On the Origin of Species by Means of Natural Selection Part 12

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.

But whether it be species belonging to the same or to a distinct cla.s.s, which yield their places to other species which have been modified and improved, a few of the sufferers may often long be preserved, from being fitted to some peculiar line of life, or from inhabiting some distant and isolated station, where they have escaped severe compet.i.tion. For instance, a single species of Trigonia, a great genus of sh.e.l.ls in the secondary formations, survives 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 {322} 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 a new area, they will have exterminated in a correspondingly rapid manner many of the old inhabitants; and the forms which thus yield their places will commonly be allied, for they will partake of some inferiority in common.

Thus, as it seems to me, the manner in which single species and whole groups of species become extinct, accords well with the theory of natural selection. We need not marvel at extinction; if we must marvel, let it be at our presumption in imagining for a moment that we understand the many complex contingencies, on which the existence of each species depends. If we forget for an instant, that each species tends to increase inordinately, and that some check is always in action, yet seldom perceived by us, the whole economy of nature will be utterly obscured. Whenever we can precisely say why this species is more abundant in individuals than that; why this species and not another can be naturalised in a given country; then, and not till then, we may justly feel surprise why we cannot account for the extinction of this particular species or group of species.

_On the Forms of Life changing almost simultaneously throughout the World._--Scarcely any palaeontological discovery is more striking than the fact, that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant parts of the world, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North {323} America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the organic remains in certain beds present an unmistakeable degree of resemblance to those of the Chalk. It is not that the same species are met with; for in some cases not one species is identically the same, but they belong to the same families, genera, and sections of genera, and sometimes are similarly characterised in such trifling points as mere superficial sculpture.

Moreover other forms, which are not found in the Chalk of Europe, but which occur in the formations either above or below, are similarly absent at these distant points of the world. In the several successive palaeozoic formations of Russia, Western Europe and North America, a similar parallelism in the forms of life has been observed by several authors: so it is, according to Lyell, with the several European and North American tertiary deposits. Even if the few fossil species which are common to the Old and New Worlds be kept wholly out of view, the general parallelism in the successive forms of life, in the stages of the widely separated palaeozoic and tertiary periods, would still be manifest, and the several formations could be easily correlated.

These observations, however, relate to the marine inhabitants of distant parts of the world: we have not sufficient data to judge whether the productions of the land and of fresh water change at distant points in the same parallel manner. We may doubt whether they have thus changed: if the Megatherium, Mylodon, Macrauchenia, and Toxodon had been brought to Europe from La Plata, without any information in regard to their geological position, no one would have suspected that they had co-existed with still living sea-sh.e.l.ls; but as these anomalous monsters co-existed with the {324} Mastodon and Horse, it might at least have been inferred that they had lived during one of the later tertiary stages.

When the marine forms of life are spoken of as having changed simultaneously throughout the world, it must not be supposed that this expression relates to the same thousandth or hundred-thousandth year, or even that it has a very strict geological sense; for if all the marine animals which live at the present day in Europe, and all those that lived in Europe during the pleistocene period (an enormously remote period as measured by years, including the whole glacial epoch), were to be compared with those now living in South America or in Australia, the most skilful naturalist would hardly be able to say whether the existing or the pleistocene inhabitants of Europe resembled most closely those of the southern hemisphere. So, again, several highly competent observers believe that the existing productions of the United States are more closely related to those which lived in Europe during certain later tertiary stages, than to those which now live here; and if this be so, it is evident that fossiliferous beds deposited at the present day on the sh.o.r.es of North America would hereafter be liable to be cla.s.sed with somewhat older European beds. Nevertheless, looking to a remotely future epoch, there can, I think, be little doubt that all the more modern _marine_ formations, namely, the upper pliocene, the pleistocene and strictly modern beds, of Europe, North and South America, and Australia, from containing fossil remains in some degree allied, and from not including those forms which are only found in the older underlying deposits, would be correctly ranked as simultaneous in a geological sense.

The fact of the forms of life changing simultaneously, in the above large sense, at distant parts of the world, has greatly struck those admirable observers, MM. {325} de Verneuil and d'Archiac. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe, they add, "If struck by this strange sequence, we turn our attention to North America, and there discover a series of a.n.a.logous phenomena, it will appear certain that all these modifications of species, their extinction, and the introduction of new ones, cannot be owing to mere changes in marine currents or other causes more or less local and temporary, but depend on general laws which govern the whole animal kingdom." M. Barrande has made forcible remarks to precisely the same effect. It is, indeed, quite futile to look to changes of currents, climate, or other physical conditions, as the cause of these great mutations in the forms of life throughout the world, under the most different climates. We must, as Barrande has remarked, look to some special law. We shall see this more clearly when we treat of the present distribution of organic beings, and find how slight is the relation between the physical conditions of various countries, and the nature of their inhabitants.

This great fact of the parallel succession of the forms of life throughout the world, is explicable on the theory of natural selection. New species are formed by new varieties arising, which have some advantage over older forms; and those forms, which are already dominant, or have some advantage over the other forms in their own country, would naturally oftenest give rise to new varieties or incipient species; for these latter must be victorious in a still higher degree in order to be preserved and to survive. We have distinct evidence on this head, in the plants which are dominant, that is, which are commonest in their own homes, and are most widely diffused, having produced the greatest number of new varieties. It is also natural that the {326} dominant, varying, and far-spreading species, which already have invaded to a certain extent the territories of other species, should be those which would have the best chance of spreading still further, and of giving rise in new countries to new varieties and species. The process of diffusion may often be very slow, being dependent on climatal and geographical changes, or on strange accidents, but in the long run the dominant forms will generally succeed in spreading. The diffusion would, it is probable, be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. We might therefore expect to find, as we apparently do find, a less strict degree of parallel succession in the productions of the land than of the sea.

Dominant species spreading from any region might encounter still more dominant species, and then their triumphant course, or even their existence, would cease. We know not at all precisely what are all the conditions most favourable for the multiplication of new and dominant species; but we can, I think, clearly see that a number of individuals, from giving a better chance of the appearance of favourable variations, and that severe compet.i.tion with many already existing forms, would be highly favourable, as would be the power of spreading into new territories. A certain amount of isolation, recurring at long intervals of time, would probably be also favourable, as before explained. One quarter of the world may have been most favourable for the production of new and dominant species on the land, and another for those in the waters of the sea. If two great regions had been for a long period favourably circ.u.mstanced in an equal degree, whenever their inhabitants met, the battle would be prolonged and severe; and some from one birthplace and some from the other might be victorious. But in the course of time, the {327} forms dominant in the highest degree, wherever produced, would tend everywhere to prevail. As they prevailed, they would cause the extinction of other and inferior forms; and as these inferior forms would be allied in groups by inheritance, whole groups would tend slowly to disappear; though here and there a single member might long be enabled to survive.

Thus, as it seems to me, the parallel, and, taken in a large sense, simultaneous, succession of the same forms of life throughout the world, accords well with the principle of new species having been formed by dominant species spreading widely and varying; the new species thus produced being themselves dominant owing to inheritance, and to having already had some advantage over their parents or over other species; these again spreading, varying, and producing new species. The forms which are beaten and which yield their places to the new and victorious forms, will generally be allied in groups, from inheriting some inferiority in common; and therefore as new and improved groups spread throughout the world, old groups will disappear from the world; and the succession of forms in both ways will everywhere tend to correspond.

There is one other remark connected with this subject worth making. I have given my reasons for believing that all our greater fossiliferous formations were deposited during periods of subsidence; and that blank intervals of vast duration occurred during the periods when the bed of the sea was either stationary or rising, and likewise when sediment was not thrown down quickly enough to embed and preserve organic remains. During these long and blank intervals I suppose that the inhabitants of each region underwent a considerable amount of modification and extinction, and that there was much migration from {328} other parts of the world. As we have reason to believe that large areas are affected by the same movement, it is probable that strictly contemporaneous formations have often been acc.u.mulated over very wide s.p.a.ces in the same quarter of the world; but we are far from having any right to conclude that this has invariably been the case, and that large areas have invariably been affected by the same movements. When two formations have been deposited in two regions during nearly, but not exactly the same period, we should find in both, from the causes explained in the foregoing paragraphs, the same general succession in the forms of life; but the species would not exactly correspond; for there will have been a little more time in the one region than in the other for modification, extinction, and immigration.

I suspect that cases of this nature occur in Europe. Mr. Prestwich, in his admirable Memoirs on the eocene deposits of England and France, is able to draw a close general parallelism between the successive stages in the two countries; but when he compares certain stages in England with those in France, although he finds in both a curious accordance in the numbers of the species belonging to the same genera, yet the species themselves differ in a manner very difficult to account for, considering the proximity of the two areas,--unless, indeed, it be a.s.sumed that an isthmus separated two seas inhabited by distinct, but contemporaneous, faunas. Lyell has made similar observations on some of the later tertiary formations. Barrande, also, shows that there is a striking general parallelism in the successive Silurian deposits of Bohemia and Scandinavia; nevertheless he finds a surprising amount of difference in the species. If the several formations in these regions have not been deposited during the same exact {329} periods,--a formation in one region often corresponding with a blank interval in the other,--and if in both regions the species have gone on slowly changing during the acc.u.mulation of the several formations and during the long intervals of time between them; in this case, the several formations in the two regions could be arranged in the same order, in accordance with the general succession of the form of life, and the order would falsely appear to be strictly parallel; nevertheless the species would not all be the same in the apparently corresponding stages in the two regions.

_On the Affinities of extinct Species to each other, and to living forms._--Let us now look to the mutual affinities of extinct and living species. They all fall into one grand natural system; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, all fossils can be cla.s.sed either in still existing groups, or between them. That the extinct forms of life help to fill up the wide intervals between existing genera, families, and orders, cannot be disputed. For if we confine our attention either to the living or to the extinct alone, the series is far less perfect than if we combine both into one general system. With respect to the Vertebrata, whole pages could be filled with striking ill.u.s.trations from our great palaeontologist, Owen, showing how extinct animals fall in between existing groups. Cuvier ranked the Ruminants and Pachyderms, as the two most distinct orders of mammals; but Owen has discovered so many fossil links, that he has had to alter the whole cla.s.sification of these two orders; and has placed certain pachyderms in the same sub-order with ruminants: for example, he dissolves by fine gradations the apparently {330} wide difference between the pig and the camel. In regard to the Invertebrata, Barrande, and a higher authority could not be named, a.s.serts that he is every day taught that Palaeozoic animals, though belonging to the same orders, families, or genera with those living at the present day, were not at this early epoch limited in such distinct groups as they now are.

Some writers have objected to any extinct species or group of species being considered as intermediate between living species or groups. If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms, the objection is probably valid. But I apprehend that in a perfectly natural cla.s.sification many fossil species would have to stand between living species, and some extinct genera between living genera, even between genera belonging to distinct families. The most common case, especially with respect to very distinct groups, such as fish and reptiles, seems to be, that supposing them to be distinguished at the present day from each other by a dozen characters, the ancient members of the same two groups would be distinguished by a somewhat lesser number of characters, so that the two groups, though formerly quite distinct, at that period made some small approach to each other.

It is a common belief that the more ancient a form is, by so much the more it tends to connect by some of its characters groups now widely separated from each other. This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages; and it would be difficult to prove the truth of the proposition, for every now and then even a living animal, as the Lepidosiren, is discovered having affinities directed towards very distinct groups. Yet if we compare the older Reptiles and {331} Batrachians, the older Fish, the older Cephalopods, and the eocene Mammals, with the more recent members of the same cla.s.ses, we must admit that there is some truth in the remark.

Let us see how far these several facts and inferences accord with the theory of descent with modification. As the subject is somewhat complex, I must request the reader to turn to the diagram in the fourth chapter. We may suppose that the numbered letters represent genera, and the dotted lines diverging from them the species in each genus. The diagram is much too simple, too few genera and too few species being given, but this is unimportant for us. The horizontal lines may represent successive geological formations, and all the forms beneath the uppermost line may be considered as extinct. The three existing genera, a^{14}, q^{14}, p^{14}, will form a small family; b^{14} and f^{14} a closely allied family or sub-family; and o^{14}, e^{14}, m^{14}, a third family. These three families, together with the many extinct genera on the several lines of descent diverging from the parent-form (A), will form an order; for all will have inherited something in common from their ancient and common progenitor. On the principle of the continued tendency to divergence of character, which was formerly ill.u.s.trated by this diagram, the more recent any form is, the more it will generally differ from its ancient progenitor.

Hence we can understand the rule that the most ancient fossils differ most from existing forms. We must not, however, a.s.sume that divergence of character is a necessary contingency; it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature. Therefore it is quite possible, as we have seen in the case of some Silurian forms, that a species might go on being slightly modified in relation to its slightly altered conditions of {332} life, and yet retain throughout a vast period the same general characteristics. This is represented in the diagram by the letter F^{14}.

All the many forms, extinct and recent, descended from (A), make, as before remarked, one order; and this order, from the continued effects of extinction and divergence of character, has become divided into several sub-families and families, some of which are supposed to have perished at different periods, and some to have endured to the present day.

By looking at the diagram we can see that if many of the extinct forms, supposed to be embedded in the successive formations, were discovered at several points low down in the series, the three existing families on the uppermost line would be rendered less distinct from each other. If, for instance, the genera a^1, a^5, a^{10}, f^8, m^3, m^6, m^9, were disinterred, these three families would be so closely linked together that they probably would have to be united into one great family, in nearly the same manner as has occurred with ruminants and pachyderms. Yet he who objected to call the extinct genera, which thus linked the living genera of three families together, intermediate in character, would be justified, as they are intermediate, not directly, but only by a long and circuitous course through many widely different forms. If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations --for instance, above No. VI.--but none from beneath this line, then only the two families on the left hand (namely, a^{14}, &c, and b^{14},&c.) would have to be united into one family; and the two other families (namely, a^{14} to f^{14} now including five genera, and o^{14} to m^{14}) would yet remain distinct. These two families, however, would be less distinct from each other than they were before the discovery of the fossils. If, for instance, we suppose the existing genera of the two families to differ from each {333} other by a dozen characters, in this case the genera, at the early period marked VI., would differ by a lesser number of characters; for at this early stage of descent they have not diverged in character from the common progenitor of the order, nearly so much as they subsequently diverged. Thus it comes that ancient and extinct genera are often in some slight degree intermediate in character between their modified descendants, or between their collateral relations.

In nature the case will be far more complicated than is represented in the diagram; for the groups will have been more numerous, they will have endured for extremely unequal lengths of time, and will have been modified in various degrees. As we possess only the last volume of the geological record, and that in a very broken condition, we have no right to expect, except in very rare cases, to fill up wide intervals in the natural system, and thus unite distinct families or orders. All that we have a right to expect, is that those groups, which have within known geological periods undergone much modification, should in the older formations make some slight approach to each other; so that the older members should differ less from each other in some of their characters than do the existing members of the same groups; and this by the concurrent evidence of our best palaeontologists seems frequently to be the case.

Thus, on the theory of descent with modification, the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms, seem to me explained in a satisfactory manner. And they are wholly inexplicable on any other view.

On this same theory, it is evident that the fauna of any great period in the earth's history will be intermediate in general character between that which preceded and that which succeeded it. Thus, the species which lived at the sixth great stage of descent in the {334} diagram are the modified offspring of those which lived at the fifth stage, and are the parents of those which became still more modified at the seventh stage; hence they could hardly fail to be nearly intermediate in character between the forms of life above and below. We must, however, allow for the entire extinction of some preceding forms, and in any one region for the immigration of new forms from other regions, and for a large amount of modification, during the long and blank intervals between the successive formations. Subject to these allowances, the fauna of each geological period undoubtedly is intermediate in character, between the preceding and succeeding faunas. I need give only one instance, namely, the manner in which the fossils of the Devonian system, when this system was first discovered, were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous, and underlying Silurian system. But each fauna is not necessarily exactly intermediate, as unequal intervals of time have elapsed between consecutive formations.