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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.