On the Origin of Species by Means of Natural Selection

tion of the Geo

ed from the rate of deposition and of denudation-On the poorness of our pal?ontological collections-On the intermittence of geological formations-On the absence

ant 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

red 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 e

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

d 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 competi

t 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 clas

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 incomprehensively 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 differen

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 a

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. In the Cordillera I estimated one pile of conglomerate at ten thousand feet in thickness. Let the observer remember Lyell's profound remark that the thickness and extent of sedimenta

e

a (not includin

,1

dary

,1

ary s

,

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 M

y 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,-thos

ow 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 pi

ght 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

f 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 p

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

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,

ered by Sir C. Lyell and Dr. Dawson in the carboniferous strata of North America, of which shell several specimens have now been collected. 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 pres

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

or even a short geological period. Along the whole west coast, which is inhabited by a peculiar marine fauna, tertiary beds are so poorly 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

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 ove

te 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

ide 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 accumulate

e in the truth of these views, for they are in strict accordance with the general principles incul

ring 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,

een 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

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 w

ighbouring 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, wit

this whole time. But we have seen that a thick fossiliferous formation can only be accumulated during a period of subsidence; and to keep the depth approximately the same, which is necessary in order to enable the same species to live on the same space, the supply of sediment must nearly have counterbalanced the amount of subsidence. But this same movement of subsidence will often tend to sink th

nd which must have required an enormous period for their accumulation; yet no one ignorant of this fact would have suspected the vast lapse of time represented by the thinner formation. Many cases could be given of the lower beds of a formation having been upraised, denuded, submerged, and then re-covered by the upper beds of the same formation,-facts, showing what wide, yet easily overlooked, intervals have occurred in its accumulation. In other cases we have the plainest evidence in great fossilised trees, still standing upright as they grew, of many long intervals of time and changes of level during the process of deposition, which would never even have been suspected, had not the trees chanced to have been preserved: thus Messrs

ffect in any one geological section. Supposing B and C to be two species, and a third, A, to be found in an underlying bed; even if A were strictly intermediate between B and C, it would simply be ranked as a third and distinct species, unless at the same time it could be most closely connected with either one or both forms by intermediate varieties. Nor should it be forgotten, as before explained, that A might be the actual

ne species of D'Orbigny and others into the rank of varieties; and on this view we do find the kind of evidence of change which on my theory we ought to find. Moreover, if we look to rather wider intervals, namely, to distinct but consecutive stages of the same great formati

ording to this view, the chance of discovering in a formation in any one country all the early stages of transition between any two forms, is small, for the successive changes are supposed to have been local or confined to some one spot. Most marine animals have a wide range; and we have seen that with plants it is those which have the widest range, that oftenest present varieties; so

our being enabled to connect species by numerous, fine, intermediate, fossil links, by asking ourselves whether, for instance, geologists at some future period will be able to prove, that our different breeds of cattle, sheep, horses, and dogs have descended from a single stock or from several aboriginal stocks; or, again, whether certain sea-shells inhabiting the shores of North America

which may be urged against my views. Hence it will be worth while to sum up the foregoing remarks, under an imaginary illustration. The Malay Archipelago is of about the size of Europe from the North Cape to the Mediterranean, and from Britain to Russia; and therefore equals all the geological formations which have been examined with any accuracy, excepting those of the United States of America. I fully agree with Mr. Godwin-Austen, that t

lating. I suspect that not many of the strictly littoral animals, or of those which lived on naked submarine rocks, would be embedded; and those embedded in gravel or sand, would not endure to a

ce would be separated from each other by enormous intervals, during which the area would be either stationary or rising; whilst rising, each fossiliferous formation would be destroyed, almost as soon as accumulated, by the incessant coast-action,

for the preservation of all the transitional gradations between any two or more species. If such gradations were not fully preserved, transitional varieties would merely appear as so many distinct species. It is, also, probable that each great period of subsidence would be interrup

varieties would at first generally be local or confined to one place, but if possessed of any decided advantage, or when further modified and improved, they would slowly spread and supplant their parent-forms. When such varieties returned t

of life. We ought only to look for a few links, some more closely, some more distantly related to each other; and these links, let them be ever so close, if found in different stages of the same formation, would, by most pal?ontologists, be ranked as distinct species. But I do not pretend that I should

ent of a group of forms, all of which have descended from some one progenitor, must have been an extremely slow process; and the progenitors must have lived long ages before their modified descendants. But we continually over-rate the perfection of the geological record, and falsely infer, because certain genera or families have not been found beneath a certain stage, that they did not exist before that stage. We continually forget how large the world is, compared with the area over which our geological formations have been carefully examined; we forget that gro

ife, for instance to fly through the air; but that when this had been effected, and a few species had thus acquired a great advantage over other organis

dstone at nearly the commencement of this great series. Cuvier used to urge that no monkey occurred in any tertiary stratum; but now extinct species have been discovered in India, South America, and in Europe even as far back as the eocene stage. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States, who would have ventured to suppose that, besides reptiles, no less than at least thirty kinds of birds, some of gigantic size, existed during that period? Not a fragment of bone has been discovered in these

lve can be recognised; from all these circumstances, I inferred that had sessile cirripedes existed during the secondary periods, they would certainly have been preserved and discovered; and as not one species had then been discovered in beds of this age, I concluded that this great group had been suddenly developed at the commencement of the tertiary series. This was a sore trouble to me, adding as I thought one more instance of the abrupt appearance of a great group of species. But my work had hardly been published, when a skilful pal?ontologis

ghly remarkable; but I cannot see that it would be an insuperable difficulty on my theory, unless it could likewise be shown that the species of this group appeared suddenly and simultaneously throughout the world at this same period. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator; and by running through Pictet's Pal?ontology it will be seen that very few species are known from several formations in Europe. Some few families of fish now have a confined range; the teleostean fish might formerly have had a similarly confined range, and after having been largely

r pal?ontological ideas on many points, which the discoveries of even the last dozen years have effected, it seems to me to be about as rash in us to dogmatize on the succession of organ

ne progenitor, apply with nearly equal force to the earliest known species. For instance, I cannot doubt that all the Silurian trilobites have descended from some one crustacean, which must have lived long before the Silurian age, and which probably differed greatly from any known animal. Some of the most ancient Silurian animals, as the Nautilus, Lingula, &c., do not differ much from living spec

ng periods elapsed, as long as, or probably far longer than, the whole interval from the Silurian age to the pre

ower stage to the Silurian system, abounding with new and peculiar species. Traces of life have been detected in the Longmynd beds, beneath Barrande's so-called primordial zone. The presence of phosphatic nodules and bituminous matter in some of the lowest azoic rocks, probably indicates the former existence of life at these periods. But the difficulty of understanding the absence of vast piles of fossiliferous strata, which on my theory no doubt were somewhere accumulated before the Silurian epoch, is very grea

e several formations of Europe and of the United States; and from the amount of sediment, miles in thickness, of which the formations are composed, we may infer that from first to last large islands or tracts of land, whence the sediment was derived, occurred in the neighbourhood of the existing continents of Europe

, that where continents now exist, large tracts of land have existed, subjected no doubt to great oscillations of level, since the earliest silurian period. The coloured map appended to my volume on Coral Reefs, led me to conclude that the great oceans are still mainly areas of subsidence, the great archipelagoes still areas of oscillations of level, and the continents areas of elevation. But have we any right to assume that things have thus remained from the beginning of this world? Our continents seem to have been formed by a preponderance, during many oscillations of level, of the force of elevation; but may not the areas of preponderant movement have changed in the lapse of ages? At a period immeasurably antecedent to the silurian epoch, continents may have existed where oceans are now spread out; and clear and open oceans may have existed where our continents now stand. Nor sho

have unanimously, often vehemently, maintained the immutability of species. But I have reason to believe that one great authority, Sir Charles Lyell, from further reflexion entertains grave doubts on this subject. I feel how rash it is to differ from these authorities, to whom, with others, we owe all our knowledge. Those who think the natural geological record in any degree perfect, and who do not attach much weight to the facts and arguments of other kinds given in this volume, will undoubtedly at once reject my theory. For my part, following out Lyell's metaphor, I look at the natural geological record, as a history of the world i