Chapter IX
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. Prof. 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
Prof. 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 Prof. 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.
We may, I think, safely conclude that sediment must be accumulated in
extremely thick, solid, or extensive masses, in order to withstand the
incessant action of the waves, when first upraised and during subsequent
oscillations of level. Such thick and extensive accumulations of sediment
may be formed in two ways; either, in profound depths of the sea, in which
case, judging from the researches of E. Forbes, we may conclude that the
bottom will be inhabited by extremely few animals, and the mass when
upraised will give a most imperfect record of the forms of life which then
existed; or, sediment may be accumulated to any thickness and extent over a
shallow bottom, if it continue slowly to subside. In this latter case, as
long as the rate of subsidence and supply of sediment nearly balance each
other, the sea will remain shallow and favourable for life, and thus a
fossiliferous formation thick enough, when upraised, to resist any amount
of degradation, may be formed.
I am convinced that all our ancient formations, which are rich in fossils,
have thus been formed during subsidence. Since publishing my views on this
subject in 1845, I have watched the progress of Geology, and have been
surprised to note how author after author, in treating of this or that
great formation, has come to the conclusion that it was accumulated during
subsidence. I may add, that the only ancient tertiary formation on the
west coast of South America, which has been bulky enough to resist such
degradation as it has as yet suffered, but which will hardly last to a
distant geological age, was certainly deposited during a downward
oscillation of level, and thus gained considerable thickness.
All geological facts tell us plainly that each area has undergone numerous
slow oscillations of level, and apparently these oscillations have affected
wide spaces. Consequently formations rich in fossils and sufficiently
thick and extensive to resist subsequent degradation, may have been formed
over wide spaces during periods of subsidence, but only where the supply of
sediment was sufficient to keep the sea shallow and to embed and preserve
the remains before they had time to decay. On the other hand, as long as
the bed of the sea remained stationary, thick deposits could not have been
accumulated in the shallow parts, which are the most favourable to life.
Still less could this have happened during the alternate periods of
elevation; or, to speak more accurately, the beds which were then
accumulated will have been destroyed by being upraised and brought within
the limits of the coast-action.
Thus the geological record will almost necessarily be rendered
intermittent. I feel much confidence in the truth of these views, for they
are in strict accordance with the general principles inculcated by Sir C.
Lyell; and E. Forbes independently arrived at a similar conclusion.
One remark is here worth a passing notice. During periods of elevation the
area of the land and of the adjoining shoal parts of the sea will be
increased, and new stations will often be formed;--all circumstances most
favourable, as previously explained, for the formation of new varieties and
species; but during such periods there will generally be a blank in the
geological record. On the other hand, during subsidence, the inhabited
area and number of inhabitants will decrease (excepting the productions on
the shores of a continent when first broken up into an archipelago), and
consequently during subsidence, though there will be much extinction, fewer
new varieties or species will be formed; and it is during these very
periods of subsidence, that our great deposits rich in fossils have been
accumulated. Nature may almost be said to have guarded against the
frequent discovery of her transitional or linking forms.
From the foregoing considerations it cannot be doubted that the geological
record, viewed as a whole, is extremely imperfect; but if we confine our
attention to any one formation, it becomes more difficult to understand,
why we do not therein find closely graduated varieties between the allied
species which lived at its commencement and at its close. Some cases are
on record of the same species presenting distinct varieties in the upper
and lower parts of the same formation, but, as they are rare, they may be
here passed over. Although each formation has indisputably required a vast
number of years for its deposition, I can see several reasons why each
should not include a graduated series of links between the species which
then lived; but I can by no means pretend to assign due proportional weight
to the following considerations.
Although each formation may mark a very long lapse of years, each perhaps
is short compared with the period requisite to change one species into
another. I am aware that two palaeontologists, whose opinions are worthy
of much deference, namely Bronn and Woodward, have concluded that the
average duration of each formation is twice or thrice as long as the
average duration of specific forms. But insuperable difficulties, as it
seems to me, prevent us coming to any just conclusion on this head. When
we see a species first appearing in the middle of any formation, it would
be rash in the extreme to infer that it had not elsewhere previously
existed. So again when we find a species disappearing before the uppermost
layers have been deposited, it would be equally rash to suppose that it
then became wholly extinct. We forget how small the area of Europe is
compared with the rest of the world; nor have the several stages of the
same formation throughout Europe been correlated with perfect accuracy.
With marine animals of all kinds, we may safely infer a large amount of
migration during climatal and other changes; and when we see a species
first appearing in any formation, the probability is that it only then
first immigrated into that area. It is well known, for instance, that
several species appeared somewhat earlier in the palaeozoic beds of North
America than in those of Europe; time having apparently been required for
their migration from the American to the European seas. In examining the
latest deposits of various quarters of the world, it has everywhere been
noted, that some few still existing species are common in the deposit, but
have become extinct in the immediately surrounding sea; or, conversely,
that some are now abundant in the neighbouring sea, but are rare or absent
in this particular deposit. It is an excellent lesson to reflect on the
ascertained amount of migration of the inhabitants of Europe during the
Glacial period, which forms only a part of one whole geological period; and
likewise to reflect on the great changes of level, on the inordinately
great change of climate, on the prodigious lapse of time, all included
within this same glacial period. Yet it may be doubted whether in any
quarter of the world, sedimentary deposits, including fossil remains, have
gone on accumulating within the same area during the whole of this period.
It is not, for instance, probable that sediment was deposited during the
whole of the glacial period near the mouth of the Mississippi, within that
limit of depth at which marine animals can flourish; for we know what vast
geographical changes occurred in other parts of America during this space
of time. When such beds as were deposited in shallow water near the mouth
of the Mississippi during some part of the glacial period shall have been
upraised, organic remains will probably first appear and disappear at
different levels, owing to the migration of species and to geographical
changes. And in the distant future, a geologist examining these beds,
might be tempted to conclude that the average duration of life of the
embedded fossils had been less than that of the glacial period, instead of
having been really far greater, that is extending from before the glacial
epoch to the present day.
In order to get a perfect gradation between two forms in the upper and
lower parts of the same formation, the deposit must have gone on
accumulating for a very long period, in order to have given sufficient time
for the slow process of variation; hence the deposit will generally have to
be a very thick one; and the species undergoing modification will have had
to live on the same area throughout this whole time. But we have seen that
a thick fossiliferous formation can only be accumulated during a period of
subsidence; and to keep the depth approximately the same, which is
necessary in order to enable the same species to live on the same space,
the supply of sediment must nearly have counterbalanced the amount of
subsidence. But this same movement of subsidence will often tend to sink
the area whence the sediment is derived, and thus diminish the supply
whilst the downward movement continues. In fact, this nearly exact
balancing between the supply of sediment and the amount of subsidence is
probably a rare contingency; for it has been observed by more than one
palaeontologist, that very thick deposits are usually barren of organic
remains, except near their upper or lower limits.
It would seem that each separate formation, like the whole pile of
formations in any country, has generally been intermittent in its
accumulation. When we see, as is so often the case, a formation composed
of beds of different mineralogical composition, we may reasonably suspect
that the process of deposition has been much interrupted, as a change in
the currents of the sea and a supply of sediment of a different nature will
generally have been due to geographical changes requiring much time. Nor
will the closest inspection of a formation give any idea of the time which
its deposition has consumed. Many instances could be given of beds only a
few feet in thickness, representing formations, elsewhere thousands of feet
in thickness, and which must have required an enormous period for their
accumulation; yet no one ignorant of this fact would have suspected the
vast lapse of time represented by the thinner formation. Many cases could
be given of the lower beds of a formation having been upraised, denuded,
submerged, and then re-covered by the upper beds of the same
formation,--facts, showing what wide, yet easily overlooked, intervals have
occurred in its accumulation. In other cases we have the plainest evidence
in great fossilised trees, still standing upright as they grew, of many
long intervals of time and changes of level during the process of
deposition, which would never even have been suspected, had not the trees
chanced to have been preserved: thus, Messrs. Lyell and Dawson found
carboniferous beds 1400 feet thick in Nova Scotia, with ancient
root-bearing strata, one above the other, at no less than sixty-eight
different levels. Hence, when the same species occur at the bottom,
middle, and top of a formation, the probability is that they have not lived
on the same spot during the whole period of deposition, but have
disappeared and reappeared, perhaps many times, during the same geological
period. So that if such species were to undergo a considerable amount of
modification during any one geological period, a section would not probably
include all the fine intermediate gradations which must on my theory have
existed between them, but abrupt, though perhaps very slight, changes of
form.
It is all-important to remember that naturalists have no golden rule by
which to distinguish species and varieties; they grant some little
variability to each species, but when they meet with a somewhat greater
amount of difference between any two forms, they rank both as species,
unless they are enabled to connect them together by close intermediate
gradations. And this from the reasons just assigned we can seldom hope to
effect in any one geological section. Supposing B and C to be two species,
and a third, A, to be found in an underlying bed; even if A were strictly
intermediate between B and C, it would simply be ranked as a third and
distinct species, unless at the same time it could be most closely
connected with either one or both forms by intermediate varieties. Nor
should it be forgotten, as before explained, that A might be the actual
progenitor of B and C, and yet might not at all necessarily be strictly
intermediate between them in all points of structure. So that we might
obtain the parent-species and its several modified descendants from the
lower and upper beds of a formation, and unless we obtained numerous
transitional gradations, we should not recognise their relationship, and
should consequently be compelled to rank them all as distinct species.
It is notorious on what excessively slight differences many
palaeontologists have founded their species; and they do this the more
readily if the specimens come from different sub-stages of the same
formation. Some experienced conchologists are now sinking many of the very
fine species of D'Orbigny and others into the rank of varieties; and on
this view we do find the kind of evidence of change which on my theory we
ought to find. Moreover, if we look to rather wider intervals, namely, to
distinct but consecutive stages of the same great formation, we find that
the embedded fossils, though almost universally ranked as specifically
different, yet are far more closely allied to each other than are the
species found in more widely separated formations; but to this subject I
shall have to return in the following chapter.
One other consideration is worth notice: with animals and plants that can
propagate rapidly and are not highly locomotive, there is reason to
suspect, as we have formerly seen, that their varieties are generally at
first local; and that such local varieties do not spread widely and
supplant their parent-forms until they have been modified and perfected in
some considerable degree. According to this view, the chance of
discovering in a formation in any one country all the early stages of
transition between any two forms, is small, for the successive changes are
supposed to have been local or confined to some one spot. Most marine
animals have a wide range; and we have seen that with plants it is those
which have the widest range, that oftenest present varieties; so that with
shells and other marine animals, it is probably those which have had the
widest range, far exceeding the limits of the known geological formations
of Europe, which have oftenest given rise, first to local varieties and
ultimately to new species; and this again would greatly lessen the chance
of our being able to trace the stages of transition in any one geological
formation.
It should not be forgotten, that at the present day, with perfect specimens
for examination, two forms can seldom be connected by intermediate
varieties and thus proved to be the same species, until many specimens have
been collected from many places; and in the case of fossil species this
could rarely be effected by palaeontologists. We shall, perhaps, best
perceive the improbability of our being enabled to connect species by
numerous, fine, intermediate, fossil links, by asking ourselves whether,
for instance, geologists at some future period will be able to prove, that
our different breeds of cattle, sheep, horses, and dogs have descended from
a single stock or from several aboriginal stocks; or, again, whether
certain sea-shells inhabiting the shores of North America, which are ranked
by some conchologists as distinct species from their European
representatives, and by other conchologists as only varieties, are really
varieties or are, as it is called, specifically distinct. This could be
effected only by the future geologist discovering in a fossil state
numerous intermediate gradations; and such success seems to me improbable
in the highest degree.
Geological research, though it has added numerous species to existing and
extinct genera, and has made the intervals between some few groups less
wide than they otherwise would have been, yet has done scarcely anything in
breaking down the distinction between species, by connecting them together
by numerous, fine, intermediate varieties; and this not having been
effected, is probably the gravest and most obvious of all the many
objections which may be urged against my views. Hence it will be worth
while to sum up the foregoing remarks, under an imaginary illustration.
The Malay Archipelago is of about the size of Europe from the North Cape to
the Mediterranean, and from Britain to Russia; and therefore equals all the
geological formations which have been examined with any accuracy, excepting
those of the United States of America. I fully agree with Mr.
Godwin-Austen, that the present condition of the Malay Archipelago, with
its numerous large islands separated by wide and shallow seas, probably
represents the former state of Europe, when most of our formations were
accumulating. The Malay Archipelago is one of the richest regions of the
whole world in organic beings; yet if all the species were to be collected
which have ever lived there, how imperfectly would they represent the
natural history of the world!
But we have every reason to believe that the terrestrial productions of the
archipelago would be preserved in an excessively imperfect manner in the
formations which we suppose to be there accumulating. I suspect that not
many of the strictly littoral animals, or of those which lived on naked
submarine rocks, would be embedded; and those embedded in gravel or sand,
would not endure to a distant epoch. Wherever sediment did not accumulate
on the bed of the sea, or where it did not accumulate at a sufficient rate
to protect organic bodies from decay, no remains could be preserved.
In our archipelago, I believe that fossiliferous formations could be formed
of sufficient thickness to last to an age, as distant in futurity as the
secondary formations lie in the past, only during periods of subsidence.
These periods of subsidence would be separated from each other by enormous
intervals, during which the area would be either stationary or rising;
whilst rising, each fossiliferous formation would be destroyed, almost as
soon as accumulated, by the incessant coast-action, as we now see on the
shores of South America. During the periods of subsidence there would
probably be much extinction of life; during the periods of elevation, there
would be much variation, but the geological record would then be least
perfect.
It may be doubted whether the duration of any one great period of
subsidence over the whole or part of the archipelago, together with a
contemporaneous accumulation of sediment, would exceed the average duration
of the same specific forms; and these contingencies are indispensable for
the preservation of all the transitional gradations between any two or more
species. If such gradations were not fully preserved, transitional
varieties would merely appear as so many distinct species. It is, also,
probable that each great period of subsidence would be interrupted by
oscillations of level, and that slight climatal changes would intervene
during such lengthy periods; and in these cases the inhabitants of the
archipelago would have to migrate, and no closely consecutive record of
their modifications could be preserved in any one formation.
Very many of the marine inhabitants of the archipelago now range thousands
of miles beyond its confines; and analogy leads me to believe that it would
be chiefly these far-ranging species which would oftenest produce new
varieties; and the varieties would at first generally be local or confined
to one place, but if possessed of any decided advantage, or when further
modified and improved, they would slowly spread and supplant their
parent-forms. When such varieties returned to their ancient homes, as they
would differ from their former state, in a nearly uniform, though perhaps
extremely slight degree, they would, according to the principles followed
by many palaeontologists, be ranked as new and distinct species.
If then, there be some degree of truth in these remarks, we have no right
to expect to find in our geological formations, an infinite number of those
fine transitional forms, which on my theory assuredly have connected all
the past and present species of the same group into one long and branching
chain of life. We ought only to look for a few links, some more closely,
some more distantly related to each other; and these links, let them be
ever so close, if found in different stages of the same formation, would,
by most palaeontologists, be ranked as distinct species. But I do not
pretend that I should ever have suspected how poor a record of the
mutations of life, the best preserved geological section presented, had not
the difficulty of our not discovering innumerable transitional links
between the species which appeared at the commencement and close of each
formation, pressed so hardly on my theory.
On the sudden appearance of whole groups of Allied Species. -- The abrupt
manner in which whole groups of species suddenly appear in certain
formations, has been urged by several palaeontologists, for instance, by
Agassiz, Pictet, and by none more forcibly than by Professor Sedgwick, as a
fatal objection to the belief in the transmutation of species. If numerous
species, belonging to the same genera or families, have really started into
life all at once, the fact would be fatal to the theory of descent with
slow modification through natural selection. For the development of a
group of forms, all of which have descended from some one progenitor, must
have been an extremely slow process; and the progenitors must have lived
long ages before their modified descendants. But we continually over-rate
the perfection of the geological record, and falsely infer, because certain
genera or families have not been found beneath a certain stage, that they
did not exist before that stage. We continually forget how large the world
is, compared with the area over which our geological formations have been
carefully examined; we forget that groups of species may elsewhere have
long existed and have slowly multiplied before they invaded the ancient
archipelagoes of Europe and of the United States. We do not make due
allowance for the enormous intervals of time, which have probably elapsed
between our consecutive formations,--longer perhaps in some cases than the
time required for the accumulation of each formation. These intervals will
have given time for the multiplication of species from some one or some few
parent-forms; and in the succeeding formation such species will appear as
if suddenly created.
I may here recall a remark formerly made, namely that it might require a
long succession of ages to adapt an organism to some new and peculiar line
of life, for instance to fly through the air; but that when this had been
effected, and a few species had thus acquired a great advantage over other
organisms, a comparatively short time would be necessary to produce many
divergent forms, which would be able to spread rapidly and widely
throughout the world.
I will now give a few examples to illustrate these remarks; and to show how
liable we are to error in supposing that whole groups of species have
suddenly been produced. I may recall the well-known fact that in
geological treatises, published not many years ago, the great class of
mammals was always spoken of as having abruptly come in at the commencement
of the tertiary series. And now one of the richest known accumulations of
fossil mammals belongs to the middle of the secondary series; and one true
mammal has been discovered in the new red sandstone at nearly the
commencement of this great series. Cuvier used to urge that no monkey
occurred in any tertiary stratum; but now extinct species have been
discovered in India, South America, and in Europe even as far back as the
eocene stage. The most striking case, however, is that of the Whale
family; as these animals have huge bones, are marine, and range over the
world, the fact of not a single bone of a whale having been discovered in
any secondary formation, seemed fully to justify the belief that this great
and distinct order had been suddenly produced in the interval between the
latest secondary and earliest tertiary formation. But now we may read in
the Supplement to Lyell's 'Manual,' published in 1858, clear evidence of
the existence of whales in the upper greensand, some time before the close
of the secondary period.
I may give another instance, which from having passed under my own eyes has
much struck me. In a memoir on Fossil Sessile Cirripedes, I have stated
that, from the number of existing and extinct tertiary species; from the
extraordinary abundance of the individuals of many species all over the
world, from the Arctic regions to the equator, inhabiting various zones of
depths from the upper tidal limits to 50 fathoms; from the perfect manner
in which specimens are preserved in the oldest tertiary beds; from the ease
with which even a fragment of a valve can be recognised; from all these
circumstances, I inferred that had sessile cirripedes existed during the
secondary periods, they would certainly have been preserved and discovered;
and as not one species had been discovered in beds of this age, I concluded
that this great group had been suddenly developed at the commencement of
the tertiary series. This was a sore trouble to me, adding as I thought
one more instance of the abrupt appearance of a great group of species.
But my work had hardly been published, when a skilful palaeontologist, M.
Bosquet, sent me a drawing of a perfect specimen of an unmistakeable
sessile cirripede, which he had himself extracted from the chalk of
Belgium. And, as if to make the case as striking as possible, this sessile
cirripede was a Chthamalus, a very common, large, and ubiquitous genus, of
which not one specimen has as yet been found even in any tertiary stratum.
Hence we now positively know that sessile cirripedes existed during the
secondary period; and these cirripedes might have been the progenitors of
our many tertiary and existing species.
The case most frequently insisted on by palaeontologists of the apparently
sudden appearance of a whole group of species, is that of the teleostean
fishes, low down in the Chalk period. This group includes the large
majority of existing species. Lately, Professor Pictet has carried their
existence one sub-stage further back; and some palaeontologists believe
that certain much older fishes, of which the affinities are as yet
imperfectly known, are really teleostean. Assuming, however, that the
whole of them did appear, as Agassiz believes, at the commencement of the
chalk formation, the fact would certainly be highly remarkable; but I
cannot see that it would be an insuperable difficulty on my theory, unless
it could likewise be shown that the species of this group appeared suddenly
and simultaneously throughout the world at this same period. It is almost
superfluous to remark that hardly any fossil-fish are known from south of
the equator; and by running through Pictet's Palaeontology it will be seen
that very few species are known from several formations in Europe. Some
few families of fish now have a confined range; the teleostean fish might
formerly have had a similarly confined range, and after having been largely
developed in some one sea, might have spread widely. Nor have we any right
to suppose that the seas of the world have always been so freely open from
south to north as they are at present. Even at this day, if the Malay
Archipelago were converted into land, the tropical parts of the Indian
Ocean would form a large and perfectly enclosed basin, in which any great
group of marine animals might be multiplied; and here they would remain
confined, until some of the species became adapted to a cooler climate, and
were enabled to double the southern capes of Africa or Australia, and thus
reach other and distant seas.
From these and similar considerations, but chiefly from our ignorance of
the geology of other countries beyond the confines of Europe and the United
States; and from the revolution in our palaeontological ideas on many
points, which the discoveries of even the last dozen years have effected,
it seems to me to be about as rash in us to dogmatize on the succession of
organic beings throughout the world, as it would be for a naturalist to
land for five minutes on some one barren point in Australia, and then to
discuss the number and range of its productions.
On the sudden appearance of groups of Allied Species in the lowest known
fossiliferous strata. -- There is another and allied difficulty, which is
much graver. I allude to the manner in which numbers of species of the
same group, suddenly appear in the lowest known fossiliferous rocks. Most
of the arguments which have convinced me that all the existing species of
the same group have descended from one progenitor, apply with nearly equal
force to the earliest known species. For instance, I cannot doubt that all
the Silurian trilobites have descended from some one crustacean, which must
have lived long before the Silurian age, and which probably differed
greatly from any known animal. Some of the most ancient Silurian animals,
as the Nautilus, Lingula, &c., do not differ much from living species; and
it cannot on my theory be supposed, that these old species were the
progenitors of all the species of the orders to which they belong, for they
do not present characters in any degree intermediate between them. If,
moreover, they had been the progenitors of these orders, they would almost
certainly have been long ago supplanted and exterminated by their numerous
and improved descendants.
Consequently, if my theory be true, it is indisputable that before the
lowest Silurian stratum was deposited, long periods elapsed, as long as, or
probably far longer than, the whole interval from the Silurian age to the
present day; and that during these vast, yet quite unknown, periods of
time, the world swarmed with living creatures.
To the question why we do not find records of these vast primordial
periods, I can give no satisfactory answer. Several of the most eminent
geologists, with Sir R. Murchison at their head, are convinced that we see
in the organic remains of the lowest Silurian stratum the dawn of life on
this planet. Other highly competent judges, as Lyell and the late E.
Forbes, dispute this conclusion. We should not forget that only a small
portion of the world is known with accuracy. M. Barrande has lately added
another and lower stage to the Silurian system, abounding with new and
peculiar species. Traces of life have been detected in the Longmynd beds
beneath Barrande's so-called primordial zone. The presence of phosphatic
nodules and bituminous matter in some of the lowest azoic rocks, probably
indicates the former existence of life at these periods. But the
difficulty of understanding the absence of vast piles of fossiliferous
strata, which on my theory no doubt were somewhere accumulated before the
Silurian epoch, is very great. If these most ancient beds had been wholly
worn away by denudation, or obliterated by metamorphic action, we ought to
find only small remnants of the formations next succeeding them in age, and
these ought to be very generally in a metamorphosed condition. But the
descriptions which we now possess of the Silurian deposits over immense
territories in Russia and in North America, do not support the view, that
the older a formation is, the more it has suffered the extremity of
denudation and metamorphism.
The case at present must remain inexplicable; and may be truly urged as a
valid argument against the views here entertained. To show that it may
hereafter receive some explanation, I will give the following hypothesis.
From the nature of the organic remains, which do not appear to have
inhabited profound depths, in the several formations of Europe and of the
United States; and from the amount of sediment, miles in thickness, of
which the formations are composed, we may infer that from first to last
large islands or tracts of land, whence the sediment was derived, occurred
in the neighbourhood of the existing continents of Europe and North
America. But we do not know what was the state of things in the intervals
between the successive formations; whether Europe and the United States
during these intervals existed as dry land, or as a submarine surface near
land, on which sediment was not deposited, or again as the bed of an open
and unfathomable sea.
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 accumulated 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 subsidence, the great archipelagoes still areas of oscillations of
level, and the continents areas of elevation. But have we any right to
assume that things have thus remained from eternity? 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 assuming 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 superincumbent 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 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, Owen, Agassiz, 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 great
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.