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Books: On the Origin of Species

C >> Charles Darwin >> On the Origin of Species




Laying aside the question of fertility and sterility, in all other
respects there seems to be a general and close similarity in the
offspring of crossed species, and of crossed varieties. If we look at
species as having been specially created, and at varieties as having
been produced by secondary laws, this similarity would be an
astonishing fact. But it harmonises perfectly with the view that there
is no essential distinction between species and varieties.

SUMMARY OF CHAPTER.

First crosses between forms sufficiently distinct to be ranked as
species, and their hybrids, are very generally, but not universally,
sterile. The sterility is of all degrees, and is often so slight that
the two most careful experimentalists who have ever lived, have come
to diametrically opposite conclusions in ranking forms by this test.
The sterility is innately variable in individuals of the same species,
and is eminently susceptible of favourable and unfavourable
conditions. The degree of sterility does not strictly follow
systematic affinity, but is governed by several curious and complex
laws. It is generally different, and sometimes widely different, in
reciprocal crosses between the same two species. It is not always
equal in degree in a first cross and in the hybrid produced from this
cross.

In the same manner as in grafting trees, the capacity of one species
or variety to take on another, is incidental on generally unknown
differences in their vegetative systems, so in crossing, the greater
or less facility of one species to unite with another, is incidental
on unknown differences in their reproductive systems. There is no more
reason to think that species have been specially endowed with various
degrees of sterility to prevent them crossing and blending in nature,
than to think that trees have been specially endowed with various and
somewhat analogous degrees of difficulty in being grafted together in
order to prevent them becoming inarched in our forests.

The sterility of first crosses between pure species, which have their
reproductive systems perfect, seems to depend on several
circumstances; in some cases largely on the early death of the embryo.
The sterility of hybrids, which have their reproductive systems
imperfect, and which have had this system and their whole organisation
disturbed by being compounded of two distinct species, seems closely
allied to that sterility which so frequently affects pure species,
when their natural conditions of life have been disturbed. This view
is supported by a parallelism of another kind;--namely, that the
crossing of forms only slightly different is favourable to the vigour
and fertility of their offspring; and that slight changes in the
conditions of life are apparently favourable to the vigour and
fertility of all organic beings. It is not surprising that the degree
of difficulty in uniting two species, and the degree of sterility of
their hybrid-offspring should generally correspond, though due to
distinct causes; for both depend on the amount of difference of some
kind between the species which are crossed. Nor is it surprising that
the facility of effecting a first cross, the fertility of the hybrids
produced, and the capacity of being grafted together--though this
latter capacity evidently depends on widely different
circumstances--should all run, to a certain extent, parallel with the
systematic affinity of the forms which are subjected to experiment;
for systematic affinity attempts to express all kinds of resemblance
between all species.

First crosses between forms known to be varieties, or sufficiently
alike to be considered as varieties, and their mongrel offspring, are
very generally, but not quite universally, fertile. Nor is this nearly
general and perfect fertility surprising, when we remember how liable
we are to argue in a circle with respect to varieties in a state of
nature; and when we remember that the greater number of varieties have
been produced under domestication by the selection of mere external
differences, and not of differences in the reproductive system. In all
other respects, excluding fertility, there is a close general
resemblance between hybrids and mongrels. Finally, then, the facts
briefly given in this chapter do not seem to me opposed to, but even
rather to support the view, that there is no fundamental distinction
between species and varieties.


CHAPTER 9. ON THE IMPERFECTION OF THE GEOLOGICAL RECORD.

On the absence of intermediate varieties at the present day.
On the nature of extinct intermediate varieties; on their number.
On the vast lapse of time, as inferred from the rate of deposition and
of denudation.
On the poorness of our palaeontological collections.
On the intermittence of geological formations.
On the absence of intermediate varieties in any one formation.
On the sudden appearance of groups of species.
On their sudden appearance in the lowest known fossiliferous strata.

In the sixth chapter I enumerated the chief objections which might be
justly urged against the views maintained in this volume. Most of them
have now been discussed. One, namely the distinctness of specific
forms, and their not being blended together by innumerable
transitional links, is a very obvious difficulty. I assigned reasons
why such links do not commonly occur at the present day, under the
circumstances apparently most favourable for their presence, namely on
an extensive and continuous area with graduated physical conditions. I
endeavoured to show, that the life of each species depends in a more
important manner on the presence of other already defined organic
forms, than on climate; and, therefore, that the really governing
conditions of life do not graduate away quite insensibly like heat or
moisture. I endeavoured, also, to show that intermediate varieties,
from existing in lesser numbers than the forms which they connect,
will generally be beaten out and exterminated during the course of
further modification and improvement. The main cause, however, of
innumerable intermediate links not now occurring everywhere throughout
nature depends on the very process of natural selection, through which
new varieties continually take the places of and exterminate their
parent-forms. But just in proportion as this process of extermination
has acted on an enormous scale, so must the number of intermediate
varieties, which have formerly existed on the earth, be truly
enormous. Why then is not every geological formation and every stratum
full of such intermediate links? Geology assuredly does not reveal any
such finely graduated organic chain; and this, perhaps, is the most
obvious and gravest objection which can be urged against my theory.
The explanation lies, as I believe, in the extreme imperfection of the
geological record.

In the first place it should always be borne in mind what sort of
intermediate forms must, on my theory, have formerly existed. I have
found it difficult, when looking at any two species, to avoid
picturing to myself, forms DIRECTLY intermediate between them. But
this is a wholly false view; we should always look for forms
intermediate between each species and a common but unknown progenitor;
and the progenitor will generally have differed in some respects from
all its modified descendants. To give a simple illustration: the
fantail and pouter pigeons have both descended from the rock-pigeon;
if we possessed all the intermediate varieties which have ever
existed, we should have an extremely close series between both and the
rock-pigeon; but we should have no varieties directly intermediate
between the fantail and pouter; none, for instance, combining a tail
somewhat expanded with a crop somewhat enlarged, the characteristic
features of these two breeds. These two breeds, moreover, have become
so much modified, that if we had no historical or indirect evidence
regarding their origin, it would not have been possible to have
determined from a mere comparison of their structure with that of the
rock-pigeon, whether they had descended from this species or from some
other allied species, such as C. oenas.

So with natural species, if we look to forms very distinct, for
instance to the horse and tapir, we have no reason to suppose that
links ever existed directly intermediate between them, but between
each and an unknown common parent. The common parent will have had in
its whole organisation much general resemblance to the tapir and to
the horse; but in some points of structure may have differed
considerably from both, even perhaps more than they differ from each
other. Hence in all such cases, we should be unable to recognise the
parent-form of any two or more species, even if we closely compared
the structure of the parent with that of its modified descendants,
unless at the same time we had a nearly perfect chain of the
intermediate links.

It is just possible by my theory, that one of two living forms might
have descended from the other; for instance, a horse from a tapir; and
in this case DIRECT intermediate links will have existed between them.
But such a case would imply that one form had remained for a very long
period unaltered, whilst its descendants had undergone a vast amount
of change; and the principle of competition between organism and
organism, between child and parent, will render this a very rare
event; for in all cases the new and improved forms of life will tend
to supplant the old and unimproved forms.

By the theory of natural selection all living species have been
connected with the parent-species of each genus, by differences not
greater than we see between the varieties of the same species at the
present day; and these parent-species, now generally extinct, have in
their turn been similarly connected with more ancient species; and so
on backwards, always converging to the common ancestor of each great
class. So that the number of intermediate and transitional links,
between all living and extinct species, must have been inconceivably
great. But assuredly, if this theory be true, such have lived upon
this earth.

ON THE LAPSE OF TIME.

Independently of our not finding fossil remains of such infinitely
numerous connecting links, it may be objected, that time will not have
sufficed for so great an amount of organic change, all changes having
been effected very slowly through natural selection. It is hardly
possible for me even to recall to the reader, who may not be a
practical geologist, the facts leading the mind feebly to comprehend
the lapse of time. He who can read Sir Charles Lyell's grand work on
the Principles of Geology, which the future historian will recognise
as having produced a revolution in natural science, yet does not admit
how incomprehensibly vast have been the past periods of time, may at
once close this volume. Not that it suffices to study the Principles
of Geology, or to read special treatises by different observers on
separate formations, and to mark how each author attempts to give an
inadequate idea of the duration of each formation or even each
stratum. A man must for years examine for himself great piles of
superimposed strata, and watch the sea at work grinding down old rocks
and making fresh sediment, before he can hope to comprehend anything
of the lapse of time, the monuments of which we see around us.

It is good to wander along lines of sea-coast, when formed of
moderately hard rocks, and mark the process of degradation. The tides
in most cases reach the cliffs only for a short time twice a day, and
the waves eat into them only when they are charged with sand or
pebbles; for there is reason to believe that pure water can effect
little or nothing in wearing away rock. At last the base of the cliff
is undermined, huge fragments fall down, and these remaining fixed,
have to be worn away, atom by atom, until reduced in size they can be
rolled about by the waves, and then are more quickly ground into
pebbles, sand, or mud. But how often do we see along the bases of
retreating cliffs rounded boulders, all thickly clothed by marine
productions, showing how little they are abraded and how seldom they
are rolled about! Moreover, if we follow for a few miles any line of
rocky cliff, which is undergoing degradation, we find that it is only
here and there, along a short length or round a promontory, that the
cliffs are at the present time suffering. The appearance of the
surface and the vegetation show that elsewhere years have elapsed
since the waters washed their base.

He who most closely studies the action of the sea on our shores, will,
I believe, be most deeply impressed with the slowness with which rocky
coasts are worn away. The observations on this head by Hugh Miller,
and by that excellent observer Mr. Smith of Jordan Hill, are most
impressive. With the mind thus impressed, let any one examine beds of
conglomerate many thousand feet in thickness, which, though probably
formed at a quicker rate than many other deposits, yet, from being
formed of worn and rounded pebbles, each of which bears the stamp of
time, are good to show how slowly the mass has been accumulated. Let
him remember Lyell's profound remark, that the thickness and extent of
sedimentary formations are the result and measure of the degradation
which the earth's crust has elsewhere suffered. And what an amount of
degradation is implied by the sedimentary deposits of many countries!
Professor Ramsay has given me the maximum thickness, in most cases
from actual measurement, in a few cases from estimate, of each
formation in different parts of Great Britain; and this is the
result:--

Feet
Palaeozoic strata (not including igneous beds)..57,154.
Secondary strata................................13,190.
Tertiary strata..................................2,240.

--making altogether 72,584 feet; that is, very nearly thirteen and
three-quarters British miles. Some of these formations, which are
represented in England by thin beds, are thousands of feet in
thickness on the Continent. Moreover, between each successive
formation, we have, in the opinion of most geologists, enormously long
blank periods. So that the lofty pile of sedimentary rocks in Britain,
gives but an inadequate idea of the time which has elapsed during
their accumulation; yet what time this must have consumed! Good
observers have estimated that sediment is deposited by the great
Mississippi river at the rate of only 600 feet in a hundred thousand
years. This estimate may be quite erroneous; yet, considering over
what wide spaces very fine sediment is transported by the currents of
the sea, the process of accumulation in any one area must be extremely
slow.

But the amount of denudation which the strata have in many places
suffered, independently of the rate of accumulation of the degraded
matter, probably offers the best evidence of the lapse of time. I
remember having been much struck with the evidence of denudation, when
viewing volcanic islands, which have been worn by the waves and pared
all round into perpendicular cliffs of one or two thousand feet in
height; for the gentle slope of the lava-streams, due to their
formerly liquid state, showed at a glance how far the hard, rocky beds
had once extended into the open ocean. The same story is still more
plainly told by faults,--those great cracks along which the strata
have been upheaved on one side, or thrown down on the other, to the
height or depth of thousands of feet; for since the crust cracked, the
surface of the land has been so completely planed down by the action
of the sea, that no trace of these vast dislocations is externally
visible.

The Craven fault, for instance, extends for upwards of 30 miles, and
along this line the vertical displacement of the strata has varied
from 600 to 3000 feet. Professor Ramsay has published an account of a
downthrow in Anglesea of 2300 feet; and he informs me that he fully
believes there is one in Merionethshire of 12,000 feet; yet in these
cases there is nothing on the surface to show such prodigious
movements; the pile of rocks on the one or other side having been
smoothly swept away. The consideration of these facts impresses my
mind almost in the same manner as does the vain endeavour to grapple
with the idea of eternity.

I am tempted to give one other case, the well-known one of the
denudation of the Weald. Though it must be admitted that the
denudation of the Weald has been a mere trifle, in comparison with
that which has removed masses of our palaeozoic strata, in parts ten
thousand feet in thickness, as shown in Professor Ramsay's masterly
memoir on this subject. Yet it is an admirable lesson to stand on the
North Downs and to look at the distant South Downs; for, remembering
that at no great distance to the west the northern and southern
escarpments meet and close, one can safely picture to oneself the
great dome of rocks which must have covered up the Weald within so
limited a period as since the latter part of the Chalk formation. The
distance from the northern to the southern Downs is about 22 miles,
and the thickness of the several formations is on an average about
1100 feet, as I am informed by Professor Ramsay. But if, as some
geologists suppose, a range of older rocks underlies the Weald, on the
flanks of which the overlying sedimentary deposits might have
accumulated in thinner masses than elsewhere, the above estimate would
be erroneous; but this source of doubt probably would not greatly
affect the estimate as applied to the western extremity of the
district. If, then, we knew the rate at which the sea commonly wears
away a line of cliff of any given height, we could measure the time
requisite to have denuded the Weald. This, of course, cannot be done;
but we may, in order to form some crude notion on the subject, assume
that the sea would eat into cliffs 500 feet in height at the rate of
one inch in a century. This will at first appear much too small an
allowance; but it is the same as if we were to assume a cliff one yard
in height to be eaten back along a whole line of coast at the rate of
one yard in nearly every twenty-two years. I doubt whether any rock,
even as soft as chalk, would yield at this rate excepting on the most
exposed coasts; though no doubt the degradation of a lofty cliff would
be more rapid from the breakage of the fallen fragments. On the other
hand, I do not believe that any line of coast, ten or twenty miles in
length, ever suffers degradation at the same time along its whole
indented length; and we must remember that almost all strata contain
harder layers or nodules, which from long resisting attrition form a
breakwater at the base. Hence, under ordinary circumstances, I
conclude that for a cliff 500 feet in height, a denudation of one inch
per century for the whole length would be an ample allowance. At this
rate, on the above data, the denudation of the Weald must have
required 306,662,400 years; or say three hundred million years.

The action of fresh water on the gently inclined Wealden district,
when upraised, could hardly have been great, but it would somewhat
reduce the above estimate. On the other hand, during oscillations of
level, which we know this area has undergone, the surface may have
existed for millions of years as land, and thus have escaped the
action of the sea: when deeply submerged for perhaps equally long
periods, it would, likewise, have escaped the action of the
coast-waves. So that in all probability a far longer period than 300
million years has elapsed since the latter part of the Secondary
period.

I have made these few remarks because it is highly important for us to
gain some notion, however imperfect, of the lapse of years. During
each of these years, over the whole world, the land and the water has
been peopled by hosts of living forms. What an infinite number of
generations, which the mind cannot grasp, must have succeeded each
other in the long roll of years! Now turn to our richest geological
museums, and what a paltry display we behold!

ON THE POORNESS OF OUR PALAEONTOLOGICAL COLLECTIONS.

That our palaeontological collections are very imperfect, is admitted
by every one. The remark of that admirable palaeontologist, the late
Edward Forbes, should not be forgotten, namely, that numbers of our
fossil species are known and named from single and often broken
specimens, or from a few specimens collected on some one spot. Only a
small portion of the surface of the earth has been geologically
explored, and no part with sufficient care, as the important
discoveries made every year in Europe prove. No organism wholly soft
can be preserved. Shells and bones will decay and disappear when left
on the bottom of the sea, where sediment is not accumulating. I
believe we are continually taking a most erroneous view, when we
tacitly admit to ourselves that sediment is being deposited over
nearly the whole bed of the sea, at a rate sufficiently quick to embed
and preserve fossil remains. Throughout an enormously large proportion
of the ocean, the bright blue tint of the water bespeaks its purity.
The many cases on record of a formation conformably covered, after an
enormous interval of time, by another and later formation, without the
underlying bed having suffered in the interval any wear and tear, seem
explicable only on the view of the bottom of the sea not rarely lying
for ages in an unaltered condition. The remains which do become
embedded, if in sand or gravel, will when the beds are upraised
generally be dissolved by the percolation of rain-water. I suspect
that but few of the very many animals which live on the beach between
high and low watermark are preserved. For instance, the several
species of the Chthamalinae (a sub-family of sessile cirripedes) coat
the rocks all over the world in infinite numbers: they are all
strictly littoral, with the exception of a single Mediterranean
species, which inhabits deep water and has been found fossil in
Sicily, whereas not one other species has hitherto been found in any
tertiary formation: yet it is now known that the genus Chthamalus
existed during the chalk period. The molluscan genus Chiton offers a
partially analogous case.

With respect to the terrestrial productions which lived during the
Secondary and Palaeozoic periods, it is superfluous to state that our
evidence from fossil remains is fragmentary in an extreme degree. For
instance, not a land shell is known belonging to either of these vast
periods, with one exception discovered by Sir C. Lyell in the
carboniferous strata of North America. In regard to mammiferous
remains, a single glance at the historical table published in the
Supplement to Lyell's Manual, will bring home the truth, how
accidental and rare is their preservation, far better than pages of
detail. Nor is their rarity surprising, when we remember how large a
proportion of the bones of tertiary mammals have been discovered
either in caves or in lacustrine deposits; and that not a cave or true
lacustrine bed is known belonging to the age of our secondary or
palaeozoic formations.

But the imperfection in the geological record mainly results from
another and more important cause than any of the foregoing; namely,
from the several formations being separated from each other by wide
intervals of time. When we see the formations tabulated in written
works, or when we follow them in nature, it is difficult to avoid
believing that they are closely consecutive. But we know, for
instance, from Sir R. Murchison's great work on Russia, what wide gaps
there are in that country between the superimposed formations; so it
is in North America, and in many other parts of the world. The most
skilful geologist, if his attention had been exclusively confined to
these large territories, would never have suspected that during the
periods which were blank and barren in his own country, great piles of
sediment, charged with new and peculiar forms of life, had elsewhere
been accumulated. And if in each separate territory, hardly any idea
can be formed of the length of time which has elapsed between the
consecutive formations, we may infer that this could nowhere be
ascertained. The frequent and great changes in the mineralogical
composition of consecutive formations, generally implying great
changes in the geography of the surrounding lands, whence the sediment
has been derived, accords with the belief of vast intervals of time
having elapsed between each formation.

But we can, I think, see why the geological formations of each region
are almost invariably intermittent; that is, have not followed each
other in close sequence. Scarcely any fact struck me more when
examining many hundred miles of the South American coasts, which have
been upraised several hundred feet within the recent period, than the
absence of any recent deposits sufficiently extensive to last for even
a short geological period. Along the whole west coast, which is
inhabited by a peculiar marine fauna, tertiary beds are so scantily
developed, that no record of several successive and peculiar marine
faunas will probably be preserved to a distant age. A little
reflection will explain why along the rising coast of the western side
of South America, no extensive formations with recent or tertiary
remains can anywhere be found, though the supply of sediment must for
ages have been great, from the enormous degradation of the coast-rocks
and from muddy streams entering the sea. The explanation, no doubt,
is, that the littoral and sub-littoral deposits are continually worn
away, as soon as they are brought up by the slow and gradual rising of
the land within the grinding action of the coast-waves.