The Origin of Species by Means of Natural Selection Part 17

ON THE POORNESS OF PALAEONTOLOGICAL COLLECTIONS.

Now let us turn to our richest 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. Sh.e.l.ls and bones decay and disappear when left on the bottom of the sea, where sediment is not acc.u.mulating. We probably take a quite erroneous view, when we a.s.sume 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 carbonic 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. .h.i.therto 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 acc.u.mulation, are entirely dest.i.tute of organic remains, without our being able to a.s.sign 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 ma.s.s 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-sh.e.l.l 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-sh.e.l.ls 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 acc.u.mulated. And if, in every 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 the 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 acc.u.mulated in extremely thick, solid, or extensive ma.s.ses, in order to withstand the incessant action of the waves, when first upraised and during subsequent oscillations of level, as well as the subsequent subaerial degradation.

Such thick and extensive acc.u.mulations 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 ma.s.s when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its acc.u.mulation. 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 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 publis.h.i.+ng 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 acc.u.mulated during subsidence. I may add, that the only ancient tertiary formation on the west coast of South America, which has been bulky enough to resist such degradation as it has as yet suffered, but which will hardly last to a distant geological age, was deposited during a downward oscillation of level, and thus gained considerable thickness.

All geological facts tell us plainly that each area has undergone numerous slow oscillations of level, and apparently these oscillations have affected wide s.p.a.ces. Consequently, formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation, will have been formed over wide s.p.a.ces 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 cannot have been acc.u.mulated 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 acc.u.mulated will generally have been destroyed by being upraised and brought within the limits of the coast-action.

These remarks apply chiefly to littoral and sublittoral 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 thirty or forty to sixty 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 acc.u.mulations, 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 stripped of their covering to an enormous extent. For it is scarcely possible that such rocks could have been solidified and crystallised while 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, etc., 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 at 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 1,100 geographical miles, were examined by me, and they all belonged to this cla.s.s. 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' 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 granite rocks exceed, in the proportion of 19 to 12.5, the whole of the newer Palaeozoic formations. In many regions the metamorphic and granite 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 crystallised. 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 pa.s.sing 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 circ.u.mstances 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 sh.o.r.es 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 acc.u.mulated.

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 a.s.sign 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 acc.u.mulating 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 s.p.a.ce 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 acc.u.mulating 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 acc.u.mulate 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 s.p.a.ce, 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 acc.u.mulation. 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 acc.u.mulation; 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 acc.u.mulation. 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.

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 a.s.signed, 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-relations.h.i.+p, 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 sub-stages 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 sh.e.l.ls believed by the majority of naturalists to be identical with existing species; but some excellent naturalists, as Aga.s.siz 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. If 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-form 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 sh.e.l.ls 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 to 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-sh.e.l.ls inhabiting the sh.o.r.es 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 a.s.serted over and over again, by writers who believe in the immutability of species, that geology yields no linking forms. This a.s.sertion, 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 ill.u.s.tration. 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. G.o.dwin-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, while most of our formations were acc.u.mulating. 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 acc.u.mulating. 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 acc.u.mulate on the bed of the sea, or where it did not acc.u.mulate 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 sh.o.r.es would be destroyed, almost as soon as acc.u.mulated, by the incessant coast-action, as we now see on the sh.o.r.es of South America. Even throughout the extensive and shallow seas within the archipelago, sedimentary beds could hardly be acc.u.mulated 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 acc.u.mulation 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 climatical 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 a.n.a.logy 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 a.s.suredly 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 Aga.s.siz, Pictet, and Sedgwick, as a fatal objection to the belief in the trans.m.u.tation 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 enormous intervals of time which have elapsed between our consecutive formations, longer perhaps in many cases than the time required for the acc.u.mulation 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 ill.u.s.tration, 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 pa.s.sed; 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 ill.u.s.trate 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 acc.u.mulations 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 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 cla.s.s 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 pa.s.sed 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 fifty 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 circ.u.mstances, 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 Aga.s.siz, in the Chalk period.

This group includes the large majority of existing species. But certain Jura.s.sic and Tria.s.sic forms are now commonly admitted to be teleostean; and even some palaeozoic forms have thus been cla.s.sed 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 STRATA.

There 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, etc., 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 twenty or more than four hundred million years ago, but probably not less than ninety-eight or more than two hundred 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 sixty 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 one hundred and forty 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 a.s.sumed 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 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 azotic 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 surpa.s.s 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 cla.s.ses of animals, but is highly organised for its cla.s.s; 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 a.s.signing 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 a 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.