The Origin of Species

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by Charles Darwin

of their production, and still less with the order of their disappearance;

  for the parent rock-pigeon now lives; and many varieties between the

  rock-pigeon and the carrier have become extinct; and carriers which are

  extreme in the important character of length of beak originated earlier

  than short-beaked tumblers, which are at the opposite end of the series in

  this same respect.

  Closely connected with the statement, that the organic remains from an

  intermediate formation are in some degree intermediate in character, is the

  fact, insisted on by all palaeontologists, that fossils from two

  consecutive formations are far more closely related to each other, than are

  the fossils from two remote formations. Pictet gives as a well-known

  instance, the general resemblance of the organic remains from the several

  stages of the chalk formation, though the species are distinct in each

  stage. This fact alone, from its generality, seems to have shaken

  Professor Pictet in his firm belief in the immutability of species. He who

  is acquainted with the distribution of existing species over the globe,

  will not attempt to account for the close resemblance of the distinct

  species in closely consecutive formations, by the physical conditions of

  the ancient areas having remained nearly the same. Let it be remembered

  that the forms of life, at least those inhabiting the sea, have changed

  almost simultaneously throughout the world, and therefore under the most

  different climates and conditions. Consider the prodigious vicissitudes of

  climate during the pleistocene period, which includes the whole glacial

  period, and note how little the specific forms of the inhabitants of the

  sea have been affected.

  On the theory of descent, the full meaning of the fact of fossil remains

  from closely consecutive formations, though ranked as distinct species,

  being closely related, is obvious. As the accumulation of each formation

  has often been interrupted, and as long blank intervals have intervened

  between successive formations, we ought not to expect to find, as I

  attempted to show in the last chapter, in any one or two formations all the

  intermediate varieties between the species which appeared at the

  commencement and close of these periods; but we ought to find after

  intervals, very long as measured by years, but only moderately long as

  measured geologically, closely allied forms, or, as they have been called

  by some authors, representative species; and these we assuredly do find.

  We find, in short, such evidence of the slow and scarcely sensible mutation

  of specific forms, as we have a just right to expect to find.

  On the state of Development of Ancient Forms. -- There has been much

  discussion whether recent forms are more highly developed than ancient. I

  will not here enter on this subject, for naturalists have not as yet

  defined to each other's satisfaction what is meant by high and low forms.

  But in one particular sense the more recent forms must, on my theory, be

  higher than the more ancient; for each new species is formed by having had

  some advantage in the struggle for life over other and preceding forms. If

  under a nearly similar climate, the eocene inhabitants of one quarter of

  the world were put into competition with the existing inhabitants of the

  same or some other quarter, the eocene fauna or flora would certainly be

  beaten and exterminated; as would a secondary fauna by an eocene, and a

  palaeozoic fauna by a secondary fauna. I do not doubt that this process of

  improvement has affected in a marked and sensible manner the organisation

  of the more recent and victorious forms of life, in comparison with the

  ancient and beaten forms; but I can see no way of testing this sort of

  progress. Crustaceans, for instance, not the highest in their own class,

  may have beaten the highest molluscs. From the extraordinary manner in

  which European productions have recently spread over New Zealand, and have

  seized on places which must have been previously occupied, we may believe,

  if all the animals and plants of Great Britain were set free in New

  Zealand, that in the course of time a multitude of British forms would

  become thoroughly naturalized there, and would exterminate many of the

  natives. On the other hand, from what we see now occurring in New Zealand,

  and from hardly a single inhabitant of the southern hemisphere having

  become wild in any part of Europe, we may doubt, if all the productions of

  New Zealand were set free in Great Britain, whether any considerable number

  would be enabled to seize on places now occupied by our native plants and

  animals. Under this point of view, the productions of Great Britain may be

  said to be higher than those of New Zealand. Yet the most skilful

  naturalist from an examination of the species of the two countries could

  not have foreseen this result.

  Agassiz insists that ancient animals resemble to a certain extent the

  embryos of recent animals of the same classes; or that the geological

  succession of extinct forms is in some degree parallel to the embryological

  development of recent forms. I must follow Pictet and Huxley in thinking

  that the truth of this doctrine is very far from proved. Yet I fully

  expect to see it hereafter confirmed, at least in regard to subordinate

  groups, which have branched off from each other within comparatively recent

  times. For this doctrine of Agassiz accords well with the theory of

  natural selection. In a future chapter I shall attempt to show that the

  adult differs from its embryo, owing to variations supervening at a not

  early age, and being inherited at a corresponding age. This process,

  whilst it leaves the embryo almost unaltered, continually adds, in the

  course of successive generations, more and more difference to the adult.

  Thus the embryo comes to be left as a sort of picture, preserved by nature,

  of the ancient and less modified condition of each animal. This view may

  be true, and yet it may never be capable of full proof. Seeing, for

  instance, that the oldest known mammals, reptiles, and fish strictly belong

  to their own proper classes, though some of these old forms are in a slight

  degree less distinct from each other than are the typical members of the

  same groups at the present day, it would be vain to look for animals having

  the common embryological character of the Vertebrata, until beds far

  beneath the lowest Silurian strata are discovered--a discovery of which the

  chance is very small.

  On the Succession of the same Types within the same areas, during the later

  tertiary periods. -- Mr. Clift many years ago showed that the fossil

  mammals from the Australian caves were closely allied to the living

  marsupials of that continent. In South America, a similar relationship is

  manifest, even to an uneducated eye, in the gigantic pieces of armour like

  those of the armadillo, found in several parts of La Plata; and Professor

  Owen has shown in the most striking manner that most of the fossil mammals,

  buried there in such numbers, are related to South American types. This

  relationship is even more clearly seen i
n the wonderful collection of

  fossil bones made by MM. Lund and Clausen in the caves of Brazil. I was so

  much impressed with these facts that I strongly insisted, in 1839 and 1845,

  on this 'law of the succession of types,'--on 'this wonderful relationship

  in the same continent between the dead and the living.' Professor Owen has

  subsequently extended the same generalisation to the mammals of the Old

  World. We see the same law in this author's restorations of the extinct

  and gigantic birds of New Zealand. We see it also in the birds of the

  caves of Brazil. Mr. Woodward has shown that the same law holds good with

  sea-shells, but from the wide distribution of most genera of molluscs, it

  is not well displayed by them. Other cases could be added, as the relation

  between the extinct and living land-shells of Madeira; and between the

  extinct and living brackish-water shells of the Aralo-Caspian Sea.

  Now what does this remarkable law of the succession of the same types

  within the same areas mean? He would be a bold man, who after comparing

  the present climate of Australia and of parts of South America under the

  same latitude, would attempt to account, on the one hand, by dissimilar

  physical conditions for the dissimilarity of the inhabitants of these two

  continents, and, on the other hand, by similarity of conditions, for the

  uniformity of the same types in each during the later tertiary periods.

  Nor can it be pretended that it is an immutable law that marsupials should

  have been chiefly or solely produced in Australia; or that Edentata and

  other American types should have been solely produced in South America.

  For we know that Europe in ancient times was peopled by numerous

  marsupials; and I have shown in the publications above alluded to, that in

  America the law of distribution of terrestrial mammals was formerly

  different from what it now is. North America formerly partook strongly of

  the present character of the southern half of the continent; and the

  southern half was formerly more closely allied, than it is at present, to

  the northern half. In a similar manner we know from Falconer and Cautley's

  discoveries, that northern India was formerly more closely related in its

  mammals to Africa than it is at the present time. Analogous facts could be

  given in relation to the distribution of marine animals.

  On the theory of descent with modification, the great law of the long

  enduring, but not immutable, succession of the same types within the same

  areas, is at once explained; for the inhabitants of each quarter of the

  world will obviously tend to leave in that quarter, during the next

  succeeding period of time, closely allied though in some degree modified

  descendants. If the inhabitants of one continent formerly differed greatly

  from those of another continent, so will their modified descendants still

  differ in nearly the same manner and degree. But after very long intervals

  of time and after great geographical changes, permitting much

  inter-migration, the feebler will yield to the more dominant forms, and

  there will be nothing immutable in the laws of past and present

  distribution.

  It may be asked in ridicule, whether I suppose that the megatherium and

  other allied huge monsters have left behind them in South America the

  sloth, armadillo, and anteater, as their degenerate descendants. This

  cannot for an instant be admitted. These huge animals have become wholly

  extinct, and have left no progeny. But in the caves of Brazil, there are

  many extinct species which are closely allied in size and in other

  characters to the species still living in South America; and some of these

  fossils may be the actual progenitors of living species. It must not be

  forgotten that, on my theory, all the species of the same genus have

  descended from some one species; so that if six genera, each having eight

  species, be found in one geological formation, and in the next succeeding

  formation there be six other allied or representative genera with the same

  number of species, then we may conclude that only one species of each of

  the six older genera has left modified descendants, constituting the six

  new genera. The other seven species of the old genera have all died out

  and have left no progeny. Or, which would probably be a far commoner case,

  two or three species of two or three alone of the six older genera will

  have been the parents of the six new genera; the other old species and the

  other whole genera having become utterly extinct. In failing orders, with

  the genera and species decreasing in numbers, as apparently is the case of

  the Edentata of South America, still fewer genera and species will have

  left modified blood-descendants.

  Summary of the preceding and present Chapters -- I have attempted to show

  that the geological record is extremely imperfect; that only a small

  portion of the globe has been geologically explored with care; that only

  certain classes of organic beings have been largely preserved in a fossil

  state; that the number both of specimens and of species, preserved in our

  museums, is absolutely as nothing compared with the incalculable number of

  generations which must have passed away even during a single formation;

  that, owing to subsidence being necessary for the accumulation of

  fossiliferous deposits thick enough to resist future degradation, enormous

  intervals of time have elapsed between the successive formations; that

  there has probably been more extinction during the periods of subsidence,

  and more variation during the periods of elevation, and during the latter

  the record will have been least perfectly kept; that each single formation

  has not been continuously deposited; that the duration of each formation

  is, perhaps, short compared with the average duration of specific forms;

  that migration has played an important part in the first appearance of new

  forms in any one area and formation; that widely ranging species are those

  which have varied most, and have oftenest given rise to new species; and

  that varieties have at first often been local. All these causes taken

  conjointly, must have tended to make the geological record extremely

  imperfect, and will to a large extent explain why we do not find

  interminable varieties, connecting together all the extinct and existing

  forms of life by the finest graduated steps.

  He who rejects these views on the nature of the geological record, will

  rightly reject my whole theory. For he may ask in vain where are the

  numberless transitional links which must formerly have connected the

  closely allied or representative species, found in the several stages of

  the same great formation. He may disbelieve in the enormous intervals of

  time which have elapsed between our consecutive formations; he may overlook

  how important a part migration must have played, when the formations of any

  one great region alone, as that of Europe, are considered; he may urge the

  apparent, but often falsely apparent, sudden coming in of whole groups of

  species. He may ask where are the remains of those infinitely numerous

  organisms which must h
ave existed long before the first bed of the Silurian

  system was deposited: I can answer this latter question only

  hypothetically, by saying that as far as we can see, where our oceans now

  extend they have for an enormous period extended, and where our oscillating

  continents now stand they have stood ever since the Silurian epoch; but

  that long before that period, the world may have presented a wholly

  different aspect; and that the older continents, formed of formations older

  than any known to us, may now all be in a metamorphosed condition, or may

  lie buried under the ocean.

  Passing from these difficulties, all the other great leading facts in

  palaeontology seem to me simply to follow on the theory of descent with

  modification through natural selection. We can thus understand how it is

  that new species come in slowly and successively; how species of different

  classes do not necessarily change together, or at the same rate, or in the

  same degree; yet in the long run that all undergo modification to some

  extent. The extinction of old forms is the almost inevitable consequence

  of the production of new forms. We can understand why when a species has

  once disappeared it never reappears. Groups of species increase in numbers

  slowly, and endure for unequal periods of time; for the process of

  modification is necessarily slow, and depends on many complex

  contingencies. The dominant species of the larger dominant groups tend to

  leave many modified descendants, and thus new sub-groups and groups are

  formed. As these are formed, the species of the less vigorous groups, from

  their inferiority inherited from a common progenitor, tend to become

  extinct together, and to leave no modified offspring on the face of the

  earth. But the utter extinction of a whole group of species may often be a

  very slow process, from the survival of a few descendants, lingering in

  protected and isolated situations. When a group has once wholly

  disappeared, it does not reappear; for the link of generation has been

  broken.

  We can understand how the spreading of the dominant forms of life, which

  are those that oftenest vary, will in the long run tend to people the world

  with allied, but modified, descendants; and these will generally succeed in

  taking the places of those groups of species which are their inferiors in

  the struggle for existence. Hence, after long intervals of time, the

  productions of the world will appear to have changed simultaneously.

  We can understand how it is that all the forms of life, ancient and recent,

  make together one grand system; for all are connected by generation. We

  can understand, from the continued tendency to divergence of character, why

  the more ancient a form is, the more it generally differs from those now

  living. Why ancient and extinct forms often tend to fill up gaps between

  existing forms, sometimes blending two groups previously classed as

  distinct into one; but more commonly only bringing them a little closer

  together. The more ancient a form is, the more often, apparently, it

  displays characters in some degree intermediate between groups now

  distinct; for the more ancient a form is, the more nearly it will be

  related to, and consequently resemble, the common progenitor of groups,

  since become widely divergent. Extinct forms are seldom directly

  intermediate between existing forms; but are intermediate only by a long

  and circuitous course through many extinct and very different forms. We

  can clearly see why the organic remains of closely consecutive formations

  are more closely allied to each other, than are those of remote formations;

  for the forms are more closely linked together by generation: we can

  clearly see why the remains of an intermediate formation are intermediate

  in character.

  The inhabitants of each successive period in the world's history have

 

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