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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, their degenerate descendants. 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 old 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.
Summary of the preceding and present Chapter.-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 almost necessary for the accumulation of deposits rich in fossils and thick enough to resist future degradation, enormous intervals of time have elapsed between most of our 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. These causes taken conjointly, must have made the geological record extremely imperfect, and will to a large extent explain why—though we do find many links between the members of the same group—we do not find interminable varieties, connecting together all extinct and existing forms 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 ? 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. 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, the other great leading facts in palæontology 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 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. 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 other extinct and 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 must have beaten their predecessors in the race for life, and are, in so far, higher in the scale of nature, and their structure will generally have become more specialised ; and this may account for that ill-defined yet common sentiment, felt by many palæontologists, 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. 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. n the other hand, all the chief laws of palæontology 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.
GEOGRAPHICAL DISTRIBUTION Present distribution cannot be accounted for by differences in phy
sical conditions—Importance of barriers—Affinity of the productions of the same continent-Centres of creation-Means of dispersal, by changes of climate and of the level of the land, and by occasional means-
-Dispersal during the Glacial period co-extensive with the world. 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 the 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 species generally require ; for it is a mostrare 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 can 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