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11 - Geographical Distribution 11-02 - Importance of barriers 10 A second great fact which strikes us in our general review is, that barriers of any kind, or obstacles to free migration, are related in a close and important manner to the differences between the productions of various regions.

We see this in the great difference of nearly all the terrestrial productions of the New and Old Worlds, excepting in the northern parts, where the land almost joins, and where, under a slightly different climate, there might have been free migration for the northern temperate forms, as there now is for the strictly arctic productions.

We see the same fact in the great difference between the inhabitants of Australia, Africa, and South America under the same latitude: for these countries are almost as much isolated from each other as is possible.


Australia
Australia

South America
South America

Africa
Africa


On each continent, also, we see the same fact; for on the opposite sides of lofty and continuous mountain-ranges, and of great deserts, and sometimes even of large rivers, we find different productions; though as mountain chains, deserts, &c., are not as impassable, or likely to have endured so long as the oceans separating continents, the differences are very inferior in degree to those characteristic of distinct continents.

Turning to the sea, we find the same law.

No two marine faunas are more distinct, with hardly a fish, shell, or crab in common, than those of the eastern and western shores of South and Central America; yet these great faunas are separated only by the narrow, but impassable, isthmus of panama.

fish
fish

Sea Shell
Sea Shell

crab
crab

South America
South America

Central America
Central America

Panama
Panama


Westward of the shores of America, a wide space of open ocean extends, with not an island as a halting-place for emigrants; here we have a barrier of another kind, and as soon as this is passed we meet in the eastern islands of the Pacific, with another and totally distinct fauna.

Pacific Ocean
Pacific Ocean


So that here three marine faunas range far northward and southward, in parallel lines not far from each other, under corresponding climates; but from being separated from each other by impassable barriers, either of land or open sea, they are wholly distinct.

On the other hand, proceeding still further westward from the eastern islands of the tropical parts of the Pacific, we encounter no impassable barriers, and we have innumerable islands as halting-places, until after travelling over a hemisphere we come to the shores of Africa; and over this vast space we meet with no well-defined and distinct marine faunas.

Although hardly one shell, crab or fish is common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands, yet many fish range from the Pacific into the Indian Ocean, and many shells are common to the eastern islands of the Pacific and the eastern shores of Africa, on almost exactly opposite meridians of longitude.
11 - Geographical Distribution 11-01 - Present distribution cannot be accounted for by differences in physical conditions 10 In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the dissimilarity of the inhabitants of various regions can be accounted for by their climatal and other physical conditions.

Of late, almost every author who has studied the subject has come to this conclusion.

The case of America alone would almost suffice to prove its truth: for if we exclude the northern parts where the circumpolar land is almost continuous, all authors agree that one of the most fundamental divisions in geographical distribution is that between the New and Old Worlds; yet if we travel over the vast American continent, from the central parts of the United States to its extreme southern point, we meet with the most diversified conditions; the most humid districts, arid deserts, lofty mountains, grassy plains, forests, marshes, lakes, and great rivers, under almost every temperature.

There is hardly a climate or condition in the Old World which cannot be paralleled in the New at least as closely as the same species generally require; for it is a most rare case to find a group of organisms confined to any small spot, having conditions peculiar in only a slight degree; for instance, small areas in the Old World could be pointed out hotter than any in the New World, yet these are not inhabited by a peculiar fauna or flora.

Notwithstanding this parallelism in the conditions of the Old and New Worlds, how widely different are their living productions!

In the southern hemisphere, if we compare large tracts of land in Australia, South Africa, and western South America, between latitudes 25° and 35°, we shall find parts extremely similar in all their conditions, yet it would not be possible to point out three faunas and floras more utterly dissimilar.

Australia
Australia

South Africa
South Africa

South America
South America


Or again we may compare the productions of South America south of lat. 35° with those north of 25°, which consequently inhabit a considerably different climate, and they will be found incomparably more closely related to each other, than they are to the productions of Australia or Africa under nearly the same climate.

Analogous facts could be given with respect to the inhabitants of the sea.
10 - On The Geological Succession of Organic Beings 10-10 - Summary of preceding and present chapters 10 I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically explored with care; that only certain classes of organic beings have been largely preserved in a fossil state; that the number both of specimens and of species, preserved in our museums, is absolutely as nothing compared with the incalculable number of generations which must have passed away even during a single formation; that, owing to subsidence being necessary for the accumulation of fossiliferous deposits thick enough to resist future degradation, enormous intervals of time have elapsed between the successive formations; that there has probably been more extinction during the periods of subsidence, and more variation during the periods of elevation, and during the latter the record will have been least perfectly kept; that each single formation has not been continuously deposited; that the duration of each formation is, perhaps, short compared with the average duration of specific forms; that migration has played an important part in the first appearance of new forms in any one area and formation; that widely ranging species are those which have varied most, and have oftenest given rise to new species; and that varieties have at first often been local.

All these causes taken conjointly, must have tended to make the geological record extremely imperfect, and will to a large extent explain why we do not find interminable varieties, connecting together all the extinct and existing forms of life by the finest graduated steps.

He who rejects these views on the nature of the geological record, will rightly reject my whole theory.

For he may ask in vain where are the numberless transitional links which must formerly have connected the closely allied or representative species, found in the several stages of the same great formation.

He may disbelieve in the enormous intervals of time which have elapsed between our consecutive formations; he may overlook how important a part migration must have played, when the formations of any one great region alone, as that of Europe, are considered; he may urge the apparent, but often falsely apparent, sudden coming in of whole groups of species.

europe
europe


He may ask where are the remains of those infinitely numerous organisms which must have existed long before the first bed of the Silurian system was deposited: I can answer this latter question only hypothetically, by saying that as far as we can see, where our oceans now extend they have for an enormous period extended, and where our oscillating continents now stand they have stood ever since the Silurian epoch; but that long before that period, the world may have presented a wholly different aspect; and that the older continents, formed of formations older than any known to us, may now all be in a metamorphosed condition, or may lie buried under the ocean.

Passing from these difficulties, all the other great leading facts in palaeontology seem to me simply to follow on the theory of descent with modification through natural selection.

We can thus understand how it is that new species come in slowly and successively; how species of different classes do not necessarily change together, or at the same rate, or in the same degree; yet in the long run that all undergo modification to some extent.

The extinction of old forms is the almost inevitable consequence of the production of new forms.

We can understand why when a species has once disappeared it never reappears.

Groups of species increase in numbers slowly, and endure for unequal periods of time; for the process of modification is necessarily slow, and depends on many complex contingencies.

The dominant species of the larger dominant groups tend to leave many modified descendants, and thus new sub-groups and groups are formed.

As these are formed, the species of the less vigorous groups, from their inferiority inherited from a common progenitor, tend to become extinct together, and to leave no modified offspring on the face of the earth.

Earth
Earth


But the utter extinction of a whole group of species may often be a very slow process, from the survival of a few descendants, lingering in protected and isolated situations.

When a group has once wholly disappeared, it does not reappear; for the link of generation has been broken.

We can understand how the spreading of the dominant forms of life, which are those that oftenest vary, will in the long run tend to people the world with allied, but modified, descendants; and these will generally succeed in taking the places of those groups of species which are their inferiors in the struggle for existence.

Hence, after long intervals of time, the productions of the world will appear to have changed simultaneously.

We can understand how it is that all the forms of life, ancient and recent, make together one grand system; for all are connected by generation.

We can understand, from the continued tendency to divergence of character, why the more ancient a form is, the more it generally differs from those now living.

Why ancient and extinct forms often tend to fill up gaps between existing forms, sometimes blending two groups previously classed as distinct into one; but more commonly only bringing them a little closer together.

The more ancient a form is, the more often, apparently, it displays characters in some degree intermediate between groups now distinct; for the more ancient a form is, the more nearly it will be related to, and consequently resemble, the common progenitor of groups, since become widely divergent.

Extinct forms are seldom directly intermediate between existing forms; but are intermediate only by a long and circuitous course through many extinct and very different forms.

We can clearly see why the organic remains of closely consecutive formations are more closely allied to each other, than are those of remote formations; for the forms are more closely linked together by generation: we can clearly see why the remains of an intermediate formation are intermediate in character.

The inhabitants of each successive period in the world's history have beaten their predecessors in the race for life, and are, in so far, higher in the scale of nature; and this may account for that vague yet ill-defined sentiment, felt by many palaeontologists, that organisation on the whole has progressed.

If it should hereafter be proved that ancient animals resemble to a certain extent the embryos of more recent animals of the same class, the fact will be intelligible.

embryo
embryo


The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious, and is simply explained by inheritance.

If then the geological record be as imperfect as I believe it to be, and it may at least be asserted that the record cannot be proved to be much more perfect, the main objections to the theory of natural selection are greatly diminished or disappear.

On the other hand, all the chief laws of palaeontology plainly proclaim, as it seems to me, that species have been produced by ordinary generation: old forms having been supplanted by new and improved forms of life, produced by the laws of variation still acting round us, and preserved by Natural Selection.

fossil
fossil
10 - On The Geological Succession of Organic Beings 10-09 - On the succession of the same types within the same areas 10 Mr Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent.

Australia
Australia

mammals
mammals

marsupials
marsupials


In South America, a similar relationship is manifest, even to an uneducated eye, in the gigantic pieces of armour like those of the armadillo, found in several parts of La Plata; and Professor Owen has shown in the most striking manner that most of the fossil mammals, buried there in such numbers, are related to South American types.

South America
South America

armadillo
armadillo

La Plata
La Plata

Richard Owen
Richard Owen


This relationship is even more clearly seen in the wonderful collection of fossil bones made by MM. Lund and Clausen in the caves of Brazil.

Peter Wilhelm Lund
Peter Wilhelm Lund

Brazil
Brazil


I was so much impressed with these facts that I strongly insisted, in 1839 and 1845, on this `law of the succession of types,' on `this wonderful relationship in the same continent between the dead and the living.' Professor Owen has subsequently extended the same generalisation to the mammals of the Old World.

We see the same law in this author's restorations of the extinct and gigantic birds of New Zealand.

Extinct Birds of New Zealand
Extinct Birds of New Zealand


We see it also in the birds of the caves of Brazil.

Mr Woodward has shown that the same law holds good with sea-shells, but from the wide distribution of most genera of molluscs, it is not well displayed by them.

Sea Shell
Sea Shell

mollusca
mollusca


Other cases could be added, as the relation between the extinct and living land-shells of Madeira; and between the extinct and living brackish-water shells of the Aralo-Caspian Sea.

Land Shell
Land Shell

Madeira
Madeira

Caspian Sea
Caspian Sea


Now what does this remarkable law of the succession of the same types within the same areas mean?

He would be a bold man, who after comparing the present climate of Australia and of parts of South America under the same latitude, would attempt to account, on the one hand, by dissimilar physical conditions for the dissimilarity of the inhabitants of these two continents, and, on the other hand, by similarity of conditions, for the uniformity of the same types in each during the later tertiary periods.

Australia
Australia

South America
South America


Nor can it be pretended that it is an immutable law that marsupials should have been chiefly or solely produced in Australia; or that Edentata and other American types should have been solely produced in South America.

edentata
edentata


For we know that Europe in ancient times was peopled by numerous marsupials; and I have shown in the publications above alluded to, that in America the law of distribution of terrestrial mammals was formerly different from what it now is.

North America formerly partook strongly of the present character of the southern half of the continent; and the southern half was formerly more closely allied, than it is at present, to the northern half.

In a similar manner we know from Falconer and Cautley's discoveries, that northern India was formerly more closely related in its mammals to Africa than it is at the present time.

Hugh Falconer
Hugh Falconer

India
India

Africa
Africa


Analogous facts could be given in relation to the distribution of marine animals.

On the theory of descent with modification, the great law of the long enduring, but not immutable, succession of the same types within the same areas, is at once explained; for the inhabitants of each quarter of the world will obviously tend to leave in that quarter, during the next succeeding period of time, closely allied though in some degree modified descendants.

If the inhabitants of one continent formerly differed greatly from those of another continent, so will their modified descendants still differ in nearly the same manner and degree.

But after very long intervals of time and after great geographical changes, permitting much inter-migration, the feebler will yield to the more dominant forms, and there will be nothing immutable in the laws of past and present distribution.

It may be asked in ridicule, whether I suppose that the megatherium and other allied huge monsters have left behind them in South America the sloth, armadillo, and anteater, as their degenerate descendants.

South America
South America

sloth
sloth

armadillo
armadillo

anteater
anteater


This cannot for an instant be admitted.

These huge animals have become wholly extinct, and have left no progeny.

But in the caves of Brazil, there are many extinct species which are closely allied in size and in other characters to the species still living in South America; and some of these fossils may be the actual progenitors of living species.

It must not be forgotten that, on my theory, all the species of the same genus have descended from some one species; so that if six genera, each having eight species, be found in one geological formation, and in the next succeeding formation there be six other allied or representative genera with the same number of species, then we may conclude that only one species of each of the six older genera has left modified descendants, constituting the six new genera.

The other seven species of the old genera have all died out and have left no progeny.

Or, which would probably be a far commoner case, two or three species of two or three alone of the six older genera will have been the parents of the six new genera; the other old species and the other whole genera having become utterly extinct.

In failing orders, with the genera and species decreasing in numbers, as apparently is the case of the Edentata of South America, still fewer genera and species will have left modified blood-descendants.