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 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 nuinbers 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.
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 IMPERFECTION OF THE GEOLOGICAL
RECORD 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 IMPERFECTION OF
THE GEOLOGICAL RECORD 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 fine 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
IMPERFECTION OF THE GEOLOGICAL RECORD 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 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 IMPERFECTION OF THE
GEOLOGICAL RECORD 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 fossilferous
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, etc, 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 or 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, etc,
and all our greatest geologists, as Lyell, Murchison, Sedgwick,
etc, 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 even 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.
Peter v. Sengbusch - b-online@botanik.uni-hamburg.de