NATURAL SELECTION.
Natural Selection: its power compared with man’s selection, its power
on characters of trifling importance, its power at all ages and on both
sexes. Sexual Selection. On the generality of intercrosses between
individuals of the same species. Circumstances favourable and
unfavourable to Natural Selection, namely, intercrossing, isolation,
number of individuals. Slow action. Extinction caused by Natural
Selection. Divergence of Character, related to the diversity of
inhabitants of any small area, and to naturalisation. Action of Natural
Selection, through Divergence of Character and Extinction, on the
descendants from a common parent. Explains the Grouping of all organic
beings.
How will the struggle for existence, discussed too briefly in the last
chapter, act in regard to variation? Can the principle of selection,
which we have seen is so potent in the hands of man, apply in nature? I
think we shall see that it can act most effectually. Let it be borne in
mind in what an endless number of strange peculiarities our domestic
productions, and, in a lesser degree, those under nature, vary; and how
strong the hereditary tendency is. Under domestication, it may be truly
said that the whole organisation becomes in some degree plastic. Let it
be borne in mind how infinitely complex and close-fitting are the
mutual relations of all organic beings to each other and to their
physical conditions of life. Can it, then, be thought improbable,
seeing that variations useful to man have undoubtedly occurred, that
other variations useful in some way to each being in the great and
complex battle of life, should sometimes occur in the course of
thousands of generations? If such do occur, can we doubt (remembering
that many more individuals are born than can possibly survive) that
individuals having any advantage, however slight, over others, would
have the best chance of surviving and of procreating their kind? On the
other hand, we may feel sure that any variation in the least degree
injurious would be rigidly destroyed. This preservation of favourable
variations and the rejection of injurious variations, I call Natural
Selection. Variations neither useful nor injurious would not be
affected by natural selection, and would be left a fluctuating element,
as perhaps we see in the species called polymorphic.
We shall best understand the probable course of natural selection by
taking the case of a country undergoing some physical change, for
instance, of climate. The proportional numbers of its inhabitants would
almost immediately undergo a change, and some species might become
extinct. We may conclude, from what we have seen of the intimate and
complex manner in which the inhabitants of each country are bound
together, that any change in the numerical proportions of some of the
inhabitants, independently of the change of climate itself, would most
seriously affect many of the others. If the country were open on its
borders, new forms would certainly immigrate, and this also would
seriously disturb the relations of some of the former inhabitants. Let
it be remembered how powerful the influence of a single introduced tree
or mammal has been shown to be. But in the case of an island, or of a
country partly surrounded by barriers, into which new and better
adapted forms could not freely enter, we should then have places in the
economy of nature which would assuredly be better filled up, if some of
the original inhabitants were in some manner modified; for, had the
area been open to immigration, these same
places would have been seized on by intruders. In such case, every
slight modification, which in the course of ages chanced to arise, and
which in any way favoured the individuals of any of the species, by
better adapting them to their altered conditions, would tend to be
preserved; and natural selection would thus have free scope for the
work of improvement.
We have reason to believe, as stated in the first chapter, that a
change in the conditions of life, by specially acting on the
reproductive system, causes or increases variability; and in the
foregoing case the conditions of life are supposed to have undergone a
change, and this would manifestly be favourable to natural selection,
by giving a better chance of profitable variations occurring; and
unless profitable variations do occur, natural selection can do
nothing. Not that, as I believe, any extreme amount of variability is
necessary; as man can certainly produce great results by adding up in
any given direction mere individual differences, so could Nature, but
far more easily, from having incomparably longer time at her disposal.
Nor do I believe that any great physical change, as of climate, or any
unusual degree of isolation to check immigration, is actually necessary
to produce new and unoccupied places for natural selection to fill up
by modifying and improving some of the varying inhabitants. For as all
the inhabitants of each country are struggling together with nicely
balanced forces, extremely slight modifications in the structure or
habits of one inhabitant would often give it an advantage over others;
and still further modifications of the same kind would often still
further increase the advantage. No country can be named in which all
the native inhabitants are now so perfectly adapted to each other and
to the physical conditions under which they live, that none of
them could anyhow be improved; for in all countries, the natives have
been so far conquered by naturalised productions, that they have
allowed foreigners to take firm possession of the land. And as
foreigners have thus everywhere beaten some of the natives, we may
safely conclude that the natives might have been modified with
advantage, so as to have better resisted such intruders.
As man can produce and certainly has produced a great result by his
methodical and unconscious means of selection, what may not nature
effect? Man can act only on external and visible characters: nature
cares nothing for appearances, except in so far as they may be useful
to any being. She can act on every internal organ, on every shade of
constitutional difference, on the whole machinery of life. Man selects
only for his own good; Nature only for that of the being which she
tends. Every selected character is fully exercised by her; and the
being is placed under well-suited conditions of life. Man keeps the
natives of many climates in the same country; he seldom exercises each
selected character in some peculiar and fitting manner; he feeds a long
and a short beaked pigeon on the same food; he does not exercise a
long-backed or long-legged quadruped in any peculiar manner; he exposes
sheep with long and short wool to the same climate. He does not allow
the most vigorous males to struggle for the females. He does not
rigidly destroy all inferior animals, but protects during each varying
season, as far as lies in his power, all his productions. He often
begins his selection by some half-monstrous form; or at least by some
modification prominent enough to catch his eye, or to be plainly useful
to him. Under nature, the slightest difference of structure or
constitution may well turn the nicely-balanced scale in the
struggle for life, and so be preserved. How fleeting are the wishes and
efforts of man! how short his time! and consequently how poor will his
products be, compared with those accumulated by nature during whole
geological periods. Can we wonder, then, that nature’s productions
should be far “truer” in character than man’s productions; that they
should be infinitely better adapted to the most complex conditions of
life, and should plainly bear the stamp of far higher workmanship?
It may be said that natural selection is daily and hourly scrutinising,
throughout the world, every variation, even the slightest; rejecting
that which is bad, preserving and adding up all that is good; silently
and insensibly working, whenever and wherever opportunity offers, at
the improvement of each organic being in relation to its organic and
inorganic conditions of life. We see nothing of these slow changes in
progress, until the hand of time has marked the long lapse of ages, and
then so imperfect is our view into long past geological ages, that we
only see that the forms of life are now different from what they
formerly were.
Although natural selection can act only through and for the good of
each being, yet characters and structures, which we are apt to consider
as of very trifling importance, may thus be acted on. When we see
leaf-eating insects green, and bark-feeders mottled-grey; the alpine
ptarmigan white in winter, the red-grouse the colour of heather, and
the black-grouse that of peaty earth, we must believe that these tints
are of service to these birds and insects in preserving them from
danger. Grouse, if not destroyed at some period of their lives, would
increase in countless numbers; they are known to suffer largely from
birds of prey; and hawks are guided by eyesight to their prey,—so much
so, that on
parts of the Continent persons are warned not to keep white pigeons, as
being the most liable to destruction. Hence I can see no reason to
doubt that natural selection might be most effective in giving the
proper colour to each kind of grouse, and in keeping that colour, when
once acquired, true and constant. Nor ought we to think that the
occasional destruction of an animal of any particular colour would
produce little effect: we should remember how essential it is in a
flock of white sheep to destroy every lamb with the faintest trace of
black. In plants the down on the fruit and the colour of the flesh are
considered by botanists as characters of the most trifling importance:
yet we hear from an excellent horticulturist, Downing, that in the
United States smooth-skinned fruits suffer far more from a beetle, a
curculio, than those with down; that purple plums suffer far more from
a certain disease than yellow plums; whereas another disease attacks
yellow-fleshed peaches far more than those with other coloured flesh.
If, with all the aids of art, these slight differences make a great
difference in cultivating the several varieties, assuredly, in a state
of nature, where the trees would have to struggle with other trees and
with a host of enemies, such differences would effectually settle which
variety, whether a smooth or downy, a yellow or purple fleshed fruit,
should succeed.
In looking at many small points of difference between species, which,
as far as our ignorance permits us to judge, seem to be quite
unimportant, we must not forget that climate, food, etc., probably
produce some slight and direct effect. It is, however, far more
necessary to bear in mind that there are many unknown laws of
correlation of growth, which, when one part of the organisation is
modified through variation, and the modifications are accumulated by
natural selection for
the good of the being, will cause other modifications, often of the
most unexpected nature.
As we see that those variations which under domestication appear at any
particular period of life, tend to reappear in the offspring at the
same period;—for instance, in the seeds of the many varieties of our
culinary and agricultural plants; in the caterpillar and cocoon stages
of the varieties of the silkworm; in the eggs of poultry, and in the
colour of the down of their chickens; in the horns of our sheep and
cattle when nearly adult;—so in a state of nature, natural selection
will be enabled to act on and modify organic beings at any age, by the
accumulation of profitable variations at that age, and by their
inheritance at a corresponding age. If it profit a plant to have its
seeds more and more widely disseminated by the wind, I can see no
greater difficulty in this being effected through natural selection,
than in the cotton-planter increasing and improving by selection the
down in the pods on his cotton-trees. Natural selection may modify and
adapt the larva of an insect to a score of contingencies, wholly
different from those which concern the mature insect. These
modifications will no doubt affect, through the laws of correlation,
the structure of the adult; and probably in the case of those insects
which live only for a few hours, and which never feed, a large part of
their structure is merely the correlated result of successive changes
in the structure of their larvæ. So, conversely, modifications in the
adult will probably often affect the structure of the larva; but in all
cases natural selection will ensure that modifications consequent on
other modifications at a different period of life, shall not be in the
least degree injurious: for if they became so, they would cause the
extinction of the species.
Natural selection will modify the structure of the
young in relation to the parent, and of the parent in relation to the
young. In social animals it will adapt the structure of each individual
for the benefit of the community; if each in consequence profits by the
selected change. What natural selection cannot do, is to modify the
structure of one species, without giving it any advantage, for the good
of another species; and though statements to this effect may be found
in works of natural history, I cannot find one case which will bear
investigation. A structure used only once in an animal’s whole life, if
of high importance to it, might be modified to any extent by natural
selection; for instance, the great jaws possessed by certain insects,
and used exclusively for opening the cocoon—or the hard tip to the beak
of nestling birds, used for breaking the egg. It has been asserted,
that of the best short-beaked tumbler-pigeons more perish in the egg
than are able to get out of it; so that fanciers assist in the act of
hatching. Now, if nature had to make the beak of a full-grown pigeon
very short for the bird’s own advantage, the process of modification
would be very slow, and there would be simultaneously the most rigorous
selection of the young birds within the egg, which had the most
powerful and hardest beaks, for all with weak beaks would inevitably
perish: or, more delicate and more easily broken shells might be
selected, the thickness of the shell being known to vary like every
other structure.
_Sexual Selection_.—Inasmuch as peculiarities often appear under
domestication in one sex and become hereditarily attached to that sex,
the same fact probably occurs under nature, and if so, natural
selection will be able to modify one sex in its functional relations to
the other sex, or in relation to wholly different habits of life in the
two sexes, as is sometimes the case
with insects. And this leads me to say a few words on what I call
Sexual Selection. This depends, not on a struggle for existence, but on
a struggle between the males for possession of the females; the result
is not death to the unsuccessful competitor, but few or no offspring.
Sexual selection is, therefore, less rigorous than natural selection.
Generally, the most vigorous males, those which are best fitted for
their places in nature, will leave most progeny. But in many cases,
victory will depend not on general vigour, but on having special
weapons, confined to the male sex. A hornless stag or spurless cock
would have a poor chance of leaving offspring. Sexual selection by
always allowing the victor to breed might surely give indomitable
courage, length to the spur, and strength to the wing to strike in the
spurred leg, as well as the brutal cock-fighter, who knows well that he
can improve his breed by careful selection of the best cocks. How low
in the scale of nature this law of battle descends, I know not; male
alligators have been described as fighting, bellowing, and whirling
round, like Indians in a war-dance, for the possession of the females;
male salmons have been seen fighting all day long; male stag-beetles
often bear wounds from the huge mandibles of other males. The war is,
perhaps, severest between the males of polygamous animals, and these
seem oftenest provided with special weapons. The males of carnivorous
animals are already well armed; though to them and to others, special
means of defence may be given through means of sexual selection, as the
mane to the lion, the shoulder-pad to the boar, and the hooked jaw to
the male salmon; for the shield may be as important for victory, as the
sword or spear.
Amongst birds, the contest is often of a more peaceful character. All
those who have attended to the subject,
believe that there is the severest rivalry between the males of many
species to attract by singing the females. The rock-thrush of Guiana,
birds of Paradise, and some others, congregate; and successive males
display their gorgeous plumage and perform strange antics before the
females, which standing by as spectators, at last choose the most
attractive partner. Those who have closely attended to birds in
confinement well know that they often take individual preferences and
dislikes: thus Sir R. Heron has described how one pied peacock was
eminently attractive to all his hen birds. It may appear childish to
attribute any effect to such apparently weak means: I cannot here enter
on the details necessary to support this view; but if man can in a
short time give elegant carriage and beauty to his bantams, according
to his standard of beauty, I can see no good reason to doubt that
female birds, by selecting, during thousands of generations, the most
melodious or beautiful males, according to their standard of beauty,
might produce a marked effect. I strongly suspect that some well-known
laws with respect to the plumage of male and female birds, in
comparison with the plumage of the young, can be explained on the view
of plumage having been chiefly modified by sexual selection, acting
when the birds have come to the breeding age or during the breeding
season; the modifications thus produced being inherited at
corresponding ages or seasons, either by the males alone, or by the
males and females; but I have not space here to enter on this subject.
Thus it is, as I believe, that when the males and females of any animal
have the same general habits of life, but differ in structure, colour,
or ornament, such differences have been mainly caused by sexual
selection; that is, individual males have had, in successive
generations, some slight advantage over other
males, in their weapons, means of defence, or charms; and have
transmitted these advantages to their male offspring. Yet, I would not
wish to attribute all such sexual differences to this agency: for we
see peculiarities arising and becoming attached to the male sex in our
domestic animals (as the wattle in male carriers, horn-like
protuberances in the cocks of certain fowls, etc.), which we cannot
believe to be either useful to the males in battle, or attractive to
the females. We see analogous cases under nature, for instance, the
tuft of hair on the breast of the turkey-cock, which can hardly be
either useful or ornamental to this bird;—indeed, had the tuft appeared
under domestication, it would have been called a monstrosity.
_Illustrations of the action of Natural Selection_.—In order to make it
clear how, as I believe, natural selection acts, I must beg permission
to give one or two imaginary illustrations. Let us take the case of a
wolf, which preys on various animals, securing some by craft, some by
strength, and some by fleetness; and let us suppose that the fleetest
prey, a deer for instance, had from any change in the country increased
in numbers, or that other prey had decreased in numbers, during that
season of the year when the wolf is hardest pressed for food. I can
under such circumstances see no reason to doubt that the swiftest and
slimmest wolves would have the best chance of surviving, and so be
preserved or selected,—provided always that they retained strength to
master their prey at this or at some other period of the year, when
they might be compelled to prey on other animals. I can see no more
reason to doubt this, than that man can improve the fleetness of his
greyhounds by careful and methodical selection, or by that unconscious
selection which results from each man trying
to keep the best dogs without any thought of modifying the breed.
Even without any change in the proportional numbers of the animals on
which our wolf preyed, a cub might be born with an innate tendency to
pursue certain kinds of prey. Nor can this be thought very improbable;
for we often observe great differences in the natural tendencies of our
domestic animals; one cat, for instance, taking to catch rats, another
mice; one cat, according to Mr. St. John, bringing home winged game,
another hares or rabbits, and another hunting on marshy ground and
almost nightly catching woodcocks or snipes. The tendency to catch rats
rather than mice is known to be inherited. Now, if any slight innate
change of habit or of structure benefited an individual wolf, it would
have the best chance of surviving and of leaving offspring. Some of its
young would probably inherit the same habits or structure, and by the
repetition of this process, a new variety might be formed which would
either supplant or coexist with the parent-form of wolf. Or, again, the
wolves inhabiting a mountainous district, and those frequenting the
lowlands, would naturally be forced to hunt different prey; and from
the continued preservation of the individuals best fitted for the two
sites, two varieties might slowly be formed. These varieties would
cross and blend where they met; but to this subject of intercrossing we
shall soon have to return. I may add, that, according to Mr. Pierce,
there are two varieties of the wolf inhabiting the Catskill Mountains
in the United States, one with a light greyhound-like form, which
pursues deer, and the other more bulky, with shorter legs, which more
frequently attacks the shepherd’s flocks.
Let us now take a more complex case. Certain plants excrete a sweet
juice, apparently for the sake of eliminating something injurious from
their sap: this is
effected by glands at the base of the stipules in some Leguminosæ, and
at the back of the leaf of the common laurel. This juice, though small
in quantity, is greedily sought by insects. Let us now suppose a little
sweet juice or nectar to be excreted by the inner bases of the petals
of a flower. In this case insects in seeking the nectar would get
dusted with pollen, and would certainly often transport the pollen from
one flower to the stigma of another flower. The flowers of two distinct
individuals of the same species would thus get crossed; and the act of
crossing, we have good reason to believe (as will hereafter be more
fully alluded to), would produce very vigorous seedlings, which
consequently would have the best chance of flourishing and surviving.
Some of these seedlings would probably inherit the nectar-excreting
power. Those individual flowers which had the largest glands or
nectaries, and which excreted most nectar, would be oftenest visited by
insects, and would be oftenest crossed; and so in the long-run would
gain the upper hand. Those flowers, also, which had their stamens and
pistils placed, in relation to the size and habits of the particular
insects which visited them, so as to favour in any degree the
transportal of their pollen from flower to flower, would likewise be
favoured or selected. We might have taken the case of insects visiting
flowers for the sake of collecting pollen instead of nectar; and as
pollen is formed for the sole object of fertilisation, its destruction
appears a simple loss to the plant; yet if a little pollen were
carried, at first occasionally and then habitually, by the
pollen-devouring insects from flower to flower, and a cross thus
effected, although nine-tenths of the pollen were destroyed, it might
still be a great gain to the plant; and those individuals which
produced more and more pollen, and had larger and larger anthers, would
be selected.
When our plant, by this process of the continued preservation or
natural selection of more and more attractive flowers, had been
rendered highly attractive to insects, they would, unintentionally on
their part, regularly carry pollen from flower to flower; and that they
can most effectually do this, I could easily show by many striking
instances. I will give only one—not as a very striking case, but as
likewise illustrating one step in the separation of the sexes of
plants, presently to be alluded to. Some holly-trees bear only male
flowers, which have four stamens producing rather a small quantity of
pollen, and a rudimentary pistil; other holly-trees bear only female
flowers; these have a full-sized pistil, and four stamens with
shrivelled anthers, in which not a grain of pollen can be detected.
Having found a female tree exactly sixty yards from a male tree, I put
the stigmas of twenty flowers, taken from different branches, under the
microscope, and on all, without exception, there were pollen-grains,
and on some a profusion of pollen. As the wind had set for several days
from the female to the male tree, the pollen could not thus have been
carried. The weather had been cold and boisterous, and therefore not
favourable to bees, nevertheless every female flower which I examined
had been effectually fertilised by the bees, accidentally dusted with
pollen, having flown from tree to tree in search of nectar. But to
return to our imaginary case: as soon as the plant had been rendered so
highly attractive to insects that pollen was regularly carried from
flower to flower, another process might commence. No naturalist doubts
the advantage of what has been called the “physiological division of
labour;” hence we may believe that it would be advantageous to a plant
to produce stamens alone in one flower or on one whole plant, and
pistils alone in
another flower or on another plant. In plants under culture and placed
under new conditions of life, sometimes the male organs and sometimes
the female organs become more or less impotent; now if we suppose this
to occur in ever so slight a degree under nature, then as pollen is
already carried regularly from flower to flower, and as a more complete
separation of the sexes of our plant would be advantageous on the
principle of the division of labour, individuals with this tendency
more and more increased, would be continually favoured or selected,
until at last a complete separation of the sexes would be effected.
Let us now turn to the nectar-feeding insects in our imaginary case: we
may suppose the plant of which we have been slowly increasing the
nectar by continued selection, to be a common plant; and that certain
insects depended in main part on its nectar for food. I could give many
facts, showing how anxious bees are to save time; for instance, their
habit of cutting holes and sucking the nectar at the bases of certain
flowers, which they can, with a very little more trouble, enter by the
mouth. Bearing such facts in mind, I can see no reason to doubt that an
accidental deviation in the size and form of the body, or in the
curvature and length of the proboscis, etc., far too slight to be
appreciated by us, might profit a bee or other insect, so that an
individual so characterised would be able to obtain its food more
quickly, and so have a better chance of living and leaving descendants.
Its descendants would probably inherit a tendency to a similar slight
deviation of structure. The tubes of the corollas of the common red and
incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty
glance appear to differ in length; yet the hive-bee can easily suck the
nectar out of the incarnate clover, but not out of the common red
clover, which is visited by humble-bees alone; so that whole fields of
the red clover offer in vain an abundant supply of precious nectar to
the hive-bee. Thus it might be a great advantage to the hive-bee to
have a slightly longer or differently constructed proboscis. On the
other hand, I have found by experiment that the fertility of clover
greatly depends on bees visiting and moving parts of the corolla, so as
to push the pollen on to the stigmatic surface. Hence, again, if
humble-bees were to become rare in any country, it might be a great
advantage to the red clover to have a shorter or more deeply divided
tube to its corolla, so that the hive-bee could visit its flowers. Thus
I can understand how a flower and a bee might slowly become, either
simultaneously or one after the other, modified and adapted in the most
perfect manner to each other, by the continued preservation of
individuals presenting mutual and slightly favourable deviations of
structure.
I am well aware that this doctrine of natural selection, exemplified in
the above imaginary instances, is open to the same objections which
were at first urged against Sir Charles Lyell’s noble views on “the
modern changes of the earth, as illustrative of geology;” but we now
very seldom hear the action, for instance, of the coast-waves, called a
trifling and insignificant cause, when applied to the excavation of
gigantic valleys or to the formation of the longest lines of inland
cliffs. Natural selection can act only by the preservation and
accumulation of infinitesimally small inherited modifications, each
profitable to the preserved being; and as modern geology has almost
banished such views as the excavation of a great valley by a single
diluvial wave, so will natural selection, if it be a true principle,
banish the belief of the continued creation of new organic
beings, or of any great and sudden modification in their structure.
_On the Intercrossing of Individuals_.—I must here introduce a short
digression. In the case of animals and plants with separated sexes, it
is of course obvious that two individuals must always unite for each
birth; but in the case of hermaphrodites this is far from obvious.
Nevertheless I am strongly inclined to believe that with all
hermaphrodites two individuals, either occasionally or habitually,
concur for the reproduction of their kind. This view, I may add, was
first suggested by Andrew Knight. We shall presently see its
importance; but I must here treat the subject with extreme brevity,
though I have the materials prepared for an ample discussion. All
vertebrate animals, all insects, and some other large groups of
animals, pair for each birth. Modern research has much diminished the
number of supposed hermaphrodites, and of real hermaphrodites a large
number pair; that is, two individuals regularly unite for reproduction,
which is all that concerns us. But still there are many hermaphrodite
animals which certainly do not habitually pair, and a vast majority of
plants are hermaphrodites. What reason, it may be asked, is there for
supposing in these cases that two individuals ever concur in
reproduction? As it is impossible here to enter on details, I must
trust to some general considerations alone.
In the first place, I have collected so large a body of facts, showing,
in accordance with the almost universal belief of breeders, that with
animals and plants a cross between different varieties, or between
individuals of the same variety but of another strain, gives vigour and
fertility to the offspring; and on the other hand, that _close_
interbreeding diminishes vigour and fertility; that
these facts alone incline me to believe that it is a general law of
nature (utterly ignorant though we be of the meaning of the law) that
no organic being self-fertilises itself for an eternity of generations;
but that a cross with another individual is occasionally—perhaps at
very long intervals—indispensable.
On the belief that this is a law of nature, we can, I think, understand
several large classes of facts, such as the following, which on any
other view are inexplicable. Every hybridizer knows how unfavourable
exposure to wet is to the fertilisation of a flower, yet what a
multitude of flowers have their anthers and stigmas fully exposed to
the weather! but if an occasional cross be indispensable, the fullest
freedom for the entrance of pollen from another individual will explain
this state of exposure, more especially as the plant’s own anthers and
pistil generally stand so close together that self-fertilisation seems
almost inevitable. Many flowers, on the other hand, have their organs
of fructification closely enclosed, as in the great papilionaceous or
pea-family; but in several, perhaps in all, such flowers, there is a
very curious adaptation between the structure of the flower and the
manner in which bees suck the nectar; for, in doing this, they either
push the flower’s own pollen on the stigma, or bring pollen from
another flower. So necessary are the visits of bees to papilionaceous
flowers, that I have found, by experiments published elsewhere, that
their fertility is greatly diminished if these visits be prevented.
Now, it is scarcely possible that bees should fly from flower to
flower, and not carry pollen from one to the other, to the great good,
as I believe, of the plant. Bees will act like a camel-hair pencil, and
it is quite sufficient just to touch the anthers of one flower and then
the stigma of another with the same brush to ensure fertilisation; but
it must not be
supposed that bees would thus produce a multitude of hybrids between
distinct species; for if you bring on the same brush a plant’s own
pollen and pollen from another species, the former will have such a
prepotent effect, that it will invariably and completely destroy, as
has been shown by Gärtner, any influence from the foreign pollen.
When the stamens of a flower suddenly spring towards the pistil, or
slowly move one after the other towards it, the contrivance seems
adapted solely to ensure self-fertilisation; and no doubt it is useful
for this end: but, the agency of insects is often required to cause the
stamens to spring forward, as Kölreuter has shown to be the case with
the barberry; and curiously in this very genus, which seems to have a
special contrivance for self-fertilisation, it is well known that if
very closely-allied forms or varieties are planted near each other, it
is hardly possible to raise pure seedlings, so largely do they
naturally cross. In many other cases, far from there being any aids for
self-fertilisation, there are special contrivances, as I could show
from the writings of C. C. Sprengel and from my own observations, which
effectually prevent the stigma receiving pollen from its own flower:
for instance, in Lobelia fulgens, there is a really beautiful and
elaborate contrivance by which every one of the infinitely numerous
pollen-granules are swept out of the conjoined anthers of each flower,
before the stigma of that individual flower is ready to receive them;
and as this flower is never visited, at least in my garden, by insects,
it never sets a seed, though by placing pollen from one flower on the
stigma of another, I raised plenty of seedlings; and whilst another
species of Lobelia growing close by, which is visited by bees, seeds
freely. In very many other cases, though there be no special mechanical
contrivance to prevent the stigma of a flower receiving its own pollen,
yet, as
C. C. Sprengel has shown, and as I can confirm, either the anthers
burst before the stigma is ready for fertilisation, or the stigma is
ready before the pollen of that flower is ready, so that these plants
have in fact separated sexes, and must habitually be crossed. How
strange are these facts! How strange that the pollen and stigmatic
surface of the same flower, though placed so close together, as if for
the very purpose of self-fertilisation, should in so many cases be
mutually useless to each other! How simply are these facts explained on
the view of an occasional cross with a distinct individual being
advantageous or indispensable!
If several varieties of the cabbage, radish, onion, and of some other
plants, be allowed to seed near each other, a large majority, as I have
found, of the seedlings thus raised will turn out mongrels: for
instance, I raised 233 seedling cabbages from some plants of different
varieties growing near each other, and of these only 78 were true to
their kind, and some even of these were not perfectly true. Yet the
pistil of each cabbage-flower is surrounded not only by its own six
stamens, but by those of the many other flowers on the same plant. How,
then, comes it that such a vast number of the seedlings are
mongrelized? I suspect that it must arise from the pollen of a distinct
_variety_ having a prepotent effect over a flower’s own pollen; and
that this is part of the general law of good being derived from the
intercrossing of distinct individuals of the same species. When
distinct _species_ are crossed the case is directly the reverse, for a
plant’s own pollen is always prepotent over foreign pollen; but to this
subject we shall return in a future chapter.
In the case of a gigantic tree covered with innumerable flowers, it may
be objected that pollen could seldom be carried from tree to tree, and
at most only from flower
to flower on the same tree, and that flowers on the same tree can be
considered as distinct individuals only in a limited sense. I believe
this objection to be valid, but that nature has largely provided
against it by giving to trees a strong tendency to bear flowers with
separated sexes. When the sexes are separated, although the male and
female flowers may be produced on the same tree, we can see that pollen
must be regularly carried from flower to flower; and this will give a
better chance of pollen being occasionally carried from tree to tree.
That trees belonging to all Orders have their sexes more often
separated than other plants, I find to be the case in this country; and
at my request Dr. Hooker tabulated the trees of New Zealand, and Dr.
Asa Gray those of the United States, and the result was as I
anticipated. On the other hand, Dr. Hooker has recently informed me
that he finds that the rule does not hold in Australia; and I have made
these few remarks on the sexes of trees simply to call attention to the
subject.
Turning for a very brief space to animals: on the land there are some
hermaphrodites, as land-mollusca and earth-worms; but these all pair.
As yet I have not found a single case of a terrestrial animal which
fertilises itself. We can understand this remarkable fact, which offers
so strong a contrast with terrestrial plants, on the view of an
occasional cross being indispensable, by considering the medium in
which terrestrial animals live, and the nature of the fertilising
element; for we know of no means, analogous to the action of insects
and of the wind in the case of plants, by which an occasional cross
could be effected with terrestrial animals without the concurrence of
two individuals. Of aquatic animals, there are many self-fertilising
hermaphrodites; but here currents in the water offer an obvious means
for an occasional cross. And, as in the case of flowers, I have as yet
failed, after consultation with one of the highest authorities, namely,
Professor Huxley, to discover a single case of an hermaphrodite animal
with the organs of reproduction so perfectly enclosed within the body,
that access from without and the occasional influence of a distinct
individual can be shown to be physically impossible. Cirripedes long
appeared to me to present a case of very great difficulty under this
point of view; but I have been enabled, by a fortunate chance,
elsewhere to prove that two individuals, though both are
self-fertilising hermaphrodites, do sometimes cross.
It must have struck most naturalists as a strange anomaly that, in the
case of both animals and plants, species of the same family and even of
the same genus, though agreeing closely with each other in almost their
whole organisation, yet are not rarely, some of them hermaphrodites,
and some of them unisexual. But if, in fact, all hermaphrodites do
occasionally intercross with other individuals, the difference between
hermaphrodites and unisexual species, as far as function is concerned,
becomes very small.
From these several considerations and from the many special facts which
I have collected, but which I am not here able to give, I am strongly
inclined to suspect that, both in the vegetable and animal kingdoms, an
occasional intercross with a distinct individual is a law of nature. I
am well aware that there are, on this view, many cases of difficulty,
some of which I am trying to investigate. Finally then, we may conclude
that in many organic beings, a cross between two individuals is an
obvious necessity for each birth; in many others it occurs perhaps only
at long intervals; but in none, as I suspect, can self-fertilisation go
on for perpetuity.
_Circumstances favourable to Natural Selection_.—This
is an extremely intricate subject. A large amount of inheritable and
diversified variability is favourable, but I believe mere individual
differences suffice for the work. A large number of individuals, by
giving a better chance for the appearance within any given period of
profitable variations, will compensate for a lesser amount of
variability in each individual, and is, I believe, an extremely
important element of success. Though nature grants vast periods of time
for the work of natural selection, she does not grant an indefinite
period; for as all organic beings are striving, it may be said, to
seize on each place in the economy of nature, if any one species does
not become modified and improved in a corresponding degree with its
competitors, it will soon be exterminated.
In man’s methodical selection, a breeder selects for some definite
object, and free intercrossing will wholly stop his work. But when many
men, without intending to alter the breed, have a nearly common
standard of perfection, and all try to get and breed from the best
animals, much improvement and modification surely but slowly follow
from this unconscious process of selection, notwithstanding a large
amount of crossing with inferior animals. Thus it will be in nature;
for within a confined area, with some place in its polity not so
perfectly occupied as might be, natural selection will always tend to
preserve all the individuals varying in the right direction, though in
different degrees, so as better to fill up the unoccupied place. But if
the area be large, its several districts will almost certainly present
different conditions of life; and then if natural selection be
modifying and improving a species in the several districts, there will
be intercrossing with the other individuals of the same species on the
confines of each. And in this case the effects of intercrossing can
hardly be counterbalanced
by natural selection always tending to modify all the individuals in
each district in exactly the same manner to the conditions of each; for
in a continuous area, the conditions will generally graduate away
insensibly from one district to another. The intercrossing will most
affect those animals which unite for each birth, which wander much, and
which do not breed at a very quick rate. Hence in animals of this
nature, for instance in birds, varieties will generally be confined to
separated countries; and this I believe to be the case. In
hermaphrodite organisms which cross only occasionally, and likewise in
animals which unite for each birth, but which wander little and which
can increase at a very rapid rate, a new and improved variety might be
quickly formed on any one spot, and might there maintain itself in a
body, so that whatever intercrossing took place would be chiefly
between the individuals of the same new variety. A local variety when
once thus formed might subsequently slowly spread to other districts.
On the above principle, nurserymen always prefer getting seed from a
large body of plants of the same variety, as the chance of
intercrossing with other varieties is thus lessened.
Even in the case of slow-breeding animals, which unite for each birth,
we must not overrate the effects of intercrosses in retarding natural
selection; for I can bring a considerable catalogue of facts, showing
that within the same area, varieties of the same animal can long remain
distinct, from haunting different stations, from breeding at slightly
different seasons, or from varieties of the same kind preferring to
pair together.
Intercrossing plays a very important part in nature in keeping the
individuals of the same species, or of the same variety, true and
uniform in character. It will obviously thus act far more efficiently
with those animals
which unite for each birth; but I have already attempted to show that
we have reason to believe that occasional intercrosses take place with
all animals and with all plants. Even if these take place only at long
intervals, I am convinced that the young thus produced will gain so
much in vigour and fertility over the offspring from long-continued
self-fertilisation, that they will have a better chance of surviving
and propagating their kind; and thus, in the long run, the influence of
intercrosses, even at rare intervals, will be great. If there exist
organic beings which never intercross, uniformity of character can be
retained amongst them, as long as their conditions of life remain the
same, only through the principle of inheritance, and through natural
selection destroying any which depart from the proper type; but if
their conditions of life change and they undergo modification,
uniformity of character can be given to their modified offspring,
solely by natural selection preserving the same favourable variations.
Isolation, also, is an important element in the process of natural
selection. In a confined or isolated area, if not very large, the
organic and inorganic conditions of life will generally be in a great
degree uniform; so that natural selection will tend to modify all the
individuals of a varying species throughout the area in the same manner
in relation to the same conditions. Intercrosses, also, with the
individuals of the same species, which otherwise would have inhabited
the surrounding and differently circumstanced districts, will be
prevented. But isolation probably acts more efficiently in checking the
immigration of better adapted organisms, after any physical change,
such as of climate or elevation of the land, etc.; and thus new places
in the natural economy of the country are left open for the old
inhabitants to struggle for, and become adapted to, through
modifications
in their structure and constitution. Lastly, isolation, by checking
immigration and consequently competition, will give time for any new
variety to be slowly improved; and this may sometimes be of importance
in the production of new species. If, however, an isolated area be very
small, either from being surrounded by barriers, or from having very
peculiar physical conditions, the total number of the individuals
supported on it will necessarily be very small; and fewness of
individuals will greatly retard the production of new species through
natural selection, by decreasing the chance of the appearance of
favourable variations.
If we turn to nature to test the truth of these remarks, and look at
any small isolated area, such as an oceanic island, although the total
number of the species inhabiting it, will be found to be small, as we
shall see in our chapter on geographical distribution; yet of these
species a very large proportion are endemic,—that is, have been
produced there, and nowhere else. Hence an oceanic island at first
sight seems to have been highly favourable for the production of new
species. But we may thus greatly deceive ourselves, for to ascertain
whether a small isolated area, or a large open area like a continent,
has been most favourable for the production of new organic forms, we
ought to make the comparison within equal times; and this we are
incapable of doing.
Although I do not doubt that isolation is of considerable importance in
the production of new species, on the whole I am inclined to believe
that largeness of area is of more importance, more especially in the
production of species, which will prove capable of enduring for a long
period, and of spreading widely. Throughout a great and open area, not
only will there be a better chance of favourable variations arising
from the large number of individuals of the same species
there supported, but the conditions of life are infinitely complex from
the large number of already existing species; and if some of these many
species become modified and improved, others will have to be improved
in a corresponding degree or they will be exterminated. Each new form,
also, as soon as it has been much improved, will be able to spread over
the open and continuous area, and will thus come into competition with
many others. Hence more new places will be formed, and the competition
to fill them will be more severe, on a large than on a small and
isolated area. Moreover, great areas, though now continuous, owing to
oscillations of level, will often have recently existed in a broken
condition, so that the good effects of isolation will generally, to a
certain extent, have concurred. Finally, I conclude that, although
small isolated areas probably have been in some respects highly
favourable for the production of new species, yet that the course of
modification will generally have been more rapid on large areas; and
what is more important, that the new forms produced on large areas,
which already have been victorious over many competitors, will be those
that will spread most widely, will give rise to most new varieties and
species, and will thus play an important part in the changing history
of the organic world.
We can, perhaps, on these views, understand some facts which will be
again alluded to in our chapter on geographical distribution; for
instance, that the productions of the smaller continent of Australia
have formerly yielded, and apparently are now yielding, before those of
the larger Europæo-Asiatic area. Thus, also, it is that continental
productions have everywhere become so largely naturalised on islands.
On a small island, the race for life will have been less severe, and
there will have been less modification and less extermination.
Hence, perhaps, it comes that the flora of Madeira, according to Oswald
Heer, resembles the extinct tertiary flora of Europe. All fresh-water
basins, taken together, make a small area compared with that of the sea
or of the land; and, consequently, the competition between fresh-water
productions will have been less severe than elsewhere; new forms will
have been more slowly formed, and old forms more slowly exterminated.
And it is in fresh water that we find seven genera of Ganoid fishes,
remnants of a once preponderant order: and in fresh water we find some
of the most anomalous forms now known in the world, as the
Ornithorhynchus and Lepidosiren, which, like fossils, connect to a
certain extent orders now widely separated in the natural scale. These
anomalous forms may almost be called living fossils; they have endured
to the present day, from having inhabited a confined area, and from
having thus been exposed to less severe competition.
To sum up the circumstances favourable and unfavourable to natural
selection, as far as the extreme intricacy of the subject permits. I
conclude, looking to the future, that for terrestrial productions a
large continental area, which will probably undergo many oscillations
of level, and which consequently will exist for long periods in a
broken condition, will be the most favourable for the production of
many new forms of life, likely to endure long and to spread widely. For
the area will first have existed as a continent, and the inhabitants,
at this period numerous in individuals and kinds, will have been
subjected to very severe competition. When converted by subsidence into
large separate islands, there will still exist many individuals of the
same species on each island: intercrossing on the confines of the range
of each species will thus be checked: after physical changes of any
kind, immigration will be prevented,
so that new places in the polity of each island will have to be filled
up by modifications of the old inhabitants; and time will be allowed
for the varieties in each to become well modified and perfected. When,
by renewed elevation, the islands shall be re-converted into a
continental area, there will again be severe competition: the most
favoured or improved varieties will be enabled to spread: there will be
much extinction of the less improved forms, and the relative
proportional numbers of the various inhabitants of the renewed
continent will again be changed; and again there will be a fair field
for natural selection to improve still further the inhabitants, and
thus produce new species.
That natural selection will always act with extreme slowness, I fully
admit. Its action depends on there being places in the polity of
nature, which can be better occupied by some of the inhabitants of the
country undergoing modification of some kind. The existence of such
places will often depend on physical changes, which are generally very
slow, and on the immigration of better adapted forms having been
checked. But the action of natural selection will probably still
oftener depend on some of the inhabitants becoming slowly modified; the
mutual relations of many of the other inhabitants being thus disturbed.
Nothing can be effected, unless favourable variations occur, and
variation itself is apparently always a very slow process. The process
will often be greatly retarded by free intercrossing. Many will exclaim
that these several causes are amply sufficient wholly to stop the
action of natural selection. I do not believe so. On the other hand, I
do believe that natural selection will always act very slowly, often
only at long intervals of time, and generally on only a very few of the
inhabitants of the same region at the same time. I further believe,
that this very slow, intermittent
action of natural selection accords perfectly well with what geology
tells us of the rate and manner at which the inhabitants of this world
have changed.
Slow though the process of selection may be, if feeble man can do much
by his powers of artificial selection, I can see no limit to the amount
of change, to the beauty and infinite complexity of the coadaptations
between all organic beings, one with another and with their physical
conditions of life, which may be effected in the long course of time by
nature’s power of selection.
_Extinction_.—This subject will be more fully discussed in our chapter
on Geology; but it must be here alluded to from being intimately
connected with natural selection. Natural selection acts solely through
the preservation of variations in some way advantageous, which
consequently endure. But as from the high geometrical powers of
increase of all organic beings, each area is already fully stocked with
inhabitants, it follows that as each selected and favoured form
increases in number, so will the less favoured forms decrease and
become rare. Rarity, as geology tells us, is the precursor to
extinction. We can, also, see that any form represented by few
individuals will, during fluctuations in the seasons or in the number
of its enemies, run a good chance of utter extinction. But we may go
further than this; for as new forms are continually and slowly being
produced, unless we believe that the number of specific forms goes on
perpetually and almost indefinitely increasing, numbers inevitably must
become extinct. That the number of specific forms has not indefinitely
increased, geology shows us plainly; and indeed we can see reason why
they should not have thus increased, for the number of places in the
polity of nature is not indefinitely great,—not that we
have any means of knowing that any one region has as yet got its
maximum of species. Probably no region is as yet fully stocked, for at
the Cape of Good Hope, where more species of plants are crowded
together than in any other quarter of the world, some foreign plants
have become naturalised, without causing, as far as we know, the
extinction of any natives.
Furthermore, the species which are most numerous in individuals will
have the best chance of producing within any given period favourable
variations. We have evidence of this, in the facts given in the second
chapter, showing that it is the common species which afford the
greatest number of recorded varieties, or incipient species. Hence,
rare species will be less quickly modified or improved within any given
period, and they will consequently be beaten in the race for life by
the modified descendants of the commoner species.
From these several considerations I think it inevitably follows, that
as new species in the course of time are formed through natural
selection, others will become rarer and rarer, and finally extinct. The
forms which stand in closest competition with those undergoing
modification and improvement, will naturally suffer most. And we have
seen in the chapter on the Struggle for Existence that it is the most
closely-allied forms,—varieties of the same species, and species of the
same genus or of related genera,—which, from having nearly the same
structure, constitution, and habits, generally come into the severest
competition with each other. Consequently, each new variety or species,
during the progress of its formation, will generally press hardest on
its nearest kindred, and tend to exterminate them. We see the same
process of extermination amongst our domesticated productions, through
the selection of improved forms by man. Many curious
instances could be given showing how quickly new breeds of cattle,
sheep, and other animals, and varieties of flowers, take the place of
older and inferior kinds. In Yorkshire, it is historically known that
the ancient black cattle were displaced by the long-horns, and that
these “were swept away by the short-horns” (I quote the words of an
agricultural writer) “as if by some murderous pestilence.”
_Divergence of Character_.—The principle, which I have designated by
this term, is of high importance on my theory, and explains, as I
believe, several important facts. In the first place, varieties, even
strongly-marked ones, though having somewhat of the character of
species—as is shown by the hopeless doubts in many cases how to rank
them—yet certainly differ from each other far less than do good and
distinct species. Nevertheless, according to my view, varieties are
species in the process of formation, or are, as I have called them,
incipient species. How, then, does the lesser difference between
varieties become augmented into the greater difference between species?
That this does habitually happen, we must infer from most of the
innumerable species throughout nature presenting well-marked
differences; whereas varieties, the supposed prototypes and parents of
future well-marked species, present slight and ill-defined differences.
Mere chance, as we may call it, might cause one variety to differ in
some character from its parents, and the offspring of this variety
again to differ from its parent in the very same character and in a
greater degree; but this alone would never account for so habitual and
large an amount of difference as that between varieties of the same
species and species of the same genus.
As has always been my practice, let us seek light on
this head from our domestic productions. We shall here find something
analogous. A fancier is struck by a pigeon having a slightly shorter
beak; another fancier is struck by a pigeon having a rather longer
beak; and on the acknowledged principle that “fanciers do not and will
not admire a medium standard, but like extremes,” they both go on (as
has actually occurred with tumbler-pigeons) choosing and breeding from
birds with longer and longer beaks, or with shorter and shorter beaks.
Again, we may suppose that at an early period one man preferred swifter
horses; another stronger and more bulky horses. The early differences
would be very slight; in the course of time, from the continued
selection of swifter horses by some breeders, and of stronger ones by
others, the differences would become greater, and would be noted as
forming two sub-breeds; finally, after the lapse of centuries, the
sub-breeds would become converted into two well-established and
distinct breeds. As the differences slowly become greater, the inferior
animals with intermediate characters, being neither very swift nor very
strong, will have been neglected, and will have tended to disappear.
Here, then, we see in man’s productions the action of what may be
called the principle of divergence, causing differences, at first
barely appreciable, steadily to increase, and the breeds to diverge in
character both from each other and from their common parent.
But how, it may be asked, can any analogous principle apply in nature?
I believe it can and does apply most efficiently, from the simple
circumstance that the more diversified the descendants from any one
species become in structure, constitution, and habits, by so much will
they be better enabled to seize on many and widely diversified places
in the polity of nature, and so be enabled to increase in numbers.
We can clearly see this in the case of animals with simple habits. Take
the case of a carnivorous quadruped, of which the number that can be
supported in any country has long ago arrived at its full average. If
its natural powers of increase be allowed to act, it can succeed in
increasing (the country not undergoing any change in its conditions)
only by its varying descendants seizing on places at present occupied
by other animals: some of them, for instance, being enabled to feed on
new kinds of prey, either dead or alive; some inhabiting new stations,
climbing trees, frequenting water, and some perhaps becoming less
carnivorous. The more diversified in habits and structure the
descendants of our carnivorous animal became, the more places they
would be enabled to occupy. What applies to one animal will apply
throughout all time to all animals—that is, if they vary—for otherwise
natural selection can do nothing. So it will be with plants. It has
been experimentally proved, that if a plot of ground be sown with one
species of grass, and a similar plot be sown with several distinct
genera of grasses, a greater number of plants and a greater weight of
dry herbage can thus be raised. The same has been found to hold good
when first one variety and then several mixed varieties of wheat have
been sown on equal spaces of ground. Hence, if any one species of grass
were to go on varying, and those varieties were continually selected
which differed from each other in at all the same manner as distinct
species and genera of grasses differ from each other, a greater number
of individual plants of this species of grass, including its modified
descendants, would succeed in living on the same piece of ground. And
we well know that each species and each variety of grass is annually
sowing almost countless seeds; and thus, as it may be said, is striving
its utmost to increase its numbers. Consequently,
I cannot doubt that in the course of many thousands of generations, the
most distinct varieties of any one species of grass would always have
the best chance of succeeding and of increasing in numbers, and thus of
supplanting the less distinct varieties; and varieties, when rendered
very distinct from each other, take the rank of species.
The truth of the principle, that the greatest amount of life can be
supported by great diversification of structure, is seen under many
natural circumstances. In an extremely small area, especially if freely
open to immigration, and where the contest between individual and
individual must be severe, we always find great diversity in its
inhabitants. For instance, I found that a piece of turf, three feet by
four in size, which had been exposed for many years to exactly the same
conditions, supported twenty species of plants, and these belonged to
eighteen genera and to eight orders, which shows how much these plants
differed from each other. So it is with the plants and insects on small
and uniform islets; and so in small ponds of fresh water. Farmers find
that they can raise most food by a rotation of plants belonging to the
most different orders: nature follows what may be called a simultaneous
rotation. Most of the animals and plants which live close round any
small piece of ground, could live on it (supposing it not to be in any
way peculiar in its nature), and may be said to be striving to the
utmost to live there; but, it is seen, that where they come into the
closest competition with each other, the advantages of diversification
of structure, with the accompanying differences of habit and
constitution, determine that the inhabitants, which thus jostle each
other most closely, shall, as a general rule, belong to what we call
different genera and orders.
The same principle is seen in the naturalisation of
plants through man’s agency in foreign lands. It might have been
expected that the plants which have succeeded in becoming naturalised
in any land would generally have been closely allied to the indigenes;
for these are commonly looked at as specially created and adapted for
their own country. It might, also, perhaps have been expected that
naturalised plants would have belonged to a few groups more especially
adapted to certain stations in their new homes. But the case is very
different; and Alph. De Candolle has well remarked in his great and
admirable work, that floras gain by naturalisation, proportionally with
the number of the native genera and species, far more in new genera
than in new species. To give a single instance: in the last edition of
Dr. Asa Gray’s ‘Manual of the Flora of the Northern United States,’ 260
naturalised plants are enumerated, and these belong to 162 genera. We
thus see that these naturalised plants are of a highly diversified
nature. They differ, moreover, to a large extent from the indigenes,
for out of the 162 genera, no less than 100 genera are not there
indigenous, and thus a large proportional addition is made to the
genera of these States.
By considering the nature of the plants or animals which have struggled
successfully with the indigenes of any country, and have there become
naturalised, we can gain some crude idea in what manner some of the
natives would have had to be modified, in order to have gained an
advantage over the other natives; and we may, I think, at least safely
infer that diversification of structure, amounting to new generic
differences, would have been profitable to them.
The advantage of diversification in the inhabitants of the same region
is, in fact, the same as that of the physiological division of labour
in the organs of the same individual body—a subject so well elucidated
by
Milne Edwards. No physiologist doubts that a stomach by being adapted
to digest vegetable matter alone, or flesh alone, draws most nutriment
from these substances. So in the general economy of any land, the more
widely and perfectly the animals and plants are diversified for
different habits of life, so will a greater number of individuals be
capable of there supporting themselves. A set of animals, with their
organisation but little diversified, could hardly compete with a set
more perfectly diversified in structure. It may be doubted, for
instance, whether the Australian marsupials, which are divided into
groups differing but little from each other, and feebly representing,
as Mr. Waterhouse and others have remarked, our carnivorous, ruminant,
and rodent mammals, could successfully compete with these
well-pronounced orders. In the Australian mammals, we see the process
of diversification in an early and incomplete stage of development.
After the foregoing discussion, which ought to have been much
amplified, we may, I think, assume that the modified descendants of any
one species will succeed by so much the better as they become more
diversified in structure, and are thus enabled to encroach on places
occupied by other beings. Now let us see how this principle of great
benefit being derived from divergence of character, combined with the
principles of natural selection and of extinction, will tend to act.
The accompanying diagram will aid us in understanding this rather
perplexing subject. Let A to L represent the species of a genus large
in its own country; these species are supposed to resemble each other
in unequal degrees, as is so generally the case in nature, and as is
represented in the diagram by the letters standing at unequal
distances. I have said a large genus, because we have seen in the
second chapter,
that on an average more of the species of large genera vary than of
small genera; and the varying species of the large genera present a
greater number of varieties. We have, also, seen that the species,
which are the commonest and the most widely-diffused, vary more than
rare species with restricted ranges. Let (A) be a common,
widely-diffused, and varying species, belonging to a genus large in its
own country. The little fan of diverging dotted lines of unequal
lengths proceeding from (A), may represent its varying offspring. The
variations are supposed to be extremely slight, but of the most
diversified nature; they are not supposed all to appear simultaneously,
but often after long intervals of time; nor are they all supposed to
endure for equal periods. Only those variations which are in some way
profitable will be preserved or naturally selected. And here the
importance of the principle of benefit being derived from divergence of
character comes in; for this will generally lead to the most different
or divergent variations (represented by the outer dotted lines) being
preserved and accumulated by natural selection. When a dotted line
reaches one of the horizontal lines, and is there marked by a small
numbered letter, a sufficient amount of variation is supposed to have
been accumulated to have formed a fairly well-marked variety, such as
would be thought worthy of record in a systematic work.
The intervals between the horizontal lines in the diagram, may
represent each a thousand generations; but it would have been better if
each had represented ten thousand generations. After a thousand
generations, species (A) is supposed to have produced two fairly
well-marked varieties, namely _a_1 and _m_1. These two varieties will
generally continue to be exposed to the same conditions which made
their parents variable,
and the tendency to variability is in itself hereditary, consequently
they will tend to vary, and generally to vary in nearly the same manner
as their parents varied. Moreover, these two varieties, being only
slightly modified forms, will tend to inherit those advantages which
made their common parent (A) more numerous than most of the other
inhabitants of the same country; they will likewise partake of those
more general advantages which made the genus to which the
parent-species belonged, a large genus in its own country. And these
circumstances we know to be favourable to the production of new
varieties.
If, then, these two varieties be variable, the most divergent of their
variations will generally be preserved during the next thousand
generations. And after this interval, variety _a_1 is supposed in the
diagram to have produced variety _a_2, which will, owing to the
principle of divergence, differ more from (A) than did variety _a_1.
Variety _m_1 is supposed to have produced two varieties, namely _m_2
and _s_2, differing from each other, and more considerably from their
common parent (A). We may continue the process by similar steps for any
length of time; some of the varieties, after each thousand generations,
producing only a single variety, but in a more and more modified
condition, some producing two or three varieties, and some failing to
produce any. Thus the varieties or modified descendants, proceeding
from the common parent (A), will generally go on increasing in number
and diverging in character. In the diagram the process is represented
up to the ten-thousandth generation, and under a condensed and
simplified form up to the fourteen-thousandth generation.
But I must here remark that I do not suppose that the process ever goes
on so regularly as is represented in the diagram, though in itself made
somewhat irregular.
I am far from thinking that the most divergent varieties will
invariably prevail and multiply: a medium form may often long endure,
and may or may not produce more than one modified descendant; for
natural selection will always act according to the nature of the places
which are either unoccupied or not perfectly occupied by other beings;
and this will depend on infinitely complex relations. But as a general
rule, the more diversified in structure the descendants from any one
species can be rendered, the more places they will be enabled to seize
on, and the more their modified progeny will be increased. In our
diagram the line of succession is broken at regular intervals by small
numbered letters marking the successive forms which have become
sufficiently distinct to be recorded as varieties. But these breaks are
imaginary, and might have been inserted anywhere, after intervals long
enough to have allowed the accumulation of a considerable amount of
divergent variation.
As all the modified descendants from a common and widely-diffused
species, belonging to a large genus, will tend to partake of the same
advantages which made their parent successful in life, they will
generally go on multiplying in number as well as diverging in
character: this is represented in the diagram by the several divergent
branches proceeding from (A). The modified offspring from the later and
more highly improved branches in the lines of descent, will, it is
probable, often take the place of, and so destroy, the earlier and less
improved branches: this is represented in the diagram by some of the
lower branches not reaching to the upper horizontal lines. In some
cases I do not doubt that the process of modification will be confined
to a single line of descent, and the number of the descendants will not
be increased; although the amount
of divergent modification may have been increased in the successive
generations. This case would be represented in the diagram, if all the
lines proceeding from (A) were removed, excepting that from _a_1 to
_a_10. In the same way, for instance, the English race-horse and
English pointer have apparently both gone on slowly diverging in
character from their original stocks, without either having given off
any fresh branches or races.
After ten thousand generations, species (A) is supposed to have
produced three forms, _a_10, _f_10, and _m_10, which, from having
diverged in character during the successive generations, will have come
to differ largely, but perhaps unequally, from each other and from
their common parent. If we suppose the amount of change between each
horizontal line in our diagram to be excessively small, these three
forms may still be only well-marked varieties; or they may have arrived
at the doubtful category of sub-species; but we have only to suppose
the steps in the process of modification to be more numerous or greater
in amount, to convert these three forms into well-defined species: thus
the diagram illustrates the steps by which the small differences
distinguishing varieties are increased into the larger differences
distinguishing species. By continuing the same process for a greater
number of generations (as shown in the diagram in a condensed and
simplified manner), we get eight species, marked by the letters between
_a_14 and _m_14, all descended from (A). Thus, as I believe, species
are multiplied and genera are formed.
In a large genus it is probable that more than one species would vary.
In the diagram I have assumed that a second species (I) has produced,
by analogous steps, after ten thousand generations, either two
well-marked varieties (_w_10 and _z_10) or two species, according to
the amount of change supposed to be represented between
the horizontal lines. After fourteen thousand generations, six new
species, marked by the letters _n_14 to _z_14, are supposed to have
been produced. In each genus, the species, which are already extremely
different in character, will generally tend to produce the greatest
number of modified descendants; for these will have the best chance of
filling new and widely different places in the polity of nature: hence
in the diagram I have chosen the extreme species (A), and the nearly
extreme species (I), as those which have largely varied, and have given
rise to new varieties and species. The other nine species (marked by
capital letters) of our original genus, may for a long period continue
transmitting unaltered descendants; and this is shown in the diagram by
the dotted lines not prolonged far upwards from want of space.
But during the process of modification, represented in the diagram,
another of our principles, namely that of extinction, will have played
an important part. As in each fully stocked country natural selection
necessarily acts by the selected form having some advantage in the
struggle for life over other forms, there will be a constant tendency
in the improved descendants of any one species to supplant and
exterminate in each stage of descent their predecessors and their
original parent. For it should be remembered that the competition will
generally be most severe between those forms which are most nearly
related to each other in habits, constitution, and structure. Hence all
the intermediate forms between the earlier and later states, that is
between the less and more improved state of a species, as well as the
original parent-species itself, will generally tend to become extinct.
So it probably will be with many whole collateral lines of descent,
which will be conquered by later and improved lines of descent. If,
however, the
modified offspring of a species get into some distinct country, or
become quickly adapted to some quite new station, in which child and
parent do not come into competition, both may continue to exist.
If then our diagram be assumed to represent a considerable amount of
modification, species (A) and all the earlier varieties will have
become extinct, having been replaced by eight new species (_a_14 to
_m_14); and (I) will have been replaced by six (_n_14 to _z_14) new
species.
But we may go further than this. The original species of our genus were
supposed to resemble each other in unequal degrees, as is so generally
the case in nature; species (A) being more nearly related to B, C, and
D, than to the other species; and species (I) more to G, H, K, L, than
to the others. These two species (A) and (I), were also supposed to be
very common and widely diffused species, so that they must originally
have had some advantage over most of the other species of the genus.
Their modified descendants, fourteen in number at the
fourteen-thousandth generation, will probably have inherited some of
the same advantages: they have also been modified and improved in a
diversified manner at each stage of descent, so as to have become
adapted to many related places in the natural economy of their country.
It seems, therefore, to me extremely probable that they will have taken
the places of, and thus exterminated, not only their parents (A) and
(I), but likewise some of the original species which were most nearly
related to their parents. Hence very few of the original species will
have transmitted offspring to the fourteen-thousandth generation. We
may suppose that only one (F), of the two species which were least
closely related to the other nine original species, has transmitted
descendants to this late stage of descent.
The new species in our diagram descended from the original eleven
species, will now be fifteen in number. Owing to the divergent tendency
of natural selection, the extreme amount of difference in character
between species _a_14 and _z_14 will be much greater than that between
the most different of the original eleven species. The new species,
moreover, will be allied to each other in a widely different manner. Of
the eight descendants from (A) the three marked _a_14, _q_14, _p_14,
will be nearly related from having recently branched off from _a_10;
_b_14 and _f_14, from having diverged at an earlier period from _a_5,
will be in some degree distinct from the three first-named species; and
lastly, _o_14, _e_14, and _m_14, will be nearly related one to the
other, but from having diverged at the first commencement of the
process of modification, will be widely different from the other five
species, and may constitute a sub-genus or even a distinct genus.
The six descendants from (I) will form two sub-genera or even genera.
But as the original species (I) differed largely from (A), standing
nearly at the extreme points of the original genus, the six descendants
from (I) will, owing to inheritance, differ considerably from the eight
descendants from (A); the two groups, moreover, are supposed to have
gone on diverging in different directions. The intermediate species,
also (and this is a very important consideration), which connected the
original species (A) and (I), have all become, excepting (F), extinct,
and have left no descendants. Hence the six new species descended from
(I), and the eight descended from (A), will have to be ranked as very
distinct genera, or even as distinct sub-families.
Thus it is, as I believe, that two or more genera are produced by
descent, with modification, from two or more species of the same genus.
And the two or more
parent-species are supposed to have descended from some one species of
an earlier genus. In our diagram, this is indicated by the broken
lines, beneath the capital letters, converging in sub-branches
downwards towards a single point; this point representing a single
species, the supposed single parent of our several new sub-genera and
genera.
It is worth while to reflect for a moment on the character of the new
species F14, which is supposed not to have diverged much in character,
but to have retained the form of (F), either unaltered or altered only
in a slight degree. In this case, its affinities to the other fourteen
new species will be of a curious and circuitous nature. Having
descended from a form which stood between the two parent-species (A)
and (I), now supposed to be extinct and unknown, it will be in some
degree intermediate in character between the two groups descended from
these species. But as these two groups have gone on diverging in
character from the type of their parents, the new species (F14) will
not be directly intermediate between them, but rather between types of
the two groups; and every naturalist will be able to bring some such
case before his mind.
In the diagram, each horizontal line has hitherto been supposed to
represent a thousand generations, but each may represent a million or
hundred million generations, and likewise a section of the successive
strata of the earth’s crust including extinct remains. We shall, when
we come to our chapter on Geology, have to refer again to this subject,
and I think we shall then see that the diagram throws light on the
affinities of extinct beings, which, though generally belonging to the
same orders, or families, or genera, with those now living, yet are
often, in some degree, intermediate in character between existing
groups; and we can understand this fact, for
the extinct species lived at very ancient epochs when the branching
lines of descent had diverged less.
I see no reason to limit the process of modification, as now explained,
to the formation of genera alone. If, in our diagram, we suppose the
amount of change represented by each successive group of diverging
dotted lines to be very great, the forms marked _a_14 to _p_14, those
marked _b_14 and _f_14, and those marked _o_14 to _m_14, will form
three very distinct genera. We shall also have two very distinct genera
descended from (I) and as these latter two genera, both from continued
divergence of character and from inheritance from a different parent,
will differ widely from the three genera descended from (A), the two
little groups of genera will form two distinct families, or even
orders, according to the amount of divergent modification supposed to
be represented in the diagram. And the two new families, or orders,
will have descended from two species of the original genus; and these
two species are supposed to have descended from one species of a still
more ancient and unknown genus.
We have seen that in each country it is the species of the larger
genera which oftenest present varieties or incipient species. This,
indeed, might have been expected; for as natural selection acts through
one form having some advantage over other forms in the struggle for
existence, it will chiefly act on those which already have some
advantage; and the largeness of any group shows that its species have
inherited from a common ancestor some advantage in common. Hence, the
struggle for the production of new and modified descendants, will
mainly lie between the larger groups, which are all trying to increase
in number. One large group will slowly conquer another large group,
reduce its numbers, and thus lessen its chance of further variation and
improvement. Within the same large
group, the later and more highly perfected sub-groups, from branching
out and seizing on many new places in the polity of Nature, will
constantly tend to supplant and destroy the earlier and less improved
sub-groups. Small and broken groups and sub-groups will finally tend to
disappear. Looking to the future, we can predict that the groups of
organic beings which are now large and triumphant, and which are least
broken up, that is, which as yet have suffered least extinction, will
for a long period continue to increase. But which groups will
ultimately prevail, no man can predict; for we well know that many
groups, formerly most extensively developed, have now become extinct.
Looking still more remotely to the future, we may predict that, owing
to the continued and steady increase of the larger groups, a multitude
of smaller groups will become utterly extinct, and leave no modified
descendants; and consequently that of the species living at any one
period, extremely few will transmit descendants to a remote futurity. I
shall have to return to this subject in the chapter on Classification,
but I may add that on this view of extremely few of the more ancient
species having transmitted descendants, and on the view of all the
descendants of the same species making a class, we can understand how
it is that there exist but very few classes in each main division of
the animal and vegetable kingdoms. Although extremely few of the most
ancient species may now have living and modified descendants, yet at
the most remote geological period, the earth may have been as well
peopled with many species of many genera, families, orders, and
classes, as at the present day.
_Summary of the Chapter_.—If during the long course of ages and under
varying conditions of life, organic beings
vary at all in the several parts of their organisation, and I think
this cannot be disputed; if there be, owing to the high geometrical
powers of increase of each species, at some age, season, or year, a
severe struggle for life, and this certainly cannot be disputed; then,
considering the infinite complexity of the relations of all organic
beings to each other and to their conditions of existence, causing an
infinite diversity in structure, constitution, and habits, to be
advantageous to them, I think it would be a most extraordinary fact if
no variation ever had occurred useful to each being’s own welfare, in
the same way as so many variations have occurred useful to man. But if
variations useful to any organic being do occur, assuredly individuals
thus characterised will have the best chance of being preserved in the
struggle for life; and from the strong principle of inheritance they
will tend to produce offspring similarly characterised. This principle
of preservation, I have called, for the sake of brevity, Natural
Selection. Natural selection, on the principle of qualities being
inherited at corresponding ages, can modify the egg, seed, or young, as
easily as the adult. Amongst many animals, sexual selection will give
its aid to ordinary selection, by assuring to the most vigorous and
best adapted males the greatest number of offspring. Sexual selection
will also give characters useful to the males alone, in their struggles
with other males.
Whether natural selection has really thus acted in nature, in modifying
and adapting the various forms of life to their several conditions and
stations, must be judged of by the general tenour and balance of
evidence given in the following chapters. But we already see how it
entails extinction; and how largely extinction has acted in the world’s
history, geology plainly declares. Natural selection, also, leads to
divergence of
character; for more living beings can be supported on the same area the
more they diverge in structure, habits, and constitution, of which we
see proof by looking at the inhabitants of any small spot or at
naturalised productions. Therefore during the modification of the
descendants of any one species, and during the incessant struggle of
all species to increase in numbers, the more diversified these
descendants become, the better will be their chance of succeeding in
the battle of life. Thus the small differences distinguishing varieties
of the same species, will steadily tend to increase till they come to
equal the greater differences between species of the same genus, or
even of distinct genera.
We have seen that it is the common, the widely-diffused, and
widely-ranging species, belonging to the larger genera, which vary
most; and these will tend to transmit to their modified offspring that
superiority which now makes them dominant in their own countries.
Natural selection, as has just been remarked, leads to divergence of
character and to much extinction of the less improved and intermediate
forms of life. On these principles, I believe, the nature of the
affinities of all organic beings may be explained. It is a truly
wonderful fact—the wonder of which we are apt to overlook from
familiarity—that all animals and all plants throughout all time and
space should be related to each other in group subordinate to group, in
the manner which we everywhere behold—namely, varieties of the same
species most closely related together, species of the same genus less
closely and unequally related together, forming sections and
sub-genera, species of distinct genera much less closely related, and
genera related in different degrees, forming sub-families, families,
orders, sub-classes, and classes. The several subordinate groups in any
class cannot be
ranked in a single file, but seem rather to be clustered round points,
and these round other points, and so on in almost endless cycles. On
the view that each species has been independently created, I can see no
explanation of this great fact in the classification of all organic
beings; but, to the best of my judgment, it is explained through
inheritance and the complex action of natural selection, entailing
extinction and divergence of character, as we have seen illustrated in
the diagram.
The affinities of all the beings of the same class have sometimes been
represented by a great tree. I believe this simile largely speaks the
truth. The green and budding twigs may represent existing species; and
those produced during each former year may represent the long
succession of extinct species. At each period of growth all the growing
twigs have tried to branch out on all sides, and to overtop and kill
the surrounding twigs and branches, in the same manner as species and
groups of species have tried to overmaster other species in the great
battle for life. The limbs divided into great branches, and these into
lesser and lesser branches, were themselves once, when the tree was
small, budding twigs; and this connexion of the former and present buds
by ramifying branches may well represent the classification of all
extinct and living species in groups subordinate to groups. Of the many
twigs which flourished when the tree was a mere bush, only two or
three, now grown into great branches, yet survive and bear all the
other branches; so with the species which lived during long-past
geological periods, very few now have living and modified descendants.
From the first growth of the tree, many a limb and branch has decayed
and dropped off; and these lost branches of various sizes may represent
those whole orders, families, and genera which have now no living
representatives, and
which are known to us only from having been found in a fossil state. As
we here and there see a thin straggling branch springing from a fork
low down in a tree, and which by some chance has been favoured and is
still alive on its summit, so we occasionally see an animal like the
Ornithorhynchus or Lepidosiren, which in some small degree connects by
its affinities two large branches of life, and which has apparently
been saved from fatal competition by having inhabited a protected
station. As buds give rise by growth to fresh buds, and these, if
vigorous, branch out and overtop on all sides many a feebler branch, so
by generation I believe it has been with the great Tree of Life, which
fills with its dead and broken branches the crust of the earth, and
covers the surface with its ever branching and beautiful ramifications.