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Charles Robert Darwin (1809–1882).  Origin of Species.
The Harvard Classics.  1909–14.
 
X. On the Imperfection of the Geological Record
 
On the Absence of Numerous Intermediate Varieties in Any Single Formation
 
 
FROM these several 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 much 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. Several cases are on record of the same species presenting varieties in the upper and lower parts of the same formation; thus, Trautschold gives a number of instances with ammonites; and Hilgendorf has described a most curious case of ten graduated forms of Planorbis multiformis in the successive beds of a fresh-water formation in Switzerland. Although each formation has indisputably required a vast number of years for its deposition, several reasons can be given why each should not commonly include a graduated series of links between the species which lived at its commencement and close; but I cannot assign due proportional weight to the following considerations.  1
  Although each formation may mark a very long lapse of years, each probably is short compared with the period requisite to change one species into another. I am aware that two palæontologists, 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 from 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 last layers have been deposited, it would be equally rash to suppose that it then became 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.  2
  We may safely infer that with marine animals of all kinds there has been a large amount of migration due to 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 appear somewhat earlier in the palæozoic 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 in 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 epoch, which forms only a part of one whole geological period; and likewise to reflect on the changes of level, on the extreme change of climate, and on the great 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 best flourish: for we know that great 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 migrations of species and to geographical changes. And in the distant future, a geologist, examining these beds, would 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.  3
  In order to get a perfect gradation between two forms in the upper and lower parts of the same formation, the deposit must have gone on continuously accumulating during a long period, sufficient for the slow process of modification; hence the deposit must be a very thick one; and the species, undergoing change must have lived in the same district throughout the whole time. But we have seen that a thick formation, fossiliferous throughout its entire thickness, can accumulate only during a period of subsidence; and to keep the depth approximately the same, which is necessary that the same marine species may live on the same space, the supply of sediment must nearly counterbalance the amount of subsidence. But this same movement of subsidence will tend to submerge the area whence the sediment is derived, and thus diminish the supply, whilst the downward movement continues. In fact, this nearly exact balancing between the supply of sediment and the amount of subsidence is probably a rare contingency; for it has been observed by more than one palæontologist, that very thick deposits are usually barren of organic remains, except near their upper or lower limits.  4
  It would seem that each separate formation, like the whole pile of formations in any country, has generally been intermittent in its accumulation. When we see, as is so often the case, a formation composed of beds of widely different mineralogical composition, we may reasonably suspect that the process of deposition has been more or less interrupted. Nor will the closest inspection of a formation give us any idea of the length of time which its deposition may have consumed. Many instances could be given of beds only a few feet in thickness, representing formations, which are elsewhere thousands of feet in thickness, and which must have required an enormous period for their accumulation; yet no one ignorant of this fact would have even 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 not have been suspected, had not the trees been preserved: thus Sir C. Lyell and Dr. Dawson found carboniferous beds 1,400 feet thick in Nova Scotia, with ancient root-bearing strata, one above the other at no less than sixty-eight different levels. Hence, when the same species occurs at the bottom, middle, and top of a formation, the probability is that it has not lived on the same spot during the whole period of deposition, but has disappeared and reappeared, perhaps many times, during the same geological period. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation, a section would not include all the fine intermediate gradations which must on our theory have existed, but abrupt, though perhaps slight, changes of form.  5
  It is all-important to remember that naturalists have no golden rule by which to distinguish species and varieties; they grant some little variability to each species, but when they meet with a somewhat greater amount of difference between any two forms, they rank both as species, unless they are enabled to connect them together by the closest intermediate gradations; and this, from the reasons just assigned, we can seldom hope to effect in any one geological section. Supposing B and C to be two species, and a third, A, to be found in an older and 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 closely connected by intermediate varieties with either one or both forms. Nor should it be forgotten, as before explained, that A might be the actual progenitor of B and C, and yet would not necessarily be strictly intermediate between them in all respects. So that we might obtain the parent-species, and its several modified descendants from the lower and upper beds of the same formation, and unless we obtained numerous transitional gradations, we should not recognise their blood-relationship, and should consequently rank them as distinct species.  6
  It is notorious on what excessively slight differences many palæontologists have founded their species; and they do this the more readily if the specimens come from different substages of the same formation. Some experienced conchologists are now sinking many of the very fine 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 the theory we ought to find. Look again at the later tertiary deposits, which include many shells believed by the majority of naturalists to be identical with existing species; but some excellent naturalists as Agassiz and Pictet, maintain that all these tertiary species are specifically distinct, though the distinction is admitted to be very slight; so that here, unless we believe that these eminent naturalists have been misled by their imaginations, and that these late tertiary species really present no difference whatever from their living. representatives, or unless we admit, in opposition to the judgment of most naturalists, that these tertiary species are all truly distinct from the recent, we have evidence of the frequent occurrence of slight modifications of the kind required. It we look to rather wider intervals of time, namely, to distinct but consecutive stages of the same great formation, we find that the embedded fossils, though universally ranked as specifically different, yet are far more closely related to each other than are the species found in more widely separated formations; so that here again we have undoubted evidence of change in the direction required by the theory; but to this latter subject I shall return in the following chapter.  7
  With animals and plants that propagate rapidly and do not wander much, there is reason to suspect, as we have formerly seen, that their varieties are generally at first local; and that such local varieties do not spread widely and supplant their parent-forms until they have been modified and perfected in some considerable degree. According 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 that, with shells and other marine animals, it is probable that those which had the widest range, far exceeding the limits of the known geological formations in Europe, have oftenest given rise, first to local varieties and ultimately to new species; and this again would greatly lessen the chance of our being able trace the stages of transition in any one geological formation.  8
  It is a more important consideration, leading to the same result, as lately insisted on by Dr. Falconer, namely, that the period during which each species underwent modification, though long as measured by years, was probably short in comparison with that during which it remained without undergoing any change.  9
  It should not be forgotten, that at the present day, with perfect specimens for examination, two forms can seldom be connected by intermediate varieties, and thus proved to be the same species, until many specimens are collected from many places; and with fossil species this can rarely be done. We shall, perhaps, best perceive the improbability of 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 are descended from a single stock or from several aboriginal stocks; or, again, whether certain sea-shells inhabiting the shores of North America, which are ranked by some conchologists as distinct species from their European representatives, and by other conchologists as only varieties, are really varieties, or are, as it is called, specifically distinct. This could be effected by the future geologist only by his discovering in a fossil state numerous intermediate gradations; and such success is improbable in the highest degree.  10
  It has been asserted over and over again, by writers who believe in the immutability of species, that geology yields no linking forms. This assertion, as we shall see in the next chapter, is certainly erroneous. As Sir J. Lubbock has remarked, “Every species is a link between other allied forms.” If we take a genus having a score of species, recent and extinct, and destroy four-fifths of them, no one doubts that the remainder will stand much more distinct from each other. If the extreme forms in the genus happen to have been thus destroyed, the genus itself will stand more distinct from other allied genera. What geological research has not revealed, is the former existence of infinitely numerous gradations, as fine as existing varieties, connecting together nearly all existing and extinct species. But this ought not to be expected; yet this has been repeatedly advanced as a most serious objection against my views.  11
  It may be worth while to sum up the foregoing remarks on the causes of the imperfection of the geological record under an imaginary illustration. The Malay Archipelago is 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 the present condition of the Malay Archipelago, with its numerous large islands separated by wide and shallow seas, probably represents the former state of Europe, whilst most of our formations were accumulating. The Malay Archipelago is one of the richest regions in organic beings; yet if all the species were to be collected which have ever lived there, how imperfectly would they represent the natural history of the world!  12
  But we have every reason to believe that the terrestrial productions of the archipelago would be preserved in an extremely imperfect manner in the formations which we suppose to be there accumulating. 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 distant epoch. Wherever sediment did not accumulate on the bed of the sea, or where it did not accumulate at a sufficient rate to protect organic bodies from decay, no remains could be preserved.  13
  Formations rich in fossils of many kinds, and of thickness sufficient to last to an age as distant in futurity as the secondary formations lie in the past, would generally be formed in the archipelago only during periods of subsidence. These periods of subsidence would be separated from each other by immense intervals of time, during which the area would be either stationary or rising; whilst rising, the fossiliferous formations on the steeper shores would be destroyed, almost as soon as accumulated, by the incessant coast-action, as we now see on the shores of South America. Even throughout the extensive and shallow seas within the archipelago, sedimentary beds could hardly be accumulated of great thickness during the periods of elevation, or become capped and protected by subsequent deposits, so as to have a good chance of enduring to a very distant future. During the periods of subsidence, there would probably be much extinction of life; during the periods of elevation, there would be much variation, but the geological record would then be less perfect.  14
  It may be doubted whether the duration of any one great period of subsidence over the whole or part of the archipelago, together with a contemporaneous accumulation of sediment, would exceed the average duration of the same specific forms; and these contingencies are indispensable for the preservation of all the transitional gradations between any two or more species. If such gradations were not all fully preserved, transitional varieties would merely appear as so many new, though closely allied species. It is also probable that each great period of subsidence would be interrupted by oscillations of level, and that slight climatal changes would intervene during such lengthy periods; and in these cases the inhabitants of the archipelago would migrate, and no closely consecutive record of their modifications could be preserved in any one formation.  15
  Very many of the marine inhabitants of the archipelago now range thousands of miles beyond its confines; and analogy plainly leads to the belief that it would be chiefly these far ranging species, though only some of them, which would oftenest produce new varieties; and the varieties would at first 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 to their ancient homes, as they would differ from their former state in a nearly uniform, though perhaps extremely slight degree, and as they would be found embedded in slightly different sub-stages of the same formation, they would, according to the principles followed by many palæontologists, be ranked as new and distinct species.  16
  If then there be some degree of truth in these remarks, we have no right to expect to find, in our geological formations, an infinite number of those fine transitional forms which, on our theory, have connected all the past and present species of the same group into one long and branching chain of life. We ought only to look for a few links, and such assuredly we do find—some more distantly, some more closely, related to each other; and these links, let them be ever so close, if found in different stages of the same formation, would, by many palæontologists, be ranked as distinct species. But I do not pretend that I should ever have suspected how poor was the record in the best preserved geological sections, had not the absence of innumerable transitional links between the species which lived at the commencement and close of each formation, pressed so hardly on my theory.  17
 

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