plants and animals of the several great European mountain ranges, one with another, though many of the species remain identically the same, some exist as varieties, some as doubtful forms or sub-species and some as distinct yet closely allied species representing each other on the several ranges.
In the foregoing illustration, I have assumed that at the commencement of our imaginary Glacial period, the arctic productions were as uniform round the polar regions as they are at the present day. But it is also necessary to assume that many sub-arctic and some few temperate forms were the same round the world, for some of the species which now exist on the lower mountain slopes and on the plains of North America and Europe are the same; and it may be asked how I account for this degree of uniformity of the sub-arctic and temperate forms round the world, at the commencement of the real Glacial period. At the present day, the sub-arctic and northern temperate productions of the Old and New Worlds are separated from each other by the whole Atlantic Ocean and by the northern part of the Pacific. During the Glacial period, when the inhabitants of the Old and New Worlds lived further southwards than they do at present, they must have been still more completely separated from each other by wider spaces of ocean; so that it may well be asked how the same species could then or previously have entered the two continents. The explanation, I believe, lies in the nature of the climate before the commencement of the Glacial period. At this, the newer Pliocene period, the majority of the inhabitants of the world were specifically the same as now, and we have good reason to believe that the climate was warmer than at the present day. Hence, we may suppose that the organisms which now live under latitude 60 degrees, lived during the Pliocene period further north, under the Polar Circle, in latitude 66-67 degrees; and that the present arctic productions then lived on the broken land still nearer to the pole. Now, if we look at a terrestrial globe, we see under the Polar Circle that there is almost continuous land from western Europe through Siberia, to eastern America. And this continuity of the circumpolar land, with the consequent freedom under a more favourable climate for intermigration, will account for the supposed uniformity of the sub-arctic and temperate productions of the Old and New Worlds, at a period anterior to the Glacial epoch.