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10 - On The Geological Succession of Organic Beings 10-03 - Species once lost do not reappear 10 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 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.

Fantail Pigeon
Fantail Pigeon

Rock Pigeon
Rock Pigeon
10 - On The Geological Succession of Organic Beings 10-04 - Groups of species follow the same general rules in their appearance and disappearance as do single species 10 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.

Edward Forbes
Edward Forbes


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.

lingula
lingula


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 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 space 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.
10 - On The Geological Succession of Organic Beings 10-05 - On Extinction 10 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.

Elie de Beaumont
Elie de Beaumont

Roderick Murchison
Roderick Murchison

Joachim Barrande
Joachim Barrande


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 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 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.

La Plata
La Plata

horse
horse

mastodon
mastodon

megatherium
megatherium

toxodon
toxodon

Land Shell
Land Shell

South America
South America


But how utterly groundless was my astonishment! Professor Owen soon perceived that the tooth, though so like that of the existing horse, belonged to an extinct species.

Richard Owen
Richard Owen


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.

elephant
elephant


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 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 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.

Death
Death


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.

cattle
cattle


Thus the appearance of new forms and the disappearance of old forms, both natural and artificial, are bound together.

In certain flourishing groups, the number of new specific forms which have been produced within a given time is probably greater than that of the old 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 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 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 class, 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 competition.

For instance, a single species of Trigonia, a great genus of shells 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.

trigonia
trigonia


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.

trilobite
trilobite

ammonite
ammonite


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.
10 - On The Geological Succession of Organic Beings 10-06 - On simultaneous changes in the forms of life throughout the world 10 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 America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India.

The Chalk Formation Needles
The Chalk Formation Needles

North America
North America

South America
South America

Tierra del Fuego
Tierra del Fuego

Cape of Good Hope
Cape of Good Hope

India
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.

Russia
Russia

europe
europe

North America
North America

Sir Charles Lyell
Sir Charles Lyell


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 coexisted with still living sea-shells; but as these anomalous monsters coexisted with the Mastodon and Horse, it might at least have been inferred that they had lived during one of the latter tertiary stages.

megatherium
megatherium

mylodon
mylodon

macrauchenia
macrauchenia

toxodon
toxodon

europe
europe

La Plata
La Plata

Sea Shell
Sea Shell

mastodon
mastodon

europe
europe

horse
horse


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.

europe
europe

South America
South America

Australia
Australia


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 shores of North America would hereafter be liable to be classed with somewhat older European beds.

United States
United States

europe
europe

North America
North America


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. de Verneuil and d'Archiac.

Adolphe dArchiac
Adolphe dArchiac


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 analogous 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.

Joachim Barrande
Joachim Barrande


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 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 competition 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 circumstanced 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 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 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 accumulated over very wide spaces 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 have occurred 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 assumed that an isthmus separated two seas inhabited by distinct, but contemporaneous, faunas.

England
England

France
France


Lyell has made similar observations on some of the later tertiary formations.

Sir Charles Lyell
Sir Charles Lyell


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.

Joachim Barrande
Joachim Barrande

Bohemia
Bohemia

Scandinavia
Scandinavia


If the several formations in these regions have not been deposited during the same exact 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 accumulation 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.