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On the Origin of Species, 6th Edition by Charles Darwin

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intermittent results accord well with what geology tells us of the rate and
manner at which the inhabitants of the world have changed.

Slow though the process of selection may be, if feeble man can do much by
artificial selection, I can see no limit to the amount of change, to the
beauty and complexity of the coadaptations between all organic beings, one
with another and with their physical conditions of life, which may have
been effected in the long course of time through nature's power of
selection, that is by the survival of the fittest.


This subject will be more fully discussed in our chapter on Geology; but it
must here be 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. Owing to
the high geometrical rate of increase of all organic beings, each area is
already fully stocked with inhabitants, and it follows from this, that as
the favoured forms increase in number, so, generally, will the less
favoured decrease and become rare. Rarity, as geology tells us, is the
precursor to extinction. We can see that any form which is represented by
few individuals will run a good chance of utter extinction, during great
fluctuations in the nature or the seasons, or from a temporary increase in
the number of its enemies. But we may go further than this; for as new
forms are produced, unless we admit that specific forms can go on
indefinitely increasing in number, many old forms must become extinct.
That the number of specific forms has not indefinitely increased, geology
plainly tells us; and we shall presently attempt to show why it is that the
number of species throughout the world has not become immeasurably great.

We have seen that the species which are most numerous in individuals have
the best chance of producing favourable variations within any given period.
We have evidence of this, in the facts stated in the second chapter,
showing that it is the common and diffused or dominant species which offer
the greatest number of recorded varieties. Hence, rare species will be
less quickly modified or improved within any given period; they will
consequently be beaten in the race for life by the modified and improved
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
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 among 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."


The principle, which I have designated by this term, is of high importance,
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 far less from each other 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 a degree of difference as that between the species of
the same genus.

As has always been my practice, I have sought light on this head from our
domestic productions. We shall here find something analogous. It will be
admitted that the production of races so different as short-horn and
Hereford cattle, race and cart horses, the several breeds of pigeons, etc.,
could never have been effected by the mere chance accumulation of similar
variations during many successive generations. In practice, a fancier is,
for instance, 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 the sub-breeds of the tumbler-pigeon) 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 of history, the men of one nation or
district required swifter horses, while those of another required stronger
and bulkier horses. The early differences would be very slight; but, in
the course of time, from the continued selection of swifter horses in the
one case, and of stronger ones in the other, the differences would become
greater, and would be noted as forming two sub-breeds. Ultimately after
the lapse of centuries, these sub-breeds would become converted into two
well-established and distinct breeds. As the differences became greater,
the inferior animals with intermediate characters, being neither very swift
nor very strong, would not have been used for breeding, and will thus 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 (though it was a long time
before I saw how), 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 discern 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 power of increase be allowed to act, it can succeed in increasing
(the country not undergoing any change in 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 animals become, the
more places they will 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 effect 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 be raised in the latter than in the former case. The same has
been found to hold good when one variety and 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 the varieties were continually selected
which differed from each other in the same manner, though in a very slight
degree, as do the distinct species and genera of grasses, a greater number
of individual plants of this species, including its modified descendants,
would succeed in living on the same piece of ground. And we know that each
species and each variety of grass is annually sowing almost countless
seeds; and is thus striving, as it may be said, to the utmost to increase
in number. Consequently, in the course of many thousand generations, the
most distinct varieties of any one species of grass would 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 very 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: also in
small ponds of fresh water. Farmers find that they can raise more 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 its nature not to be in any way peculiar), 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, 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

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
would succeed 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 naturalised genera, no less than 100
genera are not there indigenous, and thus a large proportional addition is
made to the genera now living in the United States.

By considering the nature of the plants or animals which have in any
country struggled successfully with the indigenes, and have there become
naturalised, we may gain some crude idea in what manner some of the natives
would have had to be modified in order to gain an advantage over their
compatriots; and we may at least infer that diversification of structure,
amounting to new generic differences, would be profitable to them.

The advantage of diversification of structure 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-developed orders. In the Australian
mammals, we see the process of diversification in an early and incomplete
stage of development.


After the foregoing discussion, which has been much compressed, we may
assume that the modified descendants of any one species will succeed 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 benefit being derived from divergence of character,
combined with the principles of natural selection and of extinction, tends
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 as we saw in the second chapter, on an
average more species vary in large genera than in 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 most
widely-diffused, vary more than do the rare and restricted species. Let
(A) be a common, widely-diffused, and varying species, belonging to a genus
large in its own country. The branching and 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 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 form it into 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 or more generations. After a thousand generations, species
(A) is supposed to have produced two fairly well-marked varieties, namely
a1 and m1. These two varieties will generally still be exposed to the same
conditions which made their parents variable, and the tendency to
variability is in itself hereditary; consequently they will likewise tend
to vary, and commonly in nearly the same manner as did their parents.
Moreover, these two varieties, being only slightly modified forms, will
tend to inherit those advantages which made their parent (A) more numerous
than most of the other inhabitants of the same country; they will also
partake of those more general advantages which made the genus to which the
parent-species belonged, a large genus in its own country. And all these
circumstances are 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 a1 is supposed in the
diagram to have produced variety a2, which will, owing to the principle of
divergence, differ more from (A) than did variety a1. Variety m1 is
supposed to have produced two varieties, namely m2 and s2, 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 of 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, nor that it goes on continuously; it is far more
probable that each form remains for long periods unaltered, and then again
undergoes modification. Nor do I suppose that the most divergent varieties
are invariably preserved: 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 increase. 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 allow 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 no doubt the process of
modification will be confined to a single line of descent, and the number
of modified descendants will not be increased; although the amount of
divergent modification may have been augmented. This case would be
represented in the diagram, if all the lines proceeding from (A) were
removed, excepting that from a1 to a10. In the same way the English
racehorse 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, a10, f10, and m10, 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; 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 doubtful or at least 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 a14 and m14, 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 (w10 and z10) 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 n14 to z14,
are supposed to have been produced. In any genus, the species which are
already very different in character from each other, will generally tend to
produce the greatest number of modified descendants; for these will have
the best chance of seizing on 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 long but unequal
periods continue to transmit unaltered descendants; and this is shown in
the diagram by the dotted lines unequally prolonged upwards.

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 progenitor. 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 states of a the
same 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. If,
however, the modified offspring of a species get into some distinct
country, or become quickly adapted to some quite new station, in which
offspring and progenitor do not come into competition, both may continue to

If, then, our diagram be assumed to represent a considerable amount of
modification, species (A) and all the earlier varieties will have become
extinct, being replaced by eight new species (a14 to m14); and species (I)
will be replaced by six (n14 to z14) 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, 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 (E and F) 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
a14 and z14 will be much greater than that between the most distinct 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 a14, q14, p14, will be nearly related from having recently
branched off from a10; b14 and f14, from having diverged at an earlier
period from a5, will be in some degree distinct from the three first-named
species; and lastly, o14, e14, and m14, 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 a distinct genus.

The six descendants from (I) will form two sub-genera or genera. But as
the original species (I) differed largely from (A), standing nearly at the
extreme end of the original genus, the six descendants from (I) will, owing
to inheritance alone, 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, except (F), extinct, and have left no descendants. Hence
the six new species descended from (I), and the eight descendants from (A),
will have to be ranked as very distinct genera, or even as distinct

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 be 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 represents a species, the supposed progenitor 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. Being descended from a
form that stood between the 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 two 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 call such cases 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 more
generations; it may also represent 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, 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 various remote 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 the diagram, we suppose the amount
of change represented by each successive group of diverging dotted lines to
be great, the forms marked a14 to p14, those marked b14 and f14, and those
marked o14 to m14, will form three very distinct genera. We shall also
have two very distinct genera descended from (I), differing widely from the
descendants of (A). These two groups of genera will thus form two distinct
families, or orders, according to the amount of divergent modification
supposed to be represented in the diagram. And the two new families, or
orders, are descended from two species of the original genus; and these are
supposed to be descended from some still more ancient and unknown form.

We have seen that in each country it is the species belonging to 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 number, 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 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 have as
yet suffered least extinction, will, for a long period, continue to
increase. But which groups will ultimately prevail, no man can predict;
for we 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 as, according to this view, extremely few of the more
ancient species have transmitted descendants to the present day, and, as
all the descendants of the same species form a class, we can understand how
it is that there exist so few classes in each main division of the animal
and vegetable kingdoms. Although few of the most ancient species have left
modified descendants, yet, at remote geological periods, the earth may have
been almost as well peopled with species of many genera, families, orders
and classes, as at the present day.


Natural selection acts exclusively by the preservation and accumulation of
variations, which are beneficial under the organic and inorganic conditions
to which each creature is exposed at all periods of life. The ultimate
result is that each creature tends to become more and more improved in
relation to its conditions. This improvement inevitably leads to the
gradual advancement of the organisation of the greater number of living
beings throughout the world. But here we enter on a very intricate
subject, for naturalists have not defined to each other's satisfaction what
is meant by an advance in organisation. Among the vertebrata the degree of
intellect and an approach in structure to man clearly come into play. It
might be thought that the amount of change which the various parts and
organs pass through in their development from embryo to maturity would
suffice as a standard of comparison; but there are cases, as with certain
parasitic crustaceans, in which several parts of the structure become less
perfect, so that the mature animal cannot be called higher than its larva.
Von Baer's standard seems the most widely applicable and the best, namely,
the amount of differentiation of the parts of the same organic being, in
the adult state, as I should be inclined to add, and their specialisation
for different functions; or, as Milne Edwards would express it, the
completeness of the division of physiological labour. But we shall see how
obscure this subject is if we look, for instance, to fishes, among which
some naturalists rank those as highest which, like the sharks, approach
nearest to amphibians; while other naturalists rank the common bony or
teleostean fishes as the highest, inasmuch as they are most strictly fish-
like, and differ most from the other vertebrate classes. We see still more
plainly the obscurity of the subject by turning to plants, among which the
standard of intellect is of course quite excluded; and here some botanists
rank those plants as highest which have every organ, as sepals, petals,
stamens and pistils, fully developed in each flower; whereas other
botanists, probably with more truth, look at the plants which have their
several organs much modified and reduced in number as the highest.

If we take as the standard of high organisation, the amount of
differentiation and specialisation of the several organs in each being when
adult (and this will include the advancement of the brain for intellectual
purposes), natural selection clearly leads towards this standard: for all
physiologists admit that the specialisation of organs, inasmuch as in this
state they perform their functions better, is an advantage to each being;
and hence the accumulation of variations tending towards specialisation is
within the scope of natural selection. On the other hand, we can see,
bearing in mind that all organic beings are striving to increase at a high
ratio and to seize on every unoccupied or less well occupied place in the
economy of nature, that it is quite possible for natural selection
gradually to fit a being to a situation in which several organs would be
superfluous or useless: in such cases there would be retrogression in the
scale of organisation. Whether organisation on the whole has actually
advanced from the remotest geological periods to the present day will be
more conveniently discussed in our chapter on Geological Succession.

But it may be objected that if all organic beings thus tend to rise in the
scale, how is it that throughout the world a multitude of the lowest forms
still exist; and how is it that in each great class some forms are far more
highly developed than others? Why have not the more highly developed forms
every where supplanted and exterminated the lower? Lamarck, who believed
in an innate and inevitable tendency towards perfection in all organic
beings, seems to have felt this difficulty so strongly that he was led to
suppose that new and simple forms are continually being produced by
spontaneous generation. Science has not as yet proved the truth of this
belief, whatever the future may reveal. On our theory the continued
existence of lowly organisms offers no difficulty; for natural selection,
or the survival of the fittest, does not necessarily include progressive
development--it only takes advantage of such variations as arise and are
beneficial to each creature under its complex relations of life. And it
may be asked what advantage, as far as we can see, would it be to an
infusorian animalcule--to an intestinal worm--or even to an earth-worm, to
be highly organised. If it were no advantage, these forms would be left,
by natural selection, unimproved or but little improved, and might remain
for indefinite ages in their present lowly condition. And geology tells us
that some of the lowest forms, as the infusoria and rhizopods, have
remained for an enormous period in nearly their present state. But to
suppose that most of the many now existing low forms have not in the least
advanced since the first dawn of life would be extremely rash; for every
naturalist who has dissected some of the beings now ranked as very low in
the scale, must have been struck with their really wondrous and beautiful

Nearly the same remarks are applicable, if we look to the different grades
of organisation within the same great group; for instance, in the
vertebrata, to the co-existence of mammals and fish--among mammalia, to the
co-existence of man and the ornithorhynchus--among fishes, to the co-
existence of the shark and the lancelet (Amphioxus), which latter fish in
the extreme simplicity of its structure approaches the invertebrate
classes. But mammals and fish hardly come into competition with each
other; the advancement of the whole class of mammals, or of certain members
in this class, to the highest grade would not lead to their taking the
place of fishes. Physiologists believe that the brain must be bathed by
warm blood to be highly active, and this requires aerial respiration; so
that warm-blooded mammals when inhabiting the water lie under a
disadvantage in having to come continually to the surface to breathe. With
fishes, members of the shark family would not tend to supplant the
lancelet; for the lancelet, as I hear from Fritz Muller, has as sole
companion and competitor on the barren sandy shore of South Brazil, an
anomalous annelid. The three lowest orders of mammals, namely, marsupials,
edentata, and rodents, co-exist in South America in the same region with
numerous monkeys, and probably interfere little with each other. Although
organisation, on the whole, may have advanced and be still advancing
throughout the world, yet the scale will always present many degrees of
perfection; for the high advancement of certain whole classes, or of
certain members of each class, does not at all necessarily lead to the
extinction of those groups with which they do not enter into close
competition. In some cases, as we shall hereafter see, lowly organised
forms appear to have been preserved to the present day, from inhabiting
confined or peculiar stations, where they have been subjected to less
severe competition, and where their scanty numbers have retarded the chance
of favourable variations arising.

Finally, I believe that many lowly organised forms now exist throughout the
world, from various causes. In some cases variations or individual
differences of a favourable nature may never have arisen for natural
selection to act on and accumulate. In no case, probably, has time
sufficed for the utmost possible amount of development. In some few cases
there has been what we must call retrogression or organisation. But the
main cause lies in the fact that under very simple conditions of life a
high organisation would be of no service--possibly would be of actual
disservice, as being of a more delicate nature, and more liable to be put
out of order and injured.

Looking to the first dawn of life, when all organic beings, as we may
believe, presented the simplest structure, how, it has been asked, could
the first step in the advancement or differentiation of parts have arisen?
Mr. Herbert Spencer would probably answer that, as soon as simple
unicellular organisms came by growth or division to be compounded of
several cells, or became attached to any supporting surface, his law "that
homologous units of any order become differentiated in proportion as their
relations to incident forces become different" would come into action. But
as we have no facts to guide us, speculation on the subject is almost
useless. It is, however, an error to suppose that there would be no
struggle for existence, and, consequently, no natural selection, until many
forms had been produced: variations in a single species inhabiting an
isolated station might be beneficial, and thus the whole mass of
individuals might be modified, or two distinct forms might arise. But, as
I remarked towards the close of the introduction, no one ought to feel
surprise at much remaining as yet unexplained on the origin of species, if
we make due allowance for our profound ignorance on the mutual relations of
the inhabitants of the world at the present time, and still more so during
past ages.


Mr. H.C. Watson thinks that I have overrated the importance of divergence
of character (in which, however, he apparently believes), and that
convergence, as it may be called, has likewise played a part. If two
species belonging to two distinct though allied genera, had both produced a
large number of new and divergent forms, it is conceivable that these might
approach each other so closely that they would have all to be classed under
the same genus; and thus the descendants of two distinct genera would
converge into one. But it would in most cases be extremely rash to
attribute to convergence a close and general similarity of structure in the
modified descendants of widely distinct forms. The shape of a crystal is
determined solely by the molecular forces, and it is not surprising that
dissimilar substances should sometimes assume the same form; but with
organic beings we should bear in mind that the form of each depends on an
infinitude of complex relations, namely on the variations which have
arisen, these being due to causes far too intricate to be followed out--on
the nature of the variations which have been preserved or selected, and
this depends on the surrounding physical conditions, and in a still higher
degree on the surrounding organisms with which each being has come into
competition--and lastly, on inheritance (in itself a fluctuating element)
from innumerable progenitors, all of which have had their forms determined
through equally complex relations. It is incredible that the descendants
of two organisms, which had originally differed in a marked manner, should
ever afterwards converge so closely as to lead to a near approach to
identity throughout their whole organisation. If this had occurred, we
should meet with the same form, independently of genetic connection,
recurring in widely separated geological formations; and the balance of
evidence is opposed to any such an admission.

Mr. Watson has also objected that the continued action of natural
selection, together with divergence of character, would tend to make an
indefinite number of specific forms. As far as mere inorganic conditions
are concerned, it seems probable that a sufficient number of species would
soon become adapted to all considerable diversities of heat, moisture,
etc.; but I fully admit that the mutual relations of organic beings are
more important; and as the number of species in any country goes on
increasing, the organic conditions of life must become more and more
complex. Consequently there seems at first no limit to the amount of
profitable diversification of structure, and therefore no limit to the
number of species which might be produced. We do not know that even the
most prolific area is fully stocked with specific forms: at the Cape of
Good Hope and in Australia, which support such an astonishing number of
species, many European plants have become naturalised. But geology shows
us, that from an early part of the tertiary period the number of species of
shells, and that from the middle part of this same period, the number of
mammals has not greatly or at all increased. What then checks an
indefinite increase in the number of species? The amount of life (I do not
mean the number of specific forms) supported on an area must have a limit,
depending so largely as it does on physical conditions; therefore, if an
area be inhabited by very many species, each or nearly each species will be
represented by few individuals; and such species will be liable to
extermination from accidental fluctuations in the nature of the seasons or
in the number of their enemies. The process of extermination in such cases
would be rapid, whereas the production of new species must always be slow.
Imagine the extreme case of as many species as individuals in England, and
the first severe winter or very dry summer would exterminate thousands on
thousands of species. Rare species, and each species will become rare if
the number of species in any country becomes indefinitely increased, will,
on the principal often explained, present within a given period few
favourable variations; consequently, the process of giving birth to new
specific forms would thus be retarded. When any species becomes very rare,
close interbreeding will help to exterminate it; authors have thought that
this comes into play in accounting for the deterioration of the aurochs in
Lithuania, of red deer in Scotland and of bears in Norway, etc. Lastly,
and this I am inclined to think is the most important element, a dominant
species, which has already beaten many competitors in its own home, will
tend to spread and supplant many others. Alph. de Candolle has shown that
those species which spread widely tend generally to spread VERY widely,
consequently they will tend to supplant and exterminate several species in
several areas, and thus check the inordinate increase of specific forms
throughout the world. Dr. Hooker has recently shown that in the southeast
corner of Australia, where, apparently, there are many invaders from
different quarters of the globe, the endemic Australian species have been
greatly reduced in number. How much weight to attribute to these several
considerations I will not pretend to say; but conjointly they must limit in
each country the tendency to an indefinite augmentation of specific forms.


If under changing conditions of life organic beings present individual
differences in almost every part of their structure, and this cannot be
disputed; if there be, owing to their geometrical rate of increase, a
severe struggle for life at some age, season or year, 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
life, causing an infinite diversity in structure, constitution, and habits,
to be advantageous to them, it would be a most extraordinary fact if no
variations had ever occurred useful to each being's own welfare, in the
same manner as so many variations have occurred useful to man. But if
variations useful to any organic being ever 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, these will
tend to produce offspring similarly characterised. This principle of
preservation, or the survival of the fittest, I have called natural
selection. It leads to the improvement of each creature in relation to its
organic and inorganic conditions of life; and consequently, in most cases,
to what must be regarded as an advance in organisation. Nevertheless, low
and simple forms will long endure if well fitted for their simple
conditions of life.

Natural selection, on the principle of qualities being inherited at
corresponding ages, can modify the egg, seed, or young as easily as the
adult. Among many animals sexual selection will have given 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 or rivalry with
other males; and these characters will be transmitted to one sex or to both
sexes, according to the form of inheritance which prevails.

Whether natural selection has really thus acted in adapting the various
forms of life to their several conditions and stations, must be judged by
the general tenour and balance of evidence given in the following chapters.
But we have already seen 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 the more
organic beings diverge in structure, habits and constitution, by so much
the more can a large number be supported on the area, of which we see proof
by looking to the inhabitants of any small spot, and to the productions
naturalised in foreign lands. 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 the descendants
become, the better will be their chance of success in the battle for life.
Thus the small differences distinguishing varieties of the same species,
steadily tend to increase, till they 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 within each class, which vary most;
and these 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, the nature of the affinities, and the generally well defined
distinctions between the innumerable organic beings in each class
throughout the world, 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 groups, subordinate to groups, in the manner which we everywhere
behold--namely, varieties of the same species most closely related, species
of the same genus less closely and unequally related, 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 clustered round points,
and these round other points, and so on in almost endless cycles. If
species had been independently created, no explanation would have been
possible of this kind of classification; but 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

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 former years 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 at
all times overmastered 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 young, 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 the other branches; so with the species which lived during
long-past geological periods, very few have left living and modified
descendants. From the first growth of the tree, many a limb and branch has
decayed and dropped off; and these fallen 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 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.



Effects of changed conditions -- Use and disuse, combined with natural
selection; organs of flight and of vision -- Acclimatisation -- Correlated
variation -- Compensation and economy of growth -- False correlations --
Multiple, rudimentary, and lowly organised structures variable -- Parts
developed in an unusual manner are highly variable: specific characters
more variable than generic: secondary sexual characters variable --
Species of the same genus vary in an analogous manner -- Reversions to
long-lost characters -- Summary.

I have hitherto sometimes spoken as if the variations--so common and
multiform with organic beings under domestication, and in a lesser degree
with those under nature--were due to chance. This, of course is a wholly
incorrect expression, but it serves to acknowledge plainly our ignorance of
the cause of each particular variation. Some authors believe it to be as
much the function of the reproductive system to produce individual
differences, or slight deviations of structure, as to make the child like
its parents. But the fact of variations and monstrosities occurring much
more frequently under domestication than under nature, and the greater
variability of species having wide ranges than of those with restricted
ranges, lead to the conclusion that variability is generally related to the
conditions of life to which each species has been exposed during several
successive generations. In the first chapter I attempted to show that
changed conditions act in two ways, directly on the whole organisation or
on certain parts alone, and indirectly through the reproductive system. In
all cases there are two factors, the nature of the organism, which is much
the most important of the two, and the nature of the conditions. The
direct action of changed conditions leads to definite or indefinite
results. In the latter case the organisation seems to become plastic, and
we have much fluctuating variability. In the former case the nature of the
organism is such that it yields readily, when subjected to certain
conditions, and all, or nearly all, the individuals become modified in the
same way.

It is very difficult to decide how far changed conditions, such as of
climate, food, etc., have acted in a definite manner. There is reason to
believe that in the course of time the effects have been greater than can
be proved by clear evidence. But we may safely conclude that the
innumerable complex co-adaptations of structure, which we see throughout
nature between various organic beings, cannot be attributed simply to such
action. In the following cases the conditions seem to have produced some
slight definite effect: E. Forbes asserts that shells at their southern
limit, and when living in shallow water, are more brightly coloured than
those of the same species from further north or from a greater depth; but
this certainly does not always hold good. Mr. Gould believes that birds of
the same species are more brightly coloured under a clear atmosphere, than
when living near the coast or on islands; and Wollaston is convinced that
residence near the sea affects the colours of insects. Moquin-Tandon gives
a list of plants which, when growing near the sea-shore, have their leaves
in some degree fleshy, though not elsewhere fleshy. These slightly varying
organisms are interesting in as far as they present characters analogous to
those possessed by the species which are confined to similar conditions.

When a variation is of the slightest use to any being, we cannot tell how
much to attribute to the accumulative action of natural selection, and how
much to the definite action of the conditions of life. Thus, it is well
known to furriers that animals of the same species have thicker and better
fur the further north they live; but who can tell how much of this
difference may be due to the warmest-clad individuals having been favoured
and preserved during many generations, and how much to the action of the
severe climate? For it would appear that climate has some direct action on
the hair of our domestic quadrupeds.

Instances could be given of similar varieties being produced from the same
species under external conditions of life as different as can well be
conceived; and, on the other hand, of dissimilar varieties being produced
under apparently the same external conditions. Again, innumerable
instances are known to every naturalist, of species keeping true, or not
varying at all, although living under the most opposite climates. Such
considerations as these incline me to lay less weight on the direct action
of the surrounding conditions, than on a tendency to vary, due to causes of
which we are quite ignorant.

In one sense the conditions of life may be said, not only to cause
variability, either directly or indirectly, but likewise to include natural
selection, for the conditions determine whether this or that variety shall
survive. But when man is the selecting agent, we clearly see that the two
elements of change are distinct; variability is in some manner excited, but
it is the will of man which accumulates the variations in certain
direction; and it is this latter agency which answers to the survival of
the fittest under nature.


>From the facts alluded to in the first chapter, I think there can be no
doubt that use in our domestic animals has strengthened and enlarged
certain parts, and disuse diminished them; and that such modifications are
inherited. Under free nature we have no standard of comparison by which to
judge of the effects of long-continued use or disuse, for we know not the
parent-forms; but many animals possess structures which can be best
explained by the effects of disuse. As Professor Owen has remarked, there
is no greater anomaly in nature than a bird that cannot fly; yet there are
several in this state. The logger-headed duck of South America can only
flap along the surface of the water, and has its wings in nearly the same
condition as the domestic Aylesbury duck: it is a remarkable fact that the
young birds, according to Mr. Cunningham, can fly, while the adults have
lost this power. As the larger ground-feeding birds seldom take flight
except to escape danger, it is probable that the nearly wingless condition
of several birds, now inhabiting or which lately inhabited several oceanic
islands, tenanted by no beasts of prey, has been caused by disuse. The
ostrich indeed inhabits continents, and is exposed to danger from which it
cannot escape by flight, but it can defend itself, by kicking its enemies,
as efficiently as many quadrupeds. We may believe that the progenitor of
the ostrich genus had habits like those of the bustard, and that, as the
size and weight of its body were increased during successive generations,
its legs were used more and its wings less, until they became incapable of

Kirby has remarked (and I have observed the same fact) that the anterior
tarsi, or feet, of many male dung-feeding beetles are often broken off; he
examined seventeen specimens in his own collection, and not one had even a
relic left. In the Onites apelles the tarsi are so habitually lost that
the insect has been described as not having them. In some other genera
they are present, but in a rudimentary condition. In the Ateuchus or
sacred beetle of the Egyptians, they are totally deficient. The evidence
that accidental mutilations can be inherited is at present not decisive;
but the remarkable cases observed by Brown-Sequard in guinea-pigs, of the
inherited effects of operations, should make us cautious in denying this
tendency. Hence, it will perhaps be safest to look at the entire absence
of the anterior tarsi in Ateuchus, and their rudimentary condition in some
other genera, not as cases of inherited mutilations, but as due to the
effects of long-continued disuse; for as many dung-feeding beetles are
generally found with their tarsi lost, this must happen early in life;
therefore the tarsi cannot be of much importance or be much used by these

In some cases we might easily put down to disuse modifications of structure
which are wholly, or mainly due to natural selection. Mr. Wollaston has
discovered the remarkable fact that 200 beetles, out of the 550 species
(but more are now known) inhabiting Madeira, are so far deficient in wings
that they cannot fly; and that, of the twenty-nine endemic genera, no less
than twenty-three have all their species in this condition! Several facts,
namely, that beetles in many parts of the world are very frequently blown
to sea and perish; that the beetles in Madeira, as observed by Mr.
Wollaston, lie much concealed, until the wind lulls and the sun shines;
that the proportion of wingless beetles is larger on the exposed Desertas
than in Madeira itself; and especially the extraordinary fact, so strongly
insisted on by Mr. Wollaston, that certain large groups of beetles,
elsewhere excessively numerous, which absolutely require the use of their
wings, are here almost entirely absent. These several considerations make
me believe that the wingless condition of so many Madeira beetles is mainly
due to the action of natural selection, combined probably with disuse. For
during many successive generations each individual beetle which flew least,
either from its wings having been ever so little less perfectly developed
or from indolent habit, will have had the best chance of surviving from not
being blown out to sea; and, on the other hand, those beetles which most
readily took to flight would oftenest have been blown to sea, and thus

The insects in Madeira which are not ground-feeders, and which, as certain
flower-feeding coleoptera and lepidoptera, must habitually use their wings
to gain their subsistence, have, as Mr. Wollaston suspects, their wings not
at all reduced, but even enlarged. This is quite compatible with the
action of natural selection. For when a new insect first arrived on the
island, the tendency of natural selection to enlarge or to reduce the
wings, would depend on whether a greater number of individuals were saved
by successfully battling with the winds, or by giving up the attempt and
rarely or never flying. As with mariners shipwrecked near a coast, it
would have been better for the good swimmers if they had been able to swim
still further, whereas it would have been better for the bad swimmers if
they had not been able to swim at all and had stuck to the wreck.

The eyes of moles and of some burrowing rodents are rudimentary in size,
and in some cases are quite covered by skin and fur. This state of the
eyes is probably due to gradual reduction from disuse, but aided perhaps by
natural selection. In South America, a burrowing rodent, the tuco-tuco, or
Ctenomys, is even more subterranean in its habits than the mole; and I was
assured by a Spaniard, who had often caught them, that they were frequently
blind. One which I kept alive was certainly in this condition, the cause,
as appeared on dissection, having been inflammation of the nictitating
membrane. As frequent inflammation of the eyes must be injurious to any
animal, and as eyes are certainly not necessary to animals having
subterranean habits, a reduction in their size, with the adhesion of the
eyelids and growth of fur over them, might in such case be an advantage;
and if so, natural selection would aid the effects of disuse.

It is well known that several animals, belonging to the most different
classes, which inhabit the caves of Carniola and Kentucky, are blind. In
some of the crabs the foot-stalk for the eye remains, though the eye is
gone; the stand for the telescope is there, though the telescope with its
glasses has been lost. As it is difficult to imagine that eyes, though
useless, could be in any way injurious to animals living in darkness, their
loss may be attributed to disuse. In one of the blind animals, namely, the
cave-rat (Neotoma), two of which were captured by Professor Silliman at
above half a mile distance from the mouth of the cave, and therefore not in
the profoundest depths, the eyes were lustrous and of large size; and these
animals, as I am informed by Professor Silliman, after having been exposed
for about a month to a graduated light, acquired a dim perception of

It is difficult to imagine conditions of life more similar than deep
limestone caverns under a nearly similar climate; so that, in accordance
with the old view of the blind animals having been separately created for
the American and European caverns, very close similarity in their
organisation and affinities might have been expected. This is certainly
not the case if we look at the two whole faunas; with respect to the
insects alone, Schiodte has remarked: "We are accordingly prevented from
considering the entire phenomenon in any other light than something purely
local, and the similarity which is exhibited in a few forms between the
Mammoth Cave (in Kentucky) and the caves in Carniola, otherwise than as a
very plain expression of that analogy which subsists generally between the
fauna of Europe and of North America." On my view we must suppose that
American animals, having in most cases ordinary powers of vision, slowly
migrated by successive generations from the outer world into the deeper and
deeper recesses of the Kentucky caves, as did European animals into the
caves of Europe. We have some evidence of this gradation of habit; for, as
Schiodte remarks: "We accordingly look upon the subterranean faunas as
small ramifications which have penetrated into the earth from the
geographically limited faunas of the adjacent tracts, and which, as they
extended themselves into darkness, have been accommodated to surrounding
circumstances. Animals not far remote from ordinary forms, prepare the
transition from light to darkness. Next follow those that are constructed
for twilight; and, last of all, those destined for total darkness, and
whose formation is quite peculiar." These remarks of Schiodte's it should
be understood, apply not to the same, but to distinct species. By the time
that an animal had reached, after numberless generations, the deepest
recesses, disuse will on this view have more or less perfectly obliterated
its eyes, and natural selection will often have effected other changes,
such as an increase in the length of the antennae or palpi, as a
compensation for blindness. Notwithstanding such modifications, we might
expect still to see in the cave-animals of America, affinities to the other
inhabitants of that continent, and in those of Europe to the inhabitants of
the European continent. And this is the case with some of the American
cave-animals, as I hear from Professor Dana; and some of the European
cave-insects are very closely allied to those of the surrounding country.
It would be difficult to give any rational explanation of the affinities of
the blind cave-animals to the other inhabitants of the two continents on
the ordinary view of their independent creation. That several of the
inhabitants of the caves of the Old and New Worlds should be closely
related, we might expect from the well-known relationship of most of their
other productions. As a blind species of Bathyscia is found in abundance
on shady rocks far from caves, the loss of vision in the cave species of
this one genus has probably had no relation to its dark habitation; for it
is natural that an insect already deprived of vision should readily become
adapted to dark caverns. Another blind genus (Anophthalmus) offers this
remarkable peculiarity, that the species, as Mr. Murray observes, have not
as yet been found anywhere except in caves; yet those which inhabit the
several caves of Europe and America are distinct; but it is possible that
the progenitors of these several species, while they were furnished with
eyes, may formerly have ranged over both continents, and then have become
extinct, excepting in their present secluded abodes. Far from feeling
surprise that some of the cave-animals should be very anomalous, as Agassiz
has remarked in regard to the blind fish, the Amblyopsis, and as is the
case with the blind Proteus, with reference to the reptiles of Europe, I am
only surprised that more wrecks of ancient life have not been preserved,
owing to the less severe competition to which the scanty inhabitants of
these dark abodes will have been exposed.


Habit is hereditary with plants, as in the period of flowering, in the time
of sleep, in the amount of rain requisite for seeds to germinate, etc., and
this leads me to say a few words on acclimatisation. As it is extremely
common for distinct species belonging to the same genus to inhabit hot and
cold countries, if it be true that all the species of the same genus are
descended from a single parent-form, acclimatisation must be readily
effected during a long course of descent. It is notorious that each
species is adapted to the climate of its own home: species from an arctic
or even from a temperate region cannot endure a tropical climate, or
conversely. So again, many succulent plants cannot endure a damp climate.
But the degree of adaptation of species to the climates under which they
live is often overrated. We may infer this from our frequent inability to
predict whether or not an imported plant will endure our climate, and from
the number of plants and animals brought from different countries which are
here perfectly healthy. We have reason to believe that species in a state
of nature are closely limited in their ranges by the competition of other
organic beings quite as much as, or more than, by adaptation to particular
climates. But whether or not this adaptation is in most cases very close,
we have evidence with some few plants, of their becoming, to a certain
extent, naturally habituated to different temperatures; that is, they
become acclimatised: thus the pines and rhododendrons, raised from seed
collected by Dr. Hooker from the same species growing at different heights
on the Himalayas, were found to possess in this country different
constitutional powers of resisting cold. Mr. Thwaites informs me that he
has observed similar facts in Ceylon; analogous observations have been made
by Mr. H.C. Watson on European species of plants brought from the Azores to
England; and I could give other cases. In regard to animals, several
authentic instances could be adduced of species having largely extended,
within historical times, their range from warmer to colder latitudes, and
conversely; but we do not positively know that these animals were strictly
adapted to their native climate, though in all ordinary cases we assume
such to be the case; nor do we know that they have subsequently become
specially acclimatised to their new homes, so as to be better fitted for
them than they were at first.

As we may infer that our domestic animals were originally chosen by
uncivilised man because they were useful, and because they bred readily
under confinement, and not because they were subsequently found capable of
far-extended transportation, the common and extraordinary capacity in our
domestic animals of not only withstanding the most different climates, but
of being perfectly fertile (a far severer test) under them, may be used as
an argument that a large proportion of other animals now in a state of
nature could easily be brought to bear widely different climates. We must
not, however, push the foregoing argument too far, on account of the
probable origin of some of our domestic animals from several wild stocks:
the blood, for instance, of a tropical and arctic wolf may perhaps be
mingled in our domestic breeds. The rat and mouse cannot be considered as
domestic animals, but they have been transported by man to many parts of
the world, and now have a far wider range than any other rodent; for they
live under the cold climate of Faroe in the north and of the Falklands in
the south, and on many an island in the torrid zones. Hence adaptation to
any special climate may be looked at as a quality readily grafted on an
innate wide flexibility of constitution, common to most animals. On this
view, the capacity of enduring the most different climates by man himself
and by his domestic animals, and the fact of the extinct elephant and
rhinoceros having formerly endured a glacial climate, whereas the living
species are now all tropical or sub-tropical in their habits, ought not to
be looked at as anomalies, but as examples of a very common flexibility of
constitution, brought, under peculiar circumstances, into action.

How much of the acclimatisation of species to any peculiar climate is due
to mere habit, and how much to the natural selection of varieties having
different innate constitutions, and how much to both means combined, is an
obscure question. That habit or custom has some influence, I must believe,
both from analogy and from the incessant advice given in agricultural
works, even in the ancient Encyclopaedias of China, to be very cautious in
transporting animals from one district to another. And as it is not likely
that man should have succeeded in selecting so many breeds and sub-breeds
with constitutions specially fitted for their own districts, the result
must, I think, be due to habit. On the other hand, natural selection would
inevitably tend to preserve those individuals which were born with
constitutions best adapted to any country which they inhabited. In
treatises on many kinds of cultivated plants, certain varieties are said to
withstand certain climates better than others; this is strikingly shown in
works on fruit-trees published in the United States, in which certain
varieties are habitually recommended for the northern and others for the
southern states; and as most of these varieties are of recent origin, they
cannot owe their constitutional differences to habit. The case of the
Jerusalem artichoke, which is never propagated in England by seed, and of
which, consequently, new varieties have not been produced, has even been
advanced, as proving that acclimatisation cannot be effected, for it is now
as tender as ever it was! The case, also, of the kidney-bean has been
often cited for a similar purpose, and with much greater weight; but until
some one will sow, during a score of generations, his kidney-beans so early
that a very large proportion are destroyed by frost, and then collect seed
from the few survivors, with care to prevent accidental crosses, and then
again get seed from these seedlings, with the same precautions, the
experiment cannot be said to have been even tried. Nor let it be supposed
that differences in the constitution of seedling kidney-beans never appear,
for an account has been published how much more hardy some seedlings are
than others; and of this fact I have myself observed striking instances.

On the whole, we may conclude that habit, or use and disuse, have, in some
cases, played a considerable part in the modification of the constitution
and structure; but that the effects have often been largely combined with,
and sometimes overmastered by, the natural selection of innate variations.


I mean by this expression that the whole organisation is so tied together,
during its growth and development, that when slight variations in any one
part occur and are accumulated through natural selection, other parts
become modified. This is a very important subject, most imperfectly
understood, and no doubt wholly different classes of facts may be here
easily confounded together. We shall presently see that simple inheritance
often gives the false appearance of correlation. One of the most obvious
real cases is, that variations of structure arising in the young or larvae
naturally tend to affect the structure of the mature animal. The several
parts which are homologous, and which, at an early embryonic period, are
identical in structure, and which are necessarily exposed to similar
conditions, seem eminently liable to vary in a like manner: we see this in
the right and left sides of the body varying in the same manner; in the
front and hind legs, and even in the jaws and limbs, varying together, for
the lower jaw is believed by some anatomists to be homologous with the
limbs. These tendencies, I do not doubt, may be mastered more or less
completely by natural selection: thus a family of stags once existed with
an antler only on one side; and if this had been of any great use to the
breed, it might probably have been rendered permanent by natural selection.

Homologous parts, as has been remarked by some authors, tend to cohere;
this is often seen in monstrous plants: and nothing is more common than
the union of homologous parts in normal structures, as in the union of the
petals into a tube. Hard parts seem to affect the form of adjoining soft
parts; it is believed by some authors that with birds the diversity in the
shape of the pelvis causes the remarkable diversity in the shape of the
kidneys. Others believe that the shape of the pelvis in the human mother
influences by pressure the shape of the head of the child. In snakes,
according to Schlegel, the shape of the body and the manner of swallowing
determine the position and form of several of the most important viscera.

The nature of the bond is frequently quite obscure. M. Is. Geoffroy St.
Hilaire has forcibly remarked that certain malconformations frequently, and
that others rarely, coexist without our being able to assign any reason.
What can be more singular than the relation in cats between complete
whiteness and blue eyes with deafness, or between the tortoise-shell colour
and the female sex; or in pigeons, between their feathered feet and skin
betwixt the outer toes, or between the presence of more or less down on the
young pigeon when first hatched, with the future colour of its plumage; or,
again, the relation between the hair and the teeth in the naked Turkish
dog, though here no doubt homology comes into play? With respect to this
latter case of correlation, I think it can hardly be accidental that the
two orders of mammals which are most abnormal in their dermal covering,
viz., Cetacea (whales) and Edentata (armadilloes, scaly ant-eaters, etc.),
are likewise on the whole the most abnormal in their teeth, but there are
so many exceptions to this rule, as Mr. Mivart has remarked, that it has
little value.

I know of no case better adapted to show the importance of the laws of
correlation and variation, independently of utility, and therefore of
natural selection, than that of the difference between the outer and inner
flowers in some Compositous and Umbelliferous plants. Everyone is familiar
with the difference between the ray and central florets of, for instance,
the daisy, and this difference is often accompanied with the partial or
complete abortion of the reproductive organs. But in some of these plants
the seeds also differ in shape and sculpture. These differences have
sometimes been attributed to the pressure of the involucra on the florets,
or to their mutual pressure, and the shape of the seeds in the ray-florets
of some Compositae countenances this idea; but with the Umbelliferae it is
by no means, as Dr. Hooker informs me, the species with the densest heads
which most frequently differ in their inner and outer flowers. It might
have been thought that the development of the ray-petals, by drawing
nourishment from the reproductive organs causes their abortion; but this
can hardly be the sole case, for in some Compositae the seeds of the outer
and inner florets differ, without any difference in the corolla. Possibly
these several differences may be connected with the different flow of
nutriment towards the central and external flowers. We know, at least,
that with irregular flowers those nearest to the axis are most subject to
peloria, that is to become abnormally symmetrical. I may add, as an
instance of this fact, and as a striking case of correlation, that in many
pelargoniums the two upper petals in the central flower of the truss often
lose their patches of darker colour; and when this occurs, the adherent
nectary is quite aborted, the central flower thus becoming peloric or
regular. When the colour is absent from only one of the two upper petals,
the nectary is not quite aborted but is much shortened.

With respect to the development of the corolla, Sprengel's idea that the
ray-florets serve to attract insects, whose agency is highly advantageous,
or necessary for the fertilisation of these plants, is highly probable; and
if so, natural selection may have come into play. But with respect to the
seeds, it seems impossible that their differences in shape, which are not
always correlated with any difference in the corolla, can be in any way
beneficial; yet in the Umbelliferae these differences are of such apparent
importance--the seeds being sometimes orthospermous in the exterior flowers
and coelospermous in the central flowers--that the elder De Candolle
founded his main divisions in the order on such characters. Hence
modifications of structure, viewed by systematists as of high value, may be
wholly due to the laws of variation and correlation, without being, as far
as we can judge, of the slightest service to the species.

We may often falsely attribute to correlated variation structures which are
common to whole groups of species, and which in truth are simply due to
inheritance; for an ancient progenitor may have acquired through natural
selection some one modification in structure, and, after thousands of
generations, some other and independent modification; and these two
modifications, having been transmitted to a whole group of descendants with
diverse habits, would naturally be thought to be in some necessary manner
correlated. Some other correlations are apparently due to the manner in
which natural selection can alone act. For instance, Alph. De Candolle has
remarked that winged seeds are never found in fruits which do not open; I
should explain this rule by the impossibility of seeds gradually becoming
winged through natural selection, unless the capsules were open; for in
this case alone could the seeds, which were a little better adapted to be
wafted by the wind, gain an advantage over others less well fitted for wide


The elder Geoffroy and Goethe propounded, at about the same time, their law
of compensation or balancement of growth; or, as Goethe expressed it, "in
order to spend on one side, nature is forced to economise on the other
side." I think this holds true to a certain extent with our domestic
productions: if nourishment flows to one part or organ in excess, it
rarely flows, at least in excess, to another part; thus it is difficult to
get a cow to give much milk and to fatten readily. The same varieties of
the cabbage do not yield abundant and nutritious foliage and a copious
supply of oil-bearing seeds. When the seeds in our fruits become
atrophied, the fruit itself gains largely in size and quality. In our
poultry, a large tuft of feathers on the head is generally accompanied by a
diminished comb, and a large beard by diminished wattles. With species in
a state of nature it can hardly be maintained that the law is of universal
application; but many good observers, more especially botanists, believe in
its truth. I will not, however, here give any instances, for I see hardly
any way of distinguishing between the effects, on the one hand, of a part
being largely developed through natural selection and another and adjoining
part being reduced by the same process or by disuse, and, on the other
hand, the actual withdrawal of nutriment from one part owing to the excess
of growth in another and adjoining part.

I suspect, also, that some of the cases of compensation which have been
advanced, and likewise some other facts, may be merged under a more general
principle, namely, that natural selection is continually trying to
economise in every part of the organisation. If under changed conditions
of life a structure, before useful, becomes less useful, its diminution
will be favoured, for it will profit the individual not to have its
nutriment wasted in building up a useless structure. I can thus only
understand a fact with which I was much struck when examining cirripedes,
and of which many other instances could be given: namely, that when a
cirripede is parasitic within another cirripede and is thus protected, it
loses more or less completely its own shell or carapace. This is the case
with the male Ibla, and in a truly extraordinary manner with the
Proteolepas: for the carapace in all other cirripedes consists of the
three highly important anterior segments of the head enormously developed,
and furnished with great nerves and muscles; but in the parasitic and
protected Proteolepas, the whole anterior part of the head is reduced to
the merest rudiment attached to the bases of the prehensile antennae. Now
the saving of a large and complex structure, when rendered superfluous,
would be a decided advantage to each successive individual of the species;
for in the struggle for life to which every animal is exposed, each would
have a better chance of supporting itself, by less nutriment being wasted.

Thus, as I believe, natural selection will tend in the long run to reduce
any part of the organisation, as soon as it becomes, through changed
habits, superfluous, without by any means causing some other part to be
largely developed in a corresponding degree. And conversely, that natural
selection may perfectly well succeed in largely developing an organ without
requiring as a necessary compensation the reduction of some adjoining part.


It seems to be a rule, as remarked by Is. Geoffroy St. Hilaire, both with
varieties and species, that when any part or organ is repeated many times
in the same individual (as the vertebrae in snakes, and the stamens in
polyandrous flowers) the number is variable; whereas the number of the same
part or organ, when it occurs in lesser numbers, is constant. The same
author as well as some botanists, have further remarked that multiple parts
are extremely liable to vary in structure. As "vegetative repetition," to
use Professor Owen's expression, is a sign of low organisation; the
foregoing statements accord with the common opinion of naturalists, that
beings which stand low in the scale of nature are more variable than those
which are higher. I presume that lowness here means that the several parts
of the organisation have been but little specialised for particular
functions; and as long as the same part has to perform diversified work, we
can perhaps see why it should remain variable, that is, why natural
selection should not have preserved or rejected each little deviation of
form so carefully as when the part has to serve for some one special
purpose. In the same way that a knife which has to cut all sorts of things
may be of almost any shape; whilst a tool for some particular purpose must
be of some particular shape. Natural selection, it should never be
forgotten, can act solely through and for the advantage of each being.

Rudimentary parts, as is generally admitted, are apt to be highly variable.
We shall have to recur to this subject; and I will here only add that their
variability seems to result from their uselessness, and consequently from
natural selection having had no power to check deviations in their


Several years ago I was much struck by a remark to the above effect made by
Mr. Waterhouse. Professor Owen, also, seems to have come to a nearly
similar conclusion. It is hopeless to attempt to convince any one of the
truth of the above proposition without giving the long array of facts which
I have collected, and which cannot possibly be here introduced. I can only
state my conviction that it is a rule of high generality. I am aware of
several causes of error, but I hope that I have made due allowances for
them. It should be understood that the rule by no means applies to any
part, however unusually developed, unless it be unusually developed in one
species or in a few species in comparison with the same part in many
closely allied species. Thus, the wing of the bat is a most abnormal
structure in the class of mammals; but the rule would not apply here,
because the whole group of bats possesses wings; it would apply only if
some one species had wings developed in a remarkable manner in comparison
with the other species of the same genus. The rule applies very strongly
in the case of secondary sexual characters, when displayed in any unusual
manner. The term, secondary sexual characters, used by Hunter, relates to
characters which are attached to one sex, but are not directly connected
with the act of reproduction. The rule applies to males and females; but
more rarely to females, as they seldom offer remarkable secondary sexual
characters. The rule being so plainly applicable in the case of secondary
sexual characters, may be due to the great variability of these characters,
whether or not displayed in any unusual manner--of which fact I think there
can be little doubt. But that our rule is not confined to secondary sexual
characters is clearly shown in the case of hermaphrodite cirripedes; I
particularly attended to Mr. Waterhouse's remark, whilst investigating this
order, and I am fully convinced that the rule almost always holds good. I
shall, in a future work, give a list of all the more remarkable cases. I
will here give only one, as it illustrates the rule in its largest
application. The opercular valves of sessile cirripedes (rock barnacles)
are, in every sense of the word, very important structures, and they differ
extremely little even in distinct genera; but in the several species of one
genus, Pyrgoma, these valves present a marvellous amount of
diversification; the homologous valves in the different species being
sometimes wholly unlike in shape; and the amount of variation in the
individuals of the same species is so great that it is no exaggeration to
state that the varieties of the same species differ more from each other in
the characters derived from these important organs, than do the species
belonging to other distinct genera.

As with birds the individuals of the same species, inhabiting the same
country, vary extremely little, I have particularly attended to them; and
the rule certainly seems to hold good in this class. I cannot make out
that it applies to plants, and this would have seriously shaken my belief
in its truth, had not the great variability in plants made it particularly
difficult to compare their relative degrees of variability.

When we see any part or organ developed in a remarkable degree or manner in
a species, the fair presumption is that it is of high importance to that
species: nevertheless it is in this case eminently liable to variation.
Why should this be so? On the view that each species has been
independently created, with all its parts as we now see them, I can see no
explanation. But on the view that groups of species are descended from
some other species, and have been modified through natural selection, I
think we can obtain some light. First let me make some preliminary
remarks. If, in our domestic animals, any part or the whole animal be
neglected, and no selection be applied, that part (for instance, the comb
in the Dorking fowl) or the whole breed will cease to have a uniform
character: and the breed may be said to be degenerating. In rudimentary
organs, and in those which have been but little specialised for any
particular purpose, and perhaps in polymorphic groups, we see a nearly
parallel case; for in such cases natural selection either has not or cannot
come into full play, and thus the organisation is left in a fluctuating
condition. But what here more particularly concerns us is, that those
points in our domestic animals, which at the present time are undergoing
rapid change by continued selection, are also eminently liable to
variation. Look at the individuals of the same breed of the pigeon; and
see what a prodigious amount of difference there is in the beak of
tumblers, in the beak and wattle of carriers, in the carriage and tail of
fantails, etc., these being the points now mainly attended to by English
fanciers. Even in the same sub-breed, as in that of the short-faced
tumbler, it is notoriously difficult to breed nearly perfect birds, many
departing widely from the standard. There may truly be said to be a
constant struggle going on between, on the one hand, the tendency to
reversion to a less perfect state, as well as an innate tendency to new
variations, and, on the other hand, the power of steady selection to keep
the breed true. In the long run selection gains the day, and we do not
expect to fail so completely as to breed a bird as coarse as a common
tumbler pigeon from a good short-faced strain. But as long as selection is
rapidly going on, much variability in the parts undergoing modification may
always be expected.

Now let us turn to nature. When a part has been developed in an
extraordinary manner in any one species, compared with the other species of
the same genus, we may conclude that this part has undergone an
extraordinary amount of modification since the period when the several
species branched off from the common progenitor of the genus. This period
will seldom be remote in any extreme degree, as species rarely endure for
more than one geological period. An extraordinary amount of modification
implies an unusually large and long-continued amount of variability, which
has continually been accumulated by natural selection for the benefit of
the species. But as the variability of the extraordinarily developed part
or organ has been so great and long-continued within a period not
excessively remote, we might, as a general rule, still expect to find more
variability in such parts than in other parts of the organisation which
have remained for a much longer period nearly constant. And this, I am
convinced, is the case. That the struggle between natural selection on the
one hand, and the tendency to reversion and variability on the other hand,
will in the course of time cease; and that the most abnormally developed
organs may be made constant, I see no reason to doubt. Hence, when an
organ, however abnormal it may be, has been transmitted in approximately
the same condition to many modified descendants, as in the case of the wing
of the bat, it must have existed, according to our theory, for an immense
period in nearly the same state; and thus it has come not to be more
variable than any other structure. It is only in those cases in which the
modification has been comparatively recent and extraordinarily great that
we ought to find the GENERATIVE VARIABILITY, as it may be called, still
present in a high degree. For in this case the variability will seldom as
yet have been fixed by the continued selection of the individuals varying
in the required manner and degree, and by the continued rejection of those
tending to revert to a former and less modified condition.


The principle discussed under the last heading may be applied to our
present subject. It is notorious that specific characters are more
variable than generic. To explain by a simple example what is meant: if
in a large genus of plants some species had blue flowers and some had red,
the colour would be only a specific character, and no one would be
surprised at one of the blue species varying into red, or conversely; but
if all the species had blue flowers, the colour would become a generic
character, and its variation would be a more unusual circumstance. I have
chosen this example because the explanation which most naturalists would
advance is not here applicable, namely, that specific characters are more
variable than generic, because they are taken from parts of less
physiological importance than those commonly used for classing genera. I
believe this explanation is partly, yet only indirectly, true; I shall,
however, have to return to this point in the chapter on Classification. It
would be almost superfluous to adduce evidence in support of the statement,
that ordinary specific characters are more variable than generic; but with
respect to important characters, I have repeatedly noticed in works on
natural history, that when an author remarks with surprise that some
important organ or part, which is generally very constant throughout a
large group of species, DIFFERS considerably in closely-allied species, it
is often VARIABLE in the individuals of the same species. And this fact
shows that a character, which is generally of generic value, when it sinks
in value and becomes only of specific value, often becomes variable, though
its physiological importance may remain the same. Something of the same
kind applies to monstrosities: at least Is. Geoffroy St. Hilaire
apparently entertains no doubt, that the more an organ normally differs in
the different species of the same group, the more subject it is to
anomalies in the individuals.

On the ordinary view of each species having been independently created, why
should that part of the structure, which differs from the same part in
other independently created species of the same genus, be more variable
than those parts which are closely alike in the several species? I do not
see that any explanation can be given. But on the view that species are
only strongly marked and fixed varieties, we might expect often to find
them still continuing to vary in those parts of their structure which have
varied within a moderately recent period, and which have thus come to
differ. Or to state the case in another manner: the points in which all
the species of a genus resemble each other, and in which they differ from
allied genera, are called generic characters; and these characters may be
attributed to inheritance from a common progenitor, for it can rarely have
happened that natural selection will have modified several distinct
species, fitted to more or less widely different habits, in exactly the
same manner: and as these so-called generic characters have been inherited
from before the period when the several species first branched off from
their common progenitor, and subsequently have not varied or come to differ
in any degree, or only in a slight degree, it is not probable that they
should vary at the present day. On the other hand, the points in which
species differ from other species of the same genus are called specific
characters; and as these specific characters have varied and come to differ
since the period when the species branched off from a common progenitor, it
is probable that they should still often be in some degree variable--at
least more variable than those parts of the organisation which have for a
very long period remained constant.


I think it will be admitted by naturalists, without my entering on details,
that secondary sexual characters are highly variable. It will also be
admitted that species of the same group differ from each other more widely
in their secondary sexual characters, than in other parts of their
organisation; compare, for instance, the amount of difference between the
males of gallinaceous birds, in which secondary sexual characters are
strongly displayed, with the amount of difference between the females. The
cause of the original variability of these characters is not manifest; but
we can see why they should not have been rendered as constant and uniform
as others, for they are accumulated by sexual selection, which is less
rigid in its action than ordinary selection, as it does not entail death,
but only gives fewer offspring to the less favoured males. Whatever the
cause may be of the variability of secondary sexual characters, as they are
highly variable, sexual selection will have had a wide scope for action,
and may thus have succeeded in giving to the species of the same group a
greater amount of difference in these than in other respects.

It is a remarkable fact, that the secondary differences between the two
sexes of the same species are generally displayed in the very same parts of
the organisation in which the species of the same genus differ from each
other. Of this fact I will give in illustration the first two instances
which happen to stand on my list; and as the differences in these cases are
of a very unusual nature, the relation can hardly be accidental. The same
number of joints in the tarsi is a character common to very large groups of
beetles, but in the Engidae, as Westwood has remarked, the number varies
greatly and the number likewise differs in the two sexes of the same
species. Again in the fossorial hymenoptera, the neuration of the wings is
a character of the highest importance, because common to large groups; but
in certain genera the neuration differs in the different species, and
likewise in the two sexes of the same species. Sir J. Lubbock has recently
remarked, that several minute crustaceans offer excellent illustrations of
this law. "In Pontella, for instance, the sexual characters are afforded
mainly by the anterior antennae and by the fifth pair of legs: the
specific differences also are principally given by these organs." This
relation has a clear meaning on my view: I look at all the species of the
same genus as having as certainly descended from the same progenitor, as
have the two sexes of any one species. Consequently, whatever part of the
structure of the common progenitor, or of its early descendants, became
variable; variations of this part would, it is highly probable, be taken
advantage of by natural and sexual selection, in order to fit the several
places in the economy of nature, and likewise to fit the two sexes of the
same species to each other, or to fit the males to struggle with other
males for the possession of the females.

Finally, then, I conclude that the greater variability of specific
characters, or those which distinguish species from species, than of
generic characters, or those which are possessed by all the species; that
the frequent extreme variability of any part which is developed in a
species in an extraordinary manner in comparison with the same part in its
congeners; and the slight degree of variability in a part, however
extraordinarily it may be developed, if it be common to a whole group of
species; that the great variability of secondary sexual characters and
their great difference in closely allied species; that secondary sexual and
ordinary specific differences are generally displayed in the same parts of
the organisation, are all principles closely connected together. All being
mainly due to the species of the same group being the descendants of a
common progenitor, from whom they have inherited much in common, to parts
which have recently and largely varied being more likely still to go on
varying than parts which have long been inherited and have not varied, to
natural selection having more or less completely, according to the lapse of
time, overmastered the tendency to reversion and to further variability, to
sexual selection being less rigid than ordinary selection, and to
variations in the same parts having been accumulated by natural and sexual
selection, and thus having been adapted for secondary sexual, and for
ordinary purposes.


These propositions will be most readily understood by looking to our
domestic races. The most distinct breeds of the pigeon, in countries
widely apart, present sub-varieties with reversed feathers on the head, and
with feathers on the feet, characters not possessed by the aboriginal
rock-pigeon; these then are analogous variations in two or more distinct
races. The frequent presence of fourteen or even sixteen tail-feathers in
the pouter may be considered as a variation representing the normal
structure of another race, the fantail. I presume that no one will doubt
that all such analogous variations are due to the several races of the
pigeon having inherited from a common parent the same constitution and
tendency to variation, when acted on by similar unknown influences. In the
vegetable kingdom we have a case of analogous variation, in the enlarged
stems, or as commonly called roots, of the Swedish turnip and ruta-baga,
plants which several botanists rank as varieties produced by cultivation
from a common parent: if this be not so, the case will then be one of
analogous variation in two so-called distinct species; and to these a third
may be added, namely, the common turnip. According to the ordinary view of
each species having been independently created, we should have to attribute
this similarity in the enlarged stems of these three plants, not to the
vera causa of community of descent, and a consequent tendency to vary in a
like manner, but to three separate yet closely related acts of creation.
Many similar cases of analogous variation have been observed by Naudin in
the great gourd family, and by various authors in our cereals. Similar
cases occurring with insects under natural conditions have lately been
discussed with much ability by Mr. Walsh, who has grouped them under his
law of equable variability.

With pigeons, however, we have another case, namely, the occasional
appearance in all the breeds, of slaty-blue birds with two black bars on
the wings, white loins, a bar at the end of the tail, with the outer
feathers externally edged near their bases with white. As all these marks
are characteristic of the parent rock-pigeon, I presume that no one will
doubt that this is a case of reversion, and not of a new yet analogous
variation appearing in the several breeds. We may, I think, confidently
come to this conclusion, because, as we have seen, these coloured marks are
eminently liable to appear in the crossed offspring of two distinct and
differently coloured breeds; and in this case there is nothing in the
external conditions of life to cause the reappearance of the slaty-blue,
with the several marks, beyond the influence of the mere act of crossing on
the laws of inheritance.

No doubt it is a very surprising fact that characters should reappear after
having been lost for many, probably for hundreds of generations. But when
a breed has been crossed only once by some other breed, the offspring
occasionally show for many generations a tendency to revert in character to
the foreign breed--some say, for a dozen or even a score of generations.
After twelve generations, the proportion of blood, to use a common
expression, from one ancestor, is only 1 in 2048; and yet, as we see, it is
generally believed that a tendency to reversion is retained by this remnant
of foreign blood. In a breed which has not been crossed, but in which BOTH
parents have lost some character which their progenitor possessed, the
tendency, whether strong or weak, to reproduce the lost character might, as
was formerly remarked, for all that we can see to the contrary, be
transmitted for almost any number of generations. When a character which
has been lost in a breed, reappears after a great number of generations,
the most probable hypothesis is, not that one individual suddenly takes
after an ancestor removed by some hundred generations, but that in each
successive generation the character in question has been lying latent, and
at last, under unknown favourable conditions, is developed. With the
barb-pigeon, for instance, which very rarely produces a blue bird, it is
probable that there is a latent tendency in each generation to produce blue
plumage. The abstract improbability of such a tendency being transmitted
through a vast number of generations, is not greater than that of quite
useless or rudimentary organs being similarly transmitted. A mere tendency
to produce a rudiment is indeed sometimes thus inherited.

As all the species of the same genus are supposed to be descended from a
common progenitor, it might be expected that they would occasionally vary
in an analogous manner; so that the varieties of two or more species would
resemble each other, or that a variety of one species would resemble in
certain characters another and distinct species, this other species being,
according to our view, only a well-marked and permanent variety. But
characters exclusively due to analogous variation would probably be of an
unimportant nature, for the preservation of all functionally important
characters will have been determined through natural selection, in
accordance with the different habits of the species. It might further be
expected that the species of the same genus would occasionally exhibit
reversions to long-lost characters. As, however, we do not know the common
ancestor of any natural group, we cannot distinguish between reversionary
and analogous characters. If, for instance, we did not know that the
parent rock-pigeon was not feather-footed or turn-crowned, we could not
have told, whether such characters in our domestic breeds were reversions
or only analogous variations; but we might have inferred that the blue
colour was a case of reversion from the number of the markings, which are
correlated with this tint, and which would not probably have all appeared
together from simple variation. More especially we might have inferred
this from the blue colour and the several marks so often appearing when
differently coloured breeds are crossed. Hence, although under nature it
must generally be left doubtful, what cases are reversions to formerly
existing characters, and what are new but analogous variations, yet we
ought, on our theory, sometimes to find the varying offspring of a species
assuming characters which are already present in other members of the same
group. And this undoubtedly is the case.

The difficulty in distinguishing variable species is largely due to the
varieties mocking, as it were, other species of the same genus. A
considerable catalogue, also, could be given of forms intermediate between
two other forms, which themselves can only doubtfully be ranked as species;
and this shows, unless all these closely allied forms be considered as
independently created species, that they have in varying assumed some of
the characters of the others. But the best evidence of analogous
variations is afforded by parts or organs which are generally constant in
character, but which occasionally vary so as to resemble, in some degree,
the same part or organ in an allied species. I have collected a long list
of such cases; but here, as before, I lie under the great disadvantage of
not being able to give them. I can only repeat that such cases certainly
occur, and seem to me very remarkable.

I will, however, give one curious and complex case, not indeed as affecting
any important character, but from occurring in several species of the same
genus, partly under domestication and partly under nature. It is a case
almost certainly of reversion. The ass sometimes has very distinct
transverse bars on its legs, like those on the legs of a zebra. It has
been asserted that these are plainest in the foal, and from inquiries which
I have made, I believe this to be true. The stripe on the shoulder is
sometimes double, and is very variable in length and outline. A white ass,
but NOT an albino, has been described without either spinal or shoulder
stripe; and these stripes are sometimes very obscure, or actually quite
lost, in dark-coloured asses. The koulan of Pallas is said to have been
seen with a double shoulder-stripe. Mr. Blyth has seen a specimen of the
hemionus with a distinct shoulder-stripe, though it properly has none; and
I have been informed by Colonel Poole that foals of this species are
generally striped on the legs and faintly on the shoulder. The quagga,
though so plainly barred like a zebra over the body, is without bars on the
legs; but Dr. Gray has figured one specimen with very distinct zebra-like
bars on the hocks.

With respect to the horse, I have collected cases in England of the spinal
stripe in horses of the most distinct breeds, and of ALL colours;
transverse bars on the legs are not rare in duns, mouse-duns, and in one
instance in a chestnut; a faint shoulder-stripe may sometimes be seen in
duns, and I have seen a trace in a bay horse. My son made a careful
examination and sketch for me of a dun Belgian cart-horse with a double
stripe on each shoulder and with leg-stripes. I have myself seen a dun
Devonshire pony, and a small dun Welsh pony has been carefully described to
me, both with THREE parallel stripes on each shoulder.

In the northwest part of India the Kattywar breed of horses is so generally
striped, that, as I hear from Colonel Poole, who examined this breed for
the Indian Government, a horse without stripes is not considered as purely
bred. The spine is always striped; the legs are generally barred; and the
shoulder-stripe, which is sometimes double and sometimes treble, is common;
the side of the face, moreover, is sometimes striped. The stripes are
often plainest in the foal; and sometimes quite disappear in old horses.
Colonel Poole has seen both gray and bay Kattywar horses striped when first
foaled. I have also reason to suspect, from information given me by Mr.
W.W. Edwards, that with the English race-horse the spinal stripe is much
commoner in the foal than in the full-grown animal. I have myself recently
bred a foal from a bay mare (offspring of a Turkoman horse and a Flemish
mare) by a bay English race-horse. This foal, when a week old, was marked
on its hinder quarters and on its forehead with numerous very narrow, dark,
zebra-like bars, and its legs were feebly striped. All the stripes soon
disappeared completely. Without here entering on further details I may
state that I have collected cases of leg and shoulder stripes in horses of
very different breeds in various countries from Britain to Eastern China;
and from Norway in the north to the Malay Archipelago in the south. In all
parts of the world these stripes occur far oftenest in duns and mouse-duns;
by the term dun a large range of colour is included, from one between brown
and black to a close approach to cream colour.

I am aware that Colonel Hamilton Smith, who has written on this subject,
believes that the several breeds of the horse are descended from several
aboriginal species, one of which, the dun, was striped; and that the
above-described appearances are all due to ancient crosses with the dun
stock. But this view may be safely rejected, for it is highly improbable
that the heavy Belgian cart-horse, Welsh ponies, Norwegian cobs, the lanky
Kattywar race, etc., inhabiting the most distant parts of the world, should
have all have been crossed with one supposed aboriginal stock.

Now let us turn to the effects of crossing the several species of the horse
genus. Rollin asserts that the common mule from the ass and horse is
particularly apt to have bars on its legs; according to Mr. Gosse, in
certain parts of the United States, about nine out of ten mules have
striped legs. I once saw a mule with its legs so much striped that any one
might have thought that it was a hybrid zebra; and Mr. W.C. Martin, in his
excellent treatise on the horse, has given a figure of a similar mule. In
four coloured drawings, which I have seen, of hybrids between the ass and
zebra, the legs were much more plainly barred than the rest of the body;
and in one of them there was a double shoulder-stripe. In Lord Morton's
famous hybrid, from a chestnut mare and male quagga, the hybrid and even
the pure offspring subsequently produced from the same mare by a black
Arabian sire, were much more plainly barred across the legs than is even
the pure quagga. Lastly, and this is another most remarkable case, a
hybrid has been figured by Dr. Gray (and he informs me that he knows of a
second case) from the ass and the hemionus; and this hybrid, though the ass
only occasionally has stripes on his legs and the hemionus has none and has
not even a shoulder-stripe, nevertheless had all four legs barred, and had
three short shoulder-stripes, like those on the dun Devonshire and Welsh
ponies, and even had some zebra-like stripes on the sides of its face.
With respect to this last fact, I was so convinced that not even a stripe
of colour appears from what is commonly called chance, that I was led
solely from the occurrence of the face-stripes on this hybrid from the ass
and hemionus to ask Colonel Poole whether such face-stripes ever occurred
in the eminently striped Kattywar breed of horses, and was, as we have
seen, answered in the affirmative.

What now are we to say to these several facts? We see several distinct
species of the horse genus becoming, by simple variation, striped on the
legs like a zebra, or striped on the shoulders like an ass. In the horse
we see this tendency strong whenever a dun tint appears--a tint which
approaches to that of the general colouring of the other species of the
genus. The appearance of the stripes is not accompanied by any change of
form, or by any other new character. We see this tendency to become
striped most strongly displayed in hybrids from between several of the most
distinct species. Now observe the case of the several breeds of pigeons:
they are descended from a pigeon (including two or three sub-species or
geographical races) of a bluish colour, with certain bars and other marks;
and when any breed assumes by simple variation a bluish tint, these bars
and other marks invariably reappear; but without any other change of form
or character. When the oldest and truest breeds of various colours are
crossed, we see a strong tendency for the blue tint and bars and marks to
reappear in the mongrels. I have stated that the most probable hypothesis
to account for the reappearance of very ancient characters, is--that there
is a TENDENCY in the young of each successive generation to produce the
long-lost character, and that this tendency, from unknown causes, sometimes
prevails. And we have just seen that in several species of the horse genus
the stripes are either plainer or appear more commonly in the young than in
the old. Call the breeds of pigeons, some of which have bred true for
centuries, species; and how exactly parallel is the case with that of the
species of the horse genus! For myself, I venture confidently to look back
thousands on thousands of generations, and I see an animal striped like a
zebra, but perhaps otherwise very differently constructed, the common
parent of our domestic horse (whether or not it be descended from one or
more wild stocks) of the ass, the hemionus, quagga, and zebra.

He who believes that each equine species was independently created, will, I
presume, assert that each species has been created with a tendency to vary,
both under nature and under domestication, in this particular manner, so as
often to become striped like the other species of the genus; and that each
has been created with a strong tendency, when crossed with species
inhabiting distant quarters of the world, to produce hybrids resembling in
their stripes, not their own parents, but other species of the genus. To
admit this view is, as it seems to me, to reject a real for an unreal, or
at least for an unknown cause. It makes the works of God a mere mockery
and deception; I would almost as soon believe with the old and ignorant
cosmogonists, that fossil shells had never lived, but had been created in
stone so as to mock the shells now living on the sea-shore.


Our ignorance of the laws of variation is profound. Not in one case out of
a hundred can we pretend to assign any reason why this or that part has
varied. But whenever we have the means of instituting a comparison, the
same laws appear to have acted in producing the lesser differences between
varieties of the same species, and the greater differences between species
of the same genus. Changed conditions generally induce mere fluctuating
variability, but sometimes they cause direct and definite effects; and
these may become strongly marked in the course of time, though we have not
sufficient evidence on this head. Habit in producing constitutional
peculiarities, and use in strengthening, and disuse in weakening and
diminishing organs, appear in many cases to have been potent in their
effects. Homologous parts tend to vary in the same manner, and homologous
parts tend to cohere. Modifications in hard parts and in external parts
sometimes affect softer and internal parts. When one part is largely
developed, perhaps it tends to draw nourishment from the adjoining parts;
and every part of the structure which can be saved without detriment will
be saved. Changes of structure at an early age may affect parts
subsequently developed; and many cases of correlated variation, the nature
of which we are unable to understand, undoubtedly occur. Multiple parts
are variable in number and in structure, perhaps arising from such parts
not having been closely specialised for any particular function, so that
their modifications have not been closely checked by natural selection. It
follows probably from this same cause, that organic beings low in the scale
are more variable than those standing higher in the scale, and which have
their whole organisation more specialised. Rudimentary organs, from being
useless, are not regulated by natural selection, and hence are variable.
Specific characters--that is, the characters which have come to differ
since the several species of the same genus branched off from a common
parent--are more variable than generic characters, or those which have long
been inherited, and have not differed within this same period. In these
remarks we have referred to special parts or organs being still variable,
because they have recently varied and thus come to differ; but we have also
seen in the second chapter that the same principle applies to the whole
individual; for in a district where many species of a genus are found--that
is, where there has been much former variation and differentiation, or
where the manufactory of new specific forms has been actively at work--in
that district and among these species, we now find, on an average, most
varieties. Secondary sexual characters are highly variable, and such
characters differ much in the species of the same group. Variability in
the same parts of the organisation has generally been taken advantage of in
giving secondary sexual differences to the two sexes of the same species,
and specific differences to the several species of the same genus. Any
part or organ developed to an extraordinary size or in an extraordinary
manner, in comparison with the same part or organ in the allied species,
must have gone through an extraordinary amount of modification since the
genus arose; and thus we can understand why it should often still be
variable in a much higher degree than other parts; for variation is a
long-continued and slow process, and natural selection will in such cases
not as yet have had time to overcome the tendency to further variability
and to reversion to a less modified state. But when a species with an
extraordinarily developed organ has become the parent of many modified
descendants--which on our view must be a very slow process, requiring a
long lapse of time--in this case, natural selection has succeeded in giving
a fixed character to the organ, in however extraordinary a manner it may
have been developed. Species inheriting nearly the same constitution from
a common parent, and exposed to similar influences, naturally tend to
present analogous variations, or these same species may occasionally revert
to some of the characters of their ancient progenitors. Although new and
important modifications may not arise from reversion and analogous
variation, such modifications will add to the beautiful and harmonious
diversity of nature.

Whatever the cause may be of each slight difference between the offspring
and their parents--and a cause for each must exist--we have reason to
believe that it is the steady accumulation of beneficial differences which
has given rise to all the more important modifications of structure in
relation to the habits of each species.



Difficulties of the theory of descent with modification -- Absence or
rarity of transitional varieties -- Transitions in habits of life --
Diversified habits in the same species -- Species with habits widely
different from those of their allies -- Organs of extreme perfection --
Modes of transition -- Cases of difficulty -- Natura non facit saltum --
Organs of small importance -- Organs not in all cases absolutely perfect --
The law of Unity of Type and of the Conditions of Existence embraced by the
theory of Natural Selection.

Long before the reader has arrived at this part of my work, a crowd of
difficulties will have occurred to him. Some of them are so serious that
to this day I can hardly reflect on them without being in some degree
staggered; but, to the best of my judgment, the greater number are only
apparent, and those that are real are not, I think, fatal to the theory.

These difficulties and objections may be classed under the following heads:
First, why, if species have descended from other species by fine
gradations, do we not everywhere see innumerable transitional forms? Why
is not all nature in confusion, instead of the species being, as we see
them, well defined?

Secondly, is it possible that an animal having, for instance, the structure
and habits of a bat, could have been formed by the modification of some
other animal with widely different habits and structure? Can we believe
that natural selection could produce, on the one hand, an organ of trifling
importance, such as the tail of a giraffe, which serves as a fly-flapper,
and, on the other hand, an organ so wonderful as the eye?

Thirdly, can instincts be acquired and modified through natural selection?
What shall we say to the instinct which leads the bee to make cells, and
which has practically anticipated the discoveries of profound

Fourthly, how can we account for species, when crossed, being sterile and
producing sterile offspring, whereas, when varieties are crossed, their
fertility is unimpaired?

The two first heads will be here discussed; some miscellaneous objections
in the following chapter; Instinct and Hybridism in the two succeeding


As natural selection acts solely by the preservation of profitable
modifications, each new form will tend in a fully-stocked country to take
the place of, and finally to exterminate, its own less improved parent-form
and other less-favoured forms with which it comes into competition. Thus
extinction and natural selection go hand in hand. Hence, if we look at
each species as descended from some unknown form, both the parent and all
the transitional varieties will generally have been exterminated by the
very process of the formation and perfection of the new form.

But, as by this theory innumerable transitional forms must have existed,
why do we not find them embedded in countless numbers in the crust of the
earth? It will be more convenient to discuss this question in the chapter
on the imperfection of the geological record; and I will here only state
that I believe the answer mainly lies in the record being incomparably less
perfect than is generally supposed. The crust of the earth is a vast
museum; but the natural collections have been imperfectly made, and only at
long intervals of time.

But it may be urged that when several closely allied species inhabit the
same territory, we surely ought to find at the present time many
transitional forms. Let us take a simple case: in travelling from north
to south over a continent, we generally meet at successive intervals with
closely allied or representative species, evidently filling nearly the same
place in the natural economy of the land. These representative species
often meet and interlock; and as the one becomes rarer and rarer, the other
becomes more and more frequent, till the one replaces the other. But if we
compare these species where they intermingle, they are generally as
absolutely distinct from each other in every detail of structure as are
specimens taken from the metropolis inhabited by each. By my theory these
allied species are descended from a common parent; and during the process
of modification, each has become adapted to the conditions of life of its
own region, and has supplanted and exterminated its original parent-form
and all the transitional varieties between its past and present states.

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