Part 12 (1/2)
Looking to the existing oceans, which are thrice as extensive as the land, we see them studded with many islands; but not one 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 there, palaeozoic and secondary formations would in all probability have been acc.u.mulated from sediment derived from their wear and tear; and would have been at least partially upheaved by the oscillations of level, which we may fairly conclude 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 earliest silurian period. The coloured map appended to my volume on Coral Reefs, led me to conclude that the great oceans are still mainly areas of {310} subsidence, the great archipelagoes still areas of oscillations of level, and the continents areas of elevation. But have we any right to a.s.sume that things have thus remained from the beginning of this world? Our continents seem to have been formed by a preponderance, during many oscillations of level, of the force of elevation; but may not the areas of preponderant movement have changed in the lapse of ages? At a period immeasurably antecedent to the silurian epoch, continents may have existed where oceans are now spread out; and clear and open oceans may have existed where our continents now stand. Nor should we be justified in a.s.suming that if, for instance, the bed of the Pacific Ocean were now converted into a continent, we should there find formations older than the silurian 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 superinc.u.mbent 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 bare 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 silurian epoch in a completely metamorphosed condition.
The several difficulties here discussed, namely our not finding in the successive formations infinitely numerous transitional links between the many species which now exist or have existed; the sudden manner {311} in which whole groups of species appear in our European formations; the almost entire absence, as at present known, of fossiliferous formations beneath the Silurian strata, are all undoubtedly of the gravest nature. We see this in the plainest manner by the fact that all the most eminent palaeontologists, namely Cuvier, Aga.s.siz, Barrande, Falconer, E. Forbes, &c., and all our greatest geologists, as Lyell, Murchison, Sedgwick, &c., have unanimously, often vehemently, maintained the immutability of species.
But I have reason to believe that one great authority, Sir Charles Lyell, from further reflexion entertains grave doubts on this subject. I feel how rash it is to differ from these authorities, to whom, with others, we owe all our knowledge. Those who think the natural geological record in any degree perfect, and who do not attach much weight to the facts and arguments of other kinds given in this volume, will undoubtedly at once reject my theory. For my part, following out Lyell's metaphor, I look at the natural geological record, as a history of the world 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, in which the history is supposed to be written, being more or less different in the interrupted succession of chapters, may represent the apparently abruptly changed forms of life, entombed in our consecutive, but widely separated, formations. On this view, the difficulties above discussed are greatly diminished, or even disappear.
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CHAPTER X.
ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEINGS.
On the slow and successive appearance of new species--On their different rates of change--Species once lost do not reappear--Groups of species follow the same general rules in their appearance and disappearance as do single species--On Extinction--On simultaneous changes in the forms of life throughout the world--On the affinities of extinct species to each other and to living species--On the state of development of ancient forms--On the succession of the same types within the same areas--Summary of preceding and present chapters.
Let us now see whether the several facts and rules relating to the geological succession of organic beings, better accord with the common view of the immutability of species, or with that of their slow and gradual modification, through descent and natural selection.
New species have appeared very slowly, one after another, both on the land and in the waters. Lyell has shown that it is hardly possible to resist the evidence on this head in the case of the several tertiary stages; and every year tends to fill up the blanks between them, and to make the percentage system of lost and new forms more gradual. In some of the most recent beds, though undoubtedly of high antiquity if measured by years, only one or two species are lost forms, and only one or two are new forms, having here appeared for the first time, either locally, or, as far as we know, on the face of the earth. If we may trust the observations of Philippi in Sicily, the successive changes in the marine inhabitants of that island have been many and most gradual. The secondary formations are more broken; but, as Bronn has remarked, neither the appearance {313} nor disappearance of their many now extinct species has been simultaneous in each separate formation.
Species of different genera and cla.s.ses have not changed at the same rate, or in the same degree. In the oldest tertiary beds a few living sh.e.l.ls may still be found in the midst of a mult.i.tude of extinct forms. Falconer has given a striking instance of a similar fact, in an existing crocodile a.s.sociated with many strange and lost mammals and reptiles in the sub-Himalayan deposits. The Silurian Lingula differs but little from the living species of this genus; whereas most of the other Silurian Molluscs and all the Crustaceans have changed greatly. The productions of the land seem to change at a quicker rate than those of the sea, of which a striking instance has lately been observed in Switzerland. There is some reason to believe that organisms, considered high in the scale of nature, change more quickly than those that are low: though there are exceptions to this rule.
The amount of organic change, as Pictet has remarked, does not strictly correspond with the succession of our geological formations; so that between each two consecutive formations, the forms of life have seldom changed in exactly the same degree. Yet if we compare any but the most closely related formations, all the species will be found to have undergone some change. When a species has once disappeared from the face of the earth, we have reason to believe that the same identical form never reappears. The strongest apparent exception to this latter rule, is that of the so-called ”colonies” of M. Barrande, which intrude for a period in the midst of an older formation, and then allow the pre-existing fauna to reappear; but Lyell's explanation, namely, that it is a case of temporary migration from a distinct geographical province, seems to me satisfactory.
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These several facts accord well with my theory. I believe in no fixed law of development, causing all the inhabitants of a country to change abruptly, or simultaneously, or to an equal degree. The process of modification must be extremely slow. The variability of each species is quite independent of that of all others. Whether such variability be taken advantage of by natural selection, and whether the variations be acc.u.mulated to a greater or lesser amount, thus causing a greater or lesser amount of modification in the varying species, depends on many complex contingencies,--on the variability being of a beneficial nature, on the power of intercrossing, on the rate of breeding, on the slowly changing physical conditions of the country, and more especially on the nature of the other inhabitants with which the varying species comes into compet.i.tion. Hence it is by no means surprising that one species should retain the same identical form much longer than others; or, if changing, that it should change less. We see the same fact in geographical distribution; for instance, in the land-sh.e.l.ls and coleopterous insects of Madeira having come to differ considerably from their nearest allies on the continent of Europe, whereas the marine sh.e.l.ls and birds have remained unaltered. We can perhaps understand the apparently quicker rate of change in terrestrial and in more highly organised productions compared with marine and lower productions, by the more complex relations of the higher beings to their organic and inorganic conditions of life, as explained in a former chapter. When many of the inhabitants of a country have become modified and improved, we can understand, on the principle of compet.i.tion, and on that of the many all-important relations of organism to organism, that any form which does not become in some degree modified and improved, {315} will be liable to be exterminated. Hence we can see why all the species in the same region do at last, if we look to wide enough intervals of time, become modified; for those which do not change will become extinct.
In members of the same cla.s.s the average amount of change, during long and equal periods of time, may, perhaps, be nearly the same; but as the acc.u.mulation of long-enduring fossiliferous formations depends on great ma.s.ses of sediment having been deposited on areas whilst subsiding, our formations have been almost necessarily acc.u.mulated at wide and irregularly intermittent intervals; consequently the amount of organic change exhibited by the fossils embedded in consecutive formations is not equal. Each formation, on this view, does not mark a new and complete act of creation, but only an occasional scene, taken almost at hazard, in a slowly changing drama.
We can clearly understand why a species when once lost should never reappear, even if the very same conditions of life, organic and inorganic, should recur. For though the offspring of one species might be adapted (and no doubt this has occurred in innumerable instances) to fill the exact place of another species in the economy of nature, and thus supplant it; yet the two forms--the old and the new--would not be identically the same; for both would almost certainly inherit different characters from their distinct progenitors. For instance, it is just possible, if our fantail-pigeons were all destroyed, that fanciers, by striving during long ages for the same object, might make a new breed hardly distinguishable from our present fantail; but if the parent rock-pigeon were also destroyed, and in nature we have every reason to believe that the parent-form will generally be supplanted and exterminated by its improved offspring, it is quite {316} incredible that a fantail, identical with the existing breed, could be raised from any other species of pigeon, or even from the other well-established races of the domestic pigeon, for the newly-formed fantail would be almost sure to inherit from its new progenitor some slight characteristic differences.
Groups of species, that is, genera and families, follow the same general rules in their appearance and disappearance as do single species, changing more or less quickly, and in a greater or lesser degree. A group does not reappear after it has once disappeared; or its existence, as long as it lasts, is continuous. I am aware that there are some apparent exceptions to this rule, but the exceptions are surprisingly few, so few that E. Forbes, Pictet, and Woodward (though all strongly opposed to such views as I maintain) admit its truth; and the rule strictly accords with my theory.
For as all the species of the same group have descended from some one species, it is clear that as long as any species of the group have appeared in the long succession of ages, so long must its members have continuously existed, in order to have generated either new and modified or the same old and unmodified forms. Species of the genus Lingula, for instance, must have continuously existed by an unbroken succession of generations, from the lowest Silurian stratum to the present day.
We have seen in the last chapter that the species of a group sometimes falsely appear to have come in abruptly; and I have attempted to give an explanation of this fact, which if true would have been fatal to my views.
But such cases are certainly exceptional; the general rule being a gradual increase in number, till the group reaches its maximum, and then, sooner or later, it gradually decreases. If the number of the species of a genus, or the number of {317} the genera of a family, be represented by a vertical line of varying thickness, crossing the successive geological formations in which the species are found, the line will sometimes falsely appear to begin at its lower end, not in a sharp point, but abruptly; it then gradually thickens upwards, sometimes keeping for a s.p.a.ce of equal thickness, and ultimately thins out in the upper beds, marking the decrease and final extinction of the species. This gradual increase in number of the species of a group is strictly conformable with my theory; as the species of the same genus, and the genera of the same family, can increase only slowly and progressively; for the process of modification and the production of a number of allied forms must be slow and gradual,--one species giving rise first to two or three varieties, these being slowly converted into species, which in their turn produce by equally slow steps other species, and so on, like the branching of a great tree from a single stem, till the group becomes large.
_On Extinction._--We have as yet spoken only incidentally of the disappearance of species and of groups of species. On the theory of natural selection the extinction of old forms and the production of new and improved forms are intimately connected together. The old notion of all the inhabitants of the earth having been swept away at successive periods by catastrophes, is very generally given up, even by those geologists, as Elie de Beaumont, Murchison, Barrande, &c, whose general views would naturally lead them to this conclusion. On the contrary, we have every reason to believe, from the study of the tertiary formations, that species and groups of species gradually disappear, one after another, first from one spot, then from another, and finally from the world. Both single species and whole {318} groups of species last for very unequal periods; some groups, as we have seen, having endured from the earliest known dawn of life to the present day; some having disappeared before the close of the palaeozoic period. No fixed law seems to determine the length of time during which any single species or any single genus endures. There is reason to believe that the complete extinction of the species of a group is generally a slower process than their production: if the appearance and disappearance of a group of species be represented, as before, by a vertical line of varying thickness, the line is found to taper more gradually at its upper end, which marks the progress of extermination, than at its lower end, which marks the first appearance and increase in numbers of the species. In some cases, however, the extermination of whole groups of beings, as of ammonites towards the close of the secondary period, has been wonderfully sudden.
The whole subject of the extinction of species has been involved in the most gratuitous mystery. Some authors have even supposed that as the individual has a definite length of life, so have species a definite duration. No one I think can have marvelled more at the extinction of species, than I have done. When I found in La Plata the tooth of a horse embedded with the remains of Mastodon, Megatherium, Toxodon, and other extinct monsters, which all co-existed with still living sh.e.l.ls at a very late geological period, I was filled with astonishment; for seeing that the horse, since its introduction by the Spaniards into South America, has run wild over the whole country and has increased in numbers at an unparalleled rate, I asked myself what could so recently have exterminated the former horse under conditions of life apparently so favourable. But how utterly groundless was my astonishment! {319} Professor Owen soon perceived that the tooth, though so like that of the existing horse, belonged to an extinct species. Had this horse been still living, but in some degree rare, no naturalist would have felt the least surprise at its rarity; for rarity is the attribute of a vast number of species of all cla.s.ses, in all countries. If we ask ourselves why this or that species is rare, we answer that something is unfavourable in its conditions of life; but what that something is, we can hardly ever tell. On the supposition of the fossil horse still existing as a rare species, we might have felt certain from the a.n.a.logy of all other mammals, even of the slow-breeding elephant, and from the history of the naturalisation of the domestic horse in South America, that under more favourable conditions it would in a very few years have stocked the whole continent. But we could not have told what the unfavourable conditions were which checked its increase, whether some one or several contingencies, and at what period of the horse's life, and in what degree, they severally acted. If the conditions had gone on, however slowly, becoming less and less favourable, we a.s.suredly should not have perceived the fact, yet the fossil horse would certainly have become rarer and rarer, and finally extinct;--its place being seized on by some more successful compet.i.tor.
It is most difficult always to remember that the increase of every living being is constantly being checked by unperceived injurious agencies; and that these same unperceived agencies are amply sufficient to cause rarity, and finally extinction. We see in many cases in the more recent tertiary formations, that rarity precedes extinction; and we know that this has been the progress of events with those animals which have been exterminated, either locally or wholly, through {320} man's agency. I may repeat what I published in 1845, namely, that to admit that species generally become rare before they become extinct--to feel no surprise at the rarity of a species, and yet to marvel greatly when it ceases to exist, is much the same as to admit that sickness in the individual is the forerunner of death--to feel no surprise at sickness, but when the sick man dies, to wonder and to suspect that he died by some unknown deed of violence.
The theory of natural selection is grounded on the belief that each new variety, and ultimately each new species, is produced and maintained by having some advantage over those with which it comes into compet.i.tion; and the consequent extinction of less-favoured forms almost inevitably follows.
It is the same with our domestic productions: when a new and slightly improved variety has been raised, it at first supplants the less improved varieties in the same neighbourhood; when much improved it is transported far and near, like our short-horn cattle, and takes the place of other breeds in other countries. Thus the appearance of new forms and the disappearance of old forms, both natural and artificial, are bound together. In certain flouris.h.i.+ng groups, the number of new specific forms which have been produced within a given time is probably greater than that of the old specific forms which have been exterminated; but we know that the number of species has not gone on indefinitely increasing, at least during the later geological periods, so that looking to later times we may believe that the production of new forms has caused the extinction of about the same number of old forms.
The compet.i.tion will generally be most severe, as formerly explained and ill.u.s.trated by examples, between the forms which are most like each other in all respects. {321} Hence the improved and modified descendants of a species will generally cause the extermination of the parent-species; and if many new forms have been developed from any one species, the nearest allies of that species, _i.e._ the species of the same genus, will be the most liable to extermination. Thus, as I believe, a number of new species descended from one species, that is a new genus, comes to supplant an old genus, belonging to the same family. But it must often have happened that a new species belonging to some one group will have seized on the place occupied by a species belonging to a distinct group, and thus caused its extermination; and if many allied forms be developed from the successful intruder, many will have to yield their places; and it will generally be allied forms, which will suffer from some inherited inferiority in common.