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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 but likewise 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 have existed, we do not find infinitely numerous fine transitional forms closely joining them all together ;—the sudden manner in which several whole groups of species first appeared in our European formations ;—the almost entire absence, as at present known, of fossiliferous formations beneath the Silurian strata, -are all undoubtedly of the most serious nature. We see this in the fact that the most eminent palæontologists, 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 I have reason to believe that one great authority, Sir Charles Lyell, from further reflection 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 written, being more or less different in the successive 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.
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 reappearGroups of species follow the same general rules in their appearance and disappearance as do single species-On ExtinctionOn 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 formsOn the succession of the same types within the same areas
Summary of preceding and present chapter. 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 nor disappearance of their many now extinct species has been simultaneous in each separate formation.
Species of different genera and classes have not changed at the same rate, or in the same degree. In the older tertiary beds a few living shells may still be found in the midst of a multitude of extinct forms. Falconer has given a striking instance of a similar fact, in an existing crocodile associated with many strange 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 geological formations ; 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 no reason to believe that the same identical form ever 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.
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, 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 accumulated 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 competition. 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-shells and coleopterous insects of Madeira having come to differ considerably from their nearest allies on the continent of Europe, whereas the marine shells 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 competition, 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 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 class the average amount of change, during long and equal periods of time, may, perhaps, be nearly the same; but as the accumulation of long-enduring fossiliferous formations depends on great masses of sediment having been deposited on areas whilst subsiding, our formations have been almost necessarily accumulated 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,
if the very 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 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
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 in a body; 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 the genera of a family, be represented by a vertical line of varying thickness, ascending through 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, often keeping for a space of equal thickness, and ultimately