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that widely ranging species are those which have varied most frequently, and have oftenest given rise to new species; that varieties have at first been local; and lastly, although each species must have passed through numerous transitional stages, it is probable that the periods, during which each underwent modification, though many and long as measured by years, have been short in comparison with the periods during which each remained in an unchanged condition. These causes, taken conjointly, will to a large extent explain why—though we do find many links—we do not find interminable varieties, connecting together all extinct and existing forms by the finest graduated steps. It should also be constantly borne in mind that any linking variety between two forms, which might be found, would be ranked, unless the whole chain could be perfectly restored, as a new and distinct species; for it is not pretended that we have any sure criterion by which species and varieties can be discriminated.
He who rejects this view of the imperfection of the geological record, will rightly reject the 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 successive stages of the same great formation? He may disbelieve in the immense intervals of time which must have elapsed between our consecutive formations; he may overlook how important a part migration has played, when the formations of any one great region, as those 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 Cambrian system was deposited? We now know that at least one animal did then exist; but I can answer this last question only by supposing that where our oceans now extend they have extended for an enormous period, and where our oscillating continents now stand they have stood since the commencement of the Cambrian system; but that, long before that epoch, the world presented a widely different aspect; and that the older continents, formed of formations older than any known to us, exist now only as remnants in a metamorphosed condition, or lie still buried under the ocean.
Passing from these difficulties, the other great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and 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 belonging to large and dominant groups tend to leave many modified descendants, which form new sub-groups and groups. 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 has sometimes been a 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 it is that dominant forms which spread widely and yield the greatest number of varieties tend to people the world with allied, but modified, descendants; and these will generally succeed in displacing the groups which are their inferiors in the struggle for existence. Hence, after long intervals of time, the productions of the world appear to have changed simultaneously.
We can understand how it is that all the forms of life, ancient and recent, make together a few grand classes. We can understand, from the continued tendenoy 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 bringing them only a little closer together. The more ancient a form is, the more often it stands in some degree intermediate between groups cow 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 other extinct and different forms. We can clearly see why the organic remains of closely consecutive formations are closely allied; for they are closely linked together by generation. We can clearly see why the remains of an intermediate formation are intermediate in character.
The inhabitants of the world at each successive period in its history have beaten their predecessors in the race for life, and are, in so far, higher in the scale, and their structure has generally become more specialised; and this may account for the commou belief held by so many pakeontologists, that organisation on the whole has progressed. Extinct and ancient animals resemble to a certain extent the embryos of the more recent animals belonging to the same classes, and this wonderful fact receives a simple explanation according to our views. The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious, and is intelligible on the principle of inheritance.
If then the geological record be as imperfect as many believe, 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 paleontology 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, the products of variation and the Survival of the Fittest.
Present distribution cannot be accounted for by differences in physical 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 — Alternate Glacial periods in the North and South.
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 wholly accounted for by 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 arctic and northern temperate parts, 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; 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. No doubt small areas can be pointed out in the Old World hotter than any in the New World, but these are not inhabited by a fauna different from that of the surrounding districts; for it is rare to find a group of organisms confined to a small area, of which the conditions are peculiar in only a slight degree. Notwithstanding this general 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. Or, again, we may compare the productions of South America south of lat. 35° with those north of 25°, which consequently are separated by a space of ten degrees of latitude and are exposed to considerably different conditions, yet they are 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.
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 in 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. On each continent, also, we see the same fact; for on the opposite sides of lofty and continuous mountain-ranges, of great deserts, and 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. The marine inhabitants of the eastern and western shores of South America are very distinct, with extremely few shells, Crustacea or echinodermata in common; but Dr. Giinther has recently shown that about thirty per cent, of the fishes are the same on the opposite sides of the isthmus of Panama; and this fact has led naturalists to believe that the isthmus was formerly open. 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. So that 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 almost wholly distinct. On the other hand, proceeding still farther westward from the eastern