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GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGSby@charlesdarwin
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GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGS

by Charles DarwinJanuary 20th, 2023
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We are thus brought to the question which has been largely discussed by naturalists, namely, whether species have been created at one or more points of the earth’s surface. Undoubtedly there are many cases of extreme difficulty in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points where now found. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. He who rejects it rejects the vera causa of ordinary generation with subsequent migration, and calls in the agency of a miracle. It is universally admitted that in most cases the area inhabited by a species is continuous; and that, when a plant or animal inhabits two points so distant from each other, or with an interval of such a nature, that the space could not have been easily passed over by migration, the fact is given as something remarkable and exceptional. The incapacity of migrating across a wide sea is more clear in the case of terrestrial mammals than perhaps with any other organic beings; and, accordingly, we find no inexplicable instances of the same mammals inhabiting distant points of the world. No geologist feels any difficulty in Great Britain possessing the same quadrupeds with the rest of Europe, for they were no doubt once united. But, if the same species can be produced at two separate points, why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same, so that a multitude of European animals and plants have become naturalized in America and Australia; and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres. The answer, as I believe, is, that mammals have not been able to migrate, whereas some plants, from their varied means of dispersal, have migrated across the wide and broken interspaces. The great and striking influence of barriers of all kinds is intelligible only on the view that the great majority of species have been produced on one side, and have not been able to migrate to the opposite side. Some few families, many sub-families, very many genera, and a still greater number of sections of genera, are confined to a single region: and it has been observed by several naturalists that the most natural genera, or those genera in which the species are most closely related to each other, are generally confined to the same country, or, if they have a wide range, that their range is continuous. What a strange anomaly it would be, if a directly opposite rule were to prevail, when we go down one step lower in the series, namely, to the individuals of the same species, and these had not been, at least at first, confined to some one region!
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GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGS.

We are thus brought to the question which has been largely discussed by naturalists, namely, whether species have been created at one or more points of the earth’s surface. Undoubtedly there are many cases of extreme difficulty in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points where now found. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. He who rejects it rejects the vera causa of ordinary generation with subsequent migration, and calls in the agency of a miracle. It is universally admitted that in most cases the area inhabited by a species is continuous; and that, when a plant or animal inhabits two points so distant from each other, or with an interval of such a nature, that the space could not have been easily passed over by migration, the fact is given as something remarkable and exceptional. The incapacity of migrating across a wide sea is more clear in the case of terrestrial mammals than perhaps with any other organic beings; and, accordingly, we find no inexplicable instances of the same mammals inhabiting distant points of the world. No geologist feels any difficulty in Great Britain possessing the same quadrupeds with the rest of Europe, for they were no doubt once united. But, if the same species can be produced at two separate points, why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same, so that a multitude of European animals and plants have become naturalized in America and Australia; and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres. The answer, as I believe, is, that mammals have not been able to migrate, whereas some plants, from their varied means of dispersal, have migrated across the wide and broken interspaces. The great and striking influence of barriers of all kinds is intelligible only on the view that the great majority of species have been produced on one side, and have not been able to migrate to the opposite side. Some few families, many sub-families, very many genera, and a still greater number of sections of genera, are confined to a single region: and it has been observed by several naturalists that the most natural genera, or those genera in which the species are most closely related to each other, are generally confined to the same country, or, if they have a wide range, that their range is continuous. What a strange anomaly it would be, if a directly opposite rule were to prevail, when we go down one step lower in the series, namely, to the individuals of the same species, and these had not been, at least at first, confined to some one region!

Hence it seems to me, as it has to many other naturalists, that the view of each species having been produced in one area alone, and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted, is the most probable. Undoubtedly many cases occur, in which we can not explain how the same species could have passed from one point to the other. But the geographical and climatal changes, which have certainly occurred within recent geological times, must have rendered discontinuous the formerly continuous range of many species. So that we are reduced to consider whether the exceptions to continuity of range are so numerous and of so grave a nature that we ought to give up the belief, rendered probable by general considerations, that each species has been produced within one area, and has migrated thence as far as it could.

ISOLATED CONTINENTS NEVER WERE UNITED.

Whenever it is fully admitted, as it will some day be, that each species has proceeded from a single birthplace, and when in the course of time we know something definite about the means of distribution, we shall be enabled to speculate with security on the former extension of the land. But I do not believe that it will ever be proved that within the recent period most of our continents which now stand quite separate have been continuously, or almost continuously, united with each other, and with the many existing oceanic islands. Several facts in distribution, such as the great difference in the marine faunas on the opposite sides of almost every continent, the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants, the degree of affinity between the mammals inhabiting islands with those of the nearest continent, being in part determined (as we shall hereafter see) by the depth of the intervening ocean, these and other such facts are opposed to the admission of such prodigious geographical revolutions within the recent period as are necessary on the view advanced by Forbes and admitted by his followers. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity with continents. Nor does the almost universally volcanic composition of such islands favor the admission that they are the wrecks of sunken continents; if they had originally existed as continental mountain-ranges, some at least of the islands would have been formed, like other mountain-summits, of granite, metamorphic schists, old fossiliferous and other rocks, instead of consisting of mere piles of volcanic matter.

MEANS OF DISPERSAL.

Living birds can hardly fail to be highly effective agents in the transportation of seeds. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. We may safely assume that under such circumstances their rate of flight would often be thirty-five miles an hour; and some authors have given a far higher estimate. I have never seen an instance of nutritious seeds passing through the intestines of a bird; but hard seeds of fruit pass uninjured through even the digestive organs of a turkey. In the course of two months I picked up in my garden twelve kinds of seeds out of the excrement of small birds, and these seemed perfect, and some of them, which were tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not, as I know by trial, injure in the least the germination of seeds; now, after a bird has found and devoured a large supply of food, it is positively asserted that all the grains do not pass into the gizzard for twelve or even eighteen hours. A bird in this interval might easily be blown to the distance of five hundred miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. Some hawks and owls bolt their prey whole, and, after an interval of from twelve to twenty hours, disgorge pellets, which, as I know from experiments made in the Zoölogical Gardens, include seeds capable of germination. Some seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey; and two seeds of beet grew after having been thus retained for two days and fourteen hours. Fresh-water fish, I find, eat seeds of many land and water plants: fish are frequently devoured by birds, and thus the seeds might be transported from place to place. I forced many kinds of seeds into the stomachs of dead fish, and then gave their bodies to fishing-eagles, storks, and pelicans; these birds, after an interval of many hours, either rejected the seeds in pellets or passed them in their excrement; and several of these seeds retained the power of germination. Certain seeds, however, were always killed by this process.

Locusts are sometimes blown to great distances from the land; I myself caught one three hundred and seventy miles from the coast of Africa, and have heard of others caught at greater distances.

* * * * *

As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, it can hardly be doubted that they must occasionally, as suggested by Lyell, have transported seeds from one part to another of the Arctic and Antarctic regions, and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of plants common to Europe, in comparison with the species on the other islands of the Atlantic, which stand nearer to the mainland, and (as remarked by Mr. H. C. Watson) from their somewhat northern character in comparison with the latitude, I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch.

THESE MEANS OF TRANSPORT NOT ACCIDENTAL.

These means of transport are sometimes called accidental, but this is not strictly correct; the currents of the sea are not accidental, nor is the direction of prevalent gales of wind. It should be observed that scarcely any means of transport would carry seeds for very great distances: for seeds do not retain their vitality when exposed for a great length of time to the action of sea-water; nor could they be long carried in the crops or intestines of birds. These means, however, would suffice for occasional transport across tracts of sea some hundred miles in breadth, or from island to island, or from a continent to a neighboring island, but not from one distant continent to another. The floras of distant continents would not by such means become mingled; but would remain as distinct as they now are. The currents, from their course, would never bring seeds from North America to Britain, though they might and do bring seeds from the West Indies to our western shores, where, if not killed by their very long immersion in salt-water, they could not endure our climate. Almost every year, one or two land-birds are blown across the whole Atlantic Ocean, from North America to the western shores of Ireland and England; but seeds could be transported by these rare wanderers only by one means, namely, by dirt adhering to their feet or beaks, which is in itself a rare accident. Even in this case, how small would be the chance of a seed falling on favorable soil and coming to maturity! But it would be a great error to argue that, because a well-stocked island, like Great Britain, has not, as far as is known (and it would be very difficult to prove this), received within the last few centuries, through occasional means of transport, immigrants from Europe or any other continent, a poorly-stocked island, though standing more remote from the mainland, would not receive colonists by similar means. Out of a hundred kinds of seeds or animals transported to an island, even if far less well-stocked than Britain, perhaps not more than one would be so well fitted to its new home as to become naturalized. But this is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, while the island was being upheaved, and before it had become fully stocked with inhabitants. On almost bare land, with few or no destructive insects or birds living there, nearly every seed which chanced to arrive, if fitted for the climate, would germinate and survive.

DISPERSAL DURING THE GLACIAL PERIOD.

The Glacial period is defined “as a period of great cold and of enormous extension of ice upon the surface of the earth. It is believed that glacial periods have occurred repeatedly during the geological history of the earth, but the term is generally applied to the close of the Tertiary epoch, when nearly the whole of Europe was subjected to an Arctic climate.”

* * * * *

The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one point to the other. It is indeed a remarkable fact to see so many plants of the same species living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europe; but it is far more remarkable that the plants on the White Mountains, in the United States of America, are all the same with those of Labrador, and nearly all the same, as we hear from Asa Gray, with those on the loftiest mountains of Europe. Even as long ago as 1747 such facts led Gmelin to conclude that the same species must have been independently created at many distinct points; and we might have remained in this same belief, had not Agassiz and others called vivid attention to the Glacial period, which, as we shall immediately see, affords a simple explanation of these facts. We have evidence of almost every conceivable kind, organic and inorganic, that, within a very recent geological period, Central Europe and North America suffered under an Arctic climate. The ruins of a house burned by fire do not tell their tale more plainly than do the mountains of Scotland and Wales, with their scored flanks, polished surfaces, and perched bowlders, of the icy streams with which their valleys were lately filled. So greatly has the climate of Europe changed, that in Northern Italy gigantic moraines, left by old glaciers, are now clothed by the vine and maize. Throughout a large part of the United States erratic bowlders and scored rocks plainly reveal a former cold period.

The former influence of the glacial climate on the distribution of the inhabitants of Europe, as explained by Edward Forbes, is substantially as follows. But we shall follow the changes more readily by supposing a new glacial period slowly to come on, and then pass away, as formerly occurred. As the cold came on, and as each more southern zone became fitted for the inhabitants of the north, these would take the places of the former inhabitants of the temperate regions. The latter, at the same time, would travel farther and farther southward, unless they were stopped by barriers, in which case they would perish. The mountains would become covered with snow and ice, and their former Alpine inhabitants would descend to the plains. By the time that the cold had reached its maximum, we should have an Arctic fauna and flora, covering the central parts of Europe, as far south as the Alps and Pyrenees, and even stretching into Spain. The now temperate regions of the United States would likewise be covered by Arctic plants and animals, and these would be nearly the same with those of Europe; for the present circumpolar inhabitants, which we suppose to have everywhere traveled southward, are remarkably uniform round the world.

As the warmth returned, the Arctic forms would retreat northward, closely followed up in their retreat by the productions of the more temperate regions. And, as the snow melted from the bases of the mountains, the Arctic forms would seize on the cleared and thawed ground, always ascending, as the warmth increased and the snow still further disappeared, higher and higher, while their brethren were pursuing their northern journey. Hence, when the warmth had fully returned, the same species, which had lately lived together on the European and North American lowlands, would again be found in the Arctic regions of the Old and New Worlds, and on many isolated mountain-summits far distant from each other.

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.

THE THEORY OF CREATION INADEQUATE.

As on the land, so in the waters of the sea, a slow southern migration of a marine fauna, which, during the Pliocene or even a somewhat earlier period, was nearly uniform along the continuous shores of the Polar Circle, will account, on the theory of modification, for many closely allied forms now living in marine areas completely sundered. Thus, I think, we can understand the presence of some closely allied, still existing and extinct tertiary forms on the eastern and western shores of temperate North America; and the still more striking fact of many closely allied crustaceans (as described in Dana’s admirable work), some fish and other marine animals, inhabiting the Mediterranean and the seas of Japan—these two areas being now completely separated by the breadth of a whole continent and by wide spaces of ocean.

These cases of close relationship in species either now or formerly inhabiting the seas on the eastern and western shores of North America, the Mediterranean and Japan, and the temperate lands of North America and Europe, are inexplicable on the theory of creation. We can not maintain that such species have been created alike, in correspondence with the nearly similar physical conditions of the areas; for, if we compare, for instance, certain parts of South America with parts of South Africa or Australia, we see countries closely similar in all their physical conditions, with their inhabitants utterly dissimilar.

CAUSES OF A GLACIAL CLIMATE.

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 toward 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 recur every ten to 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 two hundred and forty thousand years ago, and endured with slight alterations of climate for about one hundred and sixty thousand 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, while 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.

DIFFICULTIES NOT YET REMOVED.

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 can not be indicated. We can not say why certain species and not others have migrated; why certain species have been modified and have given rise to new forms, while others have remained unaltered. We can not hope to explain such facts, until we can say why one species and not another becomes naturalized 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.

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 can not 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 center; 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.

IDENTITY OF THE SPECIES OF ISLANDS WITH THOSE OF THE MAINLAND EXPLAINED ONLY BY THIS THEORY.

The most striking and important fact for us is the affinity of the species which inhabit islands to those of the nearest mainland, without being actually the same. Numerous instances could be given. The Galapagos Archipelago, situated under the equator, lies at the distance of between five hundred and six hundred miles from the shores of South America. Here almost every product of the land and of the water bears the unmistakable stamp of the American Continent. There are twenty-six land-birds; of these, twenty-one or perhaps twenty-three are ranked as distinct species, and would commonly be assumed to have been here created; yet the close affinity of most of these birds to American species is manifest in every character, in their habits, gestures, and tones of voice. So it is with the other animals, and with a large proportion of the plants, as shown by Dr. Hooker in his admirable Flora of this archipelago. The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, feels that he is standing on American land. Why should this be so? why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plainly the stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which closely resembles the conditions of the South American coast; in fact, there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in the climate, height, and size of the islands, between the Galapagos and Cape de Verd Archipelagos; but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape de Verd Islands are related to those of Africa, like those of the Galapagos to America. Facts such as these admit of no sort of explanation on the ordinary view of independent creation; whereas, on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists from America, whether by occasional means of transport or (though I do not believe in this doctrine) by formerly continuous land, and the Cape de Verd Islands from Africa; such colonists would be liable to modification, the principle of inheritance still betraying their original birthplace.

Many analogous facts could be given: indeed, it is an almost universal rule that the endemic productions of islands are related to those of the nearest continent, or of the nearest large island. The exceptions are few, and most of them can be explained. Thus, although Kerguelen Land stands nearer to Africa than to America, the plants are related, and that very closely, as we know from Dr. Hooker’s account, to those of America: but, on the view that this island has been mainly stocked by seeds brought with earth and stones on icebergs, drifted by the prevailing currents, this anomaly disappears. New Zealand in its endemic plants is much more closely related to Australia, the nearest mainland, than to any other region: and this is what might have been expected; but it is also plainly related to South America, which, although the next nearest continent, is so enormously remote that the fact becomes an anomaly. But this difficulty partially disappears on the view that New Zealand, South America, and the other southern lands have been stocked in part from a nearly intermediate though distant point, namely, from the Antarctic islands, when they were clothed with vegetation, during a warmer tertiary period, before the commencement of the last Glacial period. The affinity, which, though feeble, I am assured by Dr. Hooker is real, between the flora of the southwestern corner of Australia and of the Cape of Good Hope, is a far more remarkable case; but this affinity is confined to the plants, and will, no doubt, some day be explained.

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