may as well remember, however, that it really constitutes only one part
of his theory; for besides this hypothesis of the cumulative inheritance
of functionally-produced modifications--to which we may add the
inherited effects of any direct action exercised by surrounding
conditions of life,--Lamarck believed in some transcendental principle
tending to produce gradual improvement in pre-determined lines of
advance. Therefore it would really be more correct to designate the
former hypothesis by the name either of Erasmus Darwin, or, still
better, of Herbert Spencer. Nevertheless, in order to avoid confusion, I
will follow established custom, and subsequently speak of this
hypothesis as the Lamarckian hypothesis--understanding, however, that in
employing this designation I am not referring to any part or factor of
Lamarck's general theory of evolution other than the one which has just
been described--namely, the hypothesis of the cumulative transmission of
functionally-produced, or otherwise "acquired," modifications.
This, then, was the earliest hypothesis touching the causes of organic
evolution. But we may at once perceive that it is insufficient to
explain all that stands to be explained. In the first place, it refers
in chief part only to the higher animals, which are actuated to effort
by intelligence. Its explanatory power in the case of most
invertebrata--as well as in that of all plants--is extremely limited,
inasmuch as these organisms can never be moved to a greater or less use
of their several parts by any discriminating volition, such as that
which leads to the continued straining of a giraffe's neck for the
purpose of reaching foliage. In the second place, even among the higher
animals there are numberless tissues and organs which unquestionably
present a high degree of adaptive evolution, but which nevertheless
cannot be supposed to have fallen within the influence of Lamarckian
principles. Of such are the shells of crustacea, tortoises, &c., which
although undoubtedly of great use to the animals presenting them, cannot
ever have been _used_ in the sense required by Lamarck's hypothesis, i.
e. actively exercised, so as to increase a flow of nutrition to the
part. Lastly, in the third place, the validity of Lamarck's hypothesis
in any case whatsoever has of late years become a matter of serious
question, as will be fully shown and discussed in the next volume.
Meanwhile it is enough to observe that, on account of all these reasons,
the theory of Lamarck, even if it be supposed to present any truth at
all, is clearly insufficient as a full or complete theory of organic
evolution.
* * * * *
In historical order the next theory that was arrived at was the theory
of natural selection, simultaneously published by Darwin and Wallace on
July 1st, 1858.
If we may estimate the importance of an idea by the change of thought
which it effects, this idea of natural selection is unquestionably the
most important idea that has ever been conceived by the mind of man.
Yet the wonder is that it should not have been hit upon long before. Or
rather, I should say, the wonder is that its immense and immeasurable
importance should not have been previously recognised. For, since the
publication of this idea by Darwin and Wallace, it has been found that
its main features had already occurred to at least two other
minds--namely, Dr. Wells in 1813, and Mr. Patrick Matthew in 1831. But
neither of these writers perceived that in the few scattered sentences
which they had written upon the subject they had struck the key-note of
organic nature, and resolved one of the principal chords of the
universe. Still more remarkable is the fact that Mr. Herbert
Spencer--notwithstanding his great powers of abstract thought and his
great devotion of those powers to the theory of evolution, when as yet
this theory was scorned by science--still more remarkable, I say, is the
fact that Mr. Herbert Spencer should have missed what now appears so
obvious an idea. But most remarkable of all is the fact that Dr.
Whewell, with all his stores of information on the history of the
inductive sciences, and with all his acumen on the matter of scientific
method, should not only have conceived the idea of natural selection,
but expressly stated it as a logically possible explanation of the
origin of species, and yet have so stated it merely for the purpose of
dismissing it with contempt[26]. This, I think, is most remarkable,
because it serves to prove how very far men's minds at that time must
have been from entertaining, as in any way antecedently probable, the
doctrine of transmutation. In order to show this I will here quote one
passage from the writings of Whewell, and another from a distinguished
French naturalist referred to by him.
[26] For quotations, see Note A.
In 1846 Whewell wrote:--
Not only is the doctrine of the transmutation of species in itself
disproved by the best physiological reasonings, but the additional
assumptions which are requisite to enable its advocates to apply it
to the explanation of the geological and other phenomena of the
earth, are altogether gratuitous and fantastical[27].
[27] whewell, _indications of the creator_, 2nd ed., 1846.
Then he quotes with approval the following opinion:--
Against this hypothesis, which, up to the present time, I regard as
purely gratuitous, and likely to turn geologists out of the sound
and excellent road in which they now are, I willingly raise my
voice, with the most absolute conviction of being in the right[28].
[28] de blainville, _compte rendu_, 1837.
And, after displaying the proof rendered by Lyell of uniformitarianism
in geology, and cordially subscribing thereto, Whewell adds:--
We are led by our reasonings to this view, that the present order
of things was commenced by an act of creative power entirely
different to any agency which has been exerted since. None of the
influences which have modified the present races of animals and
plants since they were placed in their habitations on the earth's
surface can have had any efficacy in producing them at first. We
are necessarily driven to assume, as the beginning of the present
cycle of organic nature, an event not included in the course of
nature[29].
[29] Whewell, _ibid._, p. 162.
So much, then, for the state of the most enlightened and representative
opinions on the question of evolution before the publication of
Darwin's work; and so much, likewise, for the only reasonable
suggestions as to the causes of evolution which up to that time had been
put forward, even by those few individuals who entertained any belief in
evolution as a fact. It was the theory of natural selection that changed
all this, and created a revolution in the thought of our time, the
magnitude of which in many of its far-reaching consequences we are not
even yet in a position to appreciate; but the action of which has
already wrought a transformation in general philosophy, as well as in
the more special science of biology, that is without a parallel in the
history of mankind.
* * * * *
Although every one is now more or less well acquainted with the theory
of natural selection, it is necessary, for the sake of completeness,
that I should state the theory; and I will do so in full detail.
It is a matter of observable fact that all plants and animals are
perpetually engaged in what Darwin calls a "struggle for existence."
That is to say, in every generation of every species a great many more
individuals are born than can possibly survive; so that there is in
consequence a perpetual battle for life going on among all the
constituent individuals of any given generation. Now, in this struggle
for existence, which individuals will be victorious and live? Assuredly
those which are best fitted to live, in whatever respect, or respects,
their superiority of fitness may consist. Hence it follows that Nature,
so to speak, _selects_ the best individuals out of each generation to
live. And not only so; but as these favoured individuals transmit their
favourable qualities to their offspring, according to the fixed laws of
heredity, it further follows that the individuals composing each
successive generation have a general tendency to be better suited to
their surroundings than were their forefathers. And this follows, not
merely because in every generation it is only the "flower of the flock"
that is allowed to breed, but also because, if in any generation some
new and beneficial qualities happen to arise as slight variations from
the ancestral type, they will (other things permitting) be seized upon
by natural selection, and, being transmitted by heredity to subsequent
generations, will be added to the previously existing type. Thus the
best idea of the whole process will be gained by comparing it with the
closely analogous process whereby gardeners, fanciers, and
cattle-breeders create their wonderful productions; for just as these
men, by always "_selecting_" their best individuals to breed from,
slowly but continuously improve their stock, so Nature, by a similar
process of "_selection_" slowly but continuously makes the various
species of plants and animals better and better suited to the conditions
of their life.
Now, if this process of continuously adapting organisms to their
environment takes place in nature at all, there is no reason why we
should set any limits on the extent to which it is able to go, up to the
point at which a complete and perfect adaptation is achieved. Therefore
we might suppose that all species would eventually reach this condition
of perfect harmony with their environment, and then remain fixed. And
so, according to the theory, they would, if the environment were itself
unchanging. But forasmuch as the environment (i. e. the sum total of the
external conditions of life) of almost every organic type alters more
or less from century to century--whether from astronomical, geological,
and geographical changes, or from the immigrations and emigrations of
other species living on contiguous areas, and so on--it follows that the
process of natural selection need never reach a terminal phase. And
forasmuch as natural selection may thus continue, _ad infinitum_, slowly
to alter a specific type in adaptation to a gradually changing
environment, if in any case the alteration thus effected is sufficient
in amount to lead naturalists to name the result as a distinct species,
it follows that natural selection has transmuted one specific type into
another. Similarly, by a continuation of the process, specific types
would become transmuted into generic, generic into family types, and so
on. Thus the process is supposed to go on throughout all the countless
forms of life continuously and simultaneously--the world of organic
types being thus regarded as in a state of perpetual, though gradual,
flux.
* * * * *
Now, the first thing we have to notice about this theory is, that in all
its main elements it is merely a statement of observable facts. It is an
observable fact that in all species of plants and animals a very much
larger number of individuals are born than can possibly survive. Thus,
for example, it has been calculated that if the progeny of a single pair
of elephants--which are the slowest breeding of animals--were all
allowed to reach maturity and propagate, in 750 years there would be
living 19,000,000 descendants. Again, in the case of vegetables, if a
species of annual plant produces only two seeds a year, if these in
successive years were all allowed to reproduce their kind, in twenty
years there would be 11,000,000 plants from a single ancestor. Yet we
know that nearly all animals and plants produce many more young at a
time than in either of these two supposed cases. Indeed, as individuals
of many kinds of plants, and not a few kinds of animals, produce every
year several thousand young, we may make a rough estimate and say, that
over organic nature as a whole probably not one in a thousand young are
allowed to survive to the age of reproduction. How tremendous,
therefore, must be the struggle for existence! It is thought a terrible
thing in battle when one half the whole number of combatants perish. But
what are we to think of a battle for life where only one in a thousand
survives?
This, then, is the first fact. The second is the fact so long ago
recognised, that the battle is to the strong, the race to the swift. The
thousandth individual which does survive in the battle for
existence--which does win the race for life--is, without question, one
of the individuals best fitted to do so; that is to say, best fitted to
the conditions of its existence considered as a whole. Nature is,
therefore, always picking out, or selecting, such individuals to live
and to breed.
The third fact is, that the individuals so selected transmit their
favourable qualities to their offspring by heredity. There is no doubt
about this fact, so far as we are concerned with it. For although, as I
have already hinted, considerable doubt has of late years been cast upon
Lamarck's doctrine of the hereditary transmission of _acquired_
characters, it remains as impossible as ever it was to question the
hereditary transmission of what are called _congenital_ characters. And
this is all that Darwin's theory necessarily requires.
The fourth fact is, that although heredity as a whole produces a
wonderfully exact copy of the parent in the child, there is never a
precise reduplication. Of all the millions of human beings upon the face
of the earth, no one is so like another that we cannot see some
difference; the resemblance is everywhere specific, nowhere individual.
Now this same remark applies to all specific types. The only reason why
we notice individual differences in the case of the human type more than
we do in the case of any other types, is because our attention is here
more incessantly focussed upon these differences. We are compelled to
notice them in the case of our own species, however small they may
appear to a naturalist, because, unless we do so, we should not
recognise the members of our own family, or be able to distinguish
between a man whom we know is ready to do us an important service, and
another man whom we know is ready to cut our throats. But our common
mother Nature is able thus to distinguish between all her children. Her
eyes are much more ready to detect small individual peculiarities than
are the eyes of any naturalist. No slight variations in the cast of
feature or disposition of parts, no minute difference in the arrangement
of microscopical cells, can escape her ever vigilant attention. And,
consequently, when among all the innumerable multitudes of individual
variations any one arises which--no matter in how slight a degree--gives
to that individual a better chance of success in the struggle for life,
Nature chooses that individual to survive, and so to perpetuate the
improvement in his or her progeny.
Now I say that all these several component parts of Darwinian doctrine
are not matters of theory, but matters of fact. The only element of
theory in his doctrine of evolution by natural selection has reference
to the degree in which these observable facts, when thus brought
together, are adequate to account for the process of evolution.
* * * * *
So much, then, as a statement of the theory of natural selection. But
from this statement--i. e. from the theory of natural selection
itself--there follow certain matters of general principle which it is
important to bear in mind. These, therefore, I shall here proceed to
mention.
First of all, it is evident that the theory is applicable as an
explanation of organic changes in specific types only in so far as these
changes are of _use_, or so far as such changes endow the species with
better chances of success in the general struggle for existence. This is
the only sense in which I shall always employ the terms use, utility,
service, benefit, and so forth--that is to say, in the sense of
life-preserving.
* * * * *
Next, it must be clearly understood that the life which it is the
object, so to speak, of natural selection to preserve, is primarily the
life of the _species_; not that of the _individual_. Natural selection
preserves the life of the individual only in so far as this is conducive
to that of the species. Wherever the life-interests of the individual
clash with those of the species, that individual is sacrificed in favour
of others who happen better to subserve the interests of the species.
For example, in all organisms a greater or less amount of vigour is
wasted, so far as individual interests are concerned, in the formation
and the nourishment of progeny. In the great majority of plants and
animals an enormous amount of physiological energy is thus expended.
Look at the roe or the milt of a herring, for instance, and see what a
huge drain has been made upon the individual for the sake of its
species. Again, all unselfish instincts have been developed for the sake
of the species, and usually against the interests of the individual. An
ant which will allow her head to be slowly drawn from her body rather
than relinquish her hold upon a pupa, is clearly acting in response to
an instinct which has been developed for the benefit of the hive, though
fatal to the individual. And, in a lesser degree, the parental
instincts, wherever they occur, are more or less detrimental to the
interests of the individual, though correspondingly essential to those
of the race.
These illustrations will serve to show that natural selection always
works primarily for the life-interests of the species--and, indeed, only
works for those of the individual at all in so far as the latter happen
to coincide with the former. Or, otherwise stated, the object of natural
selection is always that of producing and maintaining specific types in
the highest degree of efficiency, no matter what may become of the
constituent individuals. Which is a striking republication by Science of
a general truth previously stated by Poetry:--
So careful of the type she seems,
So careless of the single life.
Tennyson thus noted the fact, and a few years later Darwin supplied the
explanation.
But of course in many, if not in the majority of cases, anything that
adds to the life-sustaining power of the single life thereby ministers
also to the life-sustaining power of the type; and thus we can
understand why all mechanisms and instincts which minister to the single
life have been developed--namely, because the life of the species is
made up of the lives of all its constituent individuals. It is only
where the interests of the one clash with those of the other that
natural selection works against the individual. So long as the interests
are coincident, it works in favour of both.
Natural selection, then, is a theory which seeks to explain by natural
causes the occurrence of every kind of adaptation which is to be met
with in organic nature, on the assumption that adaptations of every kind
have primary reference to the preservation of species, and therefore
also, as a general rule, to the preservation of their constituent
individuals. And from this it follows that where it is for the benefit
of a species to change its type, natural selection will effect that
change, thus leading to a specific transmutation, or the evolution of a
new species. In such cases the old species may or may not become
extinct. If the transmutation affects the species as a whole, or
throughout its entire range, of course _that_ particular type becomes
extinct, although it does so by becoming changed into a still more
suitable type in the course of successive generations. If, on the other
hand, the transmutation affects only a part of the original species, or
not throughout its entire range, then the other parts of that species
may survive for any number of ages as they originally were. In the one
case there is a ladder-like transmutation of species in time; in the
other case a possibly tree-like multiplication of species in space. But
whether the evolution of species be thus serial in time or divergent in
space, the object of natural selection, so to speak, is in either case
the same--namely, that of preserving all types which prove best suited
to the conditions of their existence.
* * * * *
Once more, the term "struggle for existence" must be understood to
comprehend, not only a competition for life among contemporary
individuals of the same species, but likewise a struggle by all such
individuals taken collectively for the continuance of their own specific
type. Thus, on the one hand, while there is a perpetual civil war being
waged between members of the same species, on the other hand there is a
foreign war being waged by the species as a whole against its world as a
whole. Hence it follows that natural selection does not secure survival
of the fittest as regards individuals only, but also survival of the
fittest as regards types. This is a most important point to remember,
because, as a general rule, these two different causes produce exactly
opposite effects. Success in the civil war, where each is fighting
against all, is determined by _individual_ fitness and _self-reliance_.
But success in the foreign war is determined by what may be termed
_tribal_ fitness and _mutual dependence_. For example, among social
insects the struggle for existence is quite as great between different
tribes or communities, as it is between different individuals of the
same community; and thus we can understand the extraordinary degree in
which not only co-operative instincts, but also largely intelligent
social habits, have here been developed[30]. Similarly, in the case of
mankind, we can understand the still more extraordinary development of
these things--culminating in the moral sense. I have heard a sermon,
preached at one of the meetings of the British Association, entirely
devoted to arguing that the moral sense could not have been evolved by
natural selection, seeing that the altruism which this sense involves is
the very opposite of selfishness, which alone ought to have been the
product of survival of the fittest in a struggle for life. And, of
course, this argument would have been perfectly sound had Darwin limited
the struggle for existence to individuals, without extending it to
communities. But if the preacher had ever read Darwin's works he would
have found that, when thus extended, the principle of natural selection
is bound to work in favour of the co-operative instincts in the case of
so highly social an animal as man; and that of these instincts
conscience is the highest imaginable exhibition.
[30] For cases, see _Animal Intelligence_, in the chapters on Ants
and Bees; and, for discussion of principles, _Mental Evolution in
Animals_, in the chapters on Instinct.
What I have called tribal fitness--in contradistinction to individual
fitness--begins with the family, developes in the community (herd, hive,
clan, &c.), and usually ends with the limits of the species. On the one
hand, however, it is but seldom that it extends so far as to embrace the
entire species; while, on the other hand, it may in some cases, and as
it were sporadically, extend beyond the species. In these latter cases
members of different species mutually assist one another, whether in the
way of what is called symbiosis, or in a variety of other ways which I
need not wait to mention. For the only point which I now desire to make
clear is, that all cases of mutual aid or co-operation, whether within
or beyond the limits of species, are cases which fall under the
explanatory sweep of the Darwinian theory[31].
[31] Prince Kropotkin in the _Nineteenth Century_ (Feb. 1888, Apr.
1891) has adduced a large and interesting body of facts, showing the
great prevalence of the principle of co-operation in organic
nature.
* * * * *
Another important point to notice is, that it constitutes no part of the
theory of natural selection to suppose that survival of the fittest must
invariably lead to _improvement_ of type, in the sense of superior
organization. On the contrary, if from change of habits or conditions of
life an organic type ceases to have any use for previously useful
organs, natural selection will not only allow these organs in successive
generations to deteriorate--by no longer placing any selective premium
upon their maintenance--but may even proceed to assist the agencies
engaged in their destruction. For, being now useless, they may become
even deleterious, by absorbing nutriment, causing weight, occupying
space, &c., without conferring any compensating benefit. Thus we can
understand why it is that parasites, for example, present the phenomena
of what is called _degeneration_, i. e. showing by their whole structure
that they have descended from a possibly very much higher type of
organization than that which they now exhibit. Having for innumerable
generations ceased to require their legs, their eyes, and so forth, all
such organs of high elaboration have either disappeared or become
vestigial, leaving the parasite as a more or less effete representative
of its ancestry.
These facts of degeneration, as we have previously seen, are of very
general occurrence, and it is evident that their importance in the field
of organic evolution as a whole has been very great. Moreover, it ought
to be particularly observed that, as just indicated, the facts may be
due either to a passive _cessation_ of selection, or to an active
_reversal_ of it. Or, more correctly, these facts are probably _always_
due to the cessation of selection, although in most cases where species
in a state of nature are concerned, the process of degeneration has been
both hastened and intensified by the super-added influence of the
reversal of selection. In the next volume I shall have occasion to recur
to this distinction, when it will be seen that it is one of no small
importance to the general theory of descent.
* * * * *
We may now proceed to consider certain misconceptions of the Darwinian
theory which are largely, not to say generally, prevalent among
supporters of the theory. These misconceptions, therefore, differ from
those which fall to be considered in the next chapter, i. e.
misconceptions which constitute grounds of objection to the theory.
* * * * *
Of all the errors connected with the theory of natural selection,
perhaps the one most frequently met with--especially among supporters of
the theory--is that of employing the theory to explain all cases of
phyletic modification (or inherited change of type) indiscriminately,
without waiting to consider whether in particular cases its application
is so much as logically possible. The term "natural selection" thus
becomes a magic word, or Sesame, at the utterance of which every closed
door is supposed to be immediately opened. Be it observed, I am not here
alluding to that merely blind faith in natural selection, which of late
years has begun dogmatically to force this principle as the sole cause
of organic evolution in every case where it is _logically possible_ that
the principle can have come into play. Such a blind faith, indeed, I
hold to be highly inimical, not only to the progress of biological
science, but even to the true interests of the natural selection theory
itself. As to this I shall have a good deal to say in the next volume.
Here, however, the point is, that the theory in question is often
invoked in cases where it is not even logically possible that it can
apply, and therefore in cases where its application betokens, not merely
an error of judgment or extravagance of dogmatism, but a fallacy of
reasoning in the nature of a logical contradiction. Almost any number of
examples might be given; but one will suffice to illustrate what is
meant. And I choose it from the writings of one of the authors of the
selection theory itself, in order to show how easy it is to be cheated
by this mere juggling with a phrase--for of course I do not doubt that a
moment's thought would have shown the writer the untenability of his
statement.
In his most recent work Mr. Wallace advances an interesting hypothesis
to the effect that differences of colour between allied species, which
are apparently too slight to serve any other purpose, may act as
"recognition marks," whereby the opposite sexes are enabled at once to
distinguish between members of their own and of closely resembling
species. Of course this hypothesis can only apply to the higher animals;
but the point here is that, supposing it to hold for them, Mr. Wallace
proceeds to argue thus:--Recognition marks "have in all probability been
acquired in the process of differentiation for the purpose of checking
the intercrossing of allied forms," because "one of the first needs of a
new species would be to keep separate from its nearest allies, and this
could be more readily done by some easily seen external mark[32]."
Now, it is clearly not so much as logically possible that these
recognition-marks (supposing them to be such) can have been acquired
by natural selection, "for the purpose of checking intercrossing of
allied forms." For the theory of natural selection, from its own
essential nature as a theory, is logically exclusive of the supposition
that survival of the fittest ever provides changes in anticipation of
future uses. Or, otherwise stated, it involves a contradiction of the
theory itself to say that the colour-changes in question were originated
by natural selection, in order to meet "one of the _first_ needs of a
_new_ species," or for the purpose of _subsequently_ preventing
intercrossing with allied forms. If it had been said that these
colour-differentiations were originated by some cause other than natural
selection (or, if by natural selection, still with regard to some
_previous_, instead of _prophetic_, "purpose"), and, when so "acquired,"
_then_ began to serve the "purpose" assigned, the argument would not
have involved the fallacy which we are now considering. But, as it
stands, the argument reverts to the teleology of pre-Darwinian days--or
the hypothesis of a "purpose" in the literal sense which sees the end
from the beginning, instead of a "purpose" in the metaphorical sense of
an adaptation that is evolved by the very modifications which subserve
it[33].
[32] _Darwinism_, pp. 218 and 227.
[33] Since the above was written Prof. Lloyd Morgan has published a
closely similar notice of the passage in question. "This language,"
he says, "seems to savour of teleology (that pitfall of the
evolutionist). The cart is put before the horse. The
recognition-marks were, I believe, not produced to prevent
intercrossing, but intercrossing has been prevented because of
preferential mating between individuals possessing special
recognition-marks. To miss this point is to miss an important
segregation-factor."--(_Animal Life and Intelligence_, p. 103.)
Again, on pp. 184-9, he furnishes an excellent discussion on the
whole subject of the fallacy alluded to in the text, and gives
illustrative quotations from other prominent Darwinians. I should
like to add that Darwin himself has nowhere fallen into this, or any
of the other fallacies, which are mentioned in the text.
* * * * *
Another very prevalent, and more deliberate, fallacy connected with the
theory of natural selection is, _that it follows deductively from the
theory itself_ that the principle of natural selection must be the sole
means of modification in all cases where modification is of an
_adaptive_ kind,--with the consequence that no other principle can ever
have been concerned in the production of structures or instincts which
are of any use to their possessors. Whether or not natural selection
actually has been the sole means of adaptive modification in the race,
as distinguished from the individual, is a question of biological
fact[34]; but it involves a grave error of reasoning to suppose that
this question can be answered deductively from the theory of natural
selection itself, as I shall show at some length in the next volume.
[34] Of course adaptive modifications produced in the individual
lifetime, and not _inherited_, do not concern the question at all.
In this and the following paragraphs, therefore, "adaptations,"
"adaptive modifications," &c., refer exclusively to such as are
hereditary, i. e. phyletic.
* * * * *
A still more extravagant, and a still more unaccountable fallacy is the
one which represents it as following deductively from the theory of
natural selection itself, that all _hereditary_ characters are
"necessarily" due to natural selection. In other words, not only all
adaptive, but likewise all non-adaptive hereditary characters, it is
said, _must_ be due to natural selection. For non-adaptive characters
are taken to be due to "correlation of growth," in connexion with some
of the adaptive ones--natural selection being thus the _indirect_ means
of producing the former _wherever_ they may occur, on account of its
being the _direct_ and the _only_ means of producing the latter. Thus it
is deduced from the theory of natural selection itself,--1st, that the
principle of natural selection is the only possible cause of adaptive
modification: 2nd, that non-adaptive modifications can only occur in the
race as correlated appendages to the adaptive: 3rd, that, consequently,
natural selection is the only possible cause of modification, whether
adaptive or non-adaptive. Here again, therefore, we must observe that
none of these sweeping generalizations can possibly be justified by
deductive reasoning from the theory of natural selection itself. Any
attempt at such deductive reasoning must necessarily end in circular
reasoning, as I shall likewise show in the second volume, where this
whole "question of utility" will be thoroughly dealt with.
* * * * *
Once more, there is an important oversight very generally committed by
the followers of Darwin. For even those who avoid the fallacies above
mentioned often fail to perceive, that natural selection can only begin
to operate if the _degree_ of adaptation is already given as
sufficiently high to count for something in the struggle for
_existence_. Any adaptations which fall below this level of importance
cannot possibly have been produced by survival of the fittest. Yet the
followers of Darwin habitually speak of adaptative characters, which _in
their own opinion_ are subservient merely to comfort or convenience, as
having been produced by such means. Clearly this is illogical; for it
belongs to the essence of Darwin's theory to suppose, that natural
selection can have no jurisdiction beyond the line where structures or
instincts already present a sufficient degree of adaptational value to
increase, in some measure, the expectation of life on the part of their
possessors. We cannot speak of adaptations as due to natural selection,
without thereby affirming that they present what I have elsewhere termed
a "selection value."
* * * * *
Lastly, as a mere matter of logical definition, it is well-nigh
self-evident that the theory of natural selection is a theory of the
origin, and cumulative development, of _adaptations_, whether these be
distinctive of species, or of genera, orders, families, classes, and
sub-kingdoms. It is only when the adaptations happen to be distinctive
of the first (or lowest) of these taxonomic divisions, that the theory
which accounts for _these_ adaptations accounts also for the forms which
present them,--i. e. becomes _also_ a theory of the origin of species.
This, however, is clearly but an accident of particular cases; and,
therefore, even in them the theory is _primarily_ a theory of
adaptations, while it is but secondarily a theory of the species which
present them. Or, otherwise stated, the theory is no more a theory of
the origin of species than it is of the origin of genera, families, and
the rest; while, on the other hand, it is _everywhere_ a theory of the
adaptive modifications whereby each of these taxonomic divisions has
been differentiated as such. Yet, sufficiently obvious as the accuracy
of this definition must appear to any one who dispassionately considers
it, several naturalists of high standing have denounced it in violent
terms. I shall therefore have to recur to the subject at somewhat
greater length hereafter. At present it is enough merely to mention the
matter, as furnishing another and a curious illustration of the not
infrequent weakness of logical perception on the part of minds well
gifted with the faculty of observation. It may be added, however, that
the definition in question is in no way hostile to the one which is
virtually given by Darwin in the title of his great work. _The Origin of
Species by means of Natural Selection_ is beyond doubt the best title
that could have been given, because at the time when the work was
published the _fact_, no less than the _method_, of organic evolution
had to be established; and hence the most important thing to be done at
that time was to prove the transmutation of species. But now that this
has been done to the satisfaction of naturalists in general, it is as I
have said, curious to find some of them denouncing a wider definition of
the principle of natural selection, merely because the narrower (or
included) definition is invested with the charm of verbal
associations[35].
[35] The question as to whether natural selection has been the only
principle concerned in the origination of species, is quite distinct
from that as to the accuracy of the above definition.
* * * * *
So much for fallacies and misconceptions touching Darwin's theory, which
are but too frequently met with in the writings of its supporters. We
must now pass on to mention some of the still greater fallacies and
misconceptions which are prevalent in the writings of its opponents.
And, in order to do this thoroughly, I shall begin by devoting the
remainder of the present chapter to a consideration of the antecedent
standing of the two theories of natural selection and supernatural
design. This having been done, in the succeeding chapters I shall deal
with the evidences for, and the objections against, the former theory.
* * * * *
Beginning, then, with the antecedent standing of these alternative
theories, the first thing to be noticed is, that they are both concerned
with the same subject-matter, which it is their common object to
explain. Moreover, this subject-matter is clearly and sharply divisible
into two great classes of facts in organic nature--namely, those of
Adaptation and those of Beauty. Darwin's theory of descent explains the
former by his doctrine of natural selection, and the latter by his
doctrine of sexual selection. In the first instance, therefore, I shall
have to deal only with the facts of adaptation, leaving for subsequent
consideration the facts of beauty.
Innumerable cases of the adaptation of organisms to their surroundings
being the facts which now stand before us to be explained either by
natural selection or by supernatural intention, we may first consider a
statement which is frequently met with--namely, that even if all such
cases of adaptation were proved to be fully explicable by the theory of
descent, this would constitute no disproof of the theory of design: all
the cases of adaptation, it is argued, might still be due to design,
even though they admit of being hypothetically accounted for by the
theory of descent. I have heard an eminent Professor tell his class that
the many instances of mechanical adaptation discovered and described by
Darwin as occurring in orchids, seemed to him to furnish better proof of
supernatural contrivance than of natural causes; and another eminent
Professor has informed me that, although he had read the _Origin of
Species_ with care, he could see in it no evidence of natural selection
which might not equally well have been adduced in favour of intelligent
design. But here we meet with a radical misconception of the whole
logical attitude of science. For, be it observed, this exception _in
limine_ to the evidence which we are about to consider does not question
that natural selection _may_ be able to do all that Darwin ascribes to
it. The objection is urged against his interpretation of the facts
merely on the ground that these facts might _equally well_ be ascribed
to intelligent design. And so undoubtedly they might, if we were all
simple enough to adopt a supernatural explanation whenever a natural one
is found sufficient to account for the facts. Once admit the irrational
principle that we may assume the operation of higher causes where the
operation of lower ones is sufficient to explain the observed phenomena,
and all our science and all our philosophy are scattered to the winds.
For the law of logic which Sir William Hamilton called the law of
parsimony--or the law which forbids us to assume the operation of higher
causes when lower ones are found sufficient to explain the observed
effects--this law constitutes the only barrier between science and
superstition. It is always possible to give a hypothetical explanation
of any phenomenon whatsoever, by referring it immediately to the
intelligence of some supernatural agent; so that the only difference
between the logic of science and the logic of superstition consists in
science recognising a validity in the law of parsimony which
superstition disregards. Therefore one can have no hesitation in saying
that this way of looking at the evidence in favour of natural selection
is not a scientific or a reasonable way of looking at it, but a purely
superstitious way. Let us take, as an illustration, a perfectly parallel
case. When Kepler was unable to explain by any known causes the paths
described by the planets, he resorted to a supernatural explanation, and
supposed that every planet was guided in its movements by some presiding
angel. But when Newton supplied a beautifully simple physical
explanation, all persons with a scientific habit of mind at once
abandoned the metaphysical one. Now, to be consistent, the
above-mentioned Professors, and all who think with them, ought still to
adhere to Kepler's hypothesis in preference to Newton's explanation;
for, excepting the law of parsimony, there is certainly no other
logical objection to the statement, that the movements of the planets
afford as good evidence of the influence of guiding angels as they do of
the influence of gravitation.
So much, then, for the illogical position that, granting the evidence in
favour of natural descent and supernatural design to be equal and
parallel, we should hesitate in our choice between the two theories.
But, of course, if the evidence is supposed _not_ to be equal and
parallel--i. e. if it is supposed that the theory of natural selection
is not so good a theory whereby to explain the facts of adaptation as is
that of supernatural design,--then the objection is no longer the one
which we are considering. It is quite another objection, and one which
is not _prima facie_ absurd. Therefore let us state clearly the distinct
question which thus arises.
Innumerable cases of adaptation of organisms to their environments are
the observed facts for which an explanation is required. To supply this
explanation, two, and only two, hypotheses are in the field. Of these
two hypotheses one is intelligent design manifested directly in special
creation; the other is natural causation operating through countless
ages of the past. Now, the adaptations in question involve an
innumerable multitude of special mechanisms, in most cases even within
the limits of any one given species; but when we consider the sum of all
these mechanisms presented by organic nature as a whole, the mind must
indeed be dull which does not feel astounded. For, be it further
observed, these mechanical contrivances[36] are, for the most part, no
merely simple arrangements, which might reasonably be supposed due,
like the phenomena of crystallization, to comparatively simple physical
causes. On the contrary, they everywhere and habitually exhibit so
deep-laid, so intricate, and often so remote an adaptation of means to
ends, that no machinery of human contrivance can properly be said to
equal their perfection from a mechanical point of view. Therefore,
without question, the hypothesis which first of all they suggest--or
suggest most readily--is the hypothesis of design. And this hypothesis
becomes virtually the only hypothesis possible, if it be assumed--as it
generally was assumed by natural theologians of the past,--that all
species of plants and animals were introduced into the world _suddenly_.
For it is quite inconceivable that any known cause, other than
intelligent design, could be competent to turn out instantaneously any
one of these intricate pieces of machinery, already adapted to the
performance of its special function. But, on the other hand, if there is
any evidence to show that one species becomes slowly transformed into
another--or that one set of adaptations becomes slowly changed into
another set as changing circumstances require,--then it becomes quite
possible to imagine that a strictly natural causation may have had
something to do with the matter. And this suggestion becomes greatly
more probable when we discover, from geological evidence and
embryological research, that in the history both of races and of
individuals the various mechanisms in question have themselves had a
history--beginning in the forms of most uniformity and simplicity,
gradually advancing to forms more varied and complex, nowhere exhibiting
any interruptions in their upward progress, until the world of organic
machinery as we now have it is seen to have been but the last phase of a
long and gradual growth, the ultimate roots of which are to be found in
the soil of undifferentiated protoplasm.
[36] It is often objected to Darwin's terminology, that it embraces
such words as "contrivance," "purpose," &c., which are strictly
applicable only to the processes or the products of thought. But
when it is understood that they are used in a neutral or
metaphorical sense, I cannot see that any harm arises from their
use.
Lastly, when there is supplied to us the suggestion of natural selection
as a cause presumably adequate to account for this continuous growth in
the number, the intricacy, and the perfection of such mechanisms, it is
only the most unphilosophical mind that can refuse to pause as between
the older hypothesis of design and the newer hypothesis of descent.
Thus it is clear that the _a priori_ standing of the rival hypotheses of
naturalism and supernaturalism in the case of all these pieces of
organic machinery, is profoundly affected by the question whether they
came into existence suddenly, or whether they did so gradually. For, if
they all came into existence suddenly, the fact would constitute
well-nigh positive proof in favour of supernaturalism, or creation by
design; whereas, if they all came into existence gradually, this fact
would in itself constitute presumptive evidence in favour of naturalism,
or of development by natural causes. And, as shown in the previous
chapters, the proof that all species of plants and animals came into
existence gradually--or the proof of evolution as a fact--is simply
overwhelming.
From a still more general point of view I may state the case in another
way, by borrowing and somewhat expanding an illustration which, I
believe, was first used by Professor Huxley. If, when the tide is out,
we see lying upon the shore a long line of detached sea-weed, marking
the level which is reached by full tide, we should be free to conclude
that the separation of the sea-weed from the sand and the stones was due
to the intelligent work of some one who intended to collect the sea-weed
for manure, or for any other purpose. But, on the other hand, we might
explain the fact by a purely physical cause--namely, the separation by
the sea-waves of the sea-weed from the sand and stones, in virtue of its
lower specific gravity. Now, thus far the fact would be explained
equally well by either hypothesis; and this fact would be the fact of
_selection_. But whether we yielded our assent to the one explanation or
to the other would depend upon a due consideration of all collateral
circumstances. The sea-weed might not be of a kind that is of any use to
man; there might be too great a quantity of it to admit of our supposing
that it had been collected by man; the fact that it was all deposited on
the high-water-mark would in itself be highly suggestive of the agency
of the sea; and so forth. Thus, in such a case any reasonable observer
would decide in favour of the physical explanation, or against the
teleological one.
Now the question whether organic evolution has been caused by physical
agencies or by intelligent design is in precisely the same predicament.
There can be no logical doubt that, theoretically at all events, the
physical agencies which the present chapter is concerned with, and which
are conveniently summed up in the term natural selection, are as
competent to produce these so-called mechanical contrivances, and the
other cases of adaptation which are to be met with in organic nature,
as intelligent design could be. Hence, our choice as between these two
hypotheses must be governed by a study of all collateral circumstances;
that is to say, by a study of the evidences in favour of the physical
explanation. To this study, therefore, we shall now address ourselves,
in the course of the following chapters.
CHAPTER VIII.
EVIDENCES OF THE THEORY OF NATURAL SELECTION.
I will now proceed to state the main arguments in favour of the theory
of natural selection, and then, in the following chapter, the main
objections which have been urged against it.
In my opinion, the main arguments in favour of the theory are three in
number.
First, it is a matter of observation that the struggle for existence in
nature does lead to the extermination of forms less fitted for the
struggle, and thus makes room for forms more fitted. This general fact
may be best observed in cases where an exotic species proves itself
better fitted to inhabit a new country than is some endemic species
which it exterminates. In Great Britain, for example, the so-called
common rat is a comparatively recent importation from Norway, and it has
so completely supplanted the original British rat, that it is now
extremely difficult to procure a single specimen of the latter: the
native black rat has been all but exterminated by the foreign brown rat.
The same thing is constantly found in the case of imported species of
plants. I have seen the river at Cambridge so choked with the inordinate
propagation of a species of water-weed which had been introduced from
America, that considerable expense had to be incurred in order to clear
the river for traffic. In New Zealand the same thing has happened with
the European water-cress, and in Australia with the common rabbit. So it
is doubtless true, as one of the natives is said to have philosophically
remarked, "the white man's rat has driven away our rat, the European fly
drives away our fly, his clover kills our grass, and so will the Maoris
disappear before the white man himself." Innumerable other cases to the
same effect might be quoted; and they all go to establish the fact that
forms less fitted to survive succumb in their competition with forms
better fitted.
* * * * *
Secondly, there is a general consideration of the largest possible
significance in the present connexion--namely, that among all the
millions of structures and instincts which are so invariably, and for
the most part so wonderfully, adapted to the needs of the species
presenting them, we cannot find a single instance, either in the
vegetable or animal kingdom, of a structure or an instinct which is
developed for the exclusive benefit of another species. Now this great
and general fact is to my mind a fact of the most enormous, not to say
overwhelming, significance. The theory of natural selection has now been
before the world for more than thirty years, and during that time it had
stood a fire of criticism such as was never encountered by any
scientific theory before. From the first Darwin invited this criticism
to adduce any single instance, either in the vegetable or animal
kingdom, of a structure or an instinct which should unquestionably be
proved to be of exclusive use to any species other than the one
presenting it. He even went so far as to say that if any one such
instance could be shown he would surrender his whole theory on the
strength of it--so assured had he become, by his own prolonged
researches, that natural selection was the true agent in the production
of adaptive structures, and, as such, could never have permitted such a
structure to occur in one species for the benefit of another. Now, as
this invitation has been before the world for so many years, and has not
yet been answered by any naturalist, we may by this time be pretty
confident that it never will be answered. How tremendous, then, is the
significance of this fact in its testimony to Darwin's theory! The
number of animal and vegetable species, both living and extinct, is to
be reckoned by millions, and every one of these species presents on an
average hundreds of adaptive structures,--at least one of which in many,
possibly in most, if not actually in all cases, is peculiar to the
species that presents it. In other words, there are millions of adaptive
structures (not to speak of instincts) which are peculiar to the species
presenting them, and also many more which are the common property of
allied species: yet, notwithstanding this inconceivable profusion of
adaptive structures in organic nature, there is no single instance that
has been pointed out of the occurrence of such a structure save for the
benefit of the species that presents it. Therefore, I say that this
immensely large and general fact speaks with literally immeasurable
force in favour of natural selection, as at all events one of the main
causes of organic evolution. For the fact is precisely what we should
expect if this theory is true, while upon no other theory can its
universality and invariability be rendered intelligible. On the
beneficent design theory, for instance, it is inexplicable that no
species should ever be found to present a structure or an instinct
having primary reference to the welfare of another species, when, _ex
hypothesi_, such an endless amount of thought has been displayed in the
creation of structures and instincts having primary reference to the
species which present them. For how magnificent a display of divine
beneficence would organic nature have afforded, if all--or even
some--species had been so inter-related as to have ministered to each
others wants. Organic species might then have been likened to a
countless multitude of voices, all singing in one great harmonious
psalm. But, as it is, we see absolutely no vestige of such
co-ordination: every species is for itself, and for itself alone--an
outcome of the always and everywhere fiercely raging struggle for life.
In order that the force of this argument may not be misapprehended, it
is necessary to bear in mind that it is in no way affected by cases
where a structure or an instinct is of primary benefit to its possessor,
and then becomes of secondary benefit to some other species on account
of the latter being able in some way or another to utilise its action.
Of course organic nature is full of cases of this kind; but they only go
to show the readiness which all species display to utilise for
themselves everything that can be turned to good account in their own
environments, and so, among other things, the structures and instincts
of other animals. For instance, it would be no answer to Darwin's
challenge if any one were to point to a hermit-crab inhabiting the
cast-off shell of a mollusk; because the shell was primarily of use to
the mollusk itself, and, so far as the mollusk is concerned, the fact of
its shell being afterwards of a secondary use to the crab is quite
immaterial. What Darwin's challenge requires is, that some structure or
instinct should be shown which is not merely of such secondary or
accidental benefit to another species, but clearly adapted to the needs
of that other species in the first instance--such, for example, as would
be the case if the tail of a rattle-snake were of no use to its
possessor, while serving to warn other animals of the proximity of a
dangerous creature; or, in the case of instincts, if it were true that a
pilot-fish accompanies a shark for the purpose of helping the shark to
discover food. Both these instances have been alleged; but both have
been shown untenable. And so it has proved of all the other cases which
thus far have been put forward.
Perhaps the most remarkable of all the allegations which ever have been
put forward in this connexion are those that were current with regard to
instincts before the publication of Darwin's work. These allegations are
the most remarkable, because they serve to show, in a degree which I do
not believe could be shown anywhere else, the warping power of
preconceived ideas. A short time ago I happened to come across the 8th
edition of the _Encyclopaedia Britannica_, and turned up the article on
"Instinct" there, in order to see what amount of change had been wrought
with regard to our views on this subject by the work of Darwin--the 8th
edition of the _Encyclopaedia Britannica_ having been published shortly
before _The Origin of Species by means of Natural Selection_. I cannot
wait to give any lengthy quotations from this representative exponent of
scientific opinion upon the subject at that time; but its general drift
may be appreciated if I transcribe merely the short concluding
paragraph, wherein he sums up his general results. Here he says:--
It thus only remains for us to regard instinct as a mental faculty,
_sui generis_, the gift of God to the lower animals, that man in
his own person, and by them, might be relieved from the meanest
drudgery of nature.
Now, here we have the most extraordinary illustration that is imaginable
of the obscuring influence of a preconceived idea. Because he started
with the belief that instincts _must_ have been implanted in animals for
the benefit of man, this writer, even when writing a purely scientific
essay, was completely blinded to the largest, the most obvious, and the
most important of the facts which the phenomena of instinct display.
For, as a matter of fact, among all the many thousands of instincts
which are known to occur in animals, there is no single one that can be
pointed to as having any special reference to man; while, on the other
hand, it is equally impossible to point to one which does not refer to
the welfare of the animal presenting it. Indeed, when the point is
suggested, it seems to me surprising how few in number are the instincts
of animals which have proved to be so much as of secondary or accidental
benefit to man, in the same way as skins, furs, and a whole host of
other animal products are thus of secondary use to him. Therefore, this
writer not only failed to perceive the most obvious truth that every
instinct, without any single exception, has reference to the animal
which presents it; but he also conceived a purely fictitious inversion
of this truth, and wrote an essay to prove a statement which all the
instincts in the animal kingdom unite in contradicting.
This example will serve to show, in a striking manner, not only the
distance that we have travelled in our interpretation of organic nature
between two successive editions of the _Encyclopaedia Britannica_, but
also the amount of verification which this fact furnishes to the theory
of natural selection. For, inasmuch as it belongs to the very essence of
this theory that all adaptive characters (whether instinctive or
structural) must have reference to their own possessors, we find
overpowering verification furnished to the theory by the fact now before
us--namely, that immediately prior to the enunciation of this theory,
the truth that all adaptive characters have reference only to the
species which present them was not perceived. In other words, it was the
testing of this theory by the facts of nature that _revealed_ to
naturalists the general law which the theory, as it were, predicted--the
general law that all adaptive characters have primary reference to the
species which present them. And when we remember that this is a kind of
verification which is furnished by millions of separate cases, the whole
mass of it taken together is, as I have before said, overwhelming.
It is somewhat remarkable that the enormous importance of this argument
in favour of natural selection as a prime factor of organic evolution
has not received the attention which it deserves. Even Darwin himself,
with his characteristic reserve, has not presented its incalculable
significance; nor do I know any of his followers who have made any
approach to an adequate use of it in their advocacy of his views. In
preparing the present chapter, therefore, I have been particularly
careful not to pitch too high my own estimate of its evidential value.
That is to say, I have considered, both in the domain of structures and
of instincts, what instances admit of being possibly adduced _per
contra_, or as standing outside the general law that adaptive structures
and instincts are of primary use only to their possessors. In the result
I can only think of two such instances. These, therefore, I will now
dispose of.
The first was pointed out, and has been fully discussed, by Darwin
himself. Certain species of ants are fond of a sweet fluid that is
secreted by aphides, and they even keep the aphides as we keep cows for
the purpose of profiting by their "milk." Now the point is, that the use
of this sweet secretion to the aphis itself has not yet been made out.
Of course, if it is of no use to the aphis, it would furnish a case
which completely meets Darwin's own challenge. But, even if this
supposition did not stand out of analogy with all the other facts of
organic nature, most of us would probably deem it prudent to hold that
the secretion must primarily be of some use to the aphis itself,
although the matter has not been sufficiently investigated to inform us
of what this use is. For, in any case, the secretion is not of any vital
importance to the ants which feed upon it: and I think but few impartial
minds would go so far to save an hypothesis as to maintain, that the
Divinity had imposed this drain upon the internal resources of one
species of insect for the sole purpose of supplying a luxury to
another. On the whole, it seems most probable that the fluid is of the
nature of an excretion, serving to carry off waste products. Such, at
all events, was the opinion at which Darwin himself arrived, as a result
of observing the facts anew, and in relation to his theory.
* * * * *
The other instance to which I have alluded as seeming at first sight
likely to answer Darwin's challenge is the formation of vegetable galls.
The great number and variety of galls agree in presenting a more or less
elaborate structure, which is not only foreign to any of the uses of
plant-life, but singularly and specially adapted to those of the
insect-life which they shelter. Yet they are produced by a growth of the
plant itself, when suitably stimulated by the insects' inoculation--or,
according to recent observations, by emanations from the bodies of the
larvae which develop from the eggs deposited in the plant by the insect.
Now, without question, this is a most remarkable fact; and if there were
many more of the like kind to be met with in organic nature, we might
seriously consider whether the formation of galls should not be held to
make against the ubiquitous agency of natural selection. But inasmuch as
the formation of galls stands out as an exception to the otherwise
universal rule of every species for itself, and for itself alone, we are
justified in regarding this one apparent exception with extreme
suspicion. Indeed, I think we are justified in regarding the peculiar
pathological effect produced in the plant by the secretions of the
insect as having been in the first instance accidentally beneficial to
the insects. Thus, if any other effect than that of a growing tumour had
been produced in the first instance, or if the needs of the insect
progeny had not been such as to have derived profit from being enclosed
in such a tumour, then, of course, the inoculating instinct of these
animals could not have been developed by natural selection. But, given
these two conditions, and it appears to me there is nothing very much
more remarkable about an accidental correlation between the effects of a
parasitic larva on a plant and the needs of that parasite, than there is
between the similarly accidental correlation between a hydated parasite
and the nutrition furnished to it by the tissues of a warm-blooded
animal. Doubtless the case of galls is somewhat more remarkable,
inasmuch as the morbid growth of the plant has more concern in the
correlation--being, in many instances, a more specialized structure on
the part of a host than occurs anywhere else, either in the animal or
vegetable world. But here I may suggest that although natural selection
cannot have acted upon the plant directly, so as to have produced galls
ever better and better adapted to the needs of the insect, it may have
so acted upon the plants indirectly _though the insects_. For it may
very well have been that natural selection would ever tend to preserve
those individual insects, the quality of whose emanations tended to
produce the form of galls best suited to nourish the insect progeny; and
thus the character of these pathological growths may have become ever
better and better adapted to the needs of the insects. Lastly, looking
to the enormous number of relations and inter-relations between all
organic species, it is scarcely to be wondered at that even so
extraordinary an instance of correlation as this should have arisen thus
by accident, and then have been perfected by such an _indirect_ agency
of natural selection as is here suggested[37].
[37] Note B.
* * * * *
The third general class of facts which tell so immensely in favour of
natural selection as an important cause of organic evolution, are those
of domestication. The art of the horticulturist, the fancier, the
cattle-breeder, &c., consists in producing greater and greater
deviations from a given wild type of plant or animal, in any particular
direction that may be desired for purposes either of use or of beauty.
Cultivated cereals, fruits, and flowers are known to have been all
derived from wild species; and, of course, the same applies to all our
domesticated varieties of animals. Yet if we compare a cabbage rose with
a wild rose, a golden pippin apple with a crab, a toy terrier with any
species of wild dog, not to mention any number of other instances, there
can be no question that, if such differences had appeared in nature, the
organisms presenting them would have been entitled to rank as distinct
species--or even, in many cases, as distinct genera. Yet we know, as a
matter of fact, that all these differences have been produced by a
process of artificial selection, or pairing, which has been continuously
practised by horticulturists and breeders through a number of
generations. It is the business of these men to note the individual
organisms which show most variation in the directions required, and then
to propagate from these individuals, in order that the progeny shall
inherit the qualities desired. The results thus become cumulative from
generation to generation, until we now have an astonishing manifestation
of useful qualities on the one hand, and of beautiful qualities on the
other, according as the organisms have been thus bred for purposes of
use or for those of beauty.
Now it is immediately obvious that in these cases the process of
artificial selection is precisely analogous to that of natural selection
(and of sexual selection which will be considered later on), in all
respects save one: the utility or the beauty which it is the aim of
artificial selection continually to enhance, is utility or beauty in
relation to the requirements or to the tastes of man; whereas the
utility or the beauty which is produced by natural selection and sexual
selection has reference only to the requirements or the tastes of the
organisms themselves. But, with the exception of this one point of
difference, the processes and the products are identical in kind.
Persevering selection by man is thus proved to be capable of creating
what are virtually new specific types, and this in any required
direction. Hence, when we remember how severe is the struggle for
existence in nature, it becomes impossible to doubt that selection by
nature is able to do at least as much as artificial selection in the way
of thus creating new types out of old ones. Artificial selection,
indeed, notwithstanding the many and marvellous results which it has
accomplished, can only be regarded as but a feeble imitation of natural
selection, which must act with so much greater vigilance and through
such immensely greater periods of time. In a word, the proved
capabilities of artificial selection furnish, in its best conceivable
form, what is called an argument _a fortiori_ in favour of natural
selection.
Or, to put it in another way, it may be said that for thousands of years
mankind has been engaged in making a gigantic experiment to test, as it
were by anticipation, the theory of natural selection. For, although
this prolonged experiment has been carried on without any such intention
on the part of the experimenters, it is none the less an experiment in
the sense that its results now furnish an overwhelming verification of
Mr. Darwin's theory. That is to say, they furnish overwhelming proof of
the efficacy of the selective principle in the modification of organic
types, when once this principle is brought steadily and continuously to
bear upon a sufficiently long series of generations.
In order to furnish ocular evidence of the value of this line of
verification, I have had the following series of drawings prepared.
Another and equally striking series might be made of the products of
artificial selection in the case of plants; but it seems to me that the
case of animals is more than sufficient for the purpose just stated.
Perhaps it is desirable to add that considerable care has been bestowed
upon the execution of these portraits; and that in every case the latter
have been taken from the most typical specimens of the artificial
variety depicted. Those of them which have not been drawn directly from
life are taken from the most authoritative sources; and, before being
submitted to the engraver, they were all examined by the best judges in
each department. In none of the groups, however, have I aimed at an
exhaustive representation of all the varieties: I have merely introduced
representatives of as many as the page would in each case accommodate.
[Illustration: FIG. 91.--Pigeons. Drawn from life (prize
specimens).]
[Illustration: FIG. 92.--Pigeons, continued. Drawn from life (prize
specimens).]
[Illustration: FIG. 93.--Fowls. Drawn from life (prize specimens).]
[Illustration: FIG. 94.--Fowls, continued. Drawn from life (prize
specimens).]
[Illustration: FIG. 95.--Pair of Japanese Fowls, long-tailed breed.
Drawn from stuffed specimens in the British Museum.]
[Illustration: FIG. 96.--Canaries. Drawn from life (prize
specimens).]
[Illustration: FIG. 97.--Sebastopol, or Frizzled Goose. Drawn from a
photograph.]
[Illustration: FIG. 98.--The Dingo, or wild dog of Australia, 1/10
nat. size. Drawn from life (_Zoological Gardens_).]
[Illustration: FIG. 99.--Dogs. Drawn from life (prize specimens).]
[Illustration: FIG. 100.--Dogs, continued. Drawn from life (prize
specimens).]
[Illustration: FIG. 101.--The Hairless Dog of Japan, 1/10 nat. size.
Drawn from a photograph kindly lent for the purpose by the
proprietor.]
[Illustration: FIG. 102.--The skull of a Bull-dog compared with that
of a Deerhound. Drawn from nature.]
[Illustration: FIG. 103. Rabbits. Drawn from life (prize
specimens).]
[Illustration: FIG. 104.--Horses. Drawn from life (prize
specimens).]
[Illustration: FIG. 105.--Sheep. The illustrations are confined to
British breeds. Drawn from life (prize specimens).]
[Illustration: FIG. 106.--Cattle. The illustrations are confined to
British breeds. Drawn from life (prize specimens).]
[Illustration: FIG. 107.--Wild Boar contrasted with a modern
Domesticated Pig. Drawn from life (_Zoological Gardens_, and prize
specimen).]
The exigencies of space have prevented, in some of the groups, strict
adherence to a uniform scale--with the result that contrasts between
different breeds in respect of size are not adequately rendered. This
remark applies especially to the dogs; for although the artist has
endeavoured to draw them in perspective, unless the distance between
those in the foreground and those in the background is understood to be
more considerable than it appears, an inadequate idea is given of the
relative differences of size. The most instructive of the groups, I
think, is that of the Canaries; because the many and great changes in
different directions must in this case have been produced by artificial
selection in so comparatively short a time--the first mention of this
bird that I can find being by Gesner, in the sixteenth century.
* * * * *
Now, it is surely unquestionable that in these typical proofs of the
efficacy of artificial selection in the modification of specific types,
we have the strongest conceivable testimony to the power of natural
selection in the same direction. For it thus appears that wherever
mankind has had occasion to operate by selection for a sufficiently long
time--that is to say, on whatever species of plant or animal he chooses
thus to operate for the purpose of modifying the type in any required
direction,--the results are always more or less the same: he finds that
all specific types lend themselves to continuous deflection in any
particulars of structure, colour, &c., that he may desire to modify.
Nevertheless, to this parallel between the known effects of artificial
selection, and the inferred effects of natural selection, two objections
have been urged. The first is, that in the case of artificial selection
the selecting agent is a voluntary intelligence, while in the case of
natural selection the selecting agent is Nature herself; and whether or
not there is any counterpart of man's voluntary intelligence in nature
is a question with which Darwinism has nothing to do. Therefore, it is
alleged, the analogy between natural selection and artificial selection
fails _ab initio_, or at the fountain-head of the causes which are taken
by the analogy to be respectively involved.
The second objection to the analogy is, that the products of artificial
selection, closely as they may resemble natural species in all other
respects, nevertheless present one conspicuous and highly important
point of difference: they rarely, if ever, present the physiological
character of mutual infertility, which is a character of extremely
general occurrence in the case of natural species, even when these are
most nearly allied.
I will deal with these two objections in the next chapter, where I shall
be concerned with the meeting of all the objections which have ever been
urged against the theory of natural selection. Meanwhile I am engaged
only in presenting the general arguments which support the theory, and
therefore mention these objections to one of them merely _en passant_.
And I do so in order to pledge myself effectually to dispose of them
later on, so that for the purposes of my present argument both these
objections may be provisionally regarded as non-existent; which means,
in other words, that we may provisionally regard the analogy between
artificial selection and natural selection as everywhere logically
intact.
* * * * *
To sum up, then, the results of the foregoing exposition thus far, what
I hold to be the three principal, or most general, arguments in favour
of the theory of natural selection, are as follows.
First, there is the _a priori_ consideration that, if on independent
grounds we believe in the theory of evolution at all, it becomes obvious
that natural selection _must_ have had _some_ part in the process. For
no one can deny the potent facts of heredity, variability, the struggle
for existence, and survival of the fittest. But to admit these facts is
to admit natural selection as a principle which must be, at any rate,
one of the factors of organic evolution, supposing such evolution to
have taken place. Next, when we turn from these _a priori_
considerations, which thus show that natural selection _must_ have been
concerned to some extent in the process of evolution, we find in organic
nature evidence _a posteriori_ of the extent to which this principle
_has_ been thus concerned. For we find that among all the countless
millions of adaptive structures which are to be met with in organic
nature, it is an invariable rule that they exist in relation to the
needs of the particular species which present them: they never have any
primary reference to the needs of other species. And as this
extraordinarily large and general fact is exactly what the theory of
natural selection would expect, the theory is verified by the fact in an
extraordinarily cogent manner. In other words, the fact goes to prove
that in _all_ cases where adaptive structures or instincts are
concerned, natural selection must have been either the sole cause at
work, or, at the least, an influence controlling the operation of all
other causes.
Lastly, an actually experimental verification of the theory has been
furnished on a gigantic scale by the operations of breeders, fanciers,
and horticulturists. For these men, by their process of selective
accumulation, have empirically proved what immense changes of type may
thus be brought about; and so have verified by anticipation, and in a
most striking manner, the theory of natural selection--which, as now so
fully explained, is nothing more than a theory of cumulative
modifications by means of selective breeding.
So much, then, by way of generalities. But perhaps the proof of natural
selection as an agency of the first importance in the transmutation of
species may be best brought home to us by considering a few of its
applications in detail. I will therefore devote the rest of the present
chapter to considering a few cases of this kind.
There are so many large fields from which such special illustrations may
be supplied, that it is difficult to decide which of them to draw upon.
For instance, the innumerable, always interesting, and often astonishing
adaptations on the part of flowers to the fertilising agency of insects,
has alone given rise to an extensive literature since the time when
Darwin himself was led to investigate the subject by the guidance of his
own theory. The same may be said of the structures and movements of
climbing plants, and in short, of all the other departments of natural
history where the theory of natural selection has led to the study of
the phenomena of adaptation. For in all these cases the theory of
natural selection, which first led to their discovery, still remains the
only scientific theory by which they can be explained. But among all the
possible fields from which evidences of this kind may be drawn, I think
the best is that which may be generically termed defensive colouring. To
this field, therefore, I will restrict myself. But, even so, the cases
to be mentioned are but mere samples taken from different divisions of
this field; and therefore it must be understood at the outset that they
could easily be multiplied a hundred-fold.
_Protective Colouring._
A vast number of animals are rendered more or less inconspicuous by
resembling the colours of the surfaces on which they habitually rest.
Such, for example, are grouse, partridges, rabbits, &c. Moreover, there
are many cases in which, if the needs of the creature be such that it
must habitually frequent surfaces of different colours, it has acquired
the power of changing its colour accordingly--e. g. cuttle-fish,
flat-fish, frogs, chameleons, &c. The physiological mechanism whereby
these adaptive changes of colour are produced differs in different
animals; but it is needless for our purposes to go into this part of the
subject. Again, there are yet other cases where protective colouring
which is admirably suited to conceal an animal through one part of the
year, would become highly conspicuous during another part of it--namely,
when the ground is covered with snow. Accordingly, in these cases the
animals change their colour in the winter months to a snowy white:
witness stoats, mountain hares, ptarmigan, &c. (Fig. 108.)
[Illustration: FIG. 108.--Seasonal changes of colour in Ptarmigan
(_Lagopus mutus_). Drawn from stuffed specimens in the British
Museum, 1/6 nat. size, with appropriate surroundings supplied.]
Now, it is sufficiently obvious that in all these classes of cases the
concealment from enemies or prey which is thus secured is of advantage
to the animals concerned; and, therefore, that in the theory of natural
selection we have a satisfactory theory whereby to explain it. And this
cannot be said of any other theory of adaptive mechanisms in nature that
has ever been propounded. The so-called Lamarckian theory, for instance,
cannot be brought to bear upon the facts at all; and on the theory of
special creation it is unintelligible why the phenomena of protective
colouring should be of such general occurrence. For, in as far as
protective colouring is of advantage to the species which present it, it
is of corresponding disadvantage to those other species against the
predatory nature of which it acts as a defence. And, of course, the
same applies to yet other species, if they serve as prey. Moreover,
the more minutely this subject is investigated in all its details,
the more exactly is it found to harmonise with the naturalistic
interpretation[38].
[38] Were it not that some of Darwin's critics have overlooked the
very point wherein the great value of protective colouring as
evidence of natural selection consists, it would be needless to
observe that it does so in the _minuteness_ of the protective
resemblance which in so many cases is presented. Of course where the
resemblance is only very general, the phenomena might be ascribed to
mere coincidence, of which the instincts of the animal have taken
advantage. But in the measure that the resemblance becomes minutely
detailed, the supposition of mere coincidence is excluded, and the
agency of some specially adaptive cause demonstrated. Again, it is
almost needless to say, no real difficulty is presented (as has been
alleged) by the cases above quoted of seasonal imitations, on the
ground that natural selection could not act alternately on the same
individual. Natural selection is not supposed to act alternately on
the same individual. It is supposed to act always in the same
manner, and if, as in the case of a regularly recurring change in
the colours of the environment, correspondingly recurrent changes
are required to appear in the colours of the animals, natural
selection sets its premium upon those individuals the constitutions
of which best lend themselves to seasonal changes of the needful
kind--probably under the influence of stimuli supplied by the
changes of external conditions (temperature, moisture, &c.).
In the first place, we always find a complete correspondence between
imitative colouring and instinctive endowment. If a caterpillar exactly
resembles the colour of a twig, it also presents the instinct of
habitually reposing in the attitude which makes it most resemble a
twig--standing out from the branch on which it rests at the same angle
as is presented by the real twigs of the tree on which it lives.
Here, again, is a bird protectively coloured so as to resemble stones
upon the rough ground where it habitually lives; and the drawing shows
the attitude in which the bird instinctively reposes, so as still
further to increase its resemblance to a stone. (Fig. 109.)
[Illustration: FIG. 109.--_Oedicnemus crepitans_, showing the
instinctive attitude of concealment. Drawn from a stuffed specimen
in the British Museum, 1/6 nat. size, with appropriate surroundings
supplied.
To take only one other instance, hares and rabbits, like grouse and
partridges--or like the plover just alluded to,--instinctively crouch
upon those surfaces the colours of which they resemble; and I have often
remarked that if, on account of any individual peculiarity of
coloration, the animal is not able thus to secure concealment, it
nevertheless exhibits the instinct of crouching which is of benefit to
all its kind, although, from the accident of its own abnormal colouring,
this instinct is then actually detrimental to the animal itself. For
example, every sportsman must have noticed that the somewhat rare
melanic variety of the common rabbit will crouch as steadily as the
normal brownish-gray type, notwithstanding that, owing to its abnormal
colour, a "nigger-rabbit" thus renders itself the most conspicuous
object in the landscape. In all such cases, of course, there has been a
deviation from the normal type in respect of colour, with the result
that the inherited instinct is no longer in tune with the other
endowments of the animal. Such a variation of colour, therefore, will
tend to be suppressed by natural selection; while any variations which
may bring the animal still more closely to resemble its habitual
surroundings will be preserved. Thus we can understand the truly
wonderful extent to which this principle of protective colouring has
been carried in many cases where the need of it has been most urgent.
Not only colour, but structure, may be profoundly modified for the
purposes of protective concealment. Thus, caterpillars which resemble
twigs do so not only in respect of colour, but also of shape; and this
even down to the most minute details in cases where the adaptation is
most complete: certain butterflies and leaf-insects so precisely
resemble the leaves upon which, or among which, they live, that it is
almost impossible to detect them in the foliage--not only the colour,
the shape, and the venation being all exactly imitated, but in some
cases even the defects to which the leaves are liable, in the way of
fungoid growths, &c. There are other insects which with similar
exactness resemble moss, lichens, and so forth. A species of fish
secures a complete resemblance to bunches of sea-weed by a frond-like
modification of all its appendages, and so on through many other
instances. Now, in all such cases where there is so precise an
imitation, both in colour and structure, it seems impossible to suggest
any other explanation of the facts than the one which is supplied by
Mr. Darwin's theory--namely, that the more perfect the resemblance is
caused to become through the continuous influence of natural selection
always picking out the best imitations, the more highly discriminative
becomes the perception of those enemies against the depredations of
which this peculiar kind of protection is developed; so that, in virtue
of this action and re-action, eventually we have a degree of imitation
which renders it almost impossible for a naturalist to detect the animal
when living in its natural environment.
[Illustration: FIG. 110.--Imitative forms and colours in insects.
Drawn from nature (_R. Coll. Surg. Mus._).]
_Warning Colours._
In strange and glaring contrast to all these cases of protective
colouring, stand other cases of conspicuous colouring. Thus, for
example, although there are numberless species of caterpillars which
present in an astonishing degree the phenomena of protective colouring,
there are numberless other species which not only fail to present these
phenomena in any degree, but actually go to the opposite extreme of
presenting colours which appear to have been developed for the sake of
their conspicuousness. At all events, these caterpillars are usually the
most conspicuous objects in their surroundings, and therefore in the
early days of Darwinism they were regarded by Darwin himself as
presenting a formidable difficulty in the way of his theory. To Mr.
Wallace belongs the merit of having cleared up this difficulty in an
extraordinarily successful manner. He virtually reasoned thus. If the
_raison d'etre_ of protective colouring be that of concealing agreeably
flavoured caterpillars from the eye-sight of birds, may not the _raison
d'etre_ of conspicuous colouring be that of protecting disagreeably
flavoured caterpillars from any possibility of being mistaken by birds?
Should this be the case, of course the more conspicuous the colouring
the better would it be for the caterpillars presenting it. Now as soon
as this suggestion was acted upon experimentally, it was found to be
borne out by facts. Birds could not be induced to eat caterpillars of
the kinds in question; and there is now no longer any doubt that their
conspicuous colouring is correlated with their distastefulness to birds,
in the same way as the inconspicuous or imitative colouring of other
caterpillars is correlated with their tastefulness to birds. Here then
is yet another instance, added to those already given, of the
verification yielded to the theory of natural selection by its proved
competency as a guide to facts in nature; for assuredly this particular
class of facts would never have been suspected but for its suggestive
agency.
As in the case of protective imitation, so in this case of warning
conspicuousness, not only colour, but structure may be greatly modified
for the purpose of securing immunity from attack. Here, of course, the
object is to assume, as far as possible, a touch-me-not appearance; so
that, although destitute of any real means of offence, the creatures in
question present a fictitiously dangerous aspect. As the
Devil's-coach-horse turns up his stingless tail when threatened by an
enemy, so in numberless ways do many harmless animals of all classes
pretend to be formidable. But the point now is that these instincts of
self-defence are often helped out by structural modifications,
expressly and exclusively adapted to this end. For example, what a
remarkable series of protective adjustments occurs in the life-history
of the Puss Moth--culminating with so comical an instance of the
particular device now under consideration as the following. I quote the
facts from Mr. E. B. Poulton's admirable book on _The Colours of
Animals_ (pp. 269-271).
[Illustration: FIG. 111.--The larva of Puss Moth (_C. vinula_)
when undisturbed; full-fed; natural size.]
The larva of the Puss Moth (Cerura vinula) is very common upon
poplar and willow. The circular dome-like eggs are laid, either
singly or in little groups of two or three, upon the upper side of
the leaf, and being of a reddish colour strongly suggest the
appearance of little galls, or the results of some other injury to
the leaf. The youngest larvae are black, and also rest upon the
upper surface of the leaf, resembling the dark patches which are
commonly seen in this position. As the larva grows, the apparent
black patch would cover too large a space, and would lead to
detection if it still occupied the whole surface of the body. The
latter gains a green ground-colour which harmonises with the leaf,
while the dark marking is chiefly confined to the back. As growth
proceeds the relative amount of green increases, and the dark mark
is thus prevented from attaining a size which would render it too
conspicuous. In the last stage of growth the green larva becomes
very large, and usually rests on the twigs of its food-plant (Fig.
111). The dark colour is still present on the back but is softened
to a purplish tint, which tends to be replaced by a combination of
white and green in many of the largest larvae. Such a larva is well
concealed by General Protective Resemblance, and one may search a
long time before finding it, although assured of its presence from
the stripped branches of the food-plant and the faeces on the ground
beneath.
[Illustration: FIG. 112.--The larva of Puss Moth in its terrifying
attitude after being disturbed; full-fed; natural size.]
As soon as a large larva is discovered and disturbed it withdraws
its head into the first body-ring, inflating the margin, which is
of a bright red colour. There are two intensely black spots on this
margin in the appropriate position for eyes, and the whole
appearance is that of a large flat face extending to the outer edge
of the red margin (see Fig. 112). The effect is an intensely
exaggerated caricature of a vertebrate face, which is probably
alarming to the vertebrate enemies of the caterpillar. The
terrifying effect is therefore mimetic. The movements entirely
depend on tactile impressions: when touched ever so lightly a
healthy larva immediately assumes the terrifying attitude, and
turns so as to present its full face towards the enemy; if touched
on the other side or on the back it instantly turns its face in the
appropriate direction. The effect is also greatly strengthened by
two pink whips which are swiftly protruded from the prongs of the
fork in which the body terminates. The prongs represent the last
pair of larval legs which have been greatly modified from their
ordinary shape and use. The end of the body is at the same time
curved forward over the back (generally much further than in Fig.
112), so that the pink filaments are brandished above the head.
_Mimicry._
Lastly, these facts as to imitative and conspicuous colouring lead on to
the yet more remarkable facts of what is called mimicry. By mimicry is
meant the imitation in form and colour of one species by another, in
order that the imitating species may be mistaken for the imitated, and
thus participate in some advantage which the latter enjoys. For
instance, if, as in the case of the conspicuously-coloured caterpillars,
it is of advantage to an ill-savoured species that it should hold out a
warning to enemies, clearly it may be of no less advantage to a
well-savoured species that it should borrow this flag, and thus be
mistaken for its ill-savoured neighbour. Now, the extent to which this
device of mimicry is carried is highly remarkable, not only in respect
of the number of its cases, but also in respect of the astonishing
accuracy which in most of these cases is exhibited by the imitation.
There need be little or virtually no zoological affinity between the
imitating and the imitated forms; that is to say, in some cases the
zoological affinity is not closer than ordinal, and therefore cannot
possibly be ascribed to kinship. Like all the other branches of the
general subject of protective resemblance in form or colouring, this
branch has already been so largely illustrated by previous writers,
that, as in the previous cases, I need only give one or two examples.
Those which I choose are chosen on account of the colours concerned not
being highly varied or brilliant, and therefore lending themselves to
less ineffectual treatment by wood-engraving than is the case where
attempts are made to render by this means even more remarkable
instances. (Figs. 113, 114, 115.)
[Illustration: FIG. 113.--Three cases of mimicry. Drawn from nature:
first two pairs nat. size, last pair 2/3 (_R. Coll. Surg. Mus._).]
[Illustration: FIG. 114.--Two further cases of mimicry; flies
resembling a wasp in the one and a bee in the other. Drawn from
nature: nat. size (_R. Coll. Surg. Mus._).]
[Illustration: FIG. 115.--A case of mimicry where a non-venomous
species of snake resembles a venomous one. Drawn from nature: 1/3
nat. size (_R. Coll. Surg. Mus._).]
It is surely apparent, without further comment, that it is impossible to
imagine stronger evidence in favour of natural selection as a true cause
in nature, than is furnished by this culminating fact in the matter of
protective resemblance, whereby it is shown that a species of one
genus, family, or even order, will accurately mimic the appearance of a
species belonging to another genus, family, or order, so as to deceive
its natural enemies into mistaking it for a creature of so totally
different a kind. And it must be added that while this fact of mimicry
is of extraordinarily frequent occurrence, there can be no possibility
of our mistaking its purpose. For the fact is never observable except in
the case of species which occupy the same area or district.
Such being what appears to me the only reasonable view of the matter, I
will now conclude this chapter on the evidences of natural selection as
at all events the main factor of organic evolution, by simply adding
illustrations of two further cases of mimicry, which are perhaps even
more remarkable than any of the foregoing examples. The first of the two
(Fig. 115) speaks for itself. The second will be rendered intelligible
by the following few words of explanation.
There are certain ants of the Amazons which present the curious instinct
of cutting off leaves from trees, and carrying them like banners over
their heads to the hive, as represented in Fig. 116, B, where one ant is
shown without a leaf, and the others each with a leaf. Their object in
thus collecting leaves is probably that of growing a fungus upon the
"soil" which is furnished by the leaves when decomposing. But, be this
as it may[39], the only point we are now concerned with is the
appearance which these ants present when engaged in their habitual
operation of carrying leaves. For it has been recently observed by Mr.
W. L. Sclater, that in the localities where these hymenopterous insects
occur, there occurs also a _homopterous_ insect which mimics the ant,
leaf and all, in a wonderfully deceptive manner. The leaf is imitated by
the thin flattened body of the insect, "which in its dorsal aspect is so
compressed laterally that it is no thicker than a leaf, and terminates
in a sharp jagged edge." The colour is exactly the same as that of a
leaf, and the brown legs show themselves beneath the green body in just
the same way as those of the ant show themselves beneath the leaf. So
that both the form and the colouring of the homopterous insect has been
brought to resemble, with singular exactness, those belonging to a
different order of insect, when the latter is engaged in its peculiar
avocation. A glance at the figure is enough to show the means employed
and the result attained. In A, an ant and its mimic are represented as
about 2-1/2 times their natural size, and both proceeding in the same
direction. It ought to be mentioned, however, that in reality the margin
of the leaf is seldom allowed to retain its natural serrations as here
depicted: the ants usually gnaw the edge of the real leaf, so that the
margin of the false one bears an even closer resemblance to it than the
illustration represents. B is a drawing from life of a group of five
ants carrying leaves, and their mimic walking beside them[40].
[39] For a full account of this instinct and its probable purpose,
see _Animal Intelligence_, pp. 93-6.
[40] Both drawings are reproduced from Mr. Poulton's paper upon the
subject (_Proc. Zool. Soc._, June 16, 1891).
[Illustration: FIG. 116 PROTECTIVE MIMICRY]
CHAPTER IX.
CRITICISMS OF THE THEORY OF NATURAL SELECTION.
I will now proceed to consider the various objections and difficulties
which have hitherto been advanced against the theory of natural
selection.
Very early in the day Owen hurled the weight of his authority against
the new theory, and this with a strength of onslaught which was only
equalled by its want of judgment. Indeed, it is painfully apparent that
he failed to apprehend the fundamental principles of the Darwinian
theory. For he says:--
Natural Selection is an explanation of the process [of
transmutation] of the same kind and value as that which has been
proffered of the mystery of "secretion." For example, a particular
mass of matter in a living animal takes certain elements out of the
blood, and rejects them as "bile." Attributes were given to the
liver which can only be predicated of the whole animal; the
"appetency" of the liver, it was said, was for the elements of
bile, and "biliosity," or the "hepatic sensation," guided the gland
to their secretion. Such figurative language, I need not say,
explains absolutely nothing of the nature of bilification[41].
[41] _Anatomy of Vertebrates_, vol. iii. p. 794.
Assuredly, it was needless for Owen to say that figurative language of
this kind explains nothing; but it was little less than puerile in him
to see no more in the theory of natural selection than such a mere
figure of speech. To say that the liver selects the elements of bile, or
that nature selects specific types, may both be equally unmeaning
re-statements of facts; but when it is explained that the term natural
selection, unlike that of "hepatic sensation," is used as a shorthand
expression for a whole group of well-known natural causes--struggle,
variation, survival, heredity,--then it becomes evidence of an almost
childish want of thought to affirm that the expression is figurative and
nothing more. The doctrine of natural selection may be a huge mistake;
but, if so, this is not because it consists of any unmeaning metaphor:
it can only be because the combination of natural causes which it
suggests is not of the same adequacy in fact as it is taken to be in
theory.
Owen further objected that the struggle for existence could only act as
a cause of the extinction of species, not of their origination--a view
of the case which again shows on his part a complete failure to grasp
the conception of Darwinism. Acting alone, the struggle for existence
could only cause extermination; but acting together with variation,
survival, and heredity, it may very well--for anything that Owen, or
others who followed in this line of criticism, show to the
contrary--have produced every species of plant and animal that has ever
appeared upon the face of the earth.
Another and closely allied objection is, that the theory of natural
selection "personifies an abstraction." Or, as the Duke of Argyll states
it, the theory is "essentially the image of mechanical necessity
concealed under the clothes, and parading in the mask, of mental
purpose. The word 'natural' suggests Matter, and the physical forces.
The word 'selection' suggests Mind, and the powers of choice." This,
however, is a mere quarrelling about words. Darwin called the principle
which he had discovered by the name natural selection in order to mark
the analogy between it and artificial selection. No doubt in this
analogy there is not necessarily supposed to be in nature any
counterpart to the mind of the breeder, nor, therefore, to his powers of
intelligent choice. But there is no need to limit the term _selection_
(_se_ and _lego_, Gr. [Greek: lego]) to powers of intelligent choice. As
previously remarked, a bank of sea-weed on the sea-shore may be said to
have been selected by the waves from all the surrounding sand and
stones. Similarly, we may say that grain is selected from chaff by the
wind in the process of winnowing corn. Or, if it be thought that there
is any ambiguity involved in such a use of the term in the case of
"Natural Selection," there is no objection to employing the phrase which
has been coined by Mr. Spencer as its equivalent--namely, "Survival of
the Fittest." The point of the theory is, that those organisms which are
best suited to their surroundings are allowed to live and to propagate,
while those which are less suited are eliminated; and whether we call
this process a process of selection, or call it by any other name, is
clearly immaterial.
A material question is raised only when it is asked whether the process
is one that can be ascribed to causation strictly natural. It is often
denied that such is the case, on the ground that natural selection does
not originate the variations which it favours, but depends upon the
variations being supplied by some other means. For, it is said, all that
natural selection does is to preserve the suitable variations _after
they have arisen_. Natural selection does not _cause_ these suitable
variations; and therefore, it is argued, Darwin and his followers are
profoundly mistaken in representing the principle as one which
_produces_ adaptations. Now, although this objection has been put
forward by some of the most intelligent minds in our generation, it
appears to me to betoken some extraordinary failure to appreciate the
very essence of Darwinian doctrine. No doubt it is perfectly true that
natural selection does not produce variations of any kind, whether
beneficial or otherwise. But if it be granted that variations of many
kinds are occurring in every generation, and that natural selection is
competent to preserve the more favourable among them, then it appears to
me unquestionable that this principle of selection deserves to be
regarded as, in the full sense of the word, a natural cause. The
variations being expressly regarded by the theory as more or less
promiscuous[42], survival of the fittest becomes the winnowing fan,
whose function it is to eliminate all the less fit in each generation,
in order to preserve the good grain, out of which to constitute the next
generation. And as this process is supposed to be continuous through
successive generations, its action is supposed to be cumulative, till
from the eye of a worm there is gradually developed the eye of an eagle.
Therefore it follows from these suppositions (which are not disputed by
the present objection), that if it had not been for the process of
selection, such development would never have been begun; and that in the
exact measure of its efficiency will the development proceed. But any
agency without the operation of which a result cannot take place may
properly be designated the cause of that result: it is the agency which,
in co-operation with all the other agencies in the cosmos, produces that
result.
[42] The degree in which variability is indefinite, or, on the
contrary, determinate, is a question which is not yet ripe for
decision--nor even, in my opinion, for discussion. But I may here
state the following general principles with regard to it.
(1) It is evident that up to some point or another variations _must_
be pre-determined in definite lines. Men do not gather grapes from
thorns, figs from thistles, nor even moss-roses from sweet-briars.
In other words, "the nature of the organism" in all cases
necessitates the limiting of variations within certain bounds.
(2) But when the question is as to what these bounds may be, we can
only answer in a general way that, according to the general theory
of evolution, they must be such as are imposed by heredity, coupled
with the degree to which external conditions of life (and possibly
also use-inheritance) are capable, in given cases, of modifying
congenital characters. These are the only causes which the theory of
descent can consistently recognise as producing variations in
determinate directions.
(3) Inasmuch as variation presupposes the existence of parts that
vary, and inasmuch as the variation of parts can only be in the
alternative directions of increase or decrease around an average, it
follows that, in the first instance at all events, every variation,
if determinate, must be so only in one or other of these two
opposite directions.
(4) In as far as variations are summated in successive generations,
so as eventually to give rise to new structures, organs, mechanisms,
&c., natural selection is theoretically competent to explain the
facts, without our having to postulate the operation of unknown
causes producing variations in determinate lines,--or not further
than is stated in paragraphs 1 and 2.
(5) Nevertheless, it does not follow that there are not such other
unknown causes; and, if there are, of course the importance of
natural selection as a cause of adaptive modification would be
limited in proportion to their number and the extent of their
operation. But it is for those who, like the late Professors Asa
Gray and Naegeli, maintain the existence of such causes, to
substantiate their belief by indicating them.
Take any analogous case. The selective agency of specific gravity which
is utilised in gold-washing does not create the original differences
between gold-dust and dust of all other kinds. But these differences
being presented by as many different bodies in nature, the gold-washer
takes advantage of the selective agency in question, and, by using it as
a cause of segregation, is enabled to separate the gold from all the
earths with which it may happen to be mixed. So far as the objects of
the gold-washer are concerned, it is immaterial with what other earths
the gold-dust may happen to be mixed. For although gold-dust may occur
in intimate association with earths of various kinds in various
proportions, and although in each case the particular admixture which
occurs must have been due to definite causes, these things, in relation
to the selective process of the washer, are what is called accidental:
that is to say, they have nothing to do with the causative action of the
selective process. Now, in precisely the same sense Darwin calls the
multitudinous variations of plants and animals accidental. By so calling
them he expressly says he does not suppose them to be accidental in the
sense of not all being due to definite causes. But they are accidental
in relation to the sifting process of natural selection: all that they
have to do is to furnish the promiscuous material on which this sifting
process acts.
Or let us take an even closer analogy. The power of selective breeding
by man is so wonderful, that in the course of successive generations all
kinds of peculiarities as to size, shape, colour, special appendages or
abortions, &c., can be produced at pleasure, as we saw in the last
chapter. Now all the promiscuous variations which are supplied to the
breeder, and out of which, by selecting only those that are suited to
his purpose, he is able to produce the required result--all those
promiscuous variations, in relation to that purpose, are accidental.
Therefore the selective agency of the breeder deserves to be regarded as
the cause of that which it produces, or of that which could not have
been produced but for the operation of such agency. But where is the
difference between artificial and natural selection in this respect?
And, if there is no difference, is not natural selection as much
entitled to be regarded as a true cause of the origin of natural
species, as artificial selection is to be regarded as a true cause of
our domesticated races? Here, as in the case of the previous
illustration, if there be any ambiguity in speaking of variations as
accidental, it arises from the incorrect or undefined manner in which
the term "accidental" is used by Darwin's critics. In its original and
philosophically-correct usage, the term "accident" signifies a property
or quality not essential to our conception of a substance: hence, it has
come to mean anything that happens as a result of unforeseen causes--or,
lastly, that which is causeless. But, as we know that nothing can happen
without causes of some kind, the term "accident" is divested of real
meaning when it is used in the last of these senses. Yet this is the
sense that is sought to be placed upon it by the objection which we are
considering. If the objectors will but understand the term in its
correct philosophical sense--or in the only sense in which it presents
any meaning at all,--they will see that Darwinians are both logically
and historically justified in employing the word "accidental" as the
word which serves most properly to convey the meaning that they
intend--namely, variations due to causes accidental to the struggle for
existence. Similarly, when it is said that variations are "spontaneous,"
or even "fortuitous," nothing further is meant than that we do not know
the causes which lead to them, and that, so far as the principle of
selection is concerned, it is immaterial what these causes may be. Or,
to revert to our former illustration, the various weights of different
kinds of earths are no doubt all due to definite causes; but, in
relation to the selective action of the gold-washer, all the different
weights of whatever kinds of earth he may happen to include in his
washing-apparatus are, _strictly speaking_, accidental. And as at
different washings he meets with different proportions of heavy earths
with light ones, and as these "variations" are immaterial to him, he may
colloquially speak of them as "fortuitous," or due to "chance," even
though he knows that at each washing they must have been determined by
definite causes.
More adequately to deal with this merely formal objection, however,
would involve more logic-chopping than is desirable on the present
occasion. But I have already dealt with it fully elsewhere,--viz. in
_The Contemporary Review_ for June, 1888, to which therefore I may refer
any one who is interested in dialectics of this kind[43].
[43] Within the last few months this objection has been presented
anew by Mr. D. Syme, whose book _On the Modification of Organisms_
exhibits a curious combination of shrewd criticisms with almost
ludicrous misunderstandings. One of the latter it is necessary to
state, because it pervades the quotation which I am about to supply.
He everywhere compares "natural selection" with "the struggle for
existence," uses them as convertible terms, and while absurdly
stating that "Darwin defines natural selection as the struggle for
existence," complains of "the liability of error, both on his own
part and on the part of his readers," which arises from his not
having everywhere adhered to this definition! (p. 8).
"Darwin has put forth two distinct and contradictory theories of the
functions of natural selection. According to the one theory natural
selection is selective or preservative, and nothing more. According
to the other theory natural selection creates the variations(!) ...
It certainly seems absurd to speak of natural selection, or the
struggle for existence, as selective or preservative, for the
struggle for existence does not preserve at all, not even the fit
variations, as both the fit and the unfit struggle for existence,
the unfit naturally more than the fit, and the fit are preserved,
not in consequence of the struggle, but in consequence of their
fitness. Suppose two varieties of the same species are driven, by an
increase of their numbers, to seek for subsistence in a colder
region than they have been accustomed to, and that one of these
varieties had a hardier constitution than the other; and let us
suppose that the former withstood the severe climate better than the
latter, and consequently survived, while the other perished. In this
case the hardier survived, not because of the struggle, but because
it had a constitution better adapted to the climate. I wish to
ascertain if a certain metal in my possession is gold or some baser
metal, and I apply the usual test; but the mere fact of my testing
this metal would not make it gold or any other kind of metal."
I have thought it worth while to quote this passage for the sake of
showing the extraordinary confusion of mind which still prevails on
the part of Darwin's critics, even with reference to the very
fundamental parts of his theory. For, as I have said, the writer of
this passage shows himself a shrewd critic in some other parts of
his essay, where he is not engaged especially on the theory of
natural selection.
I will now pass on to consider another misconception of the Darwinian
theory, which is very prevalent in the public mind. It is virtually
asked, If some species are supposed to have been improved by natural
selection, why have not all species been similarly improved? Why should
not all invertebrated animals have risen into vertebrated? Or why
should not all monkeys have become men?
The answers are manifold. In the first place, it by no means follows
that because an advance in organization has proved itself of benefit in
the case of one form of life, therefore any or every other form would
have been similarly benefited by a similar advance. The business of
natural selection is to bring this and that form of life into the
closest harmony with its environment that all the conditions of the case
permit. Sometimes it will happen that the harmony will admit of being
improved by an improvement of organization. But just as often it will
happen that it will be best secured by leaving matters as they are. If,
therefore, an organism has already been brought into a tolerably full
degree of harmony with its environment, natural selection will not try
to change it so long as the environment remains unchanged; and this, no
doubt, is the reason why some species have survived through enormous
periods of geological time without having undergone any change. Again,
as we saw in a previous chapter, there are yet other cases where, on
account of some change in the environment or even in the habits of the
organisms themselves, adaption will be best secured by an active
_reversal_ of natural selection, with the result of causing
_degeneration_.
But, it is sometimes further urged, there are cases where we cannot
doubt that improvement of organization would have been of benefit to
species; and yet such improvement has not taken place--as, for instance,
in the case all monkeys not turning into men. Here, however, we must
remember that the operation of natural selection in any case depends
upon a variety of highly complex conditions; and, therefore, that the
fact of all those conditions having been satisfied in one instance is no
reason for concluding that they must also have been satisfied in other
instances. Take, for example, the case of monkeys passing into men. The
wonder to me appears to be that this improvement should have taken place
in even one line of descent; not that, having taken place in one line,
it should not also have taken place in other lines. For how enormously
complex must have been the conditions--physical, anatomical,
physiological, psychological, sociological--which by their happy
conjunction first began to raise the inarticulate cries of an ape into
the rational speech of a man. Therefore, the more that we appreciate the
superiority of a man to an ape, the less ought we to countenance this
supposed objection to Darwin's theory--namely, that natural selection
has not effected the change in more than one line of descent.
Even in the case of two races of mankind where one has risen higher in
the scale of civilization than another, it is now generally impossible
to assign the particular causes of the difference; much more, then, must
this be impossible in the case of still more remote conditions which
have led to the divergence of species. The requisite variations may not
have arisen in the one line of descent which did arise in the other; or
if they did arise in both, some counterbalancing disadvantages may have
attended their initial development in the one case which did not obtain
in the other. In short, where so exceedingly complex a play of
conditions are concerned, the only wonder would be if two different
lines of descent _had_ happened to present two independent and yet
perfectly parallel lines of history.
These general considerations would apply equally to the great majority
of other cases where some types have made great advances upon others,
notwithstanding that we can see no reason why the latter should not in
this respect have imitated the former. But there is yet a further
consideration which must be taken into account. The struggle for
existence is always most keen between closely allied species, because,
from the similarity of their forms, habits, needs, &c., they are in
closest competition. Therefore it often happens that the mere fact of
one species having made an advance upon others of itself precludes the
others from making any similar advance: the field, so to speak, has
already been occupied as regards that particular improvement, and where
the struggle for existence is concerned possession is emphatically nine
points of the law. For example, to return to the case of apes becoming
men, the fact of one rational species having been already evolved (even
if the rational faculty were at first but dimly nascent) must make an
enormous change in the conditions as regards the possibility of any
other such species being subsequently evolved--unless, of course, it be
by way of descent from the rational one. Or, as Sir Charles Lyell has
well put it, two rational species can never _coexist_ on the globe,
although the descendants of one rational species may in time become
_transformed_ into another single rational species[44].
[44] _Principles of Geology_, vol. ii. p. 487 (11th ed.).