The Origin of Species by Means of Natural Selection
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 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 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 superincumbent 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, etc., and all our greatest geologists, as Lyell, Murchison, Sedgwick, etc., 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 my 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.
CHAPTER XI. 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 laws relating to the geological succession of organic beings accord best with the common view of the immutability of species, or with that of their slow and gradual modification, through variation 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 the stages, and to make the proportion between the lost and existing 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 extinct, and only one or two are new, having appeared there for the first time, either locally, or, as far as we know, on the face of the earth. The secondary formations are more broken; but, as Bronn has remarked, neither the appearance nor disappearance of the many species embedded in each formation has been simultaneous.
Species belonging to 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, for an existing crocodile is associated with many 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 have changed at a quicker rate than those of the sea, of which a striking instance has been observed in Switzerland. There is some reason to believe that organisms high in the scale, change more quickly than those that are low: though there are exceptions to this rule. The amount of organic change, as Pictet has remarked, is not the same in each successive so-called formation. 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 satisfactory.
These several facts accord well with our theory, which includes no fixed law of development, causing all the inhabitants of an area to change abruptly, or simultaneously, or to an equal degree. The process of modification must be slow, and will generally affect only a few species at the same time; for the variability of each species is independent of that of all others. Whether such variations or individual differences as may arise will be accumulated through natural selection in a greater or less degree, thus causing a greater or less amount of permanent modification, will depend on many complex contingencies – on the variations being of a beneficial nature, on the freedom of intercrossing, on the slowly changing physical conditions of the country, on the immigration of new colonists, and on the nature of the other inhabitants with which the varying species come 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, should change in a less degree. We find similar relations between the existing inhabitants of distinct countries; for instance, the land-shells and coleopterous insects of Madeira have 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 any area have become modified and improved, we can understand, on the principle of competition, and from the all-important relations of organism to organism in the struggle for life, that any form which did not become in some degree modified and improved, would be liable to extermination. Hence, we see why all the species in the same region do at last, if we look to long enough intervals of time, become modified; for otherwise they would 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 enduring formations, rich in fossils, depends on great masses of sediment being deposited on subsiding areas, our formations have been almost necessarily accumulated at wide and irregularly intermittent intervals of time; 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 an ever 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 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; and organisms already differing would vary in a different manner. For instance, it is possible, if all our fantail-pigeons were destroyed, that fanciers might make a new breed hardly distinguishable from the present breed; but if the parent rock-pigeon were likewise destroyed, and under nature we have every reason to believe that parent forms are generally supplanted and exterminated by their improved offspring, it is incredible that a fantail, identical with the existing breed, could be raised from any other species of pigeon, or even from any other well established race of the domestic pigeon, for the successive variations would almost certainly be in some degree different, and the newly-formed variety would probably inherit from its progenitor some 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, when it has once disappeared, never reappears; that is, 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 the theory. For all the species of the same group, however long it may have lasted, are the modified descendants one from the other, and all from a common progenitor. In the genus Lingula, for instance, the species which have successively appeared at all ages must have been connected by an unbroken series of generations, from the lowest Silurian stratum to the present day.
We have seen in the last chapter that whole groups of species sometimes falsely appear to have been abruptly developed; and I have attempted to give an explanation of this fact, which if true would be fatal to my views. But such cases are certainly exceptional; the general rule being a gradual increase in number, until the group reaches its maximum, and then, sooner or later, a gradual decrease. If the number of the species included within a genus, or the number of the genera within 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 of equal thickness for a space, 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 the theory; for the species of the same genus, and the genera of the same family, can increase only slowly and progressively; the process of modification and the production of a number of allied forms necessarily being a slow and gradual process, one species first giving rise to two or three varieties, these being slowly converted into species, which in their turn produce by equally slow steps other varieties and 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 only spoken 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 by catastrophes at successive periods is very generally given up, even by those geologists, as Elie de Beaumont, Murchison, Barrande, etc., 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. In some few cases, however, as by the breaking of an isthmus and the consequent irruption of a multitude of new inhabitants into an adjoining sea, or by the final subsidence of an island, the process of extinction may have been rapid. Both single species and whole groups of species last for very unequal periods; some groups, as we have seen, have endured from the earliest known dawn of life to the present day; some have 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 extinction of a whole group of species is generally a slower process than their production: if their appearance and disappearance 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 the early increase in number of the species. In some cases, however, the extermination of whole groups, as of ammonites, towards the close of the secondary period, has been wonderfully sudden.
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 can have marvelled more than I have done at the extinction of species. 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 shells 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 my astonishment was groundless. 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 classes, 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 analogy 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 assuredly 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 competitor.
It is most difficult always to remember that the increase of every living creature is constantly being checked by unperceived hostile agencies; and that these same unperceived agencies are amply sufficient to cause rarity, and finally extinction. So little is this subject understood, that I have heard surprise repeatedly expressed at such great monsters as the Mastodon and the more ancient Dinosaurians having become extinct; as if mere bodily strength gave victory in the battle of life. Mere size, on the contrary, would in some cases determine, as has been remarked by Owen, quicker extermination, from the greater amount of requisite food. Before man inhabited India or Africa, some cause must have checked the continued increase of the existing elephant. A highly capable judge, Dr. Falconer, believes that it is chiefly insects which, from incessantly harassing and weakening the elephant in India, check its increase; and this was Bruce's conclusion with respect to the African elephant in Abyssinia. It is certain that insects and blood-sucking bats determine the existence of the larger naturalised quadrupeds in several parts of South America.
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 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 the species 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 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 competition; 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 those naturally and artificially produced, are bound together. In flourishing groups, the number of new specific forms which have been produced within a given time has at some periods probably been greater than the number of the old specific forms which have been exterminated; but we know that species have not gone on indefinitely increasing, at least during the later geological epochs, 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 competition will generally be most severe, as formerly explained and illustrated by examples, between the forms which are most like each other in all respects. 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 has seized on the place occupied by a species belonging to a distinct group, and thus have caused its extermination. If many allied forms be developed from the successful intruder, many will have to yield their places; and it will generally be the allied forms, which will suffer from some inherited inferiority in common. But whether it be species belonging to the same or to a distinct class, which have yielded their places to other modified and improved species, a few of the sufferers may often be preserved for a long time, from being fitted to some peculiar line of life, or from inhabiting some distant and isolated station, where they will have escaped severe competition. For instance, some species of Trigonia, a great genus of shells in the secondary formations, survive in the Australian seas; and a few members of the great and almost extinct group of Ganoid fishes still inhabit our fresh waters. Therefore, the utter extinction of a group is generally, as we have seen, a slower process than its production.
With respect to the apparently sudden extermination of whole families or orders, as of Trilobites at the close of the palaeozoic period, and of Ammonites at the close of the secondary period, we must remember what has been already said on the probable wide intervals of time between our consecutive formations; and in these intervals there may have been much slow extermination. Moreover, when, by sudden immigration or by unusually rapid development, many species of a new group have taken possession of an area, many of the older species will have been exterminated in a correspondingly rapid manner; and the forms which thus yield their places will commonly be allied, for they will partake of the same inferiority in common.
