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CHANGES OF CLIMATE WHICH HAVE INFLUENCED THE DISPERSAL OF ORGANISMS: THE GLACIAL EPOCH

Proofs of the Recent Occurrence of a Glacial Epoch—Moraines—Travelled Blocks—Glacial Deposits of Scotland: the "Till"—Inferences from the Glacial Phenomena of Scotland—Glacial Phenomena of North America—Effects of the Glacial Epoch on Animal Life—Warm and Cold Periods—Palæontological Evidence of Alternate Cold and Warm Periods—Evidence of Interglacial Warm Periods on the Continent and in North America—Migrations and Extinctions of Organisms caused by the Glacial Epoch.

We have now to consider another set of physical revolutions which have profoundly affected the whole organic world. Besides the wonderful geological changes to which, as we have seen, all continents have been exposed, and which must, with extreme slowness, have brought about the greater features of the dispersal of animals and plants throughout the world, there has been also a long succession of climatal changes, which, though very slow and gradual when measured by centuries, may have sometimes been rapid as compared with the slow march of geological mutations.

These climatal changes may be divided into two classes, which have been thought to be the opposite phases of the same great phenomenon—cold or even glacial epochs in the Temperate zones on the one hand, and mild or even warm periods extending into the Arctic regions on the other. The evidence for both these changes having occurred is conclusive; and as they must be taken account of whenever we endeavour to explain the past migrations and actual distribution of the animal world, a brief outline of the more important facts and of the conclusions they lead to must be here given.

Proofs of the Recent Occurrence of a Glacial Epoch.—The phenomena that prove the recent occurrence of glacial epochs in the temperate regions are exceedingly varied, and extend over very wide areas. It will be well therefore to state, first, what those facts are as exhibited in our own country, referring afterwards to similar phenomena in other parts of the world.

Perhaps the most striking of all the evidences of glaciation are the grooved, scratched, or striated rocks. These occur abundantly in Scotland, Cumberland, and North Wales, and no rational explanation of them has ever been given except that they were formed by glaciers. In many valleys, as, for instance, that of Llanberris in North Wales, hundreds of examples may be seen, consisting of deep grooves several inches wide, smaller furrows, and striæ of extreme fineness wherever the rock is of sufficiently close and hard texture to receive such marks. These grooves or scratches are often many yards long, they are found in the bed of the valley as well as high up on its sides, and they are almost all without exception in one general direction—that of the valley itself, even though the particular surface they are upon slopes in another direction. When the native covering of turf is cleared away from the rock the grooves and striæ are often found in great perfection, and there is reason to believe that such markings cover, or have once covered, a large part of the surface. Accompanying these markings we find another, hardly less curious phenomenon, the rounding off or planing down of the hardest rocks to a smooth undulating surface. Hard crystalline schists with their strata nearly vertical, and which one would expect to find exposing jagged edges, are found ground off to a perfectly smooth but never to a flat surface. These rounded surfaces are found not only on single rocks but over whole valleys and mountain sides, and form what are termed roches moutonnées, from their often having the appearance at a distance of sheep lying down.

Now these two phenomena are actually produced by existing glaciers, while there is no other known or even conceivable cause that could have produced them. Whenever the Swiss glaciers retreat a little, as they sometimes do, the rocks in the bed of the valley they have passed over are found to be rounded, grooved, and striated just as are those of Wales and Scotland. The two sets of phenomena are so exactly identical that no one who has ever compared them can doubt that they are due to the same causes. But we have further and even more convincing evidence. Glaciers produce many other effects besides these two, and whatever effects they produce in Switzerland, in Norway, or in Greenland, we find examples of similar effects having been produced in our own country. The most striking of these are moraines and travelled blocks.

Moraines.—Almost every existing glacier carries down with it great masses of rock, stones, and earth, which fall on its surface from the precipices and mountain slopes which hem it in, or the rocky peaks which rise above it. As the glacier slowly moves downward, this débris forms long lines on each side, or on the centre whenever two glacier-streams unite, and is deposited at its termination in a huge mound called the terminal moraine. The decrease of a glacier may often be traced by successive old moraines across the valley up which it has retreated. When once seen and examined, these moraines can always be distinguished almost at a glance. Their position is most remarkable, having no apparent natural relation to the form of the valley or the surrounding slopes, so that they look like huge earthworks formed by man for purposes of defence. Their composition is equally peculiar, consisting of a mixture of earth and rocks of all sizes, usually without any arrangement, the rocks often being huge angular masses just as they had fallen from the surrounding precipices. Some of these rock masses often rest on the very top of the moraine in positions where no other natural force but that of ice could have placed them. Exactly similar mounds are found in the valleys of North Wales and Scotland, and always where the other evidences of ice-action occur abundantly.

A GLACIER WITH MORAINES.

Travelled Blocks.—The phenomenon of travelled or perched blocks is also a common one in all glacier countries, marking out very clearly the former extent of the ice. When a glacier fills a lateral valley, its foot will sometimes cross over the main valley and abut against its opposite slope, and it will deposit there some portion of its terminal moraine. But in these circumstances the end of the glacier not being confined laterally will spread out, and the moraine matter will be distributed over a large surface, so that the only well-marked token of its presence will be the larger masses of rock that may have been brought down. Such blocks are found abundantly in many of the districts of our own country where other marks of glaciation exist, and they often rest on ridges or hillocks over which the ice has passed, these elevations consisting sometimes of loose material and sometimes of rock different from that of which the blocks are composed. These are called travelled blocks, and can almost always be traced to their source in one of the higher valleys from which the glacier descended. Some of the most remarkable examples of such travelled blocks are to be found on the southern slopes of the Jura. These consist of enormous angular blocks of granite, gneiss, and other crystalline rocks, quite foreign to the Jura mountains, but exactly agreeing with those of the Alpine range fifty miles away across the great central valley of Switzerland. One of the largest of these blocks is forty feet diameter, and is situated 900 feet above the level of the Lake of Neufchatel. These blocks have been proved by Swiss geologists to have been brought by the ancient glacier of the Rhone which was fed by the whole Alpine range from Mont Blanc to the Furka Pass. This glacier must have been many thousand feet thick at the mouth of the Rhone valley near the head of the Lake of Geneva, since it spread over the whole of the great valley of Switzerland, extending from Geneva to Neufchatel, Berne, and Soleure, and even on the flanks of the Jura, reached a maximum height of 2,015 feet above the valley. The numerous blocks scattered over the Jura for a distance of about a hundred miles vary considerably in the material of which they are composed, but they are found to be each traceable to a part of the Alps corresponding to their position, on the theory that they have been brought by a glacier spreading out from the Rhone valley. Thus, all the blocks situated to the east of a central point G (see map) can be traced to the eastern side of the Rhone valley (l e d), while those found towards Geneva have all come from the west side (p h). It is also very suggestive that the highest blocks on the Jura at G have come from the eastern shoulder of Mont Blanc in the direct line h B F G. Here the glacier would naturally preserve its greatest thickness, while as it spread out eastward and westward it would become thinner. We accordingly find that the travelled blocks on either side of the central point become lower and lower, till near Soleure and Geneva they are not more than 500 feet above the valley. The evidence is altogether so conclusive that, after personal examination of the district in company with eminent Swiss geologists, Sir Charles Lyell gave up the view he had first adopted—that the blocks had been carried by floating ice during a period of submergence—as altogether untenable.^[33]

MAP SHOWING THE COURSE OF THE ANCIENT GLACIER OF THE RHONE AND THE DISTRIBUTION OF ERRATIC BLOCKS ON THE JURA.

The phenomena now described demonstrate a change of climate sufficient to cover all our higher mountains with perpetual snow, and fill the adjacent valleys with huge glaciers at least as extensive as those now found in Switzerland. But there are other phenomena, best developed in the northern part of our islands, which show that even this state of things was but the concluding phase of the glacial period, which, during its maximum development, must have reduced the northern half of our island to a condition only to be paralleled now in Greenland and the Antarctic regions. As few persons besides professed geologists are acquainted with the weight of evidence for this statement, and as it is most important for our purpose to understand the amount of the climatal changes the northern hemisphere has undergone, I will endeavour to make the evidence intelligible, referring my readers for full details to Dr. James Geikie's descriptions and illustrations.^[34]

Glacial Deposits of Scotland: the "Till."—Over almost all the lowlands and in most of the highland valleys of Scotland there are immense superficial deposits of clay, sand, gravel, or drift, which can be traced more or less directly to glacial action. Some of these are moraine matter, others are lacustrine deposits, while others again have been formed or modified by the sea during periods of submergence. But below them all, and often resting directly on the rock-surface, there are extensive layers of a very tough clayey deposit known as "till." The till is very fine in texture, very tenacious, and often of a rock-like hardness. It is always full of stones, all of which are of rude form, but with the angles rubbed off, and almost always covered with scratches and striæ often crossing each other in various directions. Sometimes the stones are so numerous that there seems to be only just enough clay to unite them into a solid mass, and they are of all sizes, from mere grit up to rocks many feet in diameter. The "till" is found chiefly in the low-lying districts, where it covers extensive areas sometimes to a depth of a hundred feet; while in the highlands it occurs in much smaller patches, but in some of the broader valleys forms terraces which have been cut through by the streams. Occasionally it is found as high as two thousand feet above the sea, in hollows or hill-sides, where it seems to have been protected from denudation.

The "till" is totally unstratified, and the rock-surfaces on which it almost always rests are invariably worn smooth, and much grooved and striated when the rock is hard; but when it is soft or jointed, it frequently shows a greatly broken surface. Its colour and texture, and the nature of the stones it contains, all correspond to the character of the rock of the district where it occurs, so that it is clearly a local formation. It is often found underneath moraines, drift, and other late glacial deposits, but never overlies them (except in special cases to be hereafter referred to), so that it is certainly an earlier deposit.

Throughout Scotland, where "till" is found, the glacial striæ, perched blocks, roches moutonnées, and other marks of glacial action, occur very high up the mountains to at least 3,000 and often even to 3,500 feet above the sea, while all lower hills and mountains are rounded and grooved on their very summits; and these grooves always radiate outwards from the highest peaks and ridges towards the valleys or the sea.

Inferences from the Glacial Phenomena of Scotland.—Now all these phenomena taken together render it certain that the whole of Scotland was once buried in a vast sea of ice, out of which only the highest mountains raised their summits. There is absolutely no escape from this conclusion; for the facts which lead to it are not local—found only in one spot or one valley—but general throughout the entire length and breadth of Scotland; and are besides supported by such a mass of detailed corroborative evidence as to amount to absolute demonstration. The weight of this vast ice-sheet, at least three thousand feet in maximum thickness, and continually moving seaward with a slow grinding motion like that of all existing glaciers, must have ground down the whole surface of the country, especially all the prominences, leaving the rounded rocks as well as the grooves and striæ we still see marking the direction of its motion. All the loose stones and rock-masses which lay on the surface would be pressed into the ice; the harder blocks would serve as scratching and grinding tools, and would thus themselves become rounded, scratched, and striated, as we see them, while all the softer masses would be ground up into impalpable mud along with the material planed off the rocky projections of the country, leaving them in the condition of roches moutonnées.

The peculiar characters of the "till," its fineness and tenacity, correspond closely with the fine matter which now issues from under all glaciers, making the streams milky white, yellow, or brown, according to the nature of the rock. The sediment from such water is a fine unctuous, sticky deposit, only needing pressure to form it into a tenacious clay; and when "till" is exposed to the action of water, it dissolves into a similar soft, sticky, unctuous mud. The present glaciers of the Alps, being confined to valleys which carry off a large quantity of drainage water, lose this mud perhaps as rapidly as it is formed; but when the ice covered the whole country, there was comparatively little drainage water, and thus the mud and stones collected in vast compact masses in all the hollows, and especially in the lower flat valleys, so that, when the ice retreated, the whole country was more or less covered with it. It was then, no doubt, rapidly denuded by rain and rivers, but, as we have seen, great quantities remain to the present day to tell the tale of its wonderful formation.^[35]

There is good evidence that, when the ice was at its maximum, it extended not only over the land, but far out to sea, covering all the Scottish islands, and stretching in one connected sheet to Ireland and Wales, where all the evidences of glaciation are as well marked as in Scotland, though the ice did not of course attain quite so great a thickness.^[36]

It is evident that the change of climate requisite to produce such marvellous effects in the British Isles could not have been local, and we accordingly find strikingly similar proofs that Scandinavia and all northern Europe have also been covered with a huge ice-sheet; while we have already seen that a similar gigantic glacier buried the Alps, carrying granitic blocks to the Jura, where it deposited them at a height of 3,450 feet above the sea; while to the south, in the plains of Italy, the terminal moraines left by the retreating glaciers have formed extensive hills, those of Ivrea the work of the great glacier from the Val d'Aosta being fifteen miles across and from 700 to 1,500 feet high.

Glacial Phenomena in North America.—In North America the marks of glaciation are even more extensive and striking than in Europe, stretching over the whole of Canada and to the south of the great lakes as far as latitude 39°. There is, in all these countries, a wide-spread deposit like the "till" of Scotland, produced by the grinding of the great ice-sheet when it was at its maximum thickness; and also extensive beds of moraine-matter, true moraines, and travelled blocks, left by the glaciers as they retreated towards the mountains and finally withdrew into the upland valleys. There are, also, in Britain, Scandinavia, and North America, proofs of the submersion of the land beneath the sea to a depth of upwards of a thousand feet; but this is a subject we need not here enter upon, as our special object is to show the reality and amount of that wonderful and comparatively recent change of climate termed the glacial epoch.

Many persons, even among scientific men, who have not given much attention to the question, look upon the whole subject of the glacial epoch as a geological theory made to explain certain phenomena which are otherwise a puzzle; and they would not be much surprised if they were some day told that it was all a delusion, and that Mr. So-and-so had explained the whole thing in a much more simple way. It is to prevent my readers being imposed upon by any such statements or doubts, that I have given this very brief and imperfect outline of the nature, extent, and completeness of the evidence on which the existence of the glacial epoch depends. There is perhaps no great conclusion in any science which rests upon a surer foundation than this; and if we are to be guided by our reason at all in deducing the unknown from the known, the past from the present, we cannot refuse our assent to the reality of the glacial epoch of the northern hemisphere in all its more important features.

Effects of the Glacial Epoch on Animal Life: Warm and Cold Periods. —It is hardly necessary to point out what an important effect this great climatal cycle must have had upon all living things. When an icy mantle crept gradually over much of the northern hemisphere till large portions of Europe and North America were reduced to the condition of Greenland now, the greater part of the animal life must have been driven southward, causing a struggle for existence which must have led to the extermination of many forms, and the migration of others into new areas. But these effects must have been greatly multiplied and intensified if, as there is very good reason to believe, the glacial epoch itself—or at least the earlier and later phases of it—consisted of two or more alternations of warm and cold periods.

The evidence that such was the case is very remarkable. The "till," as we have seen, could only have been formed when the country was entirely buried under a large ice-sheet of enormous thickness, and when it must therefore have been, in all the parts so covered, almost entirely destitute of animal and vegetable life. But in several places in Scotland fine layers of sand and gravel with beds of peaty matter, have been found resting on "till" and again covered by "till." Sometimes these intercalated beds are very thin, but in other cases they are twenty or thirty feet thick, and in them have been found remains of the extinct ox, the Irish elk, the horse, reindeer and mammoth. Here we have evidence of two distinct periods of intense cold, and an intervening milder period sufficiently prolonged for the country to become covered with vegetation and stocked with animal life. In some districts borings have proved the existence of no less than four distinct formations of "till" separated from each other by beds of sand from two to twenty feet in thickness.^[37] Facts of a similar nature have been observed in other parts of our islands. In the east of England, Mr. Skertchly (of the Geological Survey) enumerates four distinct boulder clays with intervening deposits of gravels and sands.^[38] Mr. Searles V. Wood, Jun., classes the most recent (Hessle) boulder clay as "post-glacial," but he admits an intervening warmer period, characterised by southern forms of mollusca and insects, after which glacial conditions again prevailed with northern types of mollusca.^[39] Elsewhere he says: "Looking at the presence of such fluviatile mollusca as Cyrena fluminalis and Unio littoralis and of such mammalia as the hippopotamus and other great pachyderms, and of such a littoral Lusitanian fauna as that of the Selsea bed where it is mixed up with the remains of some of those pachyderms, as well as of some other features, it has seemed to me that the climate of the earlier part of the post-glacial period in England was possibly even warmer than our present climate; and that it was succeeded by a refrigeration sufficiently severe to cause ice to form all round our coasts, and glaciers to accumulate in the valleys of the mountain districts; and that this increased severity of climate was preceded, and partially accompanied, by a limited submergence, which nowhere apparently exceeded 300 feet, and reached that amount only in the northern counties of England."^[40] This decided admission of an alternation of warm and cold climates since the height of the glacial epoch by so cautious a geologist as Mr. Wood is very important, as is his statement of an accompanying depression of the land, accompanying the increased cold, because many geologists maintain that a greater elevation of the land is the true and sufficient explanation of glacial periods.

Further evidence of this alternation is found both in the Isle of Man and in Ireland, where two distinct boulder clays have been described with intervening beds of gravels and sands.

Palæontological Evidence of Alternate Cold and Warm Periods.—Especially suggestive of a period warmer than the present, immediately following glacial conditions, is the occurrence of the hippopotamus in caves, brick-earths, and gravels of palæolithic age. Entire skeletons of this animal have been found at Leeds in a bed of dark blue clay overlaid by gravel. Further north at Kirkdale cave, in N. Lat. 54° 15′, remains of the hippopotamus occur abundantly along with those of the Elephas antiquus, Rhinoceros hemitœchus, reindeer, bear, horse, and other quadrupeds, and with countless remains of the hyænas which devoured them; while it has also been found in cave deposits in Glamorganshire, at Durdham Down near Bristol, and in the post-Pliocene drifts of England and France.

The fact of the hippopotamus having lived at 54° N. Lat. in England immediately after the glacial period seems quite inconsistent with a mere gradual amelioration of climate from that time till the present day. The entirely tropical distribution of the existing animal and the large quantity of vegetable food which it requires both indicate a much warmer climate than now prevails in any part of Europe. The problem, however, is complicated by the fact that, both in the cave-deposits and river gravels, its remains are often found associated with those of animals that imply a cold climate, such as the reindeer, the mammoth, or the woolly rhinoceros. At this time the British Isles were joined to the Continent, and a great river formed by the union of the Rhine, the Elbe and all the eastern rivers of England, flowed northward through what is now the German Ocean. The hippopotamus appears to have been abundant in Central Europe before the glacial epoch, but during the height of the cold was probably driven to the south of France, whence it may have returned by way of the Rhone valley, some of the tributaries of that river approaching those of the Rhine within a mile or two a little south-west of Mulhausen, whence it would easily reach Yorkshire. Professor Boyd Dawkins supposes that at this time our summers were warm, as in Middle Asia and the United States, while the winters were cold, and that the southern and northern animals migrated to and fro over the great plains which extended from Britain to the Continent. The following extract indicates how such a migration was calculated to bring about the peculiar association of sub-tropical and arctic forms.

"It must not, however, be supposed that the southern animals migrated from the Mediterranean area as far north as Yorkshire in the same year, or the northern as far south as the Mediterranean. There were, as we shall see presently, secular changes of climate in Pleistocene Europe, and while the cold was at its maximum the arctic animals arrived at the southern limit, and while it was at its minimum the spotted hyæna and hippopotamus and other southern animals roamed to their northern limit. Thus every part of the middle zone has been successively the frontier between the northern and southern groups, and consequently their remains are mingled together in the caverns and river-deposits, under conditions which prove them to have been contemporaries in the same region. In some of the caverns, such as that of Kirkdale, the hyæna preyed upon the reindeer at one time of the year and the hippopotamus at another. In this manner the association of northern and southern animals may be explained by their migration according to the seasons; and their association over so wide an area as the middle zone, by the secular changes of climate by which each part of the zone in turn was traversed by the advancing and retreating animals."^[41]

When we consider that remains of the hippopotamus have been found in the caves of North Wales and Bristol as well as in those of Yorkshire, associated in all with the reindeer and in some with the woolly rhinoceros or the mammoth, and that the animal must have reached these localities by means of slow-flowing rivers or flooded marshes by very circuitous routes, we shall be convinced that these long journeys from the warmer regions of South Europe could not have been made during the short summers of the glacial period. Thus the very existence of such an animal in such remote localities closely associated with those implying almost an arctic winter climate appears to afford a strong support to the argument for the existence of warm inter-glacial or post-glacial periods.

Evidence of Interglacial Warm Periods on the Continent and in North America.—Besides the evidence already adduced from our own islands, many similar facts have been noted in other countries. In Switzerland two glacial periods are distinctly recognised, between which was a warm period when vegetation was so luxuriant as to form beds of lignite sufficiently thick to be worked for coal. The plants found in these deposits are similar to those now inhabiting Switzerland—pines, oaks, birches, larch, etc., but numerous animal remains are also found, showing that the country was then inhabited by an elephant (Elephas antiquus), a rhinoceros (Rhinoceros megarhinus), the urus (Bos primigenius), the red deer (Cervus elephas), and the cave-bear, (Ursus spelœus); and there were also abundance of insects.^[42]

In Sweden also there are two "tills," the lower one having been in places partly broken up and denuded before the upper one was deposited, but no interglacial deposits have yet been found. In North America more complete evidence has been obtained. On the shores of Lake Ontario sections are exposed showing three separate beds of "till" with intervening stratified deposits, the lower one of which has yielded many plant remains and fresh-water organisms. These deposits are seen to extend continuously for more than nine miles, and the fossiliferous interglacial beds attain a thickness of 140 feet. Similar beds have been discovered near Cleveland, Ohio, consisting, first of "till" at the lake-level, secondly of about 48 feet of sand and loam, and thirdly of unstratified "till" full of striated stones—six feet thick.^[43] On the other side of the continent, in British Columbia, Mr. G. M. Dawson, geologist to the North American Boundary Commission, has discovered similar evidence of two glaciations divided from each other by a warm period.

This remarkable series of observations, spread over so wide an area, seems to afford ample proof that the glacial epoch did not consist merely of one process of change, from a temperate to a cold and arctic climate, which having reached a maximum, then passed slowly and completely away; but that there were certainly two, and probably several more alternations of arctic and temperate climates.

It is evident, however, that if there have been, not two only, but a series of such alternations of climate, we could not possibly expect to find more than the most slender indications of them, because each succeeding ice-sheet would necessarily grind down or otherwise destroy much of the superficial deposits left by its predecessors, while the torrents that must always have accompanied the melting of these huge masses of ice would wash away even such fragments as might have escaped the ice itself. It is a fortunate thing therefore, that we should find any fragments of these interglacial deposits containing animal and vegetable remains; and just as we should expect, the evidence they afford seems to show that the later phase of the cold period was less severe than the earlier. Of such deposits as were formed on land during the coming on of the glacial epoch when it was continually increasing in severity hardly a trace has been preserved, because each succeeding extension of the ice being greater and thicker than the last, destroyed what had gone before it till the maximum was reached.

Migrations and Extinction of Organisms caused by the Glacial Epoch.—Our last glacial epoch was accompanied by at least two considerable submergences and elevations of the land, and there is some reason to think, as we have already explained, that the two classes of phenomena are connected as cause and effect. We can easily see how such repeated submergences and elevations would increase and aggravate the migrations and extinctions that a glacial epoch is calculated to produce. We can therefore hardly fail to be right in attributing the wonderful changes in animal and vegetable life that have occurred in Europe and N. America between the Miocene Period and the present day, in part at least, to the two or more cold epochs that have probably intervened. These changes consist, first, in the extinction of a whole host of the higher animal forms, and secondly, in a complete change of types due to extinction and migration, leading to a much greater difference between the vegetable and animal forms of the eastern and western hemisphere than before existed. Many large and powerful mammalia lived in our own country in Pliocene times and apparently survived a part of the glacial epoch; but when it finally passed away they too had disappeared, some having become altogether extinct while others continued to exist in more southern lands. Among the first class are the sabre-toothed tiger, the extinct Siberian camel (Merycotherium), three species of elephant, two of rhinoceros, two bears, five species of deer, and the gigantic beaver; among the latter are the hyæna, bear, and lion, which are considered to be only varieties of those which once inhabited Britain. Down to Pliocene times the flora of Europe was very similar to that which now prevails in Eastern Asia and Eastern North America. The late Professor Asa Gray has pointed out that hundreds of species of trees and shrubs of peculiar genera which still flourish in those countries are now completely wanting in Europe, and there is good reason to believe that these were exterminated during the glacial period, being cut off from a southern migration, first by the Alps, and then by the Mediterranean; whereas in eastern America and Asia the mountain chains run in a north and south direction, and there is nothing to prevent the flora from having been preserved by a southward migration into a milder region.^[44]

Our next two chapters will be devoted to a discussion of the causes which brought about the glacial epoch, and that still more extraordinary climatic phenomenon—the mild climate and luxuriant vegetation of the Arctic zone. If my readers will follow me with the care and attention so difficult and interesting a problem requires and deserves, they will find that I have grappled with all the more important facts which have to be accounted for, and have offered what I believe is the first complete and sufficient explanation of them. The important influence of climatal changes on the dispersal of animals and plants is a sufficient justification for introducing such a discussion into the present volume.