CHAPTER XIII

THE GEOLOGICAL EVIDENCES OF EVOLUTION

What we may expect-The number of known species of extinct animalsCauses of the imperfection of the geological record-Geological evidences of evolution-Shells-Crocodiles-The rhinoceros tribeThe pedigree of the horse tribe-Development of deer's horns-Brain development Local relations of fossil and living animals-Cause of extinction of large animals-Indications of general progress in plants and animals—The progressive development of plants-Possible cause of sudden late appearance of exogens-Geological distribution of insects-Geological succession of vertebrata-Concluding remarks.

THE theory of evolution in the organic world necessarily implies that the forms of animals and plants have, broadly speaking, progressed from a more generalised to a more specialised structure, and from simpler to more complex forms. We know, however, that this progression has been by no means regular, but has been accompanied by repeated degradation and degeneration; while extinction on enormous scale has again and again stopped all progress in certain directions, and has often compelled a fresh start in development from some comparatively low and imperfect type.

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The enormous extension of geological research in recent times has made us acquainted with a vast number of extinct organisms, so vast that in some important groups—such as the mollusca-the fossil are more numerous than the living species; while in the mammalia they are not much less numerous, the preponderance of living species being chiefly in the smaller and in the arboreal forms which have not been so well preserved as the members of the larger groups. With such a wealth of material to illustrate the successive stages

through which animals have passed, it will naturally be expected that we should find important evidence of evolution. We should hope to learn the steps by which some isolated forms have been connected with their nearest allies, and in many cases to have the gaps filled up which now separate genus from genus, or species from species. In some cases these expectations are fulfilled, but in many other cases we seek in vain for evidence of the kind we desire; and this absence of evidence with such an apparent wealth of material is held by many persons to throw doubt on the theory of evolution itself. They urge, with much appearance of reason, that all the arguments we have hitherto adduced fall short of demonstration, and that the crucial test consists in being able to show, in a great number of cases, those connecting links which we say must have existed. Many of the gaps that still remain are so vast that it seems incredible to these writers that they could ever have been filled up by a close succession of species, since these must have spread over so many ages, and have existed in such numbers, that it seems impossible to account for their total absence from deposits in which great numbers of species belonging to other groups are preserved and have been discovered. In order to appreciate the force, or weakness, of these objections, we must inquire into the character and completeness of that record of the past life of the earth which geology has unfolded, and ascertain the nature and amount of the evidence which, under actual conditions, we may expect to find.

The Number of known Species of Extinct Animals.

When we state that the known fossil mollusca are considerably more numerous than those which now live on the earth, it appears at first sight that our knowledge is very complete, but this is far from being the case. The species have been continually changing throughout geological time, and at each period have probably been as numerous as they are now. If we divide the fossiliferous strata into twelve great divisions -the Pliocene, Miocene, Eocene, Cretaceous, Oolite, Lias, Trias, Permian, Carboniferous, Devonian, Silurian, and Cambrian, we find not only that each has a very distinct and characteristic molluscan fauna, but that the different sub

divisions often present a widely different series of species; so that although a certain number of species are common to two or more of the great divisions, the totality of the species that have lived upon the earth must be very much more than twelve times-perhaps even thirty or forty times-the number now living. In like manner, although the species of fossil mammals now recognised by more or less fragmentary fossil remains may not be much less numerous than the living species, yet the duration of existence of these was comparatively so short that they were almost completely changed, perhaps six or seven times, during the Tertiary period; and this is certainly only a fragment of the geological time during which mammalia existed on the globe.

There is also reason to believe that the higher animals were much more abundant in species during past geological epochs than now, owing to the greater equability of the climate which rendered even the arctic regions as habitable as the temperate zones are in our time.

The same equable climate would probably cause a more uniform distribution of moisture, and render what are now desert regions capable of supporting abundance of animal life. This is indicated by the number and variety of the species of large animals that have been found fossil in very limited areas which they evidently inhabited at one period.

M. Albert

Gaudry found, in the deposits of a mountain stream at Pikermi in Greece, an abundance of large mammalia such as are nowhere to be found living together at the present time. Among them were two species of Mastodon, two different rhinoceroses, a gigantic wild boar, a camel and a giraffe larger than those now living, several monkeys, carnivora ranging from martens and civets to lions and hyænas of the largest size, numerous antelopes of at least five distinct genera, and besides these many forms altogether extinct. Such were the great herds of Hipparion, an ancestral form of horse; the Helladotherium, a huge animal bigger than the giraffe; the Ancylotherium, one of the Edentata; the huge Dinotherium; the Aceratherium, allied to the rhinoceros; and the monstrous Chalicotherium, allied to the swine and ruminants, but as large as a rhinoceros; and to prey upon these, the great Machairodus or sabre-toothed tiger. And all these remains were

found in a space 300 paces long by 60 paces broad, many of the species existing in enormous quantities.

The Pikermi fossils belong to the Upper Miocene formation, but an equally rich deposit of Upper Eocene age has been discovered in South-Western France at Quercy, where M. Filhol has determined the presence of no less than forty-two species of beasts of prey alone. Equally remarkable are the various discoveries of mammalian fossils in North America, especially in the old lake bottoms now forming what are called the "bad lands" of Dakota and Nebraska, belonging to the Miocene period. Here are found an enormous assemblage of remains, often perfect skeletons, of herbivora and carnivora, as varied and interesting as those from the localities already referred to in Europe; but altogether distinct, and far exceeding, in number and variety of species of the larger animals, the whole existing fauna of North America. Very similar phenomena occur in South America and in Australia, leading us to the conclusion that the earth at the present time is impoverished as regards the larger animals, and that at each successive period of Tertiary time, at all events, it contained a far greater number of species than now inhabit it. The very richness and abundance of the remains which we find in limited areas, serve to convince us how imperfect and fragmentary must be our knowledge of the earth's fauna at any one past epoch; since we cannot believe that all, or nearly all, of the animals which inhabited any district were entombed in a single lake, or overwhelmed by the floods of a single river.

But the spots where such rich deposits occur are exceedingly few and far between when compared with the vast areas of continental land, and we have every reason to believe that in past ages, as now, numbers of curious species were rare or local, the commoner and more abundant species giving a very imperfect idea of the existing series of animal forms. Yet more important, as showing the imperfection of our knowledge, is the enormous lapse of time between the several formations in which we find organic remains in any abundance, so vast that in many cases we find ourselves almost in a new world, all the species and most of the genera of the higher animals having undergone a complete change.

Causes of the Imperfection of the Geological Record. These facts are quite in accordance with the conclusions of geologists as to the necessary imperfection of the geological record, since it requires the concurrence of a number of favourable conditions to preserve any adequate representation of the life of a given epoch. In the first place, the animals to be preserved must not die a natural death by disease, or old age, or by being the prey of other animals, but must be destroyed by some accident which shall lead to their being embedded in the soil. They must be either carried away by floods, sink into bogs or quicksands, or be enveloped in the mud or ashes of a volcanic eruption; and when thus embedded they must remain undisturbed amid all the future changes of the earth's surface.

But the chances against this are enormous, because denudation is always going on, and the rocks we now find at the earth's surface are only a small fragment of those which were originally laid down. The alternations of marine and freshwater deposits, and the frequent unconformability of strata with those which overlie them, tell us plainly of repeated elevations and depressions of the surface, and of denudation on an enormous scale. Almost every mountain

range, with its peaks, ridges, and valleys, is but the remnant of some vast plateau eaten away by sub-aerial agencies; every range of sea-cliffs tell us of long slopes of land destroyed by the waves; while almost all the older rocks which now form the surface of the earth have been once covered with newer deposits which have long since disappeared. Nowhere are the evidences of this denudation more apparent than in North and South America, where granitic or metamorphic rocks cover an area hardly less than that of all Europe. The same rocks are largely developed in Central Africa and Eastern Asia; while, besides those portions that appear exposed on the surface, areas of unknown extent are buried under strata which rest on them uncomformably, and could not, therefore, constitute the original capping under which the whole of these rocks must once have been deeply buried; because granite can only be formed, and metamorphism can only go on, deep down in the crust of the earth. What an over