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Charles Robert Darwin (1809–1882).  Origin of Species.
The Harvard Classics.  1909–14.
 
XII. Geographical Distribution
 
Alternate Glacial Periods in the North and South
 
 
BUT we must return to our more immediate subject. I am convinced that Forbes’s view may be largely extended. In Europe we meet with the plainest evidence of the Glacial period, from the western shores of Britain to the Oural range, and southward to the Pyrenees. We may infer from the frozen mammals and nature of the mountain vegetation, that Siberia was similarly affected. In the Lebanon, according to Dr. Hooker, perpetual snow formerly covered the central axis, and fed glaciers which rolled 4,000 feet down the valleys. The same observer has recently found great moraines at a low level on the Atlas range in N. Africa. Along the Himalaya, at points 900 miles apart, glaciers have left the marks of their former low descent; and in Sikkim, Dr. Hooker saw maize growing on ancient and gigantic moraines. Southward of the Asiatic continent, on the opposite side of the equator, we know, from the excellent researches of Dr. J. Haast and Dr. Hector, that in New Zealand immense glaciers formerly descended to a low level; and the same plants found by Dr. Hooker on widely separated mountains in this island tell the same story of a former cold period. From facts communicated to me by the Rev. W. B. Clarke, it appears also that there are traces of former glacial action on the mountains of the south-eastern corner of Australia.  1
  Looking to America; in the northern half, ice-borne fragments of rock have been observed on the eastern side of the continent, as far south as latitude 36°–37°, and on the shores of the Pacific, where the climate is now so different, as far south as lat. 46°. Erratic boulders have, also, been noticed on the Rocky Mountains. In the Cordillera of South America, nearly under the equator, glaciers once extended far below their present level. In Central Chile I examined a vast mound of detritus with great boulders, crossing the Portillo valley, which there can hardly be a doubt once formed a huge moraine; and Mr. D. Forbes informs me that he found in various parts of the Cordillera, from latitude 13° to 30° S., at about the height of 12,000 feet, deeply furrowed rocks, resembling those with which he was familiar in Norway, and likewise great masses of detritus, including grooved pebbles. Along this whole space of the Cordillera true glaciers do not exist even at much more considerable heights. Farther south on both sides of the continent, from lat. 41° to the southernmost extremity, we have the clearest evidence of former glacial action, in numerous immense boulders transported far from their parent source.  2
  From these several facts, namely from the glacial action having extended all round the northern and southern hemispheres—from the period having been in a geological sense recent in both hemispheres—from its having lasted in both during a great length of time, as may be inferred from the amount of work affected—and lastly from glaciers having recently descended to a low level along the whole line of the Cordillera, it at one time appeared to me that we could not avoid the conclusion that the temperature of the whole world had been simultaneously lowered during the Glacial period. But now Mr. Croll, in a series of admirable memoirs, has attempted to show that a glacial condition of climate is the result of various physical causes, brought into operation by an increase in the eccentricity of the earth’s orbit. All these causes tend towards the same end; but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. According to Mr. Croll, cold periods regularly occur every ten or fifteen thousand years; and these at long intervals are extremely severe, owing to certain contingencies, of which the most important, as Sir C. Lyell has shown, is the relative position of the land and water. Mr. Croll believes that the last great Glacial period occurred about 240,000 years ago, and endured with slight alterations of climate for about 160,000 years. With respect to more ancient Glacial periods, several geologists are convinced from direct evidence that such occurred during the Miocene and Eocene formations, not to mention still more ancient formations. But the most important result for us, arrived at by Mr. Croll, is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised, with the winters rendered much milder, chiefly through changes in the direction of the ocean currents. So conversely it will be with the northern hemisphere, whilst the southern passes through a Glacial period. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it; but I will first give the facts, which demand an explanation.  3
  In South America, Dr. Hooker has shown that besides many closely allied species, between forty and fifty of the flowering plants of Tierra del Fuego, forming no inconsiderable part of its scanty flora, are common to North America and Europe, enormously remote as these areas in opposite hemispheres are from each other. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. On the Organ mountains of Brazil, some few temperate European, some Antarctic, and some Andean genera were found by Gardner, which do not exist in the low intervening hot countries. On the Silla of Caraccas, the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera.  4
  In Africa, several forms characteristic of Europe and some few representatives of the flora of the Cape of Good Hope occur in the mountains of Abyssinia. At the Cape of Good Hope a very few European species, believed not to have been introduced by man, and on the mountains several representative European forms are found, which have not been discovered in the intertropical parts of Africa. Dr. Hooker has also lately shown that several of the plants living on the upper parts of the lofty island of Fernando Po and on the neighbouring Cameroon mountains, in the Gulf of Guinea, are closely related to those on the mountains of Abyssinia, and likewise to those of temperate Europe. It now also appears, as I hear from Dr. Hooker, that some of these same temperate plants have been discovered by the Rev. R. T. Lowe on the mountains of the Cape Verde Islands. This extension of the same temperate forms, almost under the equator, across the whole continent of Africa and to the mountains of the Cape Verde Archipelago, is one of the most astonishing facts ever recorded in the distribution of plants.  5
  On the Himalaya, and on the isolated mountain-ranges of the peninsula of India, on the heights of Ceylon, and on the volcanic cones of Java, many plants occur, either identically the same or representing each other, and at the same time representing plants of Europe, not found in the intervening hot lowlands. A list of the genera of plants collected on the loftier peaks of Java, raises a picture of a collection made on a hillock in Europe! Still more striking is the fact that peculiar Australian forms are represented by certain plants growing on the summits of the mountains of Borneo. Some of these Australian forms, as I hear from Dr. Hooker, extend along the heights of the peninsula of Malacca, and are thinly scattered on the one hand over India, and on the other hand as far north as Japan.  6
  On the southern mountains of Australia, Dr. F. Müller has discovered several European species; other species, not introduced by man, occur on the lowlands; and a long list can be given, as I am informed by Dr. Hooker, of European genera, found in Australia, but not in the intermediate torrid regions. In the admirable ‘Introduction to the Flora of New Zealand,’ by Dr. Hooker, analogous and striking facts are given in regard to the plants of that large island. Hence we see that certain plants growing on the more lofty mountains of the tropics in all parts of the world, and on the temperate plains of the north and south, are either the same species or varieties of the same species. It should, however, be observed that these plants are not strictly arctic forms; for, as Mr. H. C. Watson has remarked, “in receding from polar towards equatorial latitudes, the Alpine or mountain floras really become less and less Arctic.” Besides these identical and closely allied forms, many species inhabiting the same widely sundered areas, belong to genera not now found in the intermediate tropical lowlands.  7
  These brief remarks apply to plants alone; but some few analogous facts could be given in regard to terrestrial animals. In marine productions, similar cases likewise occur; as an example, I may quote a statement by the highest authority, Prof. Dana, that “It is certainly a wonderful fact that New Zealand should have a closer resemblance in its Crustacea to Great Britain, its antipode, than to any other part of the world.” Sir J. Richardson, also, speaks of the reappearance on the shores of New Zealand, Tasmania, &c., of northern forms of fish. Dr. Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe, but have not been found in the intermediate tropical seas.  8
  From the foregoing facts, namely, the presence of temperate forms on the highlands across the whole of equatorial Africa, and along the Peninsula of India, to Ceylon and the Malay Archipelago, and in a less well-marked manner across the wide expanse of tropical South America, it appears almost certain that at some former period, no doubt during the most severe part of a Glacial period, the lowlands of these great continents were everywhere tenanted under the equator by considerable number of temperate forms. At this period the equatorial climate at the level of the sea was probably about the same with that now experienced at the height of from five to six thousand feet under the same latitude, or perhaps even rather cooler. During this, the coldest period, the lowlands under the equator must have been clothed with a mingled tropical and temperate vegetation, like that described by Hooker as growing luxuriantly at the height of from four to five thousand feet on the lower slopes of the Himalaya, but with perhaps a still greater preponderance of temperate forms. So again in the mountainous island of Fernando Po, in the Gulf of Guinea, Mr. Mann found temperate European forms beginning to appear at the height of about five thousand feet. On the mountains of Panama, at the height of only two thousand feet, Dr. Seemann found the vegetation like that of Mexico, “with forms of the torrid zone harmoniously blended with those of the temperate.”  9
  Now let us see whether Mr. Croll’s conclusion that when the northern hemisphere suffered from the extreme cold of the great Glacial period, the southern hemisphere was actually warmer, throws any clear light on the present apparently inexplicable distribution of various organisms in the temperate parts of both hemispheres, and on the mountains of the tropics. The Glacial period, as measured by years, must have been very long; and when we remember over what vast spaces some naturalised plants and animals have spread within a few centuries, this period will have been ample for any amount of migration. As the cold became more and more intense, we know that arctic forms invaded the temperate regions; and, from the facts just given, there can hardly be a doubt that some of the more vigorous, dominant, and widest-spreading temperate forms invaded the equatorial lowlands. The inhabitants of these hot lowlands would at the same time have migrated to the tropical and subtropical regions of the south, for the southern hemisphere was at this period warmer. On the decline of the Glacial period, as both hemispheres gradually recovered their former temperatures, the northern temperate forms living on the lowlands under the equator, would have been driven to their former homes or have been destroyed, being replaced by the equatorial forms returning from the south. Some, however, of the northern temperate forms would almost certainly have ascended any adjoining high land, where, if sufficiently lofty, they would have long survived like the arctic forms on the mountains of Europe. They might have survived, even if the climate was not perfectly fitted for them, for the change of temperature must have been very slow, and plants undoubtedly possess a certain capacity for acclimatisation, as shown by their transmitting to their offspring different constitutional powers of resisting heat and cold.  10
  In the regular course of events the southern hemisphere would in its turn be subjected to a severe Glacial period, with the northern hemisphere rendered warmer; and then the southern temperate forms would invade the equatorial lowlands. The northern forms which had before been left on the mountains would now descend and mingle with the southern forms. These latter, when the warmth returned, would return to their former homes, leaving some few species on the mountains, and carrying southward with them some of the northern temperate forms which had descended from their mountain fastnesses. Thus, we should have some few species identically the same in the northern and southern temperate zones and on the mountains of the intermediate tropical regions. But the species left during a long time on these mountains, or in opposite hemispheres, would have to compete with many new forms and would be exposed to somewhat different physical conditions; hence they would be eminently liable to modification, and would generally now exist as varieties or as representative species; and this is the case. We must, also, bear in mind the occurrence in both hemispheres of former Glacial periods; for these will account, in accordance with the same principles, for the many quite distinct species inhabiting the same widely separated areas,  and belonging to genera not now found in the intermediate torrid zones.  11
  It is a remarkable fact strongly insisted on by Hooker in regard to America, and by Alph. de Candolle in regard to Australia, that many more identical or slightly modified species have migrated from the north to the south, than in a reversed direction. We see, however, a few southern forms on the mountains of Borneo and Abyssinia. I suspect that this preponderant migration from the north to the south is due to the greater extent of land in the north, and to the northern forms having existed in their own homes in greater numbers, and having consequently been advanced through natural selection and competition to a higher stage of perfection, or dominating power, than the southern forms. And thus, when the two sets became commingled in the equatorial regions, during the alternations to the Glacial periods, the northern forms were the more powerful and were able to hold their places on the mountains, and afterwards to migrate southward with the southern forms; but not so the southern in regard to the northern forms. In the same manner at the present day, we see that very many European productions cover the ground in La Plata, New Zealand, and to a lesser degree in Australia, and have beaten the natives; whereas extremely few southern forms have become naturalised in any part of the northern hemisphere, though hides, wool, and other objects likely to carry seeds have been largely imported into Europe during the last two or three centuries from La Plata and during the last forty or fifty years from Australia. The Neilgherrie mountains in India, however, offer a partial exception; for here, as I hear from Dr. Hooker, Australian forms are rapidly sowing themselves and becoming naturalised. Before the last great Glacial period, no doubt the intertropical mountains were stocked with endemic Alpine forms; but these have almost everywhere yielded to the more dominant forms generated in the larger areas and more efficient workshops of the north. In many islands the native productions are nearly equalled, or even outnumbered, by those which have become naturalised; and this is the first stage towards their extinction. Mountains are islands on the land, and their inhabitants have yielded to those produced within the larger areas of the north, just in the same way as the inhabitants of real islands have everywhere yielded and are still yielding to continental forms naturalised through man’s agency.  12
  The same principles apply to the distribution of terrestrial animals and of marine productions, in the northern and southern temperate zones, and on the intertropical mountains. When, during the height of the Glacial period, the ocean-currents were widely different to what they now are, some of the inhabitants of the temperate seas might have reached the equator; of these a few would perhaps at once be able to migrate southward, by keeping to the cooler currents, whilst others might remain and survive in the colder depths until the southern hemisphere was in its turn subjected to a glacial climate and permitted their further progress; in nearly the same manner as, according to Forbes, isolated spaces inhabited by arctic productions exist to the present day in the deeper parts of the northern temperate seas.  13
  I am far from supposing that all the difficulties in regard to the distribution and affinities of the identical and allied species, which now live so widely separated in the north and south, and sometimes on the intermediate mountain-ranges, are removed on the views above given. The exact lines of migration cannot be indicated. We cannot say why certain species and not others have migrated; why certain species have been modified and have given rise to new forms, whilst others have remained unaltered. We cannot hope to explain such facts, until we can say why one species and not another becomes naturalised by man’s agency in a foreign land; why one species ranges twice or thrice as far, and is twice or thrice as common, as another species within their own homes.  14
  Various special difficulties also remain to be solved; for instance, the occurrence, as shown by Dr. Hooker, of the same plants at points so enormously remote as Kerguelen Land, New Zealand, and Fuegia; but icebergs, as suggested by Lyell, may have been concerned in their dispersal. The existence at these and other distant points of the southern hemisphere, of species, which, though distinct, belong to genera exclusively confined to the south, is a more remarkable case. Some of these species are so distinct, that we cannot suppose that there has been time since the commencement of the last Glacial period for their migration and subsequent modification to the necessary degree. The facts seem to indicate that distinct species belonging to the same genera have migrated in radiating lines from a common centre; and I am inclined to look in the southern, as in the northern hemisphere, to a former and warmer period, before the commencement of the last Glacial period, when the Antarctic lands, now covered with ice, supported a highly peculiar and isolated flora. It may be suspected that before this flora was exterminated during the last Glacial epoch, a few forms had been already widely dispersed to various points of the southern hemisphere by occasional means of transport, and by the aid as halting-places, of now sunken islands. Thus the southern shores of America, Australia, and New Zealand may have become slightly tinted by the same peculiar forms of life.  15
  Sir C. Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alterations of climate throughout the world on geographical distribution. And we have now seen that Mr. Croll’s conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere, together with the admission of the slow modification of species, explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. The living waters have flowed during one period from the north and during another from the south, and in both cases have reached the equator; but the stream of life has flowed with greater force from the north than in the opposite direction, and has consequently more freely inundated the south. As the tide leaves its drift in horizontal lines, rising higher on the shores where the tide rises highest, so have the living waters left their living drift on our mountain summits, in a line gently rising from the arctic lowlands to a great altitude under the equator. The various beings thus left stranded may be compared with savage races of man, driven up and surviving in the mountain fastnesses of almost every land, which serves as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.  16
 

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