The Origin Of Species (1872)
Chapter X: On the Imperfection of the Geological Record
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 supplant 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, 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 serious objection which can be urged against the 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 the 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 are 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, C. livia, whether they had descended from this species or from some allied form, such as C. aenas.
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 directly intermediate between them ever existed, 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 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 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 natural and domestic 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 forms; 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 the earth.
On the Lapse of Time, as inferred from the rate of Deposition and extent of Denudation
Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected that time cannot have sufficed for so great an amount of organic change, all changes having been effected slowly. It is hardly possible for me to recall to the reader who is not 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, and yet does not admit how 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 of each stratum. We can best gain some idea of past time by knowing the agencies at work, and learning how deeply the surface of the land has been denuded, and how much sediment has been deposited. As Lyell has well remarked, the extent and thickness of our sedimentary formations are the result and the measure of the denudation which the earth's crust has elsewhere undergone. Therefore a man should examine for himself the great piles of superimposed strata, and watch the rivulets bringing down mud, and the waves wearing away the sea-cliffs, in order to comprehend something about the duration of past time, the monuments of which we see all around us.
It is good to wander along the 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 good evidence that pure water effects 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 after being reduced in size they can be rolled about by the waves, and then they 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.
We have, however, recently learnt from the observations of Ramsay, in the van of many excellent observers- of Jukes, Geikie, Croll, and others, that subaerial degradation is a much more important agency than coast-action, or the power of the waves. The whole surface of the land is exposed to the chemical action of the air and of the rain-water with its dissolved carbolic acid, and in colder countries to frost; the disintegrated matter is carried down even gentle slopes during heavy rain, and to a greater extent than might be supposed, especially in arid districts, by the wind; it is then transported by the streams and rivers, which when rapid deepen their channels, and triturate the fragments. On a rainy day, even in a gently undulating country, we see the effects of subaerial degradation in the muddy rills which flow down every slope. Messrs. Ramsay and Whitaker have shown, and the observation is a most striking one, that the great lines of escarpment in the Wealden district and those ranging across England, which formerly were looked at as ancient sea-coasts, cannot have been thus formed, for each line is composed of one and the same formation, whilst our sea-cliffs are everywhere formed by the intersection of various formations. This being the case, we are compelled to admit that the escarpments owe their origin in chief part to the rocks of which they are composed having resisted subaerial denudation better than the surrounding surface; this surface consequently has been gradually lowered, with the lines of harder rock left projecting. Nothing impresses the mind with the vast duration of time, according to our ideas of time, more forcibly than the conviction thus gained that subaerial agencies which apparently have so little power, and which seem to work so slowly, have produced great results.
When thus impressed with the slow rate at which the land is worn away through subaerial and littoral action, it is good, in order to appreciate the past duration of time, to consider, on the one hand, the masses of rock which have been removed over many extensive areas, and on the other hand the thickness of our sedimentary formations. I remember having been much struck 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 told still more plainly 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, and it makes no great difference whether the upheaval was sudden, or, as most geologists now believe, was slow and effected by many starts, the surface of the land has been so completely planed down 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 varies from 600 to 3000 feet. Professor Ramsay has published an account of a downthrow in Anglesea of 2300 feet; and he informs me that he fully believes that there is one in Merionethshire of 12,000 feet; yet in these cases there is nothing on the surface of the land to show such prodigious movements; the pile of rocks on either side of the crack having been smoothly swept away.
On the other hand, in all parts of the world the piles of sedimentary strata are of wonderful thickness. In the Cordillera I estimated one mass of conglomerate at ten thousand feet; and although conglomerates have probably been accumulated at a quicker rate than finer sediments, yet from being formed of worn and rounded pebbles, each of which bears the stamp of time, they are good to show how slowly the mass must have been heaped together. Professor Ramsay has given me the maximum thickness, from actual measurement in most cases, of the successive formations in different parts of Great Britain; and this is the result:-
Palaeozoic strata (not including igneous beds): 57,154 feet
Secondary strata: 13,190 feet
Tertiary strata: 2,249 feet
-making altogether 72,584 feet; that is, very nearly thirteen and three-quarters British miles. Some of the 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, blank periods of enormous length. So that the lofty pile of sedimentary rocks in Britain gives but an inadequate idea of the time which has elapsed during their accumulation. The consideration of these various facts impresses the mind almost in the same manner as does the vain endeavour to grapple with the idea of eternity.
Nevertheless this impression is partly false. Mr. Croll, in an interesting paper, remarks that we do not err "in forming too great a conception of the length of geological periods," but in estimating them by years. When geologists look at large and complicated phenomena, and then at the figures representing several million years, the two produce a totally different effect on the mind, and the figures are at once pronounced too small. In regard to subaerial denudation, Mr. Croll shows, by calculating the known amount of sediment annually brought down by certain rivers, relatively to their areas of drainage, that 1000 feet of solid rock, as it became gradually disintegrated, would thus be removed from the mean level of the whole area in the course of six million years. This seems an astonishing result, and some considerations lead to the suspicion that it may be too large, but even if halved or quartered it is still very surprising. Few of us, however, know what a million really means: Mr. Croll gives the following illustration: take a narrow strip of paper, 83 feet 4 inches in length, and stretch it along the wall of a large hall; then mark off at one end the tenth of an inch. This tenth of an inch will represent one hundred years, and the entire strip a million years. But let it be borne in mind, in relation to the subject of this work, what a hundred years implies, represented as it is by a measure utterly insignificant in a hall of the above dimensions. Several eminent breeders, during a single lifetime, have so largely modified some of the higher animals which propagate their kind much more slowly than most of the lower animals, that they have formed what well deserves to be called a new sub-breed. Few men have attended with due care to any one strain for more than half a century, so that a hundred years represents the work of two breeders in succession. It is not to be supposed that species in a state of nature ever change so quickly as domestic animals under the guidance of methodical selection. The comparison would be in every way fairer with the effects which follow from unconscious selection, that is the preservation of the most useful or beautiful animals, with no intention of modifying the breed; but by this process of unconscious selection, various breeds have been sensibly changed in the course of two or three centuries.
Species, however, probably change much more slowly, and within the same country only a few change at the same time. This slowness follows from all the inhabitants of the same country being already so well adapted to each other, that new places in the polity of nature do not occur until after long intervals, due to the occurrence of physical changes of some kind, or through the immigration of new forms. Moreover variations or individual differences of the right nature, by which some of the inhabitants might be better fitted to their new places under the altered circumstances, would not always occur at once. Unfortunately we have no means of determining, according to the standards of years, how long a period it takes to modify a species; but to the subject of time we must return.
On the Poorness of Palaeontological Collections
Now let us turn to our richest geological museums, and what a paltry display we behold! That our collections are imperfect is admitted by every one. The remark of that admirable palaeontologist, Edward Forbes, should never be forgotten, namely, that very many 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 decay and disappear when left on the bottom of the sea, where sediment is not accumulating. We probably take a quite erroneous view, when we assume 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 immense 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 charged with carbolic acid. Some of the many kinds of animals which live on the beach between high and low water mark seem to be rarely 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 this has been found fossil in Sicily, whereas not one other species has hitherto been found in any tertiary formation: yet it is known that the genus Chthamalus existed during the Chalk period. Lastly, many great deposits requiring a vast length of time for their accumulation, are entirely destitute of organic remains, without our being able to assign any reason: one of the most striking instances is that of the Flysch formation, which consists of shale and sandstone, several thousand, occasionally even six thousand feet in thickness, and extending for at least 300 miles from Vienna to Switzerland; and although this great mass has been most carefully searched, no fossils, except a few vegetable remains, have been found.
With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods, it is superfluous to state that our evidence is fragmentary in an extreme degree. For instance, until recently not a land-shell was known belonging to either of these vast periods, with the exception of one species discovered by Sir C. Lyell and Dr. Dawson in the carboniferous strata of North America; but now land-shells have been found in the lias. In regard to mammiferous remains, a glance at the historical table published in 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 largely 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. This doctrine has been emphatically admitted by many geologists and palaeontologists, who, like E. Forbes, entirely disbelieve in the change of species. 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 confined exclusively 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 was derived, accord with the belief of vast intervals of time having elapsed between each formation.
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 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 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, 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 successive oscillations of level as well as the subsequent subaerial degradation. Such thick and extensive accumulations of sediment may be formed in two ways; either in profound depths of the sea, in which case the bottom will not be inhabited by so many and such varied forms of life, as the more shallow seas; and the mass when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its accumulation. Or, sediment may be deposited 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 the supply of sediment nearly balance each other, the sea will remain shallow and favourable for many and varied forms, and thus a rich fossiliferous formation, thick enough, when upraised, to resist a large amount of denudation, may be formed.
I am convinced that nearly all our ancient formations, which are throughout the greater part of their thickness 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 yet suffered, but which will hardly last to a distant geological age, was deposited during a downward oscillation of level, and thus gained considerable thickness.
All geological facts tell us plainly that each area has undergone 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, will 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 remains stationary, thick deposits cannot have been accumulated in the shallow parts, which are the most favourable to life. Still less can this have happened during the alternate periods of elevation; or, to speak more accurately, the beds which were then accumulated will generally have been destroyed by being upraised and brought within the limits of the coast-action.
These remarks apply chiefly to littoral and sub-littoral deposits. In the case of an extensive and shallow sea, such as that within a large part of the Malay Archipelago, where the depth varies from 30 or 40 to 60 fathoms, a widely extended formation might be formed during a period of elevation, and yet not suffer excessively from denudation during its slow upheaval; but the thickness of the formation could not be great, for owing to the elevatory movement it would be less than the depth in which it was formed; nor would the deposit be much consolidated, nor be capped by overlying formations, so that it would run a good chance of being worn away by atmospheric degradation and by the action of the sea during subsequent oscillations of level. It has, however, been suggested by Mr. Hopkins, that if one part of the area, after rising and before being denuded, subsided, the deposit formed during the rising movement, though not thick, might afterwards become protected by fresh accumulations, and thus be preserved for a long period.
Mr. Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. But all geologists, excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe, will admit that these latter rocks have been stript of their coverings to an enormous extent. For it is scarcely possible that such rocks could have been solidified and crystallized whilst uncovered; but if the metamorphic action occurred at profound depths of the ocean, the former protecting mantle of rock may not have been very thick. Admitting then that gneiss, mica-schist, granite, diorite, &c, were once necessarily covered up, how can we account for the naked and extensive areas of such rocks in many parts of the world, except on the belief that they have subsequently been completely denuded of all overlying strata? That such extensive areas do exist cannot be doubted: the granitic region of Parime is described by Humboldt as being as least nineteen times as large as Switzerland. South of the Amazon, Boue colours an area composed of rocks of this nature as equal to that of Spain, France, Italy, part of Germany, and the British Islands, all conjoined. This region has not been carefully explored, but from the concurrent testimony of travellers, the granitic area is very large: thus, von Eschwege gives a detailed section of these rocks, stretching from Rio de Janeiro for 260 geographical miles inland in a straight line; and I travelled for 150 miles in another direction, and saw nothing but granitic rocks. Numerous specimens, collected along the whole coast from near Rio de Janeiro to the mouth of the Plata, a distance of 1100 geographical miles, were examined by me, and they all belonged to this class. Inland, along the whole northern bank of the Plata I saw, besides modern tertiary beds, only one small patch of slightly metamorphosed rock, which alone could have formed a part of the original capping of the granitic series. Turning to a well-known region, namely, to the United States and Canada, as shown in Professor H. D. Rogers's beautiful map, I have estimated the areas by cutting out and weighing the paper, and I find that the metamorphic (excluding "the semi-metamorphic") and granitic rocks exceed, in the proportion of 19 to 12.5, the whole of the newer Palaeozoic formations. In many regions the metamorphic and granitic rocks would be found much more widely extended than they appear to be, if all the sedimentary beds were removed which rest unconformably on them, and which could not have formed part of the original mantle under which they were crystallized. Hence it is probable that in some parts of the world whole formations have been completely denuded, with not a wreck left behind.
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 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 on the shores of a continent when first broken up into an archipelago), and consequently during subsidence, though there will be much extinction, few new varieties or species will be formed; and it is during these very periods of subsidence, that the deposits which are richest in fossils have been accumulated.
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.
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 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 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.
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 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 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.
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 palaeontologist, that very thick deposits are usually barren of organic remains, except near their upper or lower limits.
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 1400 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.
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.
It is notorious on what excessively slight differences many palaeontologists 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.
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.
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.
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.
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.
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!
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.
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.
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.
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 palaeontologists, be ranked as new and distinct species.
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 palaeontologists, 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.
On the sudden Appearance of whole Groups of allied Species
The abrupt manner in which whole groups of species suddenly appear in certain formations, has been urged by several palaeontologists- for instance, by Agassiz, Pictet, and Sedgwick- as a fatal objection to the belief in the transmutation of species. If numerous species, belonging to the same genera or families, have really started into life at once, the fact would be fatal to the theory of evolution through natural selection. For the development by this means of a group of forms, all of which are descended from some one progenitor, must have been an extremely slow process; and the progenitors must have lived long before their modified descendants. But we continually overrate 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. In all cases positive palaeontological evidence may be implicitly trusted; negative evidence is worthless, as experience has so often shown. We continually forget how large the world is, compared with the area over which our geological formations have been carefully examined; we forget that groups of species may elsewhere have long existed, and have slowly multiplied, before they invaded the ancient archipelagoes of Europe and the United States. We do not make due allowance for the intervals of time which have elapsed between our consecutive formations,- longer perhaps in many cases than the time required for the accumulation of each formation. These intervals will have given time for the multiplication of species from some one parent-form: and in the succeeding formation, such groups or species will appear as if suddenly created.
I may here recall a remark formerly made, namely, that it might require a long succession of ages to adapt an organism to some new and peculiar line of life, for instance, to fly through the air; and consequently that the transitional forms would often long remain confined to some one region; but that, when this adaptation had once been effected, and a few species had thus acquired a great advantage over other organisms, a comparatively short time would be necessary to produce many divergent forms, which would spread rapidly and widely, throughout the world. Professor Pictet, in his excellent review of this work, in commenting on early transitional forms, and taking birds as an illustration, cannot see how the successive modifications of the anterior limbs of a supposed prototype could possibly have been of any advantage. But look at the penguins of the Southern Ocean; have not these birds their front limbs in this precise intermediate state of "neither true arms nor true wings"? Yet these birds hold their place victoriously in the battle for life; for they exist in infinite numbers and of many kinds. I do not suppose that we here see the real transitional grades through which the wings of birds have passed; but what special difficulty is there in believing that it might profit the modified descendants of the penguin, first to become enabled to flap along the surface of the sea like the logger-headed duck, and ultimately to rise from its surface and glide through the air?
I will now give a few examples to illustrate the foregoing remarks, and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. Even in so short an interval as that between the first and second editions of Pictet's great work on Palaeontology, published in 1844-46 and in 1853-57, the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified; and a third edition would require still further changes. I may recall the well-known fact that in geological treatises, published not many years ago, mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series; and true mammals have been discovered in the new red sandstone 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, as far back as the miocene stage. Had it not been for the rare accident of the preservation of the footsteps in the new red sandstone of the United States, who would have ventured to suppose that no less than at least thirty different bird-like animals, some of gigantic size, existed during that period? Not a fragment of bone has been discovered in these beds. Not long ago, palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period; but now we know, on the authority of Professor Owen, that a bird certainly lived during the deposition of the upper greensand; and still more recently, that strange bird, the Archeopteryx, with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws, has been discovered in the oolitic slates of Solenhofen. Hardly any recent discovery shows more forcibly than this, how little we as yet know of the former inhabitants of the world.
I may give another instance, which, from having passed under my own eyes, has much struck me. In a memoir On Fossil Sessile Cirripedes, I stated that, from the large number of existing and extinct tertiary species; from the extraordinary abundance of the individuals of many species all over the world, from the Arctic regions to the equator, inhabiting various zones of depths from the upper tidal limits to 50 fathoms; from the perfect manner in which specimens are preserved in the oldest tertiary beds; from the ease with which even a fragment of a valve 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 then thought one more instance of the abrupt appearance of a great group of species. But my work had hardly been published, when a skilful palaeontologist, M. Bosquet, sent me a drawing of a perfect specimen of an unmistakable sessile cirripede, which he had himself extracted from the chalk of Belgium. And, as if to make the case as striking as possible, this cirripede was a Chthamalus, a very common, large, and ubiquitous genus, of which not one species has as yet been found even in any tertiary stratum. Still more recently, a Pyrgoma, a member of a distinct subfamily of sessile cirripedes, has been discovered by Mr. Woodward in the upper chalk; so that we now have abundant evidence of the existence of this group of animals during the secondary period.
The case most frequently insisted on by palaeontologists of the apparently sudden appearance of a whole group of species, is that of the teleostean fishes, low down, according to Agassiz, in the Chalk period. This group includes the large majority of existing species. But certain Jurassic and Triassic forms are now commonly admitted to be teleostean; and even some palaeozoic forms have thus been classed by one high authority. If the teleosteans had really appeared suddenly in the northern hemisphere at the commencement of the chalk formation the fact would have been highly remarkable; but it would not have formed an insuperable difficulty, unless it could likewise have been shown that at the same period the species were suddenly and simultaneously developed in other quarters of the world. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator; and by running through Pictet's Palaeontology 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 fishes might formerly have had a similarly confined range, and after having been largely developed in some one sea, have spread widely. Nor have we any right to suppose that the seas of the world have always been so freely open from south to north as they are at present. Even at this day, if the Malay Archipelago were converted into land, the tropical parts of the Indian Ocean would form a large and perfectly enclosed basin, in which any great group of marine animals might be multiplied: and here they would remain confined, until some of the species became adapted to a cooler climate, and were enabled to double the Southern capes of Africa or Australia, and thus reach other and distant seas.
From these considerations, from our ignorance of the geology of other countries beyond the confines of Europe and the United States, and from the revolution in our palaeontological knowledge effected by the discoveries of the last dozen years, it seems to me to be about as rash to dogmatize on the succession of organic forms throughout the world, as it would be for a naturalist to land for five minutes on a barren point in Australia, and then to discuss the number and range of its productions.
On the Sudden Appearance of Groups of allied Species in the lowest known Fossiliferous StrataThere is another and allied difficulty, which is much more serious. I allude to the manner in which species belonging to several of the main divisions of the animal kingdom suddenly appear in the lowest known fossiliferous rocks. Most of the arguments which have convinced me that all the existing species of the same group are descended from a single progenitor, apply with equal force to the earliest known species. For instance, it cannot be doubted that all the Cambrian and Silurian trilobites are descended from some one crustacean, which must have lived long before the Cambrian age, and which probably differed greatly from any known animal. Some of the most ancient animals, as the Nautilus, Lingula, &c., do not differ much from living species; and it cannot on our theory be supposed, that these old species were the progenitors of all the species belonging to the same groups which have subsequently appeared, for they are not in any degree intermediate in character.
Consequently, if the theory be true, it is indisputable that before the lowest Cambrian stratum was deposited, long periods elapsed, as long as, or probably far longer than, the whole interval from the Cambrian age to the present day; and that during these vast periods the world swarmed with living creatures. Here we encounter a formidable objection; for it seems doubtful whether the earth, in a fit state for the habitation of living creatures, has lasted long enough. Sir W. Thompson concludes that the consolidation of the crust can hardly have occurred less than 20 or more than 400 million years ago, but probably not less than 98 or more than 200 million years. These very wide limits show how doubtful the data are; and other elements may have hereafter to be introduced into the problem. Mr. Croll estimates that about 60 million years have elapsed since the Cambrian period, but this, judging from the small amount of organic change since the commencement of the Glacial epoch, appears a very short time for the many and great mutations of life, which have certainly occurred since the Cambrian formation; and the previous 140 million years can hardly be considered as sufficient for the development of the varied forms of life which already existed during the Cambrian period. It is, however, probable, as Sir William Thompson insists, that the world at a very early period was subjected to more rapid and violent changes in its physical conditions than those now occurring; and such changes would have tended to induce changes at a corresponding rate in the organisms which then existed.
To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system, I can give no satisfactory answer. Several eminent geologists, with Sir R. Murchison at their head, were until recently convinced that we beheld in the organic remains of the lowest Silurian stratum the first dawn of life. Other highly competent judges, as Lyell and E. Forbes, have disputed this conclusion. We should not forget that only a small portion of the world is known with accuracy. Not very long ago M. Barrande added another and lower stage, abounding with new and peculiar species, beneath the then known Silurian system; and now, still lower down in the Lower Cambrian formation, Mr. Hicks has found in South Wales beds rich in trilobites, and containing various molluscs and annelids. The presence of phosphatic nodules and bituminous matter, even in some of the lowest azoic rocks, probably indicates life at these periods; and the existence of the Eozoon in the Laurentian formation of Canada is generally admitted. There are three great series of strata beneath the Silurian system in Canada, in the lowest of which the Eozoon is found. Sir W. Logan states that their "united thickness may possibly far surpass that of all the succeeding rocks, from the base of the palaeozoic series to the present time. We are thus carried back to a period so remote, that the appearance of the so-called primordial fauna (of Barrande) may by some be considered as a comparatively modern event." The Eozoon belongs to the most lowly organised, of all classes of animals, but is highly organised for its class; it existed in countless numbers, and, as Dr. Dawson has remarked, certainly preyed on other minute organic beings, which must have lived in great numbers. Thus the words, which I wrote in 1859, about the existence of living beings long before the Cambrian period, and which are almost the same with those since used by Sir W. Logan, have proved true. Nevertheless, the difficulty of assigning any good reason for the absence of vast piles of strata rich in fossils beneath the Cambrian system is very great. It does not seem probable that the most ancient beds have been quite worn away by denudation, or that their fossils have been wholly obliterated by metamorphic action, for if this had been the case we should have found only small remnants of the formations next succeeding them in age, and these would always have existed in partially metamorphosed condition. But the descriptions which we possess of the Silurian deposits over immense territories in Russia and in North America, do not support the view, that the older a formation is, the more invariably it has suffered extreme denudation and metamorphism.
The case at present must remain inexplicable; and may be truly urged as a valid argument against the views here entertained. To show that it may hereafter receive some explanation, I will give the following hypothesis. From the nature of the organic remains which do not appear to have inhabited profound depths, in the 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 now existing continents of Europe and North America. This same view has since been maintained by Agassiz and others. But we do not know what was the state of things in the intervals between the several successive formations; whether Europe and the United States during these intervals existed as dry land, or as a submarine surface near land, on which sediment was not deposited, or as the bed of an open and unfathomable sea.
Looking to the existing oceans, which are thrice as extensive as the land, we see them studded with many islands; but hardly one truly oceanic island (with the exception of New Zealand, if this can be called a truly oceanic island) is as yet known to afford even a remnant of any palaeozoic or secondary formation. Hence we may perhaps infer, that during the palaeozoic and secondary periods, neither continents nor continental islands existed where our oceans now extend; for had they existed, palaeozoic and secondary formations would in all probability have been accumulated from sediment derived from their wear and tear; and these would have been at least partially upheaved by the oscillations of level, which must have intervened during these enormously long periods. If then we may infer anything from these facts, we may infer that, where our oceans now extend, oceans have extended from the remotest period of which we have any record; and on the other hand, that where continents now exist, large tracts of land have existed, subjected no doubt to great oscillations of level, since the Cambrian 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 we have no reason to assume that things have thus remained from the beginning of the 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 long antecedent to the Cambrian 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 should we be justified in assuming that if, for instance, the bed of the Pacific Ocean were now converted into a continent we should there find sedimentary formations in a recognisable condition older than the Cambrian strata, supposing such to have been formerly deposited; for it might well happen that strata which had subsided some miles nearer to the centre of the earth, and which had been pressed on by an enormous weight of super-incumbent water, might have undergone far more metamorphic action than strata which have always remained nearer to the surface. The immense areas in some parts of the world, for instance in South America, of naked metamorphic rocks, which must have been heated under great pressure, have always seemed to me to require some special explanation; and we may perhaps believe that we see in these large areas, the many formations long anterior to the Cambrian epoch in a completely metamorphosed and denuded condition.
The several difficulties here discussed, namely- that, though we find in our geological formations many links between the species which now exist and which formerly existed, we do not find infinitely numerous fine transitional forms closely joining them all together;- the sudden manner in which several groups of species first appear in our European formations;- the almost entire absence, as at present known, of formations rich in fossils beneath the Cambrian strata,- are all undoubtedly of the most serious nature. We see this in the fact that the most eminent palaeontologists, namely Cuvier, Agassiz, Barrande, Pictet, Falconer, E. Forbes, &c., and all our greatest geologists, as Lyell, Murchison, Sedgwick, &c., have unanimously, often vehemently, maintained the immutability of species. But Sir Charles Lyell now gives the support of his high authority to the opposite side; and most geologists and palaeontologists are much shaken in their former belief. Those who believe that the geological record is in any degree perfect, will undoubtedly at once reject the theory. For my part, following out Lyell's metaphor, I look at the geological record as a history of the world imperfectly kept, and written in a changing dialect; of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved; and of each page, only here and there a few lines. Each word of the slowly-changing language, more or less different in the successive chapters, may represent the forms of life, which are entombed in our consecutive formations, and which falsely appear to have been abruptly introduced. On this view, the difficulties above discussed are greatly diminished, or even disappear.