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11 - Geographical Distribution 11-06 - Dispersal during the Glacial period co-extensive with the world 10 The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where the 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 to the other.

It is indeed a remarkable fact to see so many of the same plants 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.

Pyrenees
Pyrenees

White Mountains
White Mountains

Labrador
Labrador

Asa Gray
Asa Gray

Europe
Europe


Even as long ago as 1747, such facts led Gmelin to conclude that the same species must have been independently created at several 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.

Johann Friedrich Gmelin
Johann Friedrich Gmelin

Jean Louis Rodolphe Agassiz
Jean Louis Rodolphe Agassiz


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 burnt 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 boulders, 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.

vine
vine

maize
maize


Throughout a large part of the United States, erratic boulders, and rocks scored by drifted icebergs and coast-ice, plainly reveal a former cold period.

The former influence of the glacial climate on the distribution of the inhabitants of Europe, as explained with remarkable clearness by Edward Forbes, is substantially as follows.

Edward Forbes
Edward Forbes


But we shall follow the changes more readily, by supposing a new glacial period to come slowly on, and then pass away, as formerly occurred.

As the cold came on, and as each more southern zone became fitted for arctic beings and ill-fitted for their former more temperate inhabitants, the latter would be supplanted and arctic productions would take their places.

The inhabitants of the more temperate regions would at the same time travel 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 a uniform arctic fauna and flora, covering the central parts of Europe, as far south as the Alps and Pyrenees, and even stretching into Spain.

Alps
Alps

Spain
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 travelled southward, are remarkably uniform round the world.

circumpolar
circumpolar


We may suppose that the Glacial period came on a little earlier or later in North America than in Europe, so will the southern migration there have been a little earlier or later; but this will make no difference in the final result.

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 higher and higher, as the warmth increased, whilst their brethren were pursuing their northern journey.

Hence, when the warmth had fully returned, the same arctic species, which had lately lived in a body together on the lowlands of the Old and New Worlds, would be left isolated on distant mountain-summits (having been exterminated on all lesser heights) and in the arctic regions of both hemispheres.

Thus we can understand the identity of many plants at points so immensely remote as on the mountains of the United States and 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 migration as the cold came on, and the re-migration on the returning warmth, will 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.

Hewett Cottrell Watson
Hewett Cottrell Watson


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 permitted their former migration across the low intervening tracts, since become too warm for their existence.

If the climate, since the Glacial period, has ever been in any degree warmer than at present (as some geologists in the United States believe to have been the case, chiefly from the distribution of the fossil Gnathodon), then the arctic and temperate productions will at a very late period have marched a little further north, and subsequently have retreated to their present homes; but I have met with no satisfactory evidence with respect to this intercalated slightly warmer period, since the Glacial period.

gnathodon
gnathodon


The arctic forms, during their long southern migration and re-migration northward, will have been exposed to nearly the same climate, and, as is especially to be noticed, they will have kept in a body together; consequently their mutual relations will not have been much disturbed, and, in accordance with the principles inculcated in this volume, they will not have been liable to much modification.

But with our 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 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 its coldest period will have been temporarily driven down to the plains; they will, also, have been exposed to somewhat different climatal influences.

Their mutual relations will thus have been in some degree disturbed; consequently they will have been liable to modification; and this we find has been the case; for if we compare the present Alpine plants and animals of the several great European mountain-ranges, though very many of the species are identically the same, some present varieties, some are ranked as doubtful forms, and some few are distinct yet closely allied or representative species.

In illustrating what, as I believe, actually took place during the Glacial period, I assumed that at its commencement the arctic productions were as uniform round the polar regions as they are at the present day.

But the foregoing remarks on distribution apply not only to strictly arctic forms, but also to many sub-arctic and to some few northern temperate forms, for some of these are the same on the lower mountains and on the plains of North America and Europe; and it may be reasonably asked how I account for the necessary degree of uniformity of the sub-arctic and northern temperate forms round the world, at the commencement of the 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 Atlantic Ocean and by the extreme northern part of the Pacific.

Atlantic Ocean
Atlantic Ocean

Pacific Ocean
Pacific Ocean


During the Glacial period, when the inhabitants of the Old and New Worlds lived further southwards than at present, they must have been still more completely separated by wider spaces of ocean.

I believe the above difficulty may be surmounted by looking to still earlier changes of climate of an opposite nature.

We have good reason to believe that during the newer Pliocene period, before the Glacial epoch, and whilst the majority of the inhabitants of the world were specifically the same as now, the climate was warmer than at the present day.

Hence we may suppose that the organisms now living under the climate of latitude 60°, during the Pliocene period lived further north under the Polar Circle, in latitude 66°-67°; and that the strictly arctic productions then lived on the broken land still nearer to the pole.

Now if we look at a globe, we shall see that under the Polar Circle there is almost continuous land from western Europe, through Siberia, to eastern America.

Siberia
Siberia


And to this continuity of the circumpolar land, and to the consequent freedom for intermigration under a more favourable climate, I attribute the necessary amount of uniformity in the sub-arctic and northern temperate productions of the Old and New Worlds, at a period anterior to the Glacial epoch.

Believing, from reasons before alluded to, that our continents have long remained in nearly the same relative position, though subjected to large, but partial oscillations of level, I am strongly inclined to extend the above view, and to infer that during some earlier and still warmer period, such as the older Pliocene period, a large number of the same plants and animals inhabited the almost continuous circumpolar land; and that these plants and animals, both in the Old and New Worlds, began slowly to migrate southwards as the climate became less warm, long before the commencement of the Glacial period.

We now see, as I believe, their descendants, mostly in a modified condition, in the central parts of Europe and the United States.

On this view we can understand the relationship, with very little identity, between the productions of North America and Europe, a relationship which is most remarkable, considering the distance of the two areas, and their separation by the Atlantic Ocean.

We can further understand the singular fact remarked on by several observers, that the productions of Europe and America during the later tertiary stages were more closely related to each other than they are at the present time; for during these warmer periods the northern parts of the Old and New Worlds will have been almost continuously united by land, serving as a bridge, since rendered impassable by cold, for the inter-migration of their inhabitants.

During the slowly decreasing warmth of the Pliocene period, as soon as the species in common, which inhabited the New and Old Worlds, migrated south of the Polar Circle, they must have been completely cut off from each other.

This separation, as far as the more temperate productions are concerned, took place long ages ago.

And as the plants and animals migrated southward, they will have become mingled in the one great region with the native American productions, and have had to compete with them; and in the other great region, with those of the Old World.

Consequently we have here everything favourable for much modification, for far more modification than with the Alpine productions, left isolated, within a much more recent period, on the several mountain-ranges and on the arctic lands of the two Worlds.

Hence it has come, that when we compare the now living productions of the temperate regions of the New and Old Worlds, we find very few identical species (though Asa Gray has lately shown that more plants are identical than was formerly supposed), but we find in every great class many forms, which some naturalists rank as geographical races, and others as distinct species; and a host of closely allied or representative forms which are ranked by all naturalists as specifically distinct.

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 areas completely sundered.

Thus, I think, we can understand the presence of many existing and tertiary representative forms on the eastern and western shores of temperate North America; and the still more striking case of many closely allied crustaceans (as described in Dana's admirable work), of some fish and other marine animals, in the Mediterranean and in the seas of Japan, areas now separated by a continent and by nearly a hemisphere of equatorial ocean.

crustacean
crustacean

Japan
Japan

Mediterranean
Mediterranean


These cases of relationship, without identity, of the inhabitants of seas now disjoined, and likewise of the past and present inhabitants of the temperate lands of North America and Europe, are inexplicable on the theory of creation.

We cannot say that they 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 the southern continents of the Old World, we see countries closely corresponding in all their physical conditions, but with their inhabitants utterly dissimilar.

But we must return to our more immediate subject, the Glacial period.

I am convinced that Forbes's view may be largely extended.

Edward Forbes
Edward Forbes


In Europe we have the plainest evidence of the cold 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.

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 gigantic ancient moraines.

Himalaya
Himalaya

Sikkim
Sikkim

Joseph Dalton Hooker
Joseph Dalton Hooker

maize
maize


South of the equator, we have some direct evidence of former glacial action in New Zealand; and the same plants, found on widely separated mountains in this island, tell the same story.

New Zealand
New Zealand


If one account which has been published can be trusted, we have direct evidence of glacial action in the southeastern corner of Australia.

Australia
Australia


Looking to America; in the northern half, ice-borne fragments of rock have been observed on the eastern side as far south as lat. 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.

Rocky Mountains
Rocky Mountains


In the Cordillera of Equatorial South America, glaciers once extended far below their present level.

In central Chile I was astonished at the structure of a vast mound of detritus, about 800 feet in height, crossing a valley of the Andes; and this I now feel convinced was a gigantic moraine, left far below any existing glacier.

Chile
Chile

Andes
Andes


Further south on both sides of the continent, from lat. 41° to the southernmost extremity, we have the clearest evidence of former glacial action, in huge boulders transported far from their parent source.

We do not know that the Glacial epoch was strictly simultaneous at these several far distant points on opposite sides of the world.

But we have good evidence in almost every case, that the epoch was included within the latest geological period.

We have, also, excellent evidence, that it endured for an enormous time, as measured by years, at each point.

The cold may have come on, or have ceased, earlier at one point of the globe than at another, but seeing that it endured for long at each, and that it was contemporaneous in a geological sense, it seems to me probable that it was, during a part at least of the period, actually simultaneous throughout the world.

Without some distinct evidence to the contrary, we may at least admit as probable that the glacial action was simultaneous on the eastern and western sides of North America, in the Cordillera under the equator and under the warmer temperate zones, and on both sides of the southern extremity of the continent.

If this be admitted, it is difficult to avoid believing that the temperature of the whole world was at this period simultaneously cooler.

But it would suffice for my purpose, if the temperature was at the same time lower along certain broad belts of longitude.

On this view of the whole world, or at least of broad longitudinal belts, having been simultaneously colder from pole to pole, much light can be thrown on the present distribution of identical and allied species.

In America, Dr Hooker has shown that between forty and fifty of the flowering plants of Tierra del Fuego, forming no inconsiderable part of its scanty flora, are common to Europe, enormously remote as these two points are; and there are many closely allied species.

Tierra del Fuego
Tierra del Fuego


On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur.

On the highest mountains of Brazil, some few European genera were found by Gardner, which do not exist in the wide intervening hot countries.

Brazil
Brazil


So on the Silla of Caraccas the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera.

Caracas
Caracas

Venezuela
Venezuela


On the mountains of Abyssinia, several European forms and some few representatives of the peculiar flora of the Cape of Good Hope occur.

Abyssinia
Abyssinia

Cape of Good Hope
Cape of Good Hope


At the Cape of Good Hope a very few European species, believed not to have been introduced by man, and on the mountains, some few representative European forms are found, which have not been discovered in the intertropical parts of Africa.

Africa
Africa


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.

Ceylon
Ceylon

Java
Java


A list of the genera collected on the loftier peaks of Java raises a picture of a collection made on a hill in Europe! Still more striking is the fact that southern Australian forms are clearly represented by plants growing on the summits of the mountains of Borneo.

Borneo
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 as far as Japan.

Malacca
Malacca


On the southern mountains of Australia, Dr. F. Mueller 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.

Baron Sir Ferdinand Jacob Heinrich von Mueller
Baron Sir Ferdinand Jacob Heinrich von Mueller


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 throughout the world, the plants growing on the more lofty mountains, and on the temperate lowlands of the northern and southern hemispheres, are sometimes identically the same; but they are much oftener specifically distinct, though related to each other in a most remarkable manner.

This brief abstract applies to plants alone: some strictly analogous facts could be given on the distribution of terrestrial animals.

In marine productions, similar cases occur; as an example, I may quote a remark 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.

James Dwight Dana
James Dwight Dana

Tasmania
Tasmania


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.

Algae
Algae


It should be observed that the northern species and forms found in the southern parts of the southern hemisphere, and on the mountain-ranges of the intertropical regions, are not arctic, but belong to the northern temperate zones.

As Mr. H. C. Watson has recently remarked, `In receding from polar towards equatorial latitudes, the Alpine or mountain floras really become less and less arctic.' Many of the forms living on the mountains of the warmer regions of the earth and in the southern hemisphere are of doubtful value, being ranked by some naturalists as specifically distinct, by others as varieties; but some are certainly identical, and many, though closely related to northern forms, must be ranked as distinct species.

Hewett Cottrell Watson
Hewett Cottrell Watson


Now let us see what light can be thrown on the foregoing facts, on the belief, supported as it is by a large body of geological evidence, that the whole world, or a large part of it, was during the Glacial period simultaneously much colder than at present.

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 came slowly on, all the tropical plants and other productions will have retreated from both sides towards the equator, followed in the rear by the temperate productions, and these by the arctic; but with the latter we are not now concerned.

The tropical plants probably suffered much extinction; how much no one can say; perhaps formerly the tropics supported as many species as we see at the present day crowded together at the Cape of Good Hope, and in parts of temperate Australia.

Cape of Good Hope
Cape of Good Hope


As we know that many tropical plants and animals can withstand a considerable amount of cold, many might have escaped extermination during a moderate fall of temperature, more especially by escaping into the warmest spots.

But the great fact to bear in mind is, that all tropical productions will have suffered to a certain extent.

On the other hand, the temperate productions, after migrating nearer to the equator, though they will have been placed under somewhat new conditions, will have suffered less.

And it is certain that many temperate plants, if protected from the inroads of competitors, can withstand a much warmer climate than their own.

Hence, it seems to me possible, bearing in mind that the tropical productions were in a suffering state and could not have presented a firm front against intruders, that a certain number of the more vigorous and dominant temperate forms might have penetrated the native ranks and have reached or even crossed the equator.

The invasion would, of course, have been greatly favoured by high land, and perhaps by a dry climate; for Dr. Falconer informs me that it is the damp with the heat of the tropics which is so destructive to perennial plants from a temperate climate.

Hugh Falconer
Hugh Falconer


On the other hand, the most humid and hottest districts will have afforded an asylum to the tropical natives.

The mountain-ranges north-west of the Himalaya, and the long line of the Cordillera, seem to have afforded two great lines of invasion: and it is a striking fact, lately communicated to me by Dr. Hooker, that all the flowering plants, about forty-six in number, common to Tierra del Fuego and to Europe still exist in North America, which must have lain on the line of march.

Himalaya
Himalaya

Joseph Dalton Hooker
Joseph Dalton Hooker

Tierra del Fuego
Tierra del Fuego

Europe
Europe


But I do not doubt that some temperate productions entered and crossed even the lowlands of the tropics at the period when the cold was most intense, when arctic forms had migrated some twenty-five degrees of latitude from their native country and covered the land at the foot of the Pyrenees.

Pyrenees
Pyrenees


At this period of extreme cold, I believe that the climate under the equator at the level of the sea was about the same with that now felt there at the height of six or seven thousand feet.

During this the coldest period, I suppose that large spaces of the tropical lowlands were clothed with a mingled tropical and temperate vegetation, like that now growing with strange luxuriance at the base of the Himalaya, as graphically described by Hooker.

Thus, as I believe, a considerable number of plants, a few terrestrial animals, and some marine productions, migrated during the Glacial period from the northern and southern temperate zones into the intertropical regions, and some even crossed the equator.

As the warmth returned, these temperate forms would naturally ascend the higher mountains, being exterminated on the lowlands; those which had not reached the equator, would re-migrate northward or southward towards their former homes; but the forms, chiefly northern, which had crossed the equator, would travel still further from their homes into the more temperate latitudes of the opposite hemisphere.

Although we have reason to believe from geological evidence that the whole body of arctic shells underwent scarcely any modification during their long southern migration and re-migration northward, the case may have been wholly different with those intruding forms which settled themselves on the intertropical mountains, and in the southern hemisphere.

These being surrounded by strangers will have had to compete with many new forms of life; and it is probable that selected modifications in their structure, habits, and constitutions will have profited them.

Thus many of these wanderers, though still plainly related by inheritance to their brethren of the northern or southern hemispheres, now exist in their new homes as well-marked varieties or as distinct species.

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 plants and allied forms have apparently migrated from the north to the south, than in a reversed direction.

Alphonse de Candolle
Alphonse de Candolle


We see, however, a few southern vegetable forms on the mountains of Borneo and Abyssinia.

Borneo
Borneo

Abyssinia
Abyssinia


I suspect that this preponderant migration from north to 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 they became commingled during the Glacial period, the northern forms were enabled to beat the less powerful southern forms.

Just in the same manner as we see at the present day, that very many European productions cover the ground in La Plata, and in a lesser degree in Australia, and have to a certain extent beaten the natives; whereas extremely few southern forms have become naturalised in any part of Europe, 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 thirty or forty years from Australia.

La Plata
La Plata

Australia
Australia


Something of the same kind must have occurred on the intertropical mountains: no doubt before the Glacial period they were stocked with endemic Alpine forms; but these have almost everywhere largely 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 the naturalised; and if the natives have not been actually exterminated, their numbers have been greatly reduced, and this is the first stage towards extinction.

A mountain is an island on the land; and the intertropical mountains before the Glacial period must have been completely isolated; and I believe that the productions of these islands on the land yielded to those produced within the larger areas of the north, just in the same way as the productions of real islands have everywhere lately yielded to continental forms, naturalised by man's agency.

I am far from supposing that all difficulties are removed on the view here given in regard to the range and affinities of the allied species which live in the northern and southern temperate zones and on the mountains of the intertropical regions.

Very many difficulties remain to be solved.

I do not pretend to indicate the exact lines and means of migration, or the reason why certain species and not others have migrated; why certain species have been modified and have given rise to new groups of forms, and 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 ranges twice or thrice as far, and is twice or thrice as common, as another species within their own homes.

I have said that many difficulties remain to be solved: some of the most remarkable are stated with admirable clearness by Dr. Hooker in his botanical works on the antarctic regions.

These cannot be here discussed.

I will only say that as far as regards the occurrence of identical species at points so enormously remote as Kerguelen Land, New Zealand, and Fuegia, I believe that towards the close of the Glacial period, icebergs, as suggested by Lyell, have been largely concerned in their dispersal.

Sir Charles Lyell
Sir Charles Lyell


But the existence of several quite distinct species, belonging to genera exclusively confined to the south, at these and other distant points of the southern hemisphere, is, on my theory of descent with modification, a far more remarkable case of difficulty.

For some of these species are so distinct, that we cannot suppose that there has been time since the commencement of the Glacial period for their migration, and for their subsequent modification to the necessary degree.

The facts seem to me to indicate that peculiar and very distinct species have migrated in radiating lines from some 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 Glacial period, when the antarctic lands, now covered with ice, supported a highly peculiar and isolated flora.

I suspect that before this flora was exterminated by the Glacial epoch, a few forms were widely dispersed to various points of the southern hemisphere by occasional means of transport, and by the aid, as halting-places, of existing and now sunken islands, and perhaps at the commencement of the Glacial period, by icebergs.

By these means, as I believe, the southern shores of America, Australia, New Zealand have become slightly tinted by the same peculiar forms of vegetable life.

Sir C. Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alterations of climate on geographical distribution.

I believe that the world has recently felt one of his great cycles of change; and that on this view, combined with modification through natural selection, a multitude of facts in the present distribution both of the same and of allied forms of life can be explained.

The living waters may be said to have flowed during one short period from the north and from the south, and to have crossed at the equator; but to have flowed with greater force from the north so as to have freely inundated the south.

As the tide leaves its drift in horizontal lines, though 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 height 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 serve as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.
12 - Geographical Distribution -- continued 12-10 - Distribution of fresh-water productions 10 As lakes and river-systems are separated from each other by barriers of land, it might have been thought that fresh-water productions would not have ranged widely within the same country, and as the sea is apparently a still more impassable barrier, that they never would have extended to distant countries.

But the case is exactly the reverse.

Not only have many fresh-water species, belonging to quite different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world.

I well remember, when first collecting in the fresh waters of Brazil, feeling much surprise at the similarity of the fresh-water insects, shells, &c., and at the dissimilarity of the surrounding terrestrial beings, compared with those of Britain.

Brazil
Brazil


But this power in fresh-water productions of ranging widely, though so unexpected, can, I think, in most cases be explained by their having become fitted, in a manner highly useful to them, for short and frequent migrations from pond to pond, or from stream to stream; and liability to wide dispersal would follow from this capacity as an almost necessary consequence.

We can here consider only a few cases.

In regard to fish, I believe that the same species never occur in the fresh waters of distant continents.

But on the same continent the species often range widely and almost capriciously; for two river-systems will have some fish in common and some different.

A few facts seem to favour the possibility of their occasional transport by accidental means; like that of the live fish not rarely dropped by whirlwinds in India, and the vitality of their ova when removed from the water.

India
India


But I am inclined to attribute the dispersal of fresh-water fish mainly to slight changes within the recent period in the level of the land, having caused rivers to flow into each other.

Instances, also, could be given of this having occurred during floods, without any change of level.

We have evidence in the loess of the Rhine of considerable changes of level in the land within a very recent geological period, and when the surface was peopled by existing land and fresh-water shells.

Rhine
Rhine


The wide difference of the fish on opposite sides of continuous mountain-ranges, which from an early period must have parted river-systems and completely prevented their inosculation, seems to lead to this same conclusion.

With respect to allied fresh-water fish occurring at very distant points of the world, no doubt there are many cases which cannot at present be explained: but some fresh-water fish belong to very ancient forms, and in such cases there will have been ample time for great geographical changes, and consequently time and means for much migration.

In the second place, salt-water fish can with care be slowly accustomed to live in fresh water; and, according to Valenciennes, there is hardly a single group of fishes confined exclusively to fresh water, so that we may imagine that a marine member of a fresh-water group might travel far along the shores of the sea, and subsequently become modified and adapted to the fresh waters of a distant land.

Some species of fresh-water shells have a very wide range, and allied species, which, on my theory, are descended from a common parent and must have proceeded from a single source, prevail throughout the world.

Freshwater Shells
Freshwater Shells


Their distribution at first perplexed me much, as their ova are not likely to be transported by birds, and they are immediately killed by sea water, as are the adults.

I could not even understand how some naturalised species have rapidly spread throughout the same country.

But two facts, which I have observed and no doubt many others remain to be observed throw some light on this subject.

When a duck suddenly emerges from a pond covered with duck-weed, I have twice seen these little plants adhering to its back; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have quite unintentionally stocked the one with fresh-water shells from the other.

duck
duck

duckweed (Lemnaceae)
duckweed (Lemnaceae)

aquarium
aquarium


But another agency is perhaps more effectual: I suspended a duck's feet, which might represent those of a bird sleeping in a natural pond, in an aquarium, where many ova of fresh-water shells were hatching; and I found that numbers of the extremely minute and just hatched shells crawled on the feet, and clung to them so firmly that when taken out of the water they could not be jarred off, though at a somewhat more advanced age they would voluntarily drop off.

These just hatched molluscs, though aquatic in their nature, survived on the duck's feet, in damp air, from twelve to twenty hours; and in this length of time a duck or heron might fly at least six or seven hundred miles, and would be sure to alight on a pool or rivulet, if blown across sea to an oceanic island or to any other distant point.

heron
heron


Sir Charles Lyell also informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a limpet) firmly adhering to it; and a water-beetle of the same family, a Colymbetes, once flew on board the `Beagle,' when forty-five miles distant from the nearest land: how much farther it might have flown with a favouring gale no one can tell.

Sir Charles Lyell
Sir Charles Lyell

dyticus
dyticus

ancylus
ancylus

Colymbetes
Colymbetes

HMS Beagle
HMS Beagle

beagle
beagle


With respect to plants, it has long been known what enormous ranges many fresh-water and even marsh-species have, both over continents and to the most remote oceanic islands.

This is strikingly shown, as remarked by Alph. de Candolle, in large groups of terrestrial plants, which have only a very few aquatic members; for these latter seem immediately to acquire, as if in consequence, a very wide range.

Alphonse de Candolle
Alphonse de Candolle


I think favourable means of dispersal explain this fact.

I have before mentioned that earth occasionally, though rarely, adheres in some quantity to the feet and beaks of birds.

beak
beak


Wading birds, which frequent the muddy edges of ponds, if suddenly flushed, would be the most likely to have muddy feet.

Birds of this order I can show are the greatest wanderers, and are occasionally found on the most remote and barren islands in the open ocean; they would not be likely to alight on the surface of the sea, so that the dirt would not be washed off their feet; when making land, they would be sure to fly to their natural fresh-water haunts.

I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments, but will here give only the most striking case: I took in February three table-spoonfuls of mud from three different points, beneath water, on the edge of a little pond; this mud when dry weighed only 6 3/4 ounces; I kept it covered up in my study for six months, pulling up and counting each plant as it grew; the plants were of many kinds, and were altogether 537 in number; and yet the viscid mud was all contained in a breakfast cup! Considering these facts, I think it would be an inexplicable circumstance if water-birds did not transport the seeds of fresh-water plants to vast distances, and if consequently the range of these plants was not very great.

seeds
seeds

Breakfast Cup
Breakfast Cup


The same agency may have come into play with the eggs of some of the smaller fresh-water animals.

Other and unknown agencies probably have also played a part.

I have stated that fresh-water fish eat some kinds of seeds, though they reject many other kinds after having swallowed them; even small fish swallow seeds of moderate size, as of the yellow water-lily and Potamogeton.

Yellow Water Lily
Yellow Water Lily

potamogeton
potamogeton


Herons and other birds, century after century, have gone on daily devouring fish; they then take flight and go to other waters, or are blown across the sea; and we have seen that seeds retain their power of germination, when rejected in pellets or in excrement, many hours afterwards.

heron
heron


When I saw the great size of the seeds of that fine water-lily, the Nelumbium, and remembered Alph. de Candolle's remarks on this plant, I thought that its distribution must remain quite inexplicable; but Audubon states that he found the seeds of the great southern water-lily (probably, according to Dr Hooker, the Nelumbium luteum) in a heron's stomach; although I do not know the fact, yet analogy makes me believe that a heron flying to another pond and getting a hearty meal of fish, would probably reject from its stomach a pellet containing the seeds of the Nelumbium undigested; or the seeds might be dropped by the bird whilst feeding its young, in the same way as fish are known sometimes to be dropped.

nelumbium (Lotus)
nelumbium (Lotus)

Alphonse de Candolle
Alphonse de Candolle

John James Audubon
John James Audubon

Joseph Dalton Hooker
Joseph Dalton Hooker

Nelumbium Lutea
Nelumbium Lutea

heron
heron

stomach
stomach


In considering these several means of distribution, it should be remembered that when a pond or stream is first formed, for instance, on a rising islet, it will be unoccupied; and a single seed or egg will have a good chance of succeeding.

Although there will always be a struggle for life between the individuals of the species, however few, already occupying any pond, yet as the number of kinds is small, compared with those on the land, the competition will probably be less severe between aquatic than between terrestrial species; consequently an intruder from the waters of a foreign country, would have a better chance of seizing on a place, than in the case of terrestrial colonists.

We should, also, remember that some, perhaps many, fresh-water productions are low in the scale of nature, and that we have reason to believe that such low beings change or become modified less quickly than the high; and this will give longer time than the average for the migration of the same aquatic species.

We should not forget the probability of many species having formerly ranged as continuously as fresh-water productions ever can range, over immense areas, and having subsequently become extinct in intermediate regions.

But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, I believe mainly depends on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have large powers of flight, and naturally travel from one to another and often distant piece of water.

Nature, like a careful gardener, thus takes her seeds from a bed of a particular nature, and drops them in another equally well fitted for them.

gardener
gardener
12 - Geographical Distribution -- continued 12-20 - On the inhabitants of oceanic islands 10 We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe.

I have already stated that I cannot honestly admit Forbes's view on continental extensions, which, if legitimately followed out, would lead to the belief that within the recent period all existing islands have been nearly or quite joined to some continent.

Edward Forbes
Edward Forbes


This view would remove many difficulties, but it would not, I think, explain all the facts in regard to insular productions.

In the following remarks I shall not confine myself to the mere question of dispersal; but shall consider some other facts, which bear on the truth of the two theories of independent creation and of descent with modification.

The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. de Candolle admits this for plants, and Wollaston for insects.

Alphonse de Candolle
Alphonse de Candolle


If we look to the large size and varied stations of New Zealand, extending over 780 miles of latitude, and compare its flowering plants, only 750 in number, with those on an equal area at the Cape of Good Hope or in Australia, we must, I think, admit that something quite independently of any difference in physical conditions has caused so great a difference in number.

New Zealand
New Zealand


Even the uniform county of Cambridge has 847 plants, and the little island of Anglesea 764, but a few ferns and a few introduced plants are included in these numbers, and the comparison in some other respects is not quite fair.

Cambridge County
Cambridge County

Anglesea
Anglesea


We have evidence that the barren island of Ascension aboriginally possessed under half-a-dozen flowering plants; yet many have become naturalised on it, as they have on New Zealand and on every other oceanic island which can be named.

Ascension Island
Ascension Island


In St. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions.

St. Helena
St. Helena


He who admits the doctrine of the creation of each separate species, will have to admit, that a sufficient number of the best adapted plants and animals have not been created on oceanic islands; for man has unintentionally stocked them from various sources far more fully and perfectly than has nature.

Although in oceanic islands the number of kinds of inhabitants is scanty, the proportion of endemic species (i.e. those found nowhere else in the world) is often extremely large.

If we compare, for instance, the number of the endemic land-shells in Madeira, or of the endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the islands with that of the continent, we shall see that this is true.

Land Shell
Land Shell

Madeira
Madeira

Galapagos Islands
Galapagos Islands


This fact might have been expected on my theory for, as already explained, species occasionally arriving after long intervals in a new and isolated district, and having to compete with new associates, will be eminently liable to modification, and will often produce groups of modified descendants.

But it by no means follows, that, because in an island nearly all the species of one class are peculiar, those of another class, or of another section of the same class, are peculiar; and this difference seems to depend on the species which do not become modified having immigrated with facility and in a body, so that their mutual relations have not been much disturbed.

Thus in the Galapagos Islands nearly every land-bird, but only two out of the eleven marine birds, are peculiar; and it is obvious that marine birds could arrive at these islands more easily than land-birds.

Bermuda, on the other hand, which lies at about the same distance from North America as the Galapagos Islands do from South America, and which has a very peculiar soil, does not possess one endemic land bird; and we know from Mr. J. M. Jones's admirable account of Bermuda, that very many North American birds, during their great annual migrations, visit either periodically or occasionally this island.

Bermuda
Bermuda


Madeira does not possess one peculiar bird, and many European and African birds are almost every year blown there, as I am informed by Mr. E. V. Harcourt.

Africa
Africa


So that these two islands of Bermuda and Madeira have been stocked by birds, which for long ages have struggled together in their former homes, and have become mutually adapted to each other; and when settled in their new homes, each kind will have been kept by the others to their proper places and habits, and will consequently have been little liable to modification.

Madeira, again, is inhabited by a wonderful number of peculiar land-shells, whereas not one species of sea-shell is confined to its shores: now, though we do not know how seashells are dispersed, yet we can see that their eggs or larvae, perhaps attached to seaweed or floating timber, or to the feet of wading-birds, might be transported far more easily than land-shells, across three or four hundred miles of open sea.

Sea Shell
Sea Shell


The different orders of insects in Madeira apparently present analogous facts.

Oceanic islands are sometimes deficient in certain classes, and their places are apparently occupied by the other inhabitants; in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take the place of mammals.

Extinct Birds of New Zealand
Extinct Birds of New Zealand


In the plants of the Galapagos Islands, Dr. Hooker has shown that the proportional numbers of the different orders are very different from what they are elsewhere.

Joseph Dalton Hooker
Joseph Dalton Hooker


Such cases are generally accounted for by the physical conditions of the islands; but this explanation seems to me not a little doubtful.

Facility of immigration, I believe, has been at least as important as the nature of the conditions.

Many remarkable little facts could be given with respect to the inhabitants of remote islands.

For instance, in certain islands not tenanted by mammals, some of the endemic plants have beautifully hooked seeds; yet few relations are more striking than the adaptation of hooked seeds for transportal by the wool and fur of quadrupeds.

This case presents no difficulty on my view, for a hooked seed might be transported to an island by some other means; and the plant then becoming slightly modified, but still retaining its hooked seeds, would form an endemic species, having as useless an appendage as any rudimentary organ, for instance, as the shrivelled wings under the soldered elytra of many insular beetles.

Hooked Seed
Hooked Seed

beetle
beetle


Again, islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species; now trees, as Alph. de Candolle has shown, generally have, whatever the cause may be, confined ranges.

Hence trees would be little likely to reach distant oceanic islands; and an herbaceous plant, though it would have no chance of successfully competing in stature with a fully developed tree, when established on an island and having to compete with herbaceous plants alone, might readily gain an advantage by growing taller and taller and overtopping the other plants.

If so, natural selection would often tend to add to the stature of herbaceous plants when growing on an island, to whatever order they belonged, and thus convert them first into bushes and ultimately into trees.

island
island

tree
tree
12 - Geographical Distribution -- continued 12-30 - Absence of Batrachians and of terrestrial Mammals 10 With respect to the absence of whole orders on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) have never been found on any of the many islands with which the great oceans are studded.

Bory Saint Vincent
Bory Saint Vincent

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frogs

toad
toad

newt
newt