ON THE ABSENCE OR RARITY OF TRANSITIONAL VARIETIES.
As natural selection acts solely by the preservation of profitable
modifications, each new form will tend in a fully-stocked country to
take the place of, and finally to exterminate, its own less improved
parent or other less-favoured forms with which it comes into
competition. Thus extinction and natural selection will, as we have
seen, go hand in hand. Hence, if we look at each species as descended
from some other unknown form, both the parent and all the transitional
varieties will generally have been exterminated by the very process of
formation and perfection of the new form.
But, as by this theory innumerable transitional forms must have existed,
why do we not find them embedded in countless numbers in the crust of
the earth? It will be much more convenient to discuss this question in
the chapter on the Imperfection of the geological record; and I will
here only state that I believe the answer mainly lies in the record
being incomparably less perfect than is generally supposed; the
imperfection of the record being chiefly due to organic beings not
inhabiting profound depths of the sea, and to their remains being
embedded and preserved to a future age only in masses of sediment
sufficiently thick and extensive to withstand an enormous amount of
future degradation; and such fossiliferous masses can be accumulated
only where much sediment is deposited on the shallow bed of the sea,
whilst it slowly subsides. These contingencies will concur only rarely,
and after enormously long intervals. Whilst the bed of the sea
is stationary or is rising, or when very little sediment is being
deposited, there will be blanks in our geological history. The crust of
the earth is a vast museum; but the natural collections have been made
only at intervals of time immensely remote.
But it may be urged that when several closely-allied species inhabit
the same territory we surely ought to find at the present time many
transitional forms. Let us take a simple case: in travelling from north
to south over a continent, we generally meet at successive intervals
with closely allied or representative species, evidently filling nearly
the same place in the natural economy of the land. These representative
species often meet and interlock; and as the one becomes rarer and
rarer, the other becomes more and more frequent, till the one replaces
the other. But if we compare these species where they intermingle, they
are generally as absolutely distinct from each other in every detail of
structure as are specimens taken from the metropolis inhabited by each.
By my theory these allied species have descended from a common parent;
and during the process of modification, each has become adapted to
the conditions of life of its own region, and has supplanted and
exterminated its original parent and all the transitional varieties
between its past and present states. Hence we ought not to expect at
the present time to meet with numerous transitional varieties in each
region, though they must have existed there, and may be embedded
there in a fossil condition. But in the intermediate region, having
intermediate conditions of life, why do we not now find closely-linking
intermediate varieties? This difficulty for a long time quite confounded
me. But I think it can be in large part explained.
In the first place we should be extremely cautious in inferring, because
an area is now continuous, that it has been continuous during a long
period. Geology would lead us to believe that almost every continent has
been broken up into islands even during the later tertiary periods;
and in such islands distinct species might have been separately formed
without the possibility of intermediate varieties existing in the
intermediate zones. By changes in the form of the land and of climate,
marine areas now continuous must often have existed within recent times
in a far less continuous and uniform condition than at present. But I
will pass over this way of escaping from the difficulty; for I believe
that many perfectly defined species have been formed on strictly
continuous areas; though I do not doubt that the formerly broken
condition of areas now continuous has played an important part in the
formation of new species, more especially with freely-crossing and
wandering animals.
In looking at species as they are now distributed over a wide area,
we generally find them tolerably numerous over a large territory, then
becoming somewhat abruptly rarer and rarer on the confines, and finally
disappearing. Hence the neutral territory between two representative
species is generally narrow in comparison with the territory proper to
each. We see the same fact in ascending mountains, and sometimes it
is quite remarkable how abruptly, as Alph. De Candolle has observed,
a common alpine species disappears. The same fact has been noticed by
Forbes in sounding the depths of the sea with the dredge. To those who
look at climate and the physical conditions of life as the all-important
elements of distribution, these facts ought to cause surprise, as
climate and height or depth graduate away insensibly. But when we
bear in mind that almost every species, even in its metropolis, would
increase immensely in numbers, were it not for other competing species;
that nearly all either prey on or serve as prey for others; in short,
that each organic being is either directly or indirectly related in
the most important manner to other organic beings, we must see that the
range of the inhabitants of any country by no means exclusively depends
on insensibly changing physical conditions, but in large part on the
presence of other species, on which it depends, or by which it is
destroyed, or with which it comes into competition; and as these species
are already defined objects (however they may have become so), not
blending one into another by insensible gradations, the range of any one
species, depending as it does on the range of others, will tend to be
sharply defined. Moreover, each species on the confines of its range,
where it exists in lessened numbers, will, during fluctuations in the
number of its enemies or of its prey, or in the seasons, be extremely
liable to utter extermination; and thus its geographical range will come
to be still more sharply defined.
If I am right in believing that allied or representative species, when
inhabiting a continuous area, are generally so distributed that each
has a wide range, with a comparatively narrow neutral territory between
them, in which they become rather suddenly rarer and rarer; then, as
varieties do not essentially differ from species, the same rule will
probably apply to both; and if we in imagination adapt a varying species
to a very large area, we shall have to adapt two varieties to two
large areas, and a third variety to a narrow intermediate zone. The
intermediate variety, consequently, will exist in lesser numbers from
inhabiting a narrow and lesser area; and practically, as far as I can
make out, this rule holds good with varieties in a state of nature. I
have met with striking instances of the rule in the case of varieties
intermediate between well-marked varieties in the genus Balanus. And it
would appear from information given me by Mr. Watson, Dr. Asa Gray, and
Mr. Wollaston, that generally when varieties intermediate between two
other forms occur, they are much rarer numerically than the forms which
they connect. Now, if we may trust these facts and inferences, and
therefore conclude that varieties linking two other varieties together
have generally existed in lesser numbers than the forms which they
connect, then, I think, we can understand why intermediate varieties
should not endure for very long periods;--why as a general rule they
should be exterminated and disappear, sooner than the forms which they
originally linked together.
For any form existing in lesser numbers would, as already remarked,
run a greater chance of being exterminated than one existing in large
numbers; and in this particular case the intermediate form would be
eminently liable to the inroads of closely allied forms existing on both
sides of it. But a far more important consideration, as I believe, is
that, during the process of further modification, by which two varieties
are supposed on my theory to be converted and perfected into two
distinct species, the two which exist in larger numbers from inhabiting
larger areas, will have a great advantage over the intermediate variety,
which exists in smaller numbers in a narrow and intermediate zone.
For forms existing in larger numbers will always have a better chance,
within any given period, of presenting further favourable variations for
natural selection to seize on, than will the rarer forms which exist in
lesser numbers. Hence, the more common forms, in the race for life, will
tend to beat and supplant the less common forms, for these will be
more slowly modified and improved. It is the same principle which, as
I believe, accounts for the common species in each country, as shown
in the second chapter, presenting on an average a greater number of
well-marked varieties than do the rarer species. I may illustrate what I
mean by supposing three varieties of sheep to be kept, one adapted to an
extensive mountainous region; a second to a comparatively narrow, hilly
tract; and a third to wide plains at the base; and that the inhabitants
are all trying with equal steadiness and skill to improve their stocks
by selection; the chances in this case will be strongly in favour of the
great holders on the mountains or on the plains improving their breeds
more quickly than the small holders on the intermediate narrow, hilly
tract; and consequently the improved mountain or plain breed will soon
take the place of the less improved hill breed; and thus the two breeds,
which originally existed in greater numbers, will come into close
contact with each other, without the interposition of the supplanted,
intermediate hill-variety.
To sum up, I believe that species come to be tolerably well-defined
objects, and do not at any one period present an inextricable chaos of
varying and intermediate links: firstly, because new varieties are
very slowly formed, for variation is a very slow process, and natural
selection can do nothing until favourable variations chance to occur,
and until a place in the natural polity of the country can be better
filled by some modification of some one or more of its inhabitants.
And such new places will depend on slow changes of climate, or on the
occasional immigration of new inhabitants, and, probably, in a still
more important degree, on some of the old inhabitants becoming slowly
modified, with the new forms thus produced and the old ones acting and
reacting on each other. So that, in any one region and at any one time,
we ought only to see a few species presenting slight modifications of
structure in some degree permanent; and this assuredly we do see.
Secondly, areas now continuous must often have existed within the
recent period in isolated portions, in which many forms, more especially
amongst the classes which unite for each birth and wander much, may have
separately been rendered sufficiently distinct to rank as representative
species. In this case, intermediate varieties between the several
representative species and their common parent, must formerly have
existed in each broken portion of the land, but these links will
have been supplanted and exterminated during the process of natural
selection, so that they will no longer exist in a living state.
Thirdly, when two or more varieties have been formed in different
portions of a strictly continuous area, intermediate varieties will, it
is probable, at first have been formed in the intermediate zones, but
they will generally have had a short duration. For these intermediate
varieties will, from reasons already assigned (namely from what we know
of the actual distribution of closely allied or representative species,
and likewise of acknowledged varieties), exist in the intermediate zones
in lesser numbers than the varieties which they tend to connect. From
this cause alone the intermediate varieties will be liable to accidental
extermination; and during the process of further modification through
natural selection, they will almost certainly be beaten and supplanted
by the forms which they connect; for these from existing in greater
numbers will, in the aggregate, present more variation, and thus be
further improved through natural selection and gain further advantages.
Lastly, looking not to any one time, but to all time, if my theory be
true, numberless intermediate varieties, linking most closely all the
species of the same group together, must assuredly have existed; but the
very process of natural selection constantly tends, as has been so often
remarked, to exterminate the parent forms and the intermediate links.
Consequently evidence of their former existence could be found only
amongst fossil remains, which are preserved, as we shall in a future
chapter attempt to show, in an extremely imperfect and intermittent
record.