On the Origin of Species by Means of Natural Selection
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Chapter II
Variation Under Nature
Variability -- Individual differences -- Doubtful species -- Wide ranging,
much diffused, and common species vary most -- Species of the larger genera
in any country vary more than the species of the smaller genera -- Many of
the species of the larger genera resemble varieties in being very closely,
but unequally, related to each other, and in having restricted ranges.
Before applying the principles arrived at in the last chapter to organic
beings in a state of nature, we must briefly discuss whether these latter
are subject to any variation. To treat this subject at all properly, a
long catalogue of dry facts should be given; but these I shall reserve for
my future work. Nor shall I here discuss the various definitions which
have been given of the term species. No one definition has as yet
satisfied all naturalists; yet every naturalist knows vaguely what he means
when he speaks of a species. Generally the term includes the unknown
element of a distinct act of creation. The term 'variety' is almost
equally difficult to define; but here community of descent is almost
universally implied, though it can rarely be proved. We have also what are
called monstrosities; but they graduate into varieties. By a monstrosity I
presume is meant some considerable deviation of structure in one part,
either injurious to or not useful to the species, and not generally
propagated. Some authors use the term 'variation' in a technical sense, as
implying a modification directly due to the physical conditions of life;
and 'variations' in this sense are supposed not to be inherited: but who
can say that the dwarfed condition of shells in the brackish waters of the
Baltic, or dwarfed plants on Alpine summits, or the thicker fur of an
animal from far northwards, would not in some cases be inherited for at
least some few generations? and in this case I presume that the form would
be called a variety.
Again, we have many slight differences which may be called individual
differences, such as are known frequently to appear in the offspring from
the same parents, or which may be presumed to have thus arisen, from being
frequently observed in the individuals of the same species inhabiting the
same confined locality. No one supposes that all the individuals of the
same species are cast in the very same mould. These individual differences
are highly important for us, as they afford materials for natural selection
to accumulate, in the same manner as man can accumulate in any given
direction individual differences in his domesticated productions. These
individual differences generally affect what naturalists consider
unimportant parts; but I could show by a long catalogue of facts, that
parts which must be called important, whether viewed under a physiological
or classificatory point of view, sometimes vary in the individuals of the
same species. I am convinced that the most experienced naturalist would be
surprised at the number of the cases of variability, even in important
parts of structure, which he could collect on good authority, as I have
collected, during a course of years. It should be remembered that
systematists are far from pleased at finding variability in important
characters, and that there are not many men who will laboriously examine
internal and important organs, and compare them in many specimens of the
same species. I should never have expected that the branching of the main
nerves close to the great central ganglion of an insect would have been
variable in the same species; I should have expected that changes of this
nature could have been effected only by slow degrees: yet quite recently
Mr. Lubbock has shown a degree of variability in these main nerves in
Coccus, which may almost be compared to the irregular branching of the stem
of a tree. This philosophical naturalist, I may add, has also quite
recently shown that the muscles in the larvae of certain insects are very
far from uniform. Authors sometimes argue in a circle when they state that
important organs never vary; for these same authors practically rank that
character as important (as some few naturalists have honestly confessed)
which does not vary; and, under this point of view, no instance of any
important part varying will ever be found: but under any other point of
view many instances assuredly can be given.
There is one point connected with individual differences, which seems to me
extremely perplexing: I refer to those genera which have sometimes been
called 'protean' or 'polymorphic,' in which the species present an
inordinate amount of variation; and hardly two naturalists can agree which
forms to rank as species and which as varieties. We may instance Rubus,
Rosa, and Hieracium amongst plants, several genera of insects, and several
genera of Brachiopod shells. In most polymorphic genera some of the
species have fixed and definite characters. Genera which are polymorphic
in one country seem to be, with some few exceptions, polymorphic in other
countries, and likewise, judging from Brachiopod shells, at former periods
of time. These facts seem to be very perplexing, for they seem to show
that this kind of variability is independent of the conditions of life. I
am inclined to suspect that we see in these polymorphic genera variations
in points of structure which are of no service or disservice to the
species, and which consequently have not been seized on and rendered
definite by natural selection, as hereafter will be explained.
Those forms which possess in some considerable degree the character of
species, but which are so closely similar to some other forms, or are so
closely linked to them by intermediate gradations, that naturalists do not
like to rank them as distinct species, are in several respects the most
important for us. We have every reason to believe that many of these
doubtful and closely-allied forms have permanently retained their
characters in their own country for a long time; for as long, as far as we
know, as have good and true species. Practically, when a naturalist can
unite two forms together by others having intermediate characters, he
treats the one as a variety of the other, ranking the most common, but
sometimes the one first described, as the species, and the other as the
variety. But cases of great difficulty, which I will not here enumerate,
sometimes occur in deciding whether or not to rank one form as a variety of
another, even when they are closely connected by intermediate links; nor
will the commonly-assumed hybrid nature of the intermediate links always
remove the difficulty. In very many cases, however, one form is ranked as
a variety of another, not because the intermediate links have actually been
found, but because analogy leads the observer to suppose either that they
do now somewhere exist, or may formerly have existed; and here a wide door
for the entry of doubt and conjecture is opened.
Hence, in determining whether a form should be ranked as a species or a
variety, the opinion of naturalists having sound judgment and wide
experience seems the only guide to follow. We must, however, in many
cases, decide by a majority of naturalists, for few well-marked and
well-known varieties can be named which have not been ranked as species by
at least some competent judges.
That varieties of this doubtful nature are far from uncommon cannot be
disputed. Compare the several floras of Great Britain, of France or of the
United States, drawn up by different botanists, and see what a surprising
number of forms have been ranked by one botanist as good species, and by
another as mere varieties. Mr. H. C. Watson, to whom I lie under deep
obligation for assistance of all kinds, has marked for me 182 British
plants, which are generally considered as varieties, but which have all
been ranked by botanists as species; and in making this list he has omitted
many trifling varieties, but which nevertheless have been ranked by some
botanists as species, and he has entirely omitted several highly
polymorphic genera. Under genera, including the most polymorphic forms,
Mr. Babington gives 251 species, whereas Mr. Bentham gives only 112,--a
difference of 139 doubtful forms! Amongst animals which unite for each
birth, and which are highly locomotive, doubtful forms, ranked by one
zoologist as a species and by another as a variety, can rarely be found
within the same country, but are common in separated areas. How many of
those birds and insects in North America and Europe, which differ very
slightly from each other, have been ranked by one eminent naturalist as
undoubted species, and by another as varieties, or, as they are often
called, as geographical races! Many years ago, when comparing, and seeing
others compare, the birds from the separate islands of the Galapagos
Archipelago, both one with another, and with those from the American
mainland, I was much struck how entirely vague and arbitrary is the
distinction between species and varieties. On the islets of the little
Madeira group there are many insects which are characterized as varieties
in Mr. Wollaston's admirable work, but which it cannot be doubted would be
ranked as distinct species by many entomologists. Even Ireland has a few
animals, now generally regarded as varieties, but which have been ranked as
species by some zoologists. Several most experienced ornithologists
consider our British red grouse as only a strongly-marked race of a
Norwegian species, whereas the greater number rank it as an undoubted
species peculiar to Great Britain. A wide distance between the homes of
two doubtful forms leads many naturalists to rank both as distinct species;
but what distance, it has been well asked, will suffice? if that between
America and Europe is ample, will that between the Continent and the
Azores, or Madeira, or the Canaries, or Ireland, be sufficient? It must be
admitted that many forms, considered by highly-competent judges as
varieties, have so perfectly the character of species that they are ranked
by other highly-competent judges as good and true species. But to discuss
whether they are rightly called species or varieties, before any definition
of these terms has been generally accepted, is vainly to beat the air.
Many of the cases of strongly-marked varieties or doubtful species well
deserve consideration; for several interesting lines of argument, from
geographical distribution, analogical variation, hybridism, &c., have been
brought to bear on the attempt to determine their rank. I will here give
only a single instance,--the well-known one of the primrose and cowslip, or
Primula veris and elatior. These plants differ considerably in appearance;
they have a different flavour and emit a different odour; they flower at
slightly different periods; they grow in somewhat different stations; they
ascend mountains to different heights; they have different geographical
ranges; and lastly, according to very numerous experiments made during
several years by that most careful observer Gartner, they can be crossed
only with much difficulty. We could hardly wish for better evidence of the
two forms being specifically distinct. On the other hand, they are united
by many intermediate links, and it is very doubtful whether these links are
hybrids; and there is, as it seems to me, an overwhelming amount of
experimental evidence, showing that they descend from common parents, and
consequently must be ranked as varieties.
Close investigation, in most cases, will bring naturalists to an agreement
how to rank doubtful forms. Yet it must be confessed, that it is in the
best-known countries that we find the greatest number of forms of doubtful
value. I have been struck with the fact, that if any animal or plant in a
state of nature be highly useful to man, or from any cause closely attract
his attention, varieties of it will almost universally be found recorded.
These varieties, moreover, will be often ranked by some authors as species.
Look at the common oak, how closely it has been studied; yet a German
author makes more than a dozen species out of forms, which are very
generally considered as varieties; and in this country the highest
botanical authorities and practical men can be quoted to show that the
sessile and pedunculated oaks are either good and distinct species or mere
varieties.
When a young naturalist commences the study of a group of organisms quite
unknown to him, he is at first much perplexed to determine what differences
to consider as specific, and what as varieties; for he knows nothing of the
amount and kind of variation to which the group is subject; and this shows,
at least, how very generally there is some variation. But if he confine
his attention to one class within one country, he will soon make up his
mind how to rank most of the doubtful forms. His general tendency will be
to make many species, for he will become impressed, just like the pigeon or
poultry-fancier before alluded to, with the amount of difference in the
forms which he is continually studying; and he has little general knowledge
of analogical variation in other groups and in other countries, by which to
correct his first impressions. As he extends the range of his
observations, he will meet with more cases of difficulty; for he will
encounter a greater number of closely-allied forms. But if his
observations be widely extended, he will in the end generally be enabled to
make up his own mind which to call varieties and which species; but he will
succeed in this at the expense of admitting much variation,--and the truth
of this admission will often be disputed by other naturalists. When,
moreover, he comes to study allied forms brought from countries not now
continuous, in which case he can hardly hope to find the intermediate links
between his doubtful forms, he will have to trust almost entirely to
analogy, and his difficulties will rise to a climax.
Certainly no clear line of demarcation has as yet been drawn between
species and sub-species--that is, the forms which in the opinion of some
naturalists come very near to, but do not quite arrive at the rank of
species; or, again, between sub-species and well-marked varieties, or
between lesser varieties and individual differences. These differences
blend into each other in an insensible series; and a series impresses the
mind with the idea of an actual passage.
Hence I look at individual differences, though of small interest to the
systematist, as of high importance for us, as being the first step towards
such slight varieties as are barely thought worth recording in works on
natural history. And I look at varieties which are in any degree more
distinct and permanent, as steps leading to more strongly marked and more
permanent varieties; and at these latter, as leading to sub-species, and to
species. The passage from one stage of difference to another and higher
stage may be, in some cases, due merely to the long-continued action of
different physical conditions in two different regions; but I have not much
faith in this view; and I attribute the passage of a variety, from a state
in which it differs very slightly from its parent to one in which it
differs more, to the action of natural selection in accumulating (as will
hereafter be more fully explained) differences of structure in certain
definite directions. Hence I believe a well-marked variety may be justly
called an incipient species; but whether this belief be justifiable must be
judged of by the general weight of the several facts and views given
throughout this work.
It need not be supposed that all varieties or incipient species necessarily
attain the rank of species. They may whilst in this incipient state become
extinct, or they may endure as varieties for very long periods, as has been
shown to be the case by Mr. Wollaston with the varieties of certain fossil
land-shells in Madeira. If a variety were to flourish so as to exceed in
numbers the parent species, it would then rank as the species, and the
species as the variety; or it might come to supplant and exterminate the
parent species; or both might co-exist, and both rank as independent
species. But we shall hereafter have to return to this subject.
From these remarks it will be seen that I look at the term species, as one
arbitrarily given for the sake of convenience to a set of individuals
closely resembling each other, and that it does not essentially differ from
the term variety, which is given to less distinct and more fluctuating
forms. The term variety, again, in comparison with mere individual
differences, is also applied arbitrarily, and for mere convenience sake.
Guided by theoretical considerations, I thought that some interesting
results might be obtained in regard to the nature and relations of the
species which vary most, by tabulating all the varieties in several
well-worked floras. At first this seemed a simple task; but Mr. H. C.
Watson, to whom I am much indebted for valuable advice and assistance on
this subject, soon convinced me that there were many difficulties, as did
subsequently Dr. Hooker, even in stronger terms. I shall reserve for my
future work the discussion of these difficulties, and the tables themselves
of the proportional numbers of the varying species. Dr. Hooker permits me
to add, that after having carefully read my manuscript, and examined the
tables, he thinks that the following statements are fairly well
established. The whole subject, however, treated as it necessarily here is
with much brevity, is rather perplexing, and allusions cannot be avoided to
the 'struggle for existence,' 'divergence of character,' and other
questions, hereafter to be discussed.
Alph. De Candolle and others have shown that plants which have very wide
ranges generally present varieties; and this might have been expected, as
they become exposed to diverse physical conditions, and as they come into
competition (which, as we shall hereafter see, is a far more important
circumstance) with different sets of organic beings. But my tables further
show that, in any limited country, the species which are most common, that
is abound most in individuals, and the species which are most widely
diffused within their own country (and this is a different consideration
from wide range, and to a certain extent from commonness), often give rise
to varieties sufficiently well-marked to have been recorded in botanical
works. Hence it is the most flourishing, or, as they may be called, the
dominant species,--those which range widely over the world, are the most
diffused in their own country, and are the most numerous in
individuals,--which oftenest produce well-marked varieties, or, as I
consider them, incipient species. And this, perhaps, might have been
anticipated; for, as varieties, in order to become in any degree permanent,
necessarily have to struggle with the other inhabitants of the country, the
species which are already dominant will be the most likely to yield
offspring which, though in some slight degree modified, will still inherit
those advantages that enabled their parents to become dominant over their
compatriots.
If the plants inhabiting a country and described in any Flora be divided
into two equal masses, all those in the larger genera being placed on one
side, and all those in the smaller genera on the other side, a somewhat
larger number of the very common and much diffused or dominant species will
be found on the side of the larger genera. This, again, might have been
anticipated; for the mere fact of many species of the same genus inhabiting
any country, shows that there is something in the organic or inorganic
conditions of that country favourable to the genus; and, consequently, we
might have expected to have found in the larger genera, or those including
many species, a large proportional number of dominant species. But so many
causes tend to obscure this result, that I am surprised that my tables show
even a small majority on the side of the larger genera. I will here allude
to only two causes of obscurity. Fresh-water and salt-loving plants have
generally very wide ranges and are much diffused, but this seems to be
connected with the nature of the stations inhabited by them, and has little
or no relation to the size of the genera to which the species belong.
Again, plants low in the scale of organisation are generally much more
widely diffused than plants higher in the scale; and here again there is no
close relation to the size of the genera. The cause of lowly-organised
plants ranging widely will be discussed in our chapter on geographical
distribution.
From looking at species as only strongly-marked and well-defined varieties,
I was led to anticipate that the species of the larger genera in each
country would oftener present varieties, than the species of the smaller
genera; for wherever many closely related species (i.e. species of the same
genus) have been formed, many varieties or incipient species ought, as a
general rule, to be now forming. Where many large trees grow, we expect to
find saplings. Where many species of a genus have been formed through
variation, circumstances have been favourable for variation; and hence we
might expect that the circumstances would generally be still favourable to
variation. On the other hand, if we look at each species as a special act
of creation, there is no apparent reason why more varieties should occur in
a group having many species, than in one having few.
To test the truth of this anticipation I have arranged the plants of twelve
countries, and the coleopterous insects of two districts, into two nearly
equal masses, the species of the larger genera on one side, and those of
the smaller genera on the other side, and it has invariably proved to be
the case that a larger proportion of the species on the side of the larger
genera present varieties, than on the side of the smaller genera.
Moreover, the species of the large genera which present any varieties,
invariably present a larger average number of varieties than do the species
of the small genera. Both these results follow when another division is
made, and when all the smallest genera, with from only one to four species,
are absolutely excluded from the tables. These facts are of plain
signification on the view that species are only strongly marked and
permanent varieties; for whenever many species of the same genus have been
formed, or where, if we may use the expression, the manufactory of species
has been active, we ought generally to find the manufactory still in
action, more especially as we have every reason to believe the process of
manufacturing new species to be a slow one. And this certainly is the
case, if varieties be looked at as incipient species; for my tables clearly
show as a general rule that, wherever many species of a genus have been
formed, the species of that genus present a number of varieties, that is of
incipient species, beyond the average. It is not that all large genera are
now varying much, and are thus increasing in the number of their species,
or that no small genera are now varying and increasing; for if this had
been so, it would have been fatal to my theory; inasmuch as geology plainly
tells us that small genera have in the lapse of time often increased
greatly in size; and that large genera have often come to their maxima,
declined, and disappeared. All that we want to show is, that where many
species of a genus have been formed, on an average many are still forming;
and this holds good.
There are other relations between the species of large genera and their
recorded varieties which deserve notice. We have seen that there is no
infallible criterion by which to distinguish species and well-marked
varieties; and in those cases in which intermediate links have not been
found between doubtful forms, naturalists are compelled to come to a
determination by the amount of difference between them, judging by analogy
whether or not the amount suffices to raise one or both to the rank of
species. Hence the amount of difference is one very important criterion in
settling whether two forms should be ranked as species or varieties. Now
Fries has remarked in regard to plants, and Westwood in regard to insects,
that in large genera the amount of difference between the species is often
exceedingly small. I have endeavoured to test this numerically by
averages, and, as far as my imperfect results go, they always confirm the
view. I have also consulted some sagacious and most experienced observers,
and, after deliberation, they concur in this view. In this respect,
therefore, the species of the larger genera resemble varieties, more than
do the species of the smaller genera. Or the case may be put in another
way, and it may be said, that in the larger genera, in which a number of
varieties or incipient species greater than the average are now
manufacturing, many of the species already manufactured still to a certain
extent resemble varieties, for they differ from each other by a less than
usual amount of difference.
Moreover, the species of the large genera are related to each other, in the
same manner as the varieties of any one species are related to each other.
No naturalist pretends that all the species of a genus are equally distinct
from each other; they may generally be divided into sub-genera, or
sections, or lesser groups. As Fries has well remarked, little groups of
species are generally clustered like satellites around certain other
species. And what are varieties but groups of forms, unequally related to
each other, and clustered round certain forms--that is, round their
parent-species? Undoubtedly there is one most important point of
difference between varieties and species; namely, that the amount of
difference between varieties, when compared with each other or with their
parent-species, is much less than that between the species of the same
genus. But when we come to discuss the principle, as I call it, of
Divergence of Character, we shall see how this may be explained, and how
the lesser differences between varieties will tend to increase into the
greater differences between species.
There is one other point which seems to me worth notice. Varieties
generally have much restricted ranges: this statement is indeed scarcely
more than a truism, for if a variety were found to have a wider range than
that of its supposed parent-species, their denominations ought to be
reversed. But there is also reason to believe, that those species which
are very closely allied to other species, and in so far resemble varieties,
often have much restricted ranges. For instance, Mr. H. C. Watson has
marked for me in the well-sifted London Catalogue of plants (4th edition)
63 plants which are therein ranked as species, but which he considers as so
closely allied to other species as to be of doubtful value: these 63
reputed species range on an average over 6.9 of the provinces into which
Mr. Watson has divided Great Britain. Now, in this same catalogue, 53
acknowledged varieties are recorded, and these range over 7.7 provinces;
whereas, the species to which these varieties belong range over 14.3
provinces. So that the acknowledged varieties have very nearly the same
restricted average range, as have those very closely allied forms, marked
for me by Mr. Watson as doubtful species, but which are almost universally
ranked by British botanists as good and true species.
Finally, then, varieties have the same general characters as species, for
they cannot be distinguished from species,--except, firstly, by the
discovery of intermediate linking forms, and the occurrence of such links
cannot affect the actual characters of the forms which they connect; and
except, secondly, by a certain amount of difference, for two forms, if
differing very little, are generally ranked as varieties, notwithstanding
that intermediate linking forms have not been discovered; but the amount of
difference considered necessary to give to two forms the rank of species is
quite indefinite. In genera having more than the average number of species
in any country, the species of these genera have more than the average
number of varieties. In large genera the species are apt to be closely,
but unequally, allied together, forming little clusters round certain
species. Species very closely allied to other species apparently have
restricted ranges. In all these several respects the species of large
genera present a strong analogy with varieties. And we can clearly
understand these analogies, if species have once existed as varieties, and
have thus originated: whereas, these analogies are utterly inexplicable if
each species has been independently created.
We have, also, seen that it is the most flourishing and dominant species of
the larger genera which on an average vary most; and varieties, as we shall
hereafter see, tend to become converted into new and distinct species. The
larger genera thus tend to become larger; and throughout nature the forms
of life which are now dominant tend to become still more dominant by
leaving many modified and dominant descendants. But by steps hereafter to
be explained, the larger genera also tend to break up into smaller genera.
And thus, the forms of life throughout the universe become divided into
groups subordinate to groups.
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