On the Origin of Species by Means of Natural Selection
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Chapter IX

On the Imperfection of the Geological Record

On the absence of intermediate varieties at the present day -- On the nature of extinct intermediate varieties; on their number -- On the vast lapse of time, as inferred from the rate of deposition and of denudation -- On the poorness of our palaeontological collections -- On the intermittence of geological formations -- On the absence of intermediate varieties in any one formation -- On the sudden appearance of groups of species -- On their sudden appearance in the lowest known fossiliferous strata.

In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. Most of them have now been discussed. One, namely the distinctness of specific forms, and their not being blended together by innumerable transitional links, is a very obvious difficulty. I assigned reasons why such links do not commonly occur at the present day, under the circumstances apparently most favourable for their presence, namely on an extensive and continuous area with graduated physical conditions. I endeavoured to show, that the life of each species depends in a more important manner on the presence of other already defined organic forms, than on climate; and, therefore, that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. I endeavoured, also, to show that intermediate varieties, from existing in lesser numbers than the forms which they connect, will generally be beaten out and exterminated during the course of further modification and improvement. The main cause, however, of innumerable intermediate links not now occurring everywhere throughout nature depends on the very process of natural selection, through which new varieties continually take the places of and exterminate their parent-forms. But just in proportion as this process of extermination has acted on an enormous scale, so must the number of intermediate varieties, which have formerly existed on the earth, be truly enormous. Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this, perhaps, is the most obvious and gravest objection which can be urged against my theory. The explanation lies, as I believe, in the extreme imperfection of the geological record.

In the first place it should always be borne in mind what sort of intermediate forms must, on my theory, have formerly existed. I have found it difficult, when looking at any two species, to avoid picturing to myself, forms directly intermediate between them. But this is a wholly false view; we should always look for forms intermediate between each species and a common but unknown progenitor; and the progenitor will generally have differed in some respects from all its modified descendants. To give a simple illustration: the fantail and pouter pigeons have both descended from the rock-pigeon; if we possessed all the intermediate varieties which have ever existed, we should have an extremely close series between both and the rock-pigeon; but we should have no varieties directly intermediate between the fantail and pouter; none, for instance, combining a tail somewhat expanded with a crop somewhat enlarged, the characteristic features of these two breeds. These two breeds, moreover, have become so much modified, that if we had no historical or indirect evidence regarding their origin, it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon, whether they had descended from this species or from some other allied species, such as C. oenas.

So with natural species, if we look to forms very distinct, for instance to the horse and tapir, we have no reason to suppose that links ever existed directly intermediate between them, but between each and an unknown common parent. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse; but in some points of structure may have differed considerably from both, even perhaps more than they differ from each other. Hence in all such cases, we should be unable to recognise the parent-form of any two or more species, even if we closely compared the structure of the parent with that of its modified descendants, unless at the same time we had a nearly perfect chain of the intermediate links.

It is just possible by my theory, that one of two living forms might have descended from the other; for instance, a horse from a tapir; and in this case direct intermediate links will have existed between them. But such a case would imply that one form had remained for a very long period unaltered, whilst its descendants had undergone a vast amount of change; and the principle of competition between organism and organism, between child and parent, will render this a very rare event; for in all cases the new and improved forms of life will tend to supplant the old and unimproved forms.

By the theory of natural selection all living species have been connected with the parent-species of each genus, by differences not greater than we see between the varieties of the same species at the present day; and these parent-species, now generally extinct, have in their turn been similarly connected with more ancient species; and so on backwards, always converging to the common ancestor of each great class. So that the number of intermediate and transitional links, between all living and extinct species, must have been inconceivably great. But assuredly, if this theory be true, such have lived upon this earth.

On the lapse of Time. -- Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected, that time will not have sufficed for so great an amount of organic change, all changes having been effected very slowly through natural selection. It is hardly possible for me even to recall to the reader, who may not be a practical geologist, the facts leading the mind feebly to comprehend the lapse of time. He who can read Sir Charles Lyell's grand work on the Principles of Geology, which the future historian will recognise as having produced a revolution in natural science, yet does not admit how incomprehensibly vast have been the past periods of time, may at once close this volume. Not that it suffices to study the Principles of Geology, or to read special treatises by different observers on separate formations, and to mark how each author attempts to give an inadequate idea of the duration of each formation or even each stratum. A man must for years examine for himself great piles of superimposed strata, and watch the sea at work grinding down old rocks and making fresh sediment, before he can hope to comprehend anything of the lapse of time, the monuments of which we see around us.

It is good to wander along lines of sea-coast, when formed of moderately hard rocks, and mark the process of degradation. The tides in most cases reach the cliffs only for a short time twice a day, and the waves eat into them only when they are charged with sand or pebbles; for there is reason to believe that pure water can effect little or nothing in wearing away rock. At last the base of the cliff is undermined, huge fragments fall down, and these remaining fixed, have to be worn away, atom by atom, until reduced in size they can be rolled about by the waves, and then are more quickly ground into pebbles, sand, or mud. But how often do we see along the bases of retreating cliffs rounded boulders, all thickly clothed by marine productions, showing how little they are abraded and how seldom they are rolled about! Moreover, if we follow for a few miles any line of rocky cliff, which is undergoing degradation, we find that it is only here and there, along a short length or round a promontory, that the cliffs are at the present time suffering. The appearance of the surface and the vegetation show that elsewhere years have elapsed since the waters washed their base.

He who most closely studies the action of the sea on our shores, will, I believe, be most deeply impressed with the slowness with which rocky coasts are worn away. The observations on this head by Hugh Miller, and by that excellent observer Mr. Smith of Jordan Hill, are most impressive. With the mind thus impressed, let any one examine beds of conglomerate many thousand feet in thickness, which, though probably formed at a quicker rate than many other deposits, yet, from being formed of worn and rounded pebbles, each of which bears the stamp of time, are good to show how slowly the mass has been accumulated. Let him remember Lyell's profound remark, that the thickness and extent of sedimentary formations are the result and measure of the degradation which the earth's crust has elsewhere suffered. And what an amount of degradation is implied by the sedimentary deposits of many countries! Professor Ramsay has given me the maximum thickness, in most cases from actual measurement, in a few cases from estimate, of each formation in different parts of Great Britain; and this is the result:-

Palaeozoic strata (not including igneous beds)..57,154
Secondary strata................................13,190
Tertiary strata..................................2,240

--making altogether 72,584 feet; that is, very nearly thirteen and three-quarters British miles. Some of these formations, which are represented in England by thin beds, are thousands of feet in thickness on the Continent. Moreover, between each successive formation, we have, in the opinion of most geologists, enormously long blank periods. So that the lofty pile of sedimentary rocks in Britain, gives but an inadequate idea of the time which has elapsed during their accumulation; yet what time this must have consumed! Good observers have estimated that sediment is deposited by the great Mississippi river at the rate of only 600 feet in a hundred thousand years. This estimate may be quite erroneous; yet, considering over what wide spaces very fine sediment is transported by the currents of the sea, the process of accumulation in any one area must be extremely slow.

But the amount of denudation which the strata have in many places suffered, independently of the rate of accumulation of the degraded matter, probably offers the best evidence of the lapse of time. I remember having been much struck with the evidence of denudation, when viewing volcanic islands, which have been worn by the waves and pared all round into perpendicular cliffs of one or two thousand feet in height; for the gentle slope of the lava-streams, due to their formerly liquid state, showed at a glance how far the hard, rocky beds had once extended into the open ocean. The same story is still more plainly told by faults,--those great cracks along which the strata have been upheaved on one side, or thrown down on the other, to the height or depth of thousands of feet; for since the crust cracked, the surface of the land has been so completely planed down by the action of the sea, that no trace of these vast dislocations is externally visible.

The Craven fault, for instance, extends for upwards of 30 miles, and along this line the vertical displacement of the strata has varied from 600 to 3000 feet. Prof. Ramsay has published an account of a downthrow in Anglesea of 2300 feet; and he informs me that he fully believes there is one in Merionethshire of 12,000 feet; yet in these cases there is nothing on the surface to show such prodigious movements; the pile of rocks on the one or other side having been smoothly swept away. The consideration of these facts impresses my mind almost in the same manner as does the vain endeavour to grapple with the idea of eternity.

I am tempted to give one other case, the well-known one of the denudation of the Weald. Though it must be admitted that the denudation of the Weald has been a mere trifle, in comparison with that which has removed masses of our palaeozoic strata, in parts ten thousand feet in thickness, as shown in Prof. Ramsay's masterly memoir on this subject. Yet it is an admirable lesson to stand on the North Downs and to look at the distant South Downs; for, remembering that at no great distance to the west the northern and southern escarpments meet and close, one can safely picture to oneself the great dome of rocks which must have covered up the Weald within so limited a period as since the latter part of the Chalk formation. The distance from the northern to the southern Downs is about 22 miles, and the thickness of the several formations is on an average about 1100 feet, as I am informed by Prof. Ramsay. But if, as some geologists suppose, a range of older rocks underlies the Weald, on the flanks of which the overlying sedimentary deposits might have accumulated in thinner masses than elsewhere, the above estimate would be erroneous; but this source of doubt probably would not greatly affect the estimate as applied to the western extremity of the district. If, then, we knew the rate at which the sea commonly wears away a line of cliff of any given height, we could measure the time requisite to have denuded the Weald. This, of course, cannot be done; but we may, in order to form some crude notion on the subject, assume that the sea would eat into cliffs 500 feet in height at the rate of one inch in a century. This will at first appear much too small an allowance; but it is the same as if we were to assume a cliff one yard in height to be eaten back along a whole line of coast at the rate of one yard in nearly every twenty-two years. I doubt whether any rock, even as soft as chalk, would yield at this rate excepting on the most exposed coasts; though no doubt the degradation of a lofty cliff would be more rapid from the breakage of the fallen fragments. On the other hand, I do not believe that any line of coast, ten or twenty miles in length, ever suffers degradation at the same time along its whole indented length; and we must remember that almost all strata contain harder layers or nodules, which from long resisting attrition form a breakwater at the base. Hence, under ordinary circumstances, I conclude that for a cliff 500 feet in height, a denudation of one inch per century for the whole length would be an ample allowance. At this rate, on the above data, the denudation of the Weald must have required 306,662,400 years; or say three hundred million years.

The action of fresh water on the gently inclined Wealden district, when upraised, could hardly have been great, but it would somewhat reduce the above estimate. On the other hand, during oscillations of level, which we know this area has undergone, the surface may have existed for millions of years as land, and thus have escaped the action of the sea: when deeply submerged for perhaps equally long periods, it would, likewise, have escaped the action of the coast-waves. So that in all probability a far longer period than 300 million years has elapsed since the latter part of the Secondary period.

I have made these few remarks because it is highly important for us to gain some notion, however imperfect, of the lapse of years. During each of these years, over the whole world, the land and the water has been peopled by hosts of living forms. What an infinite number of generations, which the mind cannot grasp, must have succeeded each other in the long roll of years! Now turn to our richest geological museums, and what a paltry display we behold!

On the poorness of our Palaeontological collections. -- That our palaeontological collections are very imperfect, is admitted by every one. The remark of that admirable palaeontologist, the late Edward Forbes, should not be forgotten, namely, that numbers of our fossil species are known and named from single and often broken specimens, or from a few specimens collected on some one spot. Only a small portion of the surface of the earth has been geologically explored, and no part with sufficient care, as the important discoveries made every year in Europe prove. No organism wholly soft can be preserved. Shells and bones will decay and disappear when left on the bottom of the sea, where sediment is not accumulating. I believe we are continually taking a most erroneous view, when we tacitly admit to ourselves that sediment is being deposited over nearly the whole bed of the sea, at a rate sufficiently quick to embed and preserve fossil remains. Throughout an enormously large proportion of the ocean, the bright blue tint of the water bespeaks its purity. The many cases on record of a formation conformably covered, after an enormous interval of time, by another and later formation, without the underlying bed having suffered in the interval any wear and tear, seem explicable only on the view of the bottom of the sea not rarely lying for ages in an unaltered condition. The remains which do become embedded, if in sand or gravel, will when the beds are upraised generally be dissolved by the percolation of rain-water. I suspect that but few of the very many animals which live on the beach between high and low watermark are preserved. For instance, the several species of the Chthamalinae (a sub-family of sessile cirripedes) coat the rocks all over the world in infinite numbers: they are all strictly littoral, with the exception of a single Mediterranean species, which inhabits deep water and has been found fossil in Sicily, whereas not one other species has hitherto been found in any tertiary formation: yet it is now known that the genus Chthamalus existed during the chalk period. The molluscan genus Chiton offers a partially analogous case.

With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods, it is superfluous to state that our evidence from fossil remains is fragmentary in an extreme degree. For instance, not a land shell is known belonging to either of these vast periods, with one exception discovered by Sir C. Lyell in the carboniferous strata of North America. In regard to mammiferous remains, a single glance at the historical table published in the Supplement to Lyell's Manual, will bring home the truth, how accidental and rare is their preservation, far better than pages of detail. Nor is their rarity surprising, when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits; and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations.

But the imperfection in the geological record mainly results from another and more important cause than any of the foregoing; namely, from the several formations being separated from each other by wide intervals of time. When we see the formations tabulated in written works, or when we follow them in nature, it is difficult to avoid believing that they are closely consecutive. But we know, for instance, from Sir R. Murchison's great work on Russia, what wide gaps there are in that country between the superimposed formations; so it is in North America, and in many other parts of the world. The most skilful geologist, if his attention had been exclusively confined to these large territories, would never have suspected that during the periods which were blank and barren in his own country, great piles of sediment, charged with new and peculiar forms of life, had elsewhere been accumulated. And if in each separate territory, hardly any idea can be formed of the length of time which has elapsed between the consecutive formations, we may infer that this could nowhere be ascertained. The frequent and great changes in the mineralogical composition of consecutive formations, generally implying great changes in the geography of the surrounding lands, whence the sediment has been derived, accords with the belief of vast intervals of time having elapsed between each formation.

But we can, I think, see why the geological formations of each region are almost invariably intermittent; that is, have not followed each other in close sequence. Scarcely any fact struck me more when examining many hundred miles of the South American coasts, which have been upraised several hundred feet within the recent period, than the absence of any recent deposits sufficiently extensive to last for even a short geological period. Along the whole west coast, which is inhabited by a peculiar marine fauna, tertiary beds are so scantily developed, that no record of several successive and peculiar marine faunas will probably be preserved to a distant age. A little reflection will explain why along the rising coast of the western side of South America, no extensive formations with recent or tertiary remains can anywhere be found, though the supply of sediment must for ages have been great, from the enormous degradation of the coast-rocks and from muddy streams entering the sea. The explanation, no doubt, is, that the littoral and sub-littoral deposits are continually worn away, as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves.

We may, I think, safely conclude that sediment must be accumulated in extremely thick, solid, or extensive masses, in order to withstand the incessant action of the waves, when first upraised and during subsequent oscillations of level. Such thick and extensive accumulations of sediment may be formed in two ways; either, in profound depths of the sea, in which case, judging from the researches of E. Forbes, we may conclude that the bottom will be inhabited by extremely few animals, and the mass when upraised will give a most imperfect record of the forms of life which then existed; or, sediment may be accumulated to any thickness and extent over a shallow bottom, if it continue slowly to subside. In this latter case, as long as the rate of subsidence and supply of sediment nearly balance each other, the sea will remain shallow and favourable for life, and thus a fossiliferous formation thick enough, when upraised, to resist any amount of degradation, may be formed.

I am convinced that all our ancient formations, which are rich in fossils, have thus been formed during subsidence. Since publishing my views on this subject in 1845, I have watched the progress of Geology, and have been surprised to note how author after author, in treating of this or that great formation, has come to the conclusion that it was accumulated during subsidence. I may add, that the only ancient tertiary formation on the west coast of South America, which has been bulky enough to resist such degradation as it has as yet suffered, but which will hardly last to a distant geological age, was certainly deposited during a downward oscillation of level, and thus gained considerable thickness.

All geological facts tell us plainly that each area has undergone numerous slow oscillations of level, and apparently these oscillations have affected wide spaces. Consequently formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation, may have been formed over wide spaces during periods of subsidence, but only where the supply of sediment was sufficient to keep the sea shallow and to embed and preserve the remains before they had time to decay. On the other hand, as long as the bed of the sea remained stationary, thick deposits could not have been accumulated in the shallow parts, which are the most favourable to life. Still less could this have happened during the alternate periods of elevation; or, to speak more accurately, the beds which were then accumulated will have been destroyed by being upraised and brought within the limits of the coast-action.

Thus the geological record will almost necessarily be rendered intermittent. I feel much confidence in the truth of these views, for they are in strict accordance with the general principles inculcated by Sir C. Lyell; and E. Forbes independently arrived at a similar conclusion.

One remark is here worth a passing notice. During periods of elevation the area of the land and of the adjoining shoal parts of the sea will be increased, and new stations will often be formed;--all circumstances most favourable, as previously explained, for the formation of new varieties and species; but during such periods there will generally be a blank in the geological record. On the other hand, during subsidence, the inhabited area and number of inhabitants will decrease (excepting the productions on the shores of a continent when first broken up into an archipelago), and consequently during subsidence, though there will be much extinction, fewer new varieties or species will be formed; and it is during these very periods of subsidence, that our great deposits rich in fossils have been accumulated. Nature may almost be said to have guarded against the frequent discovery of her transitional or linking forms.

From the foregoing considerations it cannot be doubted that the geological record, viewed as a whole, is extremely imperfect; but if we confine our attention to any one formation, it becomes more difficult to understand, why we do not therein find closely graduated varieties between the allied species which lived at its commencement and at its close. Some cases are on record of the same species presenting distinct varieties in the upper and lower parts of the same formation, but, as they are rare, they may be here passed over. Although each formation has indisputably required a vast number of years for its deposition, I can see several reasons why each should not include a graduated series of links between the species which then lived; but I can by no means pretend to assign due proportional weight to the following considerations.

Although each formation may mark a very long lapse of years, each perhaps is short compared with the period requisite to change one species into another. I am aware that two palaeontologists, whose opinions are worthy of much deference, namely Bronn and Woodward, have concluded that the average duration of each formation is twice or thrice as long as the average duration of specific forms. But insuperable difficulties, as it seems to me, prevent us coming to any just conclusion on this head. When we see a species first appearing in the middle of any formation, it would be rash in the extreme to infer that it had not elsewhere previously existed. So again when we find a species disappearing before the uppermost layers have been deposited, it would be equally rash to suppose that it then became wholly extinct. We forget how small the area of Europe is compared with the rest of the world; nor have the several stages of the same formation throughout Europe been correlated with perfect accuracy.

With marine animals of all kinds, we may safely infer a large amount of migration during climatal and other changes; and when we see a species first appearing in any formation, the probability is that it only then first immigrated into that area. It is well known, for instance, that several species appeared somewhat earlier in the palaeozoic beds of North America than in those of Europe; time having apparently been required for their migration from the American to the European seas. In examining the latest deposits of various quarters of the world, it has everywhere been noted, that some few still existing species are common in the deposit, but have become extinct in the immediately surrounding sea; or, conversely, that some are now abundant in the neighbouring sea, but are rare or absent in this particular deposit. It is an excellent lesson to reflect on the ascertained amount of migration of the inhabitants of Europe during the Glacial period, which forms only a part of one whole geological period; and likewise to reflect on the great changes of level, on the inordinately great change of climate, on the prodigious lapse of time, all included within this same glacial period. Yet it may be doubted whether in any quarter of the world, sedimentary deposits, including fossil remains, have gone on accumulating within the same area during the whole of this period. It is not, for instance, probable that sediment was deposited during the whole of the glacial period near the mouth of the Mississippi, within that limit of depth at which marine animals can flourish; for we know what vast geographical changes occurred in other parts of America during this space of time. When such beds as were deposited in shallow water near the mouth of the Mississippi during some part of the glacial period shall have been upraised, organic remains will probably first appear and disappear at different levels, owing to the migration of species and to geographical changes. And in the distant future, a geologist examining these beds, might be tempted to conclude that the average duration of life of the embedded fossils had been less than that of the glacial period, instead of having been really far greater, that is extending from before the glacial epoch to the present day.

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