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On the Origin of Species by Charles Darwin

Part 4 out of 9

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function of the organ may be the same, yet some fundamental difference
can generally be detected. I am inclined to believe that in nearly the
same way as two men have sometimes independently hit on the very same
invention, so natural selection, working for the good of each being
and taking advantage of analogous variations, has sometimes modified
in very nearly the same manner two parts in two organic beings, which
owe but little of their structure in common to inheritance from the
same ancestor.

Although in many cases it is most difficult to conjecture by what
transitions an organ could have arrived at its present state; yet,
considering that the proportion of living and known forms to the
extinct and unknown is very small, I have been astonished how rarely
an organ can be named, towards which no transitional grade is known to
lead. The truth of this remark is indeed shown by that old canon in
natural history of "Natura non facit saltum." We meet with this
admission in the writings of almost every experienced naturalist; or,
as Milne Edwards has well expressed it, nature is prodigal in variety,
but niggard in innovation. Why, on the theory of Creation, should this
be so? Why should all the parts and organs of many independent beings,
each supposed to have been separately created for its proper place in
nature, be so invariably linked together by graduated steps? Why
should not Nature have taken a leap from structure to structure? On
the theory of natural selection, we can clearly understand why she
should not; for natural selection can act only by taking advantage of
slight successive variations; she can never take a leap, but must
advance by the shortest and slowest steps.


As natural selection acts by life and death,--by the preservation of
individuals with any favourable variation, and by the destruction of
those with any unfavourable deviation of structure,--I have sometimes
felt much difficulty in understanding the origin of simple parts, of
which the importance does not seem sufficient to cause the
preservation of successively varying individuals. I have sometimes
felt as much difficulty, though of a very different kind, on this
head, as in the case of an organ as perfect and complex as the eye.

In the first place, we are much too ignorant in regard to the whole
economy of any one organic being, to say what slight modifications
would be of importance or not. In a former chapter I have given
instances of most trifling characters, such as the down on fruit and
the colour of the flesh, which, from determining the attacks of
insects or from being correlated with constitutional differences,
might assuredly be acted on by natural selection. The tail of the
giraffe looks like an artificially constructed fly-flapper; and it
seems at first incredible that this could have been adapted for its
present purpose by successive slight modifications, each better and
better, for so trifling an object as driving away flies; yet we should
pause before being too positive even in this case, for we know that
the distribution and existence of cattle and other animals in South
America absolutely depends on their power of resisting the attacks of
insects: so that individuals which could by any means defend
themselves from these small enemies, would be able to range into new
pastures and thus gain a great advantage. It is not that the larger
quadrupeds are actually destroyed (except in some rare cases) by the
flies, but they are incessantly harassed and their strength reduced,
so that they are more subject to disease, or not so well enabled in a
coming dearth to search for food, or to escape from beasts of prey.

Organs now of trifling importance have probably in some cases been of
high importance to an early progenitor, and, after having been slowly
perfected at a former period, have been transmitted in nearly the same
state, although now become of very slight use; and any actually
injurious deviations in their structure will always have been checked
by natural selection. Seeing how important an organ of locomotion the
tail is in most aquatic animals, its general presence and use for many
purposes in so many land animals, which in their lungs or modified
swim-bladders betray their aquatic origin, may perhaps be thus
accounted for. A well-developed tail having been formed in an aquatic
animal, it might subsequently come to be worked in for all sorts of
purposes, as a fly-flapper, an organ of prehension, or as an aid in
turning, as with the dog, though the aid must be slight, for the hare,
with hardly any tail, can double quickly enough.

In the second place, we may sometimes attribute importance to
characters which are really of very little importance, and which have
originated from quite secondary causes, independently of natural
selection. We should remember that climate, food, etc., probably have
some little direct influence on the organisation; that characters
reappear from the law of reversion; that correlation of growth will
have had a most important influence in modifying various structures;
and finally, that sexual selection will often have largely modified
the external characters of animals having a will, to give one male an
advantage in fighting with another or in charming the females.
Moreover when a modification of structure has primarily arisen from
the above or other unknown causes, it may at first have been of no
advantage to the species, but may subsequently have been taken
advantage of by the descendants of the species under new conditions of
life and with newly acquired habits.

To give a few instances to illustrate these latter remarks. If green
woodpeckers alone had existed, and we did not know that there were
many black and pied kinds, I dare say that we should have thought that
the green colour was a beautiful adaptation to hide this
tree-frequenting bird from its enemies; and consequently that it was a
character of importance and might have been acquired through natural
selection; as it is, I have no doubt that the colour is due to some
quite distinct cause, probably to sexual selection. A trailing bamboo
in the Malay Archipelago climbs the loftiest trees by the aid of
exquisitely constructed hooks clustered around the ends of the
branches, and this contrivance, no doubt, is of the highest service to
the plant; but as we see nearly similar hooks on many trees which are
not climbers, the hooks on the bamboo may have arisen from unknown
laws of growth, and have been subsequently taken advantage of by the
plant undergoing further modification and becoming a climber. The
naked skin on the head of a vulture is generally looked at as a direct
adaptation for wallowing in putridity; and so it may be, or it may
possibly be due to the direct action of putrid matter; but we should
be very cautious in drawing any such inference, when we see that the
skin on the head of the clean-feeding male turkey is likewise naked.
The sutures in the skulls of young mammals have been advanced as a
beautiful adaptation for aiding parturition, and no doubt they
facilitate, or may be indispensable for this act; but as sutures occur
in the skulls of young birds and reptiles, which have only to escape
from a broken egg, we may infer that this structure has arisen from
the laws of growth, and has been taken advantage of in the parturition
of the higher animals.

We are profoundly ignorant of the causes producing slight and
unimportant variations; and we are immediately made conscious of this
by reflecting on the differences in the breeds of our domesticated
animals in different countries,--more especially in the less civilized
countries where there has been but little artificial selection.
Careful observers are convinced that a damp climate affects the growth
of the hair, and that with the hair the horns are correlated. Mountain
breeds always differ from lowland breeds; and a mountainous country
would probably affect the hind limbs from exercising them more, and
possibly even the form of the pelvis; and then by the law of
homologous variation, the front limbs and even the head would probably
be affected. The shape, also, of the pelvis might affect by pressure
the shape of the head of the young in the womb. The laborious
breathing necessary in high regions would, we have some reason to
believe, increase the size of the chest; and again correlation would
come into play. Animals kept by savages in different countries often
have to struggle for their own subsistence, and would be exposed to a
certain extent to natural selection, and individuals with slightly
different constitutions would succeed best under different climates;
and there is reason to believe that constitution and colour are
correlated. A good observer, also, states that in cattle
susceptibility to the attacks of flies is correlated with colour, as
is the liability to be poisoned by certain plants; so that colour
would be thus subjected to the action of natural selection. But we are
far too ignorant to speculate on the relative importance of the
several known and unknown laws of variation; and I have here alluded
to them only to show that, if we are unable to account for the
characteristic differences of our domestic breeds, which nevertheless
we generally admit to have arisen through ordinary generation, we
ought not to lay too much stress on our ignorance of the precise cause
of the slight analogous differences between species. I might have
adduced for this same purpose the differences between the races of
man, which are so strongly marked; I may add that some little light
can apparently be thrown on the origin of these differences, chiefly
through sexual selection of a particular kind, but without here
entering on copious details my reasoning would appear frivolous.

The foregoing remarks lead me to say a few words on the protest lately
made by some naturalists, against the utilitarian doctrine that every
detail of structure has been produced for the good of its possessor.
They believe that very many structures have been created for beauty in
the eyes of man, or for mere variety. This doctrine, if true, would be
absolutely fatal to my theory. Yet I fully admit that many structures
are of no direct use to their possessors. Physical conditions probably
have had some little effect on structure, quite independently of any
good thus gained. Correlation of growth has no doubt played a most
important part, and a useful modification of one part will often have
entailed on other parts diversified changes of no direct use. So again
characters which formerly were useful, or which formerly had arisen
from correlation of growth, or from other unknown cause, may reappear
from the law of reversion, though now of no direct use. The effects of
sexual selection, when displayed in beauty to charm the females, can
be called useful only in rather a forced sense. But by far the most
important consideration is that the chief part of the organisation of
every being is simply due to inheritance; and consequently, though
each being assuredly is well fitted for its place in nature, many
structures now have no direct relation to the habits of life of each
species. Thus, we can hardly believe that the webbed feet of the
upland goose or of the frigate-bird are of special use to these birds;
we cannot believe that the same bones in the arm of the monkey, in the
fore leg of the horse, in the wing of the bat, and in the flipper of
the seal, are of special use to these animals. We may safely attribute
these structures to inheritance. But to the progenitor of the upland
goose and of the frigate-bird, webbed feet no doubt were as useful as
they now are to the most aquatic of existing birds. So we may believe
that the progenitor of the seal had not a flipper, but a foot with
five toes fitted for walking or grasping; and we may further venture
to believe that the several bones in the limbs of the monkey, horse,
and bat, which have been inherited from a common progenitor, were
formerly of more special use to that progenitor, or its progenitors,
than they now are to these animals having such widely diversified
habits. Therefore we may infer that these several bones might have
been acquired through natural selection, subjected formerly, as now,
to the several laws of inheritance, reversion, correlation of growth,
etc. Hence every detail of structure in every living creature (making
some little allowance for the direct action of physical conditions)
may be viewed, either as having been of special use to some ancestral
form, or as being now of special use to the descendants of this
form--either directly, or indirectly through the complex laws of

Natural selection cannot possibly produce any modification in any one
species exclusively for the good of another species; though throughout
nature one species incessantly takes advantage of, and profits by, the
structure of another. But natural selection can and does often produce
structures for the direct injury of other species, as we see in the
fang of the adder, and in the ovipositor of the ichneumon, by which
its eggs are deposited in the living bodies of other insects. If it
could be proved that any part of the structure of any one species had
been formed for the exclusive good of another species, it would
annihilate my theory, for such could not have been produced through
natural selection. Although many statements may be found in works on
natural history to this effect, I cannot find even one which seems to
me of any weight. It is admitted that the rattlesnake has a
poison-fang for its own defence and for the destruction of its prey;
but some authors suppose that at the same time this snake is furnished
with a rattle for its own injury, namely, to warn its prey to escape.
I would almost as soon believe that the cat curls the end of its tail
when preparing to spring, in order to warn the doomed mouse. But I
have not space here to enter on this and other such cases.

Natural selection will never produce in a being anything injurious to
itself, for natural selection acts solely by and for the good of each.
No organ will be formed, as Paley has remarked, for the purpose of
causing pain or for doing an injury to its possessor. If a fair
balance be struck between the good and evil caused by each part, each
will be found on the whole advantageous. After the lapse of time,
under changing conditions of life, if any part comes to be injurious,
it will be modified; or if it be not so, the being will become
extinct, as myriads have become extinct.

Natural selection tends only to make each organic being as perfect as,
or slightly more perfect than, the other inhabitants of the same
country with which it has to struggle for existence. And we see that
this is the degree of perfection attained under nature. The endemic
productions of New Zealand, for instance, are perfect one compared
with another; but they are now rapidly yielding before the advancing
legions of plants and animals introduced from Europe. Natural
selection will not produce absolute perfection, nor do we always meet,
as far as we can judge, with this high standard under nature. The
correction for the aberration of light is said, on high authority, not
to be perfect even in that most perfect organ, the eye. If our reason
leads us to admire with enthusiasm a multitude of inimitable
contrivances in nature, this same reason tells us, though we may
easily err on both sides, that some other contrivances are less
perfect. Can we consider the sting of the wasp or of the bee as
perfect, which, when used against many attacking animals, cannot be
withdrawn, owing to the backward serratures, and so inevitably causes
the death of the insect by tearing out its viscera?

If we look at the sting of the bee, as having originally existed in a
remote progenitor as a boring and serrated instrument, like that in so
many members of the same great order, and which has been modified but
not perfected for its present purpose, with the poison originally
adapted to cause galls subsequently intensified, we can perhaps
understand how it is that the use of the sting should so often cause
the insect's own death: for if on the whole the power of stinging be
useful to the community, it will fulfil all the requirements of
natural selection, though it may cause the death of some few members.
If we admire the truly wonderful power of scent by which the males of
many insects find their females, can we admire the production for this
single purpose of thousands of drones, which are utterly useless to
the community for any other end, and which are ultimately slaughtered
by their industrious and sterile sisters? It may be difficult, but we
ought to admire the savage instinctive hatred of the queen-bee, which
urges her instantly to destroy the young queens her daughters as soon
as born, or to perish herself in the combat; for undoubtedly this is
for the good of the community; and maternal love or maternal hatred,
though the latter fortunately is most rare, is all the same to the
inexorable principle of natural selection. If we admire the several
ingenious contrivances, by which the flowers of the orchis and of many
other plants are fertilised through insect agency, can we consider as
equally perfect the elaboration by our fir-trees of dense clouds of
pollen, in order that a few granules may be wafted by a chance breeze
on to the ovules?


We have in this chapter discussed some of the difficulties and
objections which may be urged against my theory. Many of them are very
grave; but I think that in the discussion light has been thrown on
several facts, which on the theory of independent acts of creation are
utterly obscure. We have seen that species at any one period are not
indefinitely variable, and are not linked together by a multitude of
intermediate gradations, partly because the process of natural
selection will always be very slow, and will act, at any one time,
only on a very few forms; and partly because the very process of
natural selection almost implies the continual supplanting and
extinction of preceding and intermediate gradations. Closely allied
species, now living on a continuous area, must often have been formed
when the area was not continuous, and when the conditions of life did
not insensibly graduate away from one part to another. When two
varieties are formed in two districts of a continuous area, an
intermediate variety will often be formed, fitted for an intermediate
zone; but from reasons assigned, the intermediate variety will usually
exist in lesser numbers than the two forms which it connects;
consequently the two latter, during the course of further
modification, from existing in greater numbers, will have a great
advantage over the less numerous intermediate variety, and will thus
generally succeed in supplanting and exterminating it.

We have seen in this chapter how cautious we should be in concluding
that the most different habits of life could not graduate into each
other; that a bat, for instance, could not have been formed by natural
selection from an animal which at first could only glide through the

We have seen that a species may under new conditions of life change
its habits, or have diversified habits, with some habits very unlike
those of its nearest congeners. Hence we can understand, bearing in
mind that each organic being is trying to live wherever it can live,
how it has arisen that there are upland geese with webbed feet, ground
woodpeckers, diving thrushes, and petrels with the habits of auks.

Although the belief that an organ so perfect as the eye could have
been formed by natural selection, is more than enough to stagger any
one; yet in the case of any organ, if we know of a long series of
gradations in complexity, each good for its possessor, then, under
changing conditions of life, there is no logical impossibility in the
acquirement of any conceivable degree of perfection through natural
selection. In the cases in which we know of no intermediate or
transitional states, we should be very cautious in concluding that
none could have existed, for the homologies of many organs and their
intermediate states show that wonderful metamorphoses in function are
at least possible. For instance, a swim-bladder has apparently been
converted into an air-breathing lung. The same organ having performed
simultaneously very different functions, and then having been
specialised for one function; and two very distinct organs having
performed at the same time the same function, the one having been
perfected whilst aided by the other, must often have largely
facilitated transitions.

We are far too ignorant, in almost every case, to be enabled to assert
that any part or organ is so unimportant for the welfare of a species,
that modifications in its structure could not have been slowly
accumulated by means of natural selection. But we may confidently
believe that many modifications, wholly due to the laws of growth, and
at first in no way advantageous to a species, have been subsequently
taken advantage of by the still further modified descendants of this
species. We may, also, believe that a part formerly of high importance
has often been retained (as the tail of an aquatic animal by its
terrestrial descendants), though it has become of such small
importance that it could not, in its present state, have been acquired
by natural selection,--a power which acts solely by the preservation
of profitable variations in the struggle for life.

Natural selection will produce nothing in one species for the
exclusive good or injury of another; though it may well produce parts,
organs, and excretions highly useful or even indispensable, or highly
injurious to another species, but in all cases at the same time useful
to the owner. Natural selection in each well-stocked country, must act
chiefly through the competition of the inhabitants one with another,
and consequently will produce perfection, or strength in the battle
for life, only according to the standard of that country. Hence the
inhabitants of one country, generally the smaller one, will often
yield, as we see they do yield, to the inhabitants of another and
generally larger country. For in the larger country there will have
existed more individuals, and more diversified forms, and the
competition will have been severer, and thus the standard of
perfection will have been rendered higher. Natural selection will not
necessarily produce absolute perfection; nor, as far as we can judge
by our limited faculties, can absolute perfection be everywhere found.

On the theory of natural selection we can clearly understand the full
meaning of that old canon in natural history, "Natura non facit
saltum." This canon, if we look only to the present inhabitants of the
world, is not strictly correct, but if we include all those of past
times, it must by my theory be strictly true.

It is generally acknowledged that all organic beings have been formed
on two great laws--Unity of Type, and the Conditions of Existence. By
unity of type is meant that fundamental agreement in structure, which
we see in organic beings of the same class, and which is quite
independent of their habits of life. On my theory, unity of type is
explained by unity of descent. The expression of conditions of
existence, so often insisted on by the illustrious Cuvier, is fully
embraced by the principle of natural selection. For natural selection
acts by either now adapting the varying parts of each being to its
organic and inorganic conditions of life; or by having adapted them
during long-past periods of time: the adaptations being aided in some
cases by use and disuse, being slightly affected by the direct action
of the external conditions of life, and being in all cases subjected
to the several laws of growth. Hence, in fact, the law of the
Conditions of Existence is the higher law; as it includes, through the
inheritance of former adaptations, that of Unity of Type.


Instincts comparable with habits, but different in their origin.
Instincts graduated.
Aphides and ants.
Instincts variable.
Domestic instincts, their origin.
Natural instincts of the cuckoo, ostrich, and parasitic bees.
Slave-making ants.
Hive-bee, its cell-making instinct.
Difficulties on the theory of the Natural Selection of instincts.
Neuter or sterile insects.

The subject of instinct might have been worked into the previous
chapters; but I have thought that it would be more convenient to treat
the subject separately, especially as so wonderful an instinct as that
of the hive-bee making its cells will probably have occurred to many
readers, as a difficulty sufficient to overthrow my whole theory. I
must premise, that I have nothing to do with the origin of the primary
mental powers, any more than I have with that of life itself. We are
concerned only with the diversities of instinct and of the other
mental qualities of animals within the same class.

I will not attempt any definition of instinct. It would be easy to
show that several distinct mental actions are commonly embraced by
this term; but every one understands what is meant, when it is said
that instinct impels the cuckoo to migrate and to lay her eggs in
other birds' nests. An action, which we ourselves should require
experience to enable us to perform, when performed by an animal, more
especially by a very young one, without any experience, and when
performed by many individuals in the same way, without their knowing
for what purpose it is performed, is usually said to be instinctive.
But I could show that none of these characters of instinct are
universal. A little dose, as Pierre Huber expresses it, of judgment or
reason, often comes into play, even in animals very low in the scale
of nature.

Frederick Cuvier and several of the older metaphysicians have compared
instinct with habit. This comparison gives, I think, a remarkably
accurate notion of the frame of mind under which an instinctive action
is performed, but not of its origin. How unconsciously many habitual
actions are performed, indeed not rarely in direct opposition to our
conscious will! yet they may be modified by the will or reason. Habits
easily become associated with other habits, and with certain periods
of time and states of the body. When once acquired, they often remain
constant throughout life. Several other points of resemblance between
instincts and habits could be pointed out. As in repeating a
well-known song, so in instincts, one action follows another by a sort
of rhythm; if a person be interrupted in a song, or in repeating
anything by rote, he is generally forced to go back to recover the
habitual train of thought: so P. Huber found it was with a
caterpillar, which makes a very complicated hammock; for if he took a
caterpillar which had completed its hammock up to, say, the sixth
stage of construction, and put it into a hammock completed up only to
the third stage, the caterpillar simply re-performed the fourth,
fifth, and sixth stages of construction. If, however, a caterpillar
were taken out of a hammock made up, for instance, to the third stage,
and were put into one finished up to the sixth stage, so that much of
its work was already done for it, far from feeling the benefit of
this, it was much embarrassed, and, in order to complete its hammock,
seemed forced to start from the third stage, where it had left off,
and thus tried to complete the already finished work. If we suppose
any habitual action to become inherited--and I think it can be shown
that this does sometimes happen--then the resemblance between what
originally was a habit and an instinct becomes so close as not to be
distinguished. If Mozart, instead of playing the pianoforte at three
years old with wonderfully little practice, had played a tune with no
practice at all, he might truly be said to have done so instinctively.
But it would be the most serious error to suppose that the greater
number of instincts have been acquired by habit in one generation, and
then transmitted by inheritance to succeeding generations. It can be
clearly shown that the most wonderful instincts with which we are
acquainted, namely, those of the hive-bee and of many ants, could not
possibly have been thus acquired.

It will be universally admitted that instincts are as important as
corporeal structure for the welfare of each species, under its present
conditions of life. Under changed conditions of life, it is at least
possible that slight modifications of instinct might be profitable to
a species; and if it can be shown that instincts do vary ever so
little, then I can see no difficulty in natural selection preserving
and continually accumulating variations of instinct to any extent that
may be profitable. It is thus, as I believe, that all the most complex
and wonderful instincts have originated. As modifications of corporeal
structure arise from, and are increased by, use or habit, and are
diminished or lost by disuse, so I do not doubt it has been with
instincts. But I believe that the effects of habit are of quite
subordinate importance to the effects of the natural selection of what
may be called accidental variations of instincts;--that is of
variations produced by the same unknown causes which produce slight
deviations of bodily structure.

No complex instinct can possibly be produced through natural
selection, except by the slow and gradual accumulation of numerous,
slight, yet profitable, variations. Hence, as in the case of corporeal
structures, we ought to find in nature, not the actual transitional
gradations by which each complex instinct has been acquired--for these
could be found only in the lineal ancestors of each species--but we
ought to find in the collateral lines of descent some evidence of such
gradations; or we ought at least to be able to show that gradations of
some kind are possible; and this we certainly can do. I have been
surprised to find, making allowance for the instincts of animals
having been but little observed except in Europe and North America,
and for no instinct being known amongst extinct species, how very
generally gradations, leading to the most complex instincts, can be
discovered. The canon of "Natura non facit saltum" applies with almost
equal force to instincts as to bodily organs. Changes of instinct may
sometimes be facilitated by the same species having different
instincts at different periods of life, or at different seasons of the
year, or when placed under different circumstances, etc.; in which
case either one or the other instinct might be preserved by natural
selection. And such instances of diversity of instinct in the same
species can be shown to occur in nature.

Again as in the case of corporeal structure, and conformably with my
theory, the instinct of each species is good for itself, but has
never, as far as we can judge, been produced for the exclusive good of
others. One of the strongest instances of an animal apparently
performing an action for the sole good of another, with which I am
acquainted, is that of aphides voluntarily yielding their sweet
excretion to ants: that they do so voluntarily, the following facts
show. I removed all the ants from a group of about a dozen aphides on
a dock-plant, and prevented their attendance during several hours.
After this interval, I felt sure that the aphides would want to
excrete. I watched them for some time through a lens, but not one
excreted; I then tickled and stroked them with a hair in the same
manner, as well as I could, as the ants do with their antennae; but
not one excreted. Afterwards I allowed an ant to visit them, and it
immediately seemed, by its eager way of running about, to be well
aware what a rich flock it had discovered; it then began to play with
its antennae on the abdomen first of one aphis and then of another;
and each aphis, as soon as it felt the antennae, immediately lifted up
its abdomen and excreted a limpid drop of sweet juice, which was
eagerly devoured by the ant. Even the quite young aphides behaved in
this manner, showing that the action was instinctive, and not the
result of experience. But as the excretion is extremely viscid, it is
probably a convenience to the aphides to have it removed; and
therefore probably the aphides do not instinctively excrete for the
sole good of the ants. Although I do not believe that any animal in
the world performs an action for the exclusive good of another of a
distinct species, yet each species tries to take advantage of the
instincts of others, as each takes advantage of the weaker bodily
structure of others. So again, in some few cases, certain instincts
cannot be considered as absolutely perfect; but as details on this and
other such points are not indispensable, they may be here passed over.

As some degree of variation in instincts under a state of nature, and
the inheritance of such variations, are indispensable for the action
of natural selection, as many instances as possible ought to have been
here given; but want of space prevents me. I can only assert, that
instincts certainly do vary--for instance, the migratory instinct,
both in extent and direction, and in its total loss. So it is with the
nests of birds, which vary partly in dependence on the situations
chosen, and on the nature and temperature of the country inhabited,
but often from causes wholly unknown to us: Audubon has given several
remarkable cases of differences in nests of the same species in the
northern and southern United States. Fear of any particular enemy is
certainly an instinctive quality, as may be seen in nestling birds,
though it is strengthened by experience, and by the sight of fear of
the same enemy in other animals. But fear of man is slowly acquired,
as I have elsewhere shown, by various animals inhabiting desert
islands; and we may see an instance of this, even in England, in the
greater wildness of all our large birds than of our small birds; for
the large birds have been most persecuted by man. We may safely
attribute the greater wildness of our large birds to this cause; for
in uninhabited islands large birds are not more fearful than small;
and the magpie, so wary in England, is tame in Norway, as is the
hooded crow in Egypt.

That the general disposition of individuals of the same species, born
in a state of nature, is extremely diversified, can be shown by a
multitude of facts. Several cases also, could be given, of occasional
and strange habits in certain species, which might, if advantageous to
the species, give rise, through natural selection, to quite new
instincts. But I am well aware that these general statements, without
facts given in detail, can produce but a feeble effect on the reader's
mind. I can only repeat my assurance, that I do not speak without good

The possibility, or even probability, of inherited variations of
instinct in a state of nature will be strengthened by briefly
considering a few cases under domestication. We shall thus also be
enabled to see the respective parts which habit and the selection of
so-called accidental variations have played in modifying the mental
qualities of our domestic animals. A number of curious and authentic
instances could be given of the inheritance of all shades of
disposition and tastes, and likewise of the oddest tricks, associated
with certain frames of mind or periods of time. But let us look to the
familiar case of the several breeds of dogs: it cannot be doubted that
young pointers (I have myself seen a striking instance) will sometimes
point and even back other dogs the very first time that they are taken
out; retrieving is certainly in some degree inherited by retrievers;
and a tendency to run round, instead of at, a flock of sheep, by
shepherd-dogs. I cannot see that these actions, performed without
experience by the young, and in nearly the same manner by each
individual, performed with eager delight by each breed, and without
the end being known,--for the young pointer can no more know that he
points to aid his master, than the white butterfly knows why she lays
her eggs on the leaf of the cabbage,--I cannot see that these actions
differ essentially from true instincts. If we were to see one kind of
wolf, when young and without any training, as soon as it scented its
prey, stand motionless like a statue, and then slowly crawl forward
with a peculiar gait; and another kind of wolf rushing round, instead
of at, a herd of deer, and driving them to a distant point, we should
assuredly call these actions instinctive. Domestic instincts, as they
may be called, are certainly far less fixed or invariable than natural
instincts; but they have been acted on by far less rigorous selection,
and have been transmitted for an incomparably shorter period, under
less fixed conditions of life.

How strongly these domestic instincts, habits, and dispositions are
inherited, and how curiously they become mingled, is well shown when
different breeds of dogs are crossed. Thus it is known that a cross
with a bull-dog has affected for many generations the courage and
obstinacy of greyhounds; and a cross with a greyhound has given to a
whole family of shepherd-dogs a tendency to hunt hares. These domestic
instincts, when thus tested by crossing, resemble natural instincts,
which in a like manner become curiously blended together, and for a
long period exhibit traces of the instincts of either parent: for
example, Le Roy describes a dog, whose great-grandfather was a wolf,
and this dog showed a trace of its wild parentage only in one way, by
not coming in a straight line to his master when called.

Domestic instincts are sometimes spoken of as actions which have
become inherited solely from long-continued and compulsory habit, but
this, I think, is not true. No one would ever have thought of
teaching, or probably could have taught, the tumbler-pigeon to
tumble,--an action which, as I have witnessed, is performed by young
birds, that have never seen a pigeon tumble. We may believe that some
one pigeon showed a slight tendency to this strange habit, and that
the long-continued selection of the best individuals in successive
generations made tumblers what they now are; and near Glasgow there
are house-tumblers, as I hear from Mr. Brent, which cannot fly
eighteen inches high without going head over heels. It may be doubted
whether any one would have thought of training a dog to point, had not
some one dog naturally shown a tendency in this line; and this is
known occasionally to happen, as I once saw in a pure terrier. When
the first tendency was once displayed, methodical selection and the
inherited effects of compulsory training in each successive generation
would soon complete the work; and unconscious selection is still at
work, as each man tries to procure, without intending to improve the
breed, dogs which will stand and hunt best. On the other hand, habit
alone in some cases has sufficed; no animal is more difficult to tame
than the young of the wild rabbit; scarcely any animal is tamer than
the young of the tame rabbit; but I do not suppose that domestic
rabbits have ever been selected for tameness; and I presume that we
must attribute the whole of the inherited change from extreme wildness
to extreme tameness, simply to habit and long-continued close

Natural instincts are lost under domestication: a remarkable instance
of this is seen in those breeds of fowls which very rarely or never
become "broody," that is, never wish to sit on their eggs. Familiarity
alone prevents our seeing how universally and largely the minds of our
domestic animals have been modified by domestication. It is scarcely
possible to doubt that the love of man has become instinctive in the
dog. All wolves, foxes, jackals, and species of the cat genus, when
kept tame, are most eager to attack poultry, sheep, and pigs; and this
tendency has been found incurable in dogs which have been brought home
as puppies from countries, such as Tierra del Fuego and Australia,
where the savages do not keep these domestic animals. How rarely, on
the other hand, do our civilised dogs, even when quite young, require
to be taught not to attack poultry, sheep, and pigs! No doubt they
occasionally do make an attack, and are then beaten; and if not cured,
they are destroyed; so that habit, with some degree of selection, has
probably concurred in civilising by inheritance our dogs. On the other
hand, young chickens have lost, wholly by habit, that fear of the dog
and cat which no doubt was originally instinctive in them, in the same
way as it is so plainly instinctive in young pheasants, though reared
under a hen. It is not that chickens have lost all fear, but fear only
of dogs and cats, for if the hen gives the danger-chuckle, they will
run (more especially young turkeys) from under her, and conceal
themselves in the surrounding grass or thickets; and this is evidently
done for the instinctive purpose of allowing, as we see in wild
ground-birds, their mother to fly away. But this instinct retained by
our chickens has become useless under domestication, for the
mother-hen has almost lost by disuse the power of flight.

Hence, we may conclude, that domestic instincts have been acquired and
natural instincts have been lost partly by habit, and partly by man
selecting and accumulating during successive generations, peculiar
mental habits and actions, which at first appeared from what we must
in our ignorance call an accident. In some cases compulsory habit
alone has sufficed to produce such inherited mental changes; in other
cases compulsory habit has done nothing, and all has been the result
of selection, pursued both methodically and unconsciously; but in most
cases, probably, habit and selection have acted together.

We shall, perhaps, best understand how instincts in a state of nature
have become modified by selection, by considering a few cases. I will
select only three, out of the several which I shall have to discuss in
my future work,--namely, the instinct which leads the cuckoo to lay
her eggs in other birds' nests; the slave-making instinct of certain
ants; and the comb-making power of the hive-bee: these two latter
instincts have generally, and most justly, been ranked by naturalists
as the most wonderful of all known instincts.

It is now commonly admitted that the more immediate and final cause of
the cuckoo's instinct is, that she lays her eggs, not daily, but at
intervals of two or three days; so that, if she were to make her own
nest and sit on her own eggs, those first laid would have to be left
for some time unincubated, or there would be eggs and young birds of
different ages in the same nest. If this were the case, the process of
laying and hatching might be inconveniently long, more especially as
she has to migrate at a very early period; and the first hatched young
would probably have to be fed by the male alone. But the American
cuckoo is in this predicament; for she makes her own nest and has eggs
and young successively hatched, all at the same time. It has been
asserted that the American cuckoo occasionally lays her eggs in other
birds' nests; but I hear on the high authority of Dr. Brewer, that
this is a mistake. Nevertheless, I could give several instances of
various birds which have been known occasionally to lay their eggs in
other birds' nests. Now let us suppose that the ancient progenitor of
our European cuckoo had the habits of the American cuckoo; but that
occasionally she laid an egg in another bird's nest. If the old bird
profited by this occasional habit, or if the young were made more
vigorous by advantage having been taken of the mistaken maternal
instinct of another bird, than by their own mother's care, encumbered
as she can hardly fail to be by having eggs and young of different
ages at the same time; then the old birds or the fostered young would
gain an advantage. And analogy would lead me to believe, that the
young thus reared would be apt to follow by inheritance the occasional
and aberrant habit of their mother, and in their turn would be apt to
lay their eggs in other birds' nests, and thus be successful in
rearing their young. By a continued process of this nature, I believe
that the strange instinct of our cuckoo could be, and has been,
generated. I may add that, according to Dr. Gray and to some other
observers, the European cuckoo has not utterly lost all maternal love
and care for her own offspring.

The occasional habit of birds laying their eggs in other birds' nests,
either of the same or of a distinct species, is not very uncommon with
the Gallinaceae; and this perhaps explains the origin of a singular
instinct in the allied group of ostriches. For several hen ostriches,
at least in the case of the American species, unite and lay first a
few eggs in one nest and then in another; and these are hatched by the
males. This instinct may probably be accounted for by the fact of the
hens laying a large number of eggs; but, as in the case of the cuckoo,
at intervals of two or three days. This instinct, however, of the
American ostrich has not as yet been perfected; for a surprising
number of eggs lie strewed over the plains, so that in one day's
hunting I picked up no less than twenty lost and wasted eggs.

Many bees are parasitic, and always lay their eggs in the nests of
bees of other kinds. This case is more remarkable than that of the
cuckoo; for these bees have not only their instincts but their
structure modified in accordance with their parasitic habits; for they
do not possess the pollen-collecting apparatus which would be
necessary if they had to store food for their own young. Some species,
likewise, of Sphegidae (wasp-like insects) are parasitic on other
species; and M. Fabre has lately shown good reason for believing that
although the Tachytes nigra generally makes its own burrow and stores
it with paralysed prey for its own larvae to feed on, yet that when
this insect finds a burrow already made and stored by another sphex,
it takes advantage of the prize, and becomes for the occasion
parasitic. In this case, as with the supposed case of the cuckoo, I
can see no difficulty in natural selection making an occasional habit
permanent, if of advantage to the species, and if the insect whose
nest and stored food are thus feloniously appropriated, be not thus


This remarkable instinct was first discovered in the Formica
(Polyerges) rufescens by Pierre Huber, a better observer even than his
celebrated father. This ant is absolutely dependent on its slaves;
without their aid, the species would certainly become extinct in a
single year. The males and fertile females do no work. The workers or
sterile females, though most energetic and courageous in capturing
slaves, do no other work. They are incapable of making their own
nests, or of feeding their own larvae. When the old nest is found
inconvenient, and they have to migrate, it is the slaves which
determine the migration, and actually carry their masters in their
jaws. So utterly helpless are the masters, that when Huber shut up
thirty of them without a slave, but with plenty of the food which they
like best, and with their larvae and pupae to stimulate them to work,
they did nothing; they could not even feed themselves, and many
perished of hunger. Huber then introduced a single slave (F. fusca),
and she instantly set to work, fed and saved the survivors; made some
cells and tended the larvae, and put all to rights. What can be more
extraordinary than these well-ascertained facts? If we had not known
of any other slave-making ant, it would have been hopeless to have
speculated how so wonderful an instinct could have been perfected.

Formica sanguinea was likewise first discovered by P. Huber to be a
slave-making ant. This species is found in the southern parts of
England, and its habits have been attended to by Mr. F. Smith, of the
British Museum, to whom I am much indebted for information on this and
other subjects. Although fully trusting to the statements of Huber and
Mr. Smith, I tried to approach the subject in a sceptical frame of
mind, as any one may well be excused for doubting the truth of so
extraordinary and odious an instinct as that of making slaves. Hence I
will give the observations which I have myself made, in some little
detail. I opened fourteen nests of F. sanguinea, and found a few
slaves in all. Males and fertile females of the slave-species are
found only in their own proper communities, and have never been
observed in the nests of F. sanguinea. The slaves are black and not
above half the size of their red masters, so that the contrast in
their appearance is very great. When the nest is slightly disturbed,
the slaves occasionally come out, and like their masters are much
agitated and defend the nest: when the nest is much disturbed and the
larvae and pupae are exposed, the slaves work energetically with their
masters in carrying them away to a place of safety. Hence, it is
clear, that the slaves feel quite at home. During the months of June
and July, on three successive years, I have watched for many hours
several nests in Surrey and Sussex, and never saw a slave either leave
or enter a nest. As, during these months, the slaves are very few in
number, I thought that they might behave differently when more
numerous; but Mr. Smith informs me that he has watched the nests at
various hours during May, June and August, both in Surrey and
Hampshire, and has never seen the slaves, though present in large
numbers in August, either leave or enter the nest. Hence he considers
them as strictly household slaves. The masters, on the other hand, may
be constantly seen bringing in materials for the nest, and food of all
kinds. During the present year, however, in the month of July, I came
across a community with an unusually large stock of slaves, and I
observed a few slaves mingled with their masters leaving the nest, and
marching along the same road to a tall Scotch-fir-tree, twenty-five
yards distant, which they ascended together, probably in search of
aphides or cocci. According to Huber, who had ample opportunities for
observation, in Switzerland the slaves habitually work with their
masters in making the nest, and they alone open and close the doors in
the morning and evening; and, as Huber expressly states, their
principal office is to search for aphides. This difference in the
usual habits of the masters and slaves in the two countries, probably
depends merely on the slaves being captured in greater numbers in
Switzerland than in England.

One day I fortunately chanced to witness a migration from one nest to
another, and it was a most interesting spectacle to behold the masters
carefully carrying, as Huber has described, their slaves in their
jaws. Another day my attention was struck by about a score of the
slave-makers haunting the same spot, and evidently not in search of
food; they approached and were vigorously repulsed by an independent
community of the slave species (F. fusca); sometimes as many as three
of these ants clinging to the legs of the slave-making F. sanguinea.
The latter ruthlessly killed their small opponents, and carried their
dead bodies as food to their nest, twenty-nine yards distant; but they
were prevented from getting any pupae to rear as slaves. I then dug up
a small parcel of the pupae of F. fusca from another nest, and put
them down on a bare spot near the place of combat; they were eagerly
seized, and carried off by the tyrants, who perhaps fancied that,
after all, they had been victorious in their late combat.

At the same time I laid on the same place a small parcel of the pupae
of another species, F. flava, with a few of these little yellow ants
still clinging to the fragments of the nest. This species is
sometimes, though rarely, made into slaves, as has been described by
Mr. Smith. Although so small a species, it is very courageous, and I
have seen it ferociously attack other ants. In one instance I found to
my surprise an independent community of F. flava under a stone beneath
a nest of the slave-making F. sanguinea; and when I had accidentally
disturbed both nests, the little ants attacked their big neighbours
with surprising courage. Now I was curious to ascertain whether F.
sanguinea could distinguish the pupae of F. fusca, which they
habitually make into slaves, from those of the little and furious F.
flava, which they rarely capture, and it was evident that they did at
once distinguish them: for we have seen that they eagerly and
instantly seized the pupae of F. fusca, whereas they were much
terrified when they came across the pupae, or even the earth from the
nest of F. flava, and quickly ran away; but in about a quarter of an
hour, shortly after all the little yellow ants had crawled away, they
took heart and carried off the pupae.

One evening I visited another community of F. sanguinea, and found a
number of these ants entering their nest, carrying the dead bodies of
F. fusca (showing that it was not a migration) and numerous pupae. I
traced the returning file burthened with booty, for about forty yards,
to a very thick clump of heath, whence I saw the last individual of F.
sanguinea emerge, carrying a pupa; but I was not able to find the
desolated nest in the thick heath. The nest, however, must have been
close at hand, for two or three individuals of F. fusca were rushing
about in the greatest agitation, and one was perched motionless with
its own pupa in its mouth on the top of a spray of heath over its
ravaged home.

Such are the facts, though they did not need confirmation by me, in
regard to the wonderful instinct of making slaves. Let it be observed
what a contrast the instinctive habits of F. sanguinea present with
those of the F. rufescens. The latter does not build its own nest,
does not determine its own migrations, does not collect food for
itself or its young, and cannot even feed itself: it is absolutely
dependent on its numerous slaves. Formica sanguinea, on the other
hand, possesses much fewer slaves, and in the early part of the summer
extremely few. The masters determine when and where a new nest shall
be formed, and when they migrate, the masters carry the slaves. Both
in Switzerland and England the slaves seem to have the exclusive care
of the larvae, and the masters alone go on slave-making expeditions.
In Switzerland the slaves and masters work together, making and
bringing materials for the nest: both, but chiefly the slaves, tend,
and milk as it may be called, their aphides; and thus both collect
food for the community. In England the masters alone usually leave the
nest to collect building materials and food for themselves, their
slaves and larvae. So that the masters in this country receive much
less service from their slaves than they do in Switzerland.

By what steps the instinct of F. sanguinea originated I will not
pretend to conjecture. But as ants, which are not slave-makers, will,
as I have seen, carry off pupae of other species, if scattered near
their nests, it is possible that pupae originally stored as food might
become developed; and the ants thus unintentionally reared would then
follow their proper instincts, and do what work they could. If their
presence proved useful to the species which had seized them--if it
were more advantageous to this species to capture workers than to
procreate them--the habit of collecting pupae originally for food
might by natural selection be strengthened and rendered permanent for
the very different purpose of raising slaves. When the instinct was
once acquired, if carried out to a much less extent even than in our
British F. sanguinea, which, as we have seen, is less aided by its
slaves than the same species in Switzerland, I can see no difficulty
in natural selection increasing and modifying the instinct--always
supposing each modification to be of use to the species--until an ant
was formed as abjectly dependent on its slaves as is the Formica


I will not here enter on minute details on this subject, but will
merely give an outline of the conclusions at which I have arrived. He
must be a dull man who can examine the exquisite structure of a comb,
so beautifully adapted to its end, without enthusiastic admiration. We
hear from mathematicians that bees have practically solved a recondite
problem, and have made their cells of the proper shape to hold the
greatest possible amount of honey, with the least possible consumption
of precious wax in their construction. It has been remarked that a
skilful workman, with fitting tools and measures, would find it very
difficult to make cells of wax of the true form, though this is
perfectly effected by a crowd of bees working in a dark hive. Grant
whatever instincts you please, and it seems at first quite
inconceivable how they can make all the necessary angles and planes,
or even perceive when they are correctly made. But the difficulty is
not nearly so great as it at first appears: all this beautiful work
can be shown, I think, to follow from a few very simple instincts.

I was led to investigate this subject by Mr. Waterhouse, who has shown
that the form of the cell stands in close relation to the presence of
adjoining cells; and the following view may, perhaps, be considered
only as a modification of his theory. Let us look to the great
principle of gradation, and see whether Nature does not reveal to us
her method of work. At one end of a short series we have humble-bees,
which use their old cocoons to hold honey, sometimes adding to them
short tubes of wax, and likewise making separate and very irregular
rounded cells of wax. At the other end of the series we have the cells
of the hive-bee, placed in a double layer: each cell, as is well
known, is an hexagonal prism, with the basal edges of its six sides
bevelled so as to join on to a pyramid, formed of three rhombs. These
rhombs have certain angles, and the three which form the pyramidal
base of a single cell on one side of the comb, enter into the
composition of the bases of three adjoining cells on the opposite
side. In the series between the extreme perfection of the cells of the
hive-bee and the simplicity of those of the humble-bee, we have the
cells of the Mexican Melipona domestica, carefully described and
figured by Pierre Huber. The Melipona itself is intermediate in
structure between the hive and humble bee, but more nearly related to
the latter: it forms a nearly regular waxen comb of cylindrical cells,
in which the young are hatched, and, in addition, some large cells of
wax for holding honey. These latter cells are nearly spherical and of
nearly equal sizes, and are aggregated into an irregular mass. But the
important point to notice, is that these cells are always made at that
degree of nearness to each other, that they would have intersected or
broken into each other, if the spheres had been completed; but this is
never permitted, the bees building perfectly flat walls of wax between
the spheres which thus tend to intersect. Hence each cell consists of
an outer spherical portion and of two, three, or more perfectly flat
surfaces, according as the cell adjoins two, three or more other
cells. When one cell comes into contact with three other cells, which,
from the spheres being nearly of the same size, is very frequently and
necessarily the case, the three flat surfaces are united into a
pyramid; and this pyramid, as Huber has remarked, is manifestly a
gross imitation of the three-sided pyramidal basis of the cell of the
hive-bee. As in the cells of the hive-bee, so here, the three plane
surfaces in any one cell necessarily enter into the construction of
three adjoining cells. It is obvious that the Melipona saves wax by
this manner of building; for the flat walls between the adjoining
cells are not double, but are of the same thickness as the outer
spherical portions, and yet each flat portion forms a part of two

Reflecting on this case, it occurred to me that if the Melipona had
made its spheres at some given distance from each other, and had made
them of equal sizes and had arranged them symmetrically in a double
layer, the resulting structure would probably have been as perfect as
the comb of the hive-bee. Accordingly I wrote to Professor Miller, of
Cambridge, and this geometer has kindly read over the following
statement, drawn up from his information, and tells me that it is
strictly correct:--

If a number of equal spheres be described with their centres placed in
two parallel layers; with the centre of each sphere at the distance of
radius x the square root of 2 or radius x 1.41421 (or at some lesser
distance), from the centres of the six surrounding spheres in the same
layer; and at the same distance from the centres of the adjoining
spheres in the other and parallel layer; then, if planes of
intersection between the several spheres in both layers be formed,
there will result a double layer of hexagonal prisms united together
by pyramidal bases formed of three rhombs; and the rhombs and the
sides of the hexagonal prisms will have every angle identically the
same with the best measurements which have been made of the cells of
the hive-bee.

Hence we may safely conclude that if we could slightly modify the
instincts already possessed by the Melipona, and in themselves not
very wonderful, this bee would make a structure as wonderfully perfect
as that of the hive-bee. We must suppose the Melipona to make her
cells truly spherical, and of equal sizes; and this would not be very
surprising, seeing that she already does so to a certain extent, and
seeing what perfectly cylindrical burrows in wood many insects can
make, apparently by turning round on a fixed point. We must suppose
the Melipona to arrange her cells in level layers, as she already does
her cylindrical cells; and we must further suppose, and this is the
greatest difficulty, that she can somehow judge accurately at what
distance to stand from her fellow-labourers when several are making
their spheres; but she is already so far enabled to judge of distance,
that she always describes her spheres so as to intersect largely; and
then she unites the points of intersection by perfectly flat surfaces.
We have further to suppose, but this is no difficulty, that after
hexagonal prisms have been formed by the intersection of adjoining
spheres in the same layer, she can prolong the hexagon to any length
requisite to hold the stock of honey; in the same way as the rude
humble-bee adds cylinders of wax to the circular mouths of her old
cocoons. By such modifications of instincts in themselves not very
wonderful,--hardly more wonderful than those which guide a bird to
make its nest,--I believe that the hive-bee has acquired, through
natural selection, her inimitable architectural powers.

But this theory can be tested by experiment. Following the example of
Mr. Tegetmeier, I separated two combs, and put between them a long,
thick, square strip of wax: the bees instantly began to excavate
minute circular pits in it; and as they deepened these little pits,
they made them wider and wider until they were converted into shallow
basins, appearing to the eye perfectly true or parts of a sphere, and
of about the diameter of a cell. It was most interesting to me to
observe that wherever several bees had begun to excavate these basins
near together, they had begun their work at such a distance from each
other, that by the time the basins had acquired the above stated width
(i.e. about the width of an ordinary cell), and were in depth about
one sixth of the diameter of the sphere of which they formed a part,
the rims of the basins intersected or broke into each other. As soon
as this occurred, the bees ceased to excavate, and began to build up
flat walls of wax on the lines of intersection between the basins, so
that each hexagonal prism was built upon the festooned edge of a
smooth basin, instead of on the straight edges of a three-sided
pyramid as in the case of ordinary cells.

I then put into the hive, instead of a thick, square piece of wax, a
thin and narrow, knife-edged ridge, coloured with vermilion. The bees
instantly began on both sides to excavate little basins near to each
other, in the same way as before; but the ridge of wax was so thin,
that the bottoms of the basins, if they had been excavated to the same
depth as in the former experiment, would have broken into each other
from the opposite sides. The bees, however, did not suffer this to
happen, and they stopped their excavations in due time; so that the
basins, as soon as they had been a little deepened, came to have flat
bottoms; and these flat bottoms, formed by thin little plates of the
vermilion wax having been left ungnawed, were situated, as far as the
eye could judge, exactly along the planes of imaginary intersection
between the basins on the opposite sides of the ridge of wax. In
parts, only little bits, in other parts, large portions of a rhombic
plate had been left between the opposed basins, but the work, from the
unnatural state of things, had not been neatly performed. The bees
must have worked at very nearly the same rate on the opposite sides of
the ridge of vermilion wax, as they circularly gnawed away and
deepened the basins on both sides, in order to have succeeded in thus
leaving flat plates between the basins, by stopping work along the
intermediate planes or planes of intersection.

Considering how flexible thin wax is, I do not see that there is any
difficulty in the bees, whilst at work on the two sides of a strip of
wax, perceiving when they have gnawed the wax away to the proper
thinness, and then stopping their work. In ordinary combs it has
appeared to me that the bees do not always succeed in working at
exactly the same rate from the opposite sides; for I have noticed
half-completed rhombs at the base of a just-commenced cell, which were
slightly concave on one side, where I suppose that the bees had
excavated too quickly, and convex on the opposed side, where the bees
had worked less quickly. In one well-marked instance, I put the comb
back into the hive, and allowed the bees to go on working for a short
time, and again examined the cell, and I found that the rhombic plate
had been completed, and had become PERFECTLY FLAT: it was absolutely
impossible, from the extreme thinness of the little rhombic plate,
that they could have effected this by gnawing away the convex side;
and I suspect that the bees in such cases stand in the opposed cells
and push and bend the ductile and warm wax (which as I have tried is
easily done) into its proper intermediate plane, and thus flatten it.

From the experiment of the ridge of vermilion wax, we can clearly see
that if the bees were to build for themselves a thin wall of wax, they
could make their cells of the proper shape, by standing at the proper
distance from each other, by excavating at the same rate, and by
endeavouring to make equal spherical hollows, but never allowing the
spheres to break into each other. Now bees, as may be clearly seen by
examining the edge of a growing comb, do make a rough, circumferential
wall or rim all round the comb; and they gnaw into this from the
opposite sides, always working circularly as they deepen each cell.
They do not make the whole three-sided pyramidal base of any one cell
at the same time, but only the one rhombic plate which stands on the
extreme growing margin, or the two plates, as the case may be; and
they never complete the upper edges of the rhombic plates, until the
hexagonal walls are commenced. Some of these statements differ from
those made by the justly celebrated elder Huber, but I am convinced of
their accuracy; and if I had space, I could show that they are
conformable with my theory.

Huber's statement that the very first cell is excavated out of a
little parallel-sided wall of wax, is not, as far as I have seen,
strictly correct; the first commencement having always been a little
hood of wax; but I will not here enter on these details. We see how
important a part excavation plays in the construction of the cells;
but it would be a great error to suppose that the bees cannot build up
a rough wall of wax in the proper position--that is, along the plane
of intersection between two adjoining spheres. I have several
specimens showing clearly that they can do this. Even in the rude
circumferential rim or wall of wax round a growing comb, flexures may
sometimes be observed, corresponding in position to the planes of the
rhombic basal plates of future cells. But the rough wall of wax has in
every case to be finished off, by being largely gnawed away on both
sides. The manner in which the bees build is curious; they always make
the first rough wall from ten to twenty times thicker than the
excessively thin finished wall of the cell, which will ultimately be
left. We shall understand how they work, by supposing masons first to
pile up a broad ridge of cement, and then to begin cutting it away
equally on both sides near the ground, till a smooth, very thin wall
is left in the middle; the masons always piling up the cut-away
cement, and adding fresh cement, on the summit of the ridge. We shall
thus have a thin wall steadily growing upward; but always crowned by a
gigantic coping. From all the cells, both those just commenced and
those completed, being thus crowned by a strong coping of wax, the
bees can cluster and crawl over the comb without injuring the delicate
hexagonal walls, which are only about one four-hundredth of an inch in
thickness; the plates of the pyramidal basis being about twice as
thick. By this singular manner of building, strength is continually
given to the comb, with the utmost ultimate economy of wax.

It seems at first to add to the difficulty of understanding how the
cells are made, that a multitude of bees all work together; one bee
after working a short time at one cell going to another, so that, as
Huber has stated, a score of individuals work even at the commencement
of the first cell. I was able practically to show this fact, by
covering the edges of the hexagonal walls of a single cell, or the
extreme margin of the circumferential rim of a growing comb, with an
extremely thin layer of melted vermilion wax; and I invariably found
that the colour was most delicately diffused by the bees--as
delicately as a painter could have done with his brush--by atoms of
the coloured wax having been taken from the spot on which it had been
placed, and worked into the growing edges of the cells all round. The
work of construction seems to be a sort of balance struck between many
bees, all instinctively standing at the same relative distance from
each other, all trying to sweep equal spheres, and then building up,
or leaving ungnawed, the planes of intersection between these spheres.
It was really curious to note in cases of difficulty, as when two
pieces of comb met at an angle, how often the bees would entirely pull
down and rebuild in different ways the same cell, sometimes recurring
to a shape which they had at first rejected.

When bees have a place on which they can stand in their proper
positions for working,--for instance, on a slip of wood, placed
directly under the middle of a comb growing downwards so that the comb
has to be built over one face of the slip--in this case the bees can
lay the foundations of one wall of a new hexagon, in its strictly
proper place, projecting beyond the other completed cells. It suffices
that the bees should be enabled to stand at their proper relative
distances from each other and from the walls of the last completed
cells, and then, by striking imaginary spheres, they can build up a
wall intermediate between two adjoining spheres; but, as far as I have
seen, they never gnaw away and finish off the angles of a cell till a
large part both of that cell and of the adjoining cells has been
built. This capacity in bees of laying down under certain
circumstances a rough wall in its proper place between two
just-commenced cells, is important, as it bears on a fact, which seems
at first quite subversive of the foregoing theory; namely, that the
cells on the extreme margin of wasp-combs are sometimes strictly
hexagonal; but I have not space here to enter on this subject. Nor
does there seem to me any great difficulty in a single insect (as in
the case of a queen-wasp) making hexagonal cells, if she work
alternately on the inside and outside of two or three cells commenced
at the same time, always standing at the proper relative distance from
the parts of the cells just begun, sweeping spheres or cylinders, and
building up intermediate planes. It is even conceivable that an insect
might, by fixing on a point at which to commence a cell, and then
moving outside, first to one point, and then to five other points, at
the proper relative distances from the central point and from each
other, strike the planes of intersection, and so make an isolated
hexagon: but I am not aware that any such case has been observed; nor
would any good be derived from a single hexagon being built, as in its
construction more materials would be required than for a cylinder.

As natural selection acts only by the accumulation of slight
modifications of structure or instinct, each profitable to the
individual under its conditions of life, it may reasonably be asked,
how a long and graduated succession of modified architectural
instincts, all tending towards the present perfect plan of
construction, could have profited the progenitors of the hive-bee? I
think the answer is not difficult: it is known that bees are often
hard pressed to get sufficient nectar; and I am informed by Mr.
Tegetmeier that it has been experimentally found that no less than
from twelve to fifteen pounds of dry sugar are consumed by a hive of
bees for the secretion of each pound of wax; so that a prodigious
quantity of fluid nectar must be collected and consumed by the bees in
a hive for the secretion of the wax necessary for the construction of
their combs. Moreover, many bees have to remain idle for many days
during the process of secretion. A large store of honey is
indispensable to support a large stock of bees during the winter; and
the security of the hive is known mainly to depend on a large number
of bees being supported. Hence the saving of wax by largely saving
honey must be a most important element of success in any family of
bees. Of course the success of any species of bee may be dependent on
the number of its parasites or other enemies, or on quite distinct
causes, and so be altogether independent of the quantity of honey
which the bees could collect. But let us suppose that this latter
circumstance determined, as it probably often does determine, the
numbers of a humble-bee which could exist in a country; and let us
further suppose that the community lived throughout the winter, and
consequently required a store of honey: there can in this case be no
doubt that it would be an advantage to our humble-bee, if a slight
modification of her instinct led her to make her waxen cells near
together, so as to intersect a little; for a wall in common even to
two adjoining cells, would save some little wax. Hence it would
continually be more and more advantageous to our humble-bee, if she
were to make her cells more and more regular, nearer together, and
aggregated into a mass, like the cells of the Melipona; for in this
case a large part of the bounding surface of each cell would serve to
bound other cells, and much wax would be saved. Again, from the same
cause, it would be advantageous to the Melipona, if she were to make
her cells closer together, and more regular in every way than at
present; for then, as we have seen, the spherical surfaces would
wholly disappear, and would all be replaced by plane surfaces; and the
Melipona would make a comb as perfect as that of the hive-bee. Beyond
this stage of perfection in architecture, natural selection could not
lead; for the comb of the hive-bee, as far as we can see, is
absolutely perfect in economising wax.

Thus, as I believe, the most wonderful of all known instincts, that of
the hive-bee, can be explained by natural selection having taken
advantage of numerous, successive, slight modifications of simpler
instincts; natural selection having by slow degrees, more and more
perfectly, led the bees to sweep equal spheres at a given distance
from each other in a double layer, and to build up and excavate the
wax along the planes of intersection. The bees, of course, no more
knowing that they swept their spheres at one particular distance from
each other, than they know what are the several angles of the
hexagonal prisms and of the basal rhombic plates. The motive power of
the process of natural selection having been economy of wax; that
individual swarm which wasted least honey in the secretion of wax,
having succeeded best, and having transmitted by inheritance its newly
acquired economical instinct to new swarms, which in their turn will
have had the best chance of succeeding in the struggle for existence.

No doubt many instincts of very difficult explanation could be opposed
to the theory of natural selection,--cases, in which we cannot see how
an instinct could possibly have originated; cases, in which no
intermediate gradations are known to exist; cases of instinct of
apparently such trifling importance, that they could hardly have been
acted on by natural selection; cases of instincts almost identically
the same in animals so remote in the scale of nature, that we cannot
account for their similarity by inheritance from a common parent, and
must therefore believe that they have been acquired by independent
acts of natural selection. I will not here enter on these several
cases, but will confine myself to one special difficulty, which at
first appeared to me insuperable, and actually fatal to my whole
theory. I allude to the neuters or sterile females in
insect-communities: for these neuters often differ widely in instinct
and in structure from both the males and fertile females, and yet,
from being sterile, they cannot propagate their kind.

The subject well deserves to be discussed at great length, but I will
here take only a single case, that of working or sterile ants. How the
workers have been rendered sterile is a difficulty; but not much
greater than that of any other striking modification of structure; for
it can be shown that some insects and other articulate animals in a
state of nature occasionally become sterile; and if such insects had
been social, and it had been profitable to the community that a number
should have been annually born capable of work, but incapable of
procreation, I can see no very great difficulty in this being effected
by natural selection. But I must pass over this preliminary
difficulty. The great difficulty lies in the working ants differing
widely from both the males and the fertile females in structure, as in
the shape of the thorax and in being destitute of wings and sometimes
of eyes, and in instinct. As far as instinct alone is concerned, the
prodigious difference in this respect between the workers and the
perfect females, would have been far better exemplified by the
hive-bee. If a working ant or other neuter insect had been an animal
in the ordinary state, I should have unhesitatingly assumed that all
its characters had been slowly acquired through natural selection;
namely, by an individual having been born with some slight profitable
modification of structure, this being inherited by its offspring,
which again varied and were again selected, and so onwards. But with
the working ant we have an insect differing greatly from its parents,
yet absolutely sterile; so that it could never have transmitted
successively acquired modifications of structure or instinct to its
progeny. It may well be asked how is it possible to reconcile this
case with the theory of natural selection?

First, let it be remembered that we have innumerable instances, both
in our domestic productions and in those in a state of nature, of all
sorts of differences of structure which have become correlated to
certain ages, and to either sex. We have differences correlated not
only to one sex, but to that short period alone when the reproductive
system is active, as in the nuptial plumage of many birds, and in the
hooked jaws of the male salmon. We have even slight differences in the
horns of different breeds of cattle in relation to an artificially
imperfect state of the male sex; for oxen of certain breeds have
longer horns than in other breeds, in comparison with the horns of the
bulls or cows of these same breeds. Hence I can see no real difficulty
in any character having become correlated with the sterile condition
of certain members of insect-communities: the difficulty lies in
understanding how such correlated modifications of structure could
have been slowly accumulated by natural selection.

This difficulty, though appearing insuperable, is lessened, or, as I
believe, disappears, when it is remembered that selection may be
applied to the family, as well as to the individual, and may thus gain
the desired end. Thus, a well-flavoured vegetable is cooked, and the
individual is destroyed; but the horticulturist sows seeds of the same
stock, and confidently expects to get nearly the same variety;
breeders of cattle wish the flesh and fat to be well marbled together;
the animal has been slaughtered, but the breeder goes with confidence
to the same family. I have such faith in the powers of selection, that
I do not doubt that a breed of cattle, always yielding oxen with
extraordinarily long horns, could be slowly formed by carefully
watching which individual bulls and cows, when matched, produced oxen
with the longest horns; and yet no one ox could ever have propagated
its kind. Thus I believe it has been with social insects: a slight
modification of structure, or instinct, correlated with the sterile
condition of certain members of the community, has been advantageous
to the community: consequently the fertile males and females of the
same community flourished, and transmitted to their fertile offspring
a tendency to produce sterile members having the same modification.
And I believe that this process has been repeated, until that
prodigious amount of difference between the fertile and sterile
females of the same species has been produced, which we see in many
social insects.

But we have not as yet touched on the climax of the difficulty;
namely, the fact that the neuters of several ants differ, not only
from the fertile females and males, but from each other, sometimes to
an almost incredible degree, and are thus divided into two or even
three castes. The castes, moreover, do not generally graduate into
each other, but are perfectly well defined; being as distinct from
each other, as are any two species of the same genus, or rather as any
two genera of the same family. Thus in Eciton, there are working and
soldier neuters, with jaws and instincts extraordinarily different: in
Cryptocerus, the workers of one caste alone carry a wonderful sort of
shield on their heads, the use of which is quite unknown: in the
Mexican Myrmecocystus, the workers of one caste never leave the nest;
they are fed by the workers of another caste, and they have an
enormously developed abdomen which secretes a sort of honey, supplying
the place of that excreted by the aphides, or the domestic cattle as
they may be called, which our European ants guard or imprison.

It will indeed be thought that I have an overweening confidence in the
principle of natural selection, when I do not admit that such
wonderful and well-established facts at once annihilate my theory. In
the simpler case of neuter insects all of one caste or of the same
kind, which have been rendered by natural selection, as I believe to
be quite possible, different from the fertile males and females,--in
this case, we may safely conclude from the analogy of ordinary
variations, that each successive, slight, profitable modification did
not probably at first appear in all the individual neuters in the same
nest, but in a few alone; and that by the long-continued selection of
the fertile parents which produced most neuters with the profitable
modification, all the neuters ultimately came to have the desired
character. On this view we ought occasionally to find neuter-insects
of the same species, in the same nest, presenting gradations of
structure; and this we do find, even often, considering how few
neuter-insects out of Europe have been carefully examined. Mr. F.
Smith has shown how surprisingly the neuters of several British ants
differ from each other in size and sometimes in colour; and that the
extreme forms can sometimes be perfectly linked together by
individuals taken out of the same nest: I have myself compared perfect
gradations of this kind. It often happens that the larger or the
smaller sized workers are the most numerous; or that both large and
small are numerous, with those of an intermediate size scanty in
numbers. Formica flava has larger and smaller workers, with some of
intermediate size; and, in this species, as Mr. F. Smith has observed,
the larger workers have simple eyes (ocelli), which though small can
be plainly distinguished, whereas the smaller workers have their
ocelli rudimentary. Having carefully dissected several specimens of
these workers, I can affirm that the eyes are far more rudimentary in
the smaller workers than can be accounted for merely by their
proportionally lesser size; and I fully believe, though I dare not
assert so positively, that the workers of intermediate size have their
ocelli in an exactly intermediate condition. So that we here have two
bodies of sterile workers in the same nest, differing not only in
size, but in their organs of vision, yet connected by some few members
in an intermediate condition. I may digress by adding, that if the
smaller workers had been the most useful to the community, and those
males and females had been continually selected, which produced more
and more of the smaller workers, until all the workers had come to be
in this condition; we should then have had a species of ant with
neuters very nearly in the same condition with those of Myrmica. For
the workers of Myrmica have not even rudiments of ocelli, though the
male and female ants of this genus have well-developed ocelli.

I may give one other case: so confidently did I expect to find
gradations in important points of structure between the different
castes of neuters in the same species, that I gladly availed myself of
Mr. F. Smith's offer of numerous specimens from the same nest of the
driver ant (Anomma) of West Africa. The reader will perhaps best
appreciate the amount of difference in these workers, by my giving not
the actual measurements, but a strictly accurate illustration: the
difference was the same as if we were to see a set of workmen building
a house of whom many were five feet four inches high, and many sixteen
feet high; but we must suppose that the larger workmen had heads four
instead of three times as big as those of the smaller men, and jaws
nearly five times as big. The jaws, moreover, of the working ants of
the several sizes differed wonderfully in shape, and in the form and
number of the teeth. But the important fact for us is, that though the
workers can be grouped into castes of different sizes, yet they
graduate insensibly into each other, as does the widely-different
structure of their jaws. I speak confidently on this latter point, as
Mr. Lubbock made drawings for me with the camera lucida of the jaws
which I had dissected from the workers of the several sizes.

With these facts before me, I believe that natural selection, by
acting on the fertile parents, could form a species which should
regularly produce neuters, either all of large size with one form of
jaw, or all of small size with jaws having a widely different
structure; or lastly, and this is our climax of difficulty, one set of
workers of one size and structure, and simultaneously another set of
workers of a different size and structure;--a graduated series having
been first formed, as in the case of the driver ant, and then the
extreme forms, from being the most useful to the community, having
been produced in greater and greater numbers through the natural
selection of the parents which generated them; until none with an
intermediate structure were produced.

Thus, as I believe, the wonderful fact of two distinctly defined
castes of sterile workers existing in the same nest, both widely
different from each other and from their parents, has originated. We
can see how useful their production may have been to a social
community of insects, on the same principle that the division of
labour is useful to civilised man. As ants work by inherited instincts
and by inherited tools or weapons, and not by acquired knowledge and
manufactured instruments, a perfect division of labour could be
effected with them only by the workers being sterile; for had they
been fertile, they would have intercrossed, and their instincts and
structure would have become blended. And nature has, as I believe,
effected this admirable division of labour in the communities of ants,
by the means of natural selection. But I am bound to confess, that,
with all my faith in this principle, I should never have anticipated
that natural selection could have been efficient in so high a degree,
had not the case of these neuter insects convinced me of the fact. I
have, therefore, discussed this case, at some little but wholly
insufficient length, in order to show the power of natural selection,
and likewise because this is by far the most serious special
difficulty, which my theory has encountered. The case, also, is very
interesting, as it proves that with animals, as with plants, any
amount of modification in structure can be effected by the
accumulation of numerous, slight, and as we must call them accidental,
variations, which are in any manner profitable, without exercise or
habit having come into play. For no amount of exercise, or habit, or
volition, in the utterly sterile members of a community could possibly
have affected the structure or instincts of the fertile members, which
alone leave descendants. I am surprised that no one has advanced this
demonstrative case of neuter insects, against the well-known doctrine
of Lamarck.


I have endeavoured briefly in this chapter to show that the mental
qualities of our domestic animals vary, and that the variations are
inherited. Still more briefly I have attempted to show that instincts
vary slightly in a state of nature. No one will dispute that instincts
are of the highest importance to each animal. Therefore I can see no
difficulty, under changing conditions of life, in natural selection
accumulating slight modifications of instinct to any extent, in any
useful direction. In some cases habit or use and disuse have probably
come into play. I do not pretend that the facts given in this chapter
strengthen in any great degree my theory; but none of the cases of
difficulty, to the best of my judgment, annihilate it. On the other
hand, the fact that instincts are not always absolutely perfect and
are liable to mistakes;--that no instinct has been produced for the
exclusive good of other animals, but that each animal takes advantage
of the instincts of others;--that the canon in natural history, of
"natura non facit saltum" is applicable to instincts as well as to
corporeal structure, and is plainly explicable on the foregoing views,
but is otherwise inexplicable,--all tend to corroborate the theory of
natural selection.

This theory is, also, strengthened by some few other facts in regard
to instincts; as by that common case of closely allied, but certainly
distinct, species, when inhabiting distant parts of the world and
living under considerably different conditions of life, yet often
retaining nearly the same instincts. For instance, we can understand
on the principle of inheritance, how it is that the thrush of South
America lines its nest with mud, in the same peculiar manner as does
our British thrush: how it is that the male wrens (Troglodytes) of
North America, build "cock-nests," to roost in, like the males of our
distinct Kitty-wrens,--a habit wholly unlike that of any other known
bird. Finally, it may not be a logical deduction, but to my
imagination it is far more satisfactory to look at such instincts as
the young cuckoo ejecting its foster-brothers,--ants making
slaves,--the larvae of ichneumonidae feeding within the live bodies of
caterpillars,--not as specially endowed or created instincts, but as
small consequences of one general law, leading to the advancement of
all organic beings, namely, multiply, vary, let the strongest live and
the weakest die.


Distinction between the sterility of first crosses and of hybrids.
Sterility various in degree, not universal, affected by close
interbreeding, removed by domestication.
Laws governing the sterility of hybrids.
Sterility not a special endowment, but incidental on other
Causes of the sterility of first crosses and of hybrids.
Parallelism between the effects of changed conditions of life and
Fertility of varieties when crossed and of their mongrel offspring not
Hybrids and mongrels compared independently of their fertility.

The view generally entertained by naturalists is that species, when
intercrossed, have been specially endowed with the quality of
sterility, in order to prevent the confusion of all organic forms.
This view certainly seems at first probable, for species within the
same country could hardly have kept distinct had they been capable of
crossing freely. The importance of the fact that hybrids are very
generally sterile, has, I think, been much underrated by some late
writers. On the theory of natural selection the case is especially
important, inasmuch as the sterility of hybrids could not possibly be
of any advantage to them, and therefore could not have been acquired
by the continued preservation of successive profitable degrees of
sterility. I hope, however, to be able to show that sterility is not a
specially acquired or endowed quality, but is incidental on other
acquired differences.

In treating this subject, two classes of facts, to a large extent
fundamentally different, have generally been confounded together;
namely, the sterility of two species when first crossed, and the
sterility of the hybrids produced from them.

Pure species have of course their organs of reproduction in a perfect
condition, yet when intercrossed they produce either few or no
offspring. Hybrids, on the other hand, have their reproductive organs
functionally impotent, as may be clearly seen in the state of the male
element in both plants and animals; though the organs themselves are
perfect in structure, as far as the microscope reveals. In the first
case the two sexual elements which go to form the embryo are perfect;
in the second case they are either not at all developed, or are
imperfectly developed. This distinction is important, when the cause
of the sterility, which is common to the two cases, has to be
considered. The distinction has probably been slurred over, owing to
the sterility in both cases being looked on as a special endowment,
beyond the province of our reasoning powers.

The fertility of varieties, that is of the forms known or believed to
have descended from common parents, when intercrossed, and likewise
the fertility of their mongrel offspring, is, on my theory, of equal
importance with the sterility of species; for it seems to make a broad
and clear distinction between varieties and species.

First, for the sterility of species when crossed and of their hybrid
offspring. It is impossible to study the several memoirs and works of
those two conscientious and admirable observers, Kolreuter and
Gartner, who almost devoted their lives to this subject, without being
deeply impressed with the high generality of some degree of sterility.
Kolreuter makes the rule universal; but then he cuts the knot, for in
ten cases in which he found two forms, considered by most authors as
distinct species, quite fertile together, he unhesitatingly ranks them
as varieties. Gartner, also, makes the rule equally universal; and he
disputes the entire fertility of Kolreuter's ten cases. But in these
and in many other cases, Gartner is obliged carefully to count the
seeds, in order to show that there is any degree of sterility. He
always compares the maximum number of seeds produced by two species
when crossed and by their hybrid offspring, with the average number
produced by both pure parent-species in a state of nature. But a
serious cause of error seems to me to be here introduced: a plant to
be hybridised must be castrated, and, what is often more important,
must be secluded in order to prevent pollen being brought to it by
insects from other plants. Nearly all the plants experimentised on by
Gartner were potted, and apparently were kept in a chamber in his
house. That these processes are often injurious to the fertility of a
plant cannot be doubted; for Gartner gives in his table about a score
of cases of plants which he castrated, and artificially fertilised
with their own pollen, and (excluding all cases such as the
Leguminosae, in which there is an acknowledged difficulty in the
manipulation) half of these twenty plants had their fertility in some
degree impaired. Moreover, as Gartner during several years repeatedly
crossed the primrose and cowslip, which we have such good reason to
believe to be varieties, and only once or twice succeeded in getting
fertile seed; as he found the common red and blue pimpernels
(Anagallis arvensis and coerulea), which the best botanists rank as
varieties, absolutely sterile together; and as he came to the same
conclusion in several other analogous cases; it seems to me that we
may well be permitted to doubt whether many other species are really
so sterile, when intercrossed, as Gartner believes.

It is certain, on the one hand, that the sterility of various species
when crossed is so different in degree and graduates away so
insensibly, and, on the other hand, that the fertility of pure species
is so easily affected by various circumstances, that for all practical
purposes it is most difficult to say where perfect fertility ends and
sterility begins. I think no better evidence of this can be required
than that the two most experienced observers who have ever lived,
namely, Kolreuter and Gartner, should have arrived at diametrically
opposite conclusions in regard to the very same species. It is also
most instructive to compare--but I have not space here to enter on
details--the evidence advanced by our best botanists on the question
whether certain doubtful forms should be ranked as species or
varieties, with the evidence from fertility adduced by different
hybridisers, or by the same author, from experiments made during
different years. It can thus be shown that neither sterility nor
fertility affords any clear distinction between species and varieties;
but that the evidence from this source graduates away, and is doubtful
in the same degree as is the evidence derived from other
constitutional and structural differences.

In regard to the sterility of hybrids in successive generations;
though Gartner was enabled to rear some hybrids, carefully guarding
them from a cross with either pure parent, for six or seven, and in
one case for ten generations, yet he asserts positively that their
fertility never increased, but generally greatly decreased. I do not
doubt that this is usually the case, and that the fertility often
suddenly decreases in the first few generations. Nevertheless I
believe that in all these experiments the fertility has been
diminished by an independent cause, namely, from close interbreeding.
I have collected so large a body of facts, showing that close
interbreeding lessens fertility, and, on the other hand, that an
occasional cross with a distinct individual or variety increases
fertility, that I cannot doubt the correctness of this almost
universal belief amongst breeders. Hybrids are seldom raised by
experimentalists in great numbers; and as the parent-species, or other
allied hybrids, generally grow in the same garden, the visits of
insects must be carefully prevented during the flowering season: hence
hybrids will generally be fertilised during each generation by their
own individual pollen; and I am convinced that this would be injurious
to their fertility, already lessened by their hybrid origin. I am
strengthened in this conviction by a remarkable statement repeatedly
made by Gartner, namely, that if even the less fertile hybrids be
artificially fertilised with hybrid pollen of the same kind, their
fertility, notwithstanding the frequent ill effects of manipulation,
sometimes decidedly increases, and goes on increasing. Now, in
artificial fertilisation pollen is as often taken by chance (as I know
from my own experience) from the anthers of another flower, as from
the anthers of the flower itself which is to be fertilised; so that a
cross between two flowers, though probably on the same plant, would be
thus effected. Moreover, whenever complicated experiments are in
progress, so careful an observer as Gartner would have castrated his
hybrids, and this would have insured in each generation a cross with
the pollen from a distinct flower, either from the same plant or from
another plant of the same hybrid nature. And thus, the strange fact of
the increase of fertility in the successive generations of
ARTIFICIALLY FERTILISED hybrids may, I believe, be accounted for by
close interbreeding having been avoided.

Now let us turn to the results arrived at by the third most
experienced hybridiser, namely, the Honourable and Reverend W.
Herbert. He is as emphatic in his conclusion that some hybrids are
perfectly fertile--as fertile as the pure parent-species--as are
Kolreuter and Gartner that some degree of sterility between distinct
species is a universal law of nature. He experimentised on some of the
very same species as did Gartner. The difference in their results may,
I think, be in part accounted for by Herbert's great horticultural
skill, and by his having hothouses at his command. Of his many
important statements I will here give only a single one as an example,
namely, that "every ovule in a pod of Crinum capense fertilised by C.
revolutum produced a plant, which (he says) I never saw to occur in a
case of its natural fecundation." So that we here have perfect, or
even more than commonly perfect, fertility in a first cross between
two distinct species.

This case of the Crinum leads me to refer to a most singular fact,
namely, that there are individual plants, as with certain species of
Lobelia, and with all the species of the genus Hippeastrum, which can
be far more easily fertilised by the pollen of another and distinct
species, than by their own pollen. For these plants have been found to
yield seed to the pollen of a distinct species, though quite sterile
with their own pollen, notwithstanding that their own pollen was found
to be perfectly good, for it fertilised distinct species. So that
certain individual plants and all the individuals of certain species
can actually be hybridised much more readily than they can be
self-fertilised! For instance, a bulb of Hippeastrum aulicum produced
four flowers; three were fertilised by Herbert with their own pollen,
and the fourth was subsequently fertilised by the pollen of a compound
hybrid descended from three other and distinct species: the result was
that "the ovaries of the three first flowers soon ceased to grow, and
after a few days perished entirely, whereas the pod impregnated by the
pollen of the hybrid made vigorous growth and rapid progress to
maturity, and bore good seed, which vegetated freely." In a letter to
me, in 1839, Mr. Herbert told me that he had then tried the experiment
during five years, and he continued to try it during several
subsequent years, and always with the same result. This result has,
also, been confirmed by other observers in the case of Hippeastrum
with its sub-genera, and in the case of some other genera, as Lobelia,
Passiflora and Verbascum. Although the plants in these experiments
appeared perfectly healthy, and although both the ovules and pollen of
the same flower were perfectly good with respect to other species, yet
as they were functionally imperfect in their mutual self-action, we
must infer that the plants were in an unnatural state. Nevertheless
these facts show on what slight and mysterious causes the lesser or
greater fertility of species when crossed, in comparison with the same
species when self-fertilised, sometimes depends.

The practical experiments of horticulturists, though not made with
scientific precision, deserve some notice. It is notorious in how
complicated a manner the species of Pelargonium, Fuchsia, Calceolaria,
Petunia, Rhododendron, etc., have been crossed, yet many of these
hybrids seed freely. For instance, Herbert asserts that a hybrid from
Calceolaria integrifolia and plantaginea, species most widely
dissimilar in general habit, "reproduced itself as perfectly as if it
had been a natural species from the mountains of Chile." I have taken
some pains to ascertain the degree of fertility of some of the complex
crosses of Rhododendrons, and I am assured that many of them are
perfectly fertile. Mr. C. Noble, for instance, informs me that he
raises stocks for grafting from a hybrid between Rhododendron Ponticum
and Catawbiense, and that this hybrid "seeds as freely as it is
possible to imagine." Had hybrids, when fairly treated, gone on
decreasing in fertility in each successive generation, as Gartner
believes to be the case, the fact would have been notorious to
nurserymen. Horticulturists raise large beds of the same hybrids, and
such alone are fairly treated, for by insect agency the several
individuals of the same hybrid variety are allowed to freely cross
with each other, and the injurious influence of close interbreeding is
thus prevented. Any one may readily convince himself of the efficiency
of insect-agency by examining the flowers of the more sterile kinds of
hybrid rhododendrons, which produce no pollen, for he will find on
their stigmas plenty of pollen brought from other flowers.

In regard to animals, much fewer experiments have been carefully tried
than with plants. If our systematic arrangements can be trusted, that
is if the genera of animals are as distinct from each other, as are
the genera of plants, then we may infer that animals more widely
separated in the scale of nature can be more easily crossed than in
the case of plants; but the hybrids themselves are, I think, more
sterile. I doubt whether any case of a perfectly fertile hybrid animal
can be considered as thoroughly well authenticated. It should,
however, be borne in mind that, owing to few animals breeding freely
under confinement, few experiments have been fairly tried: for
instance, the canary-bird has been crossed with nine other finches,
but as not one of these nine species breeds freely in confinement, we
have no right to expect that the first crosses between them and the
canary, or that their hybrids, should be perfectly fertile. Again,
with respect to the fertility in successive generations of the more
fertile hybrid animals, I hardly know of an instance in which two
families of the same hybrid have been raised at the same time from
different parents, so as to avoid the ill effects of close
interbreeding. On the contrary, brothers and sisters have usually been
crossed in each successive generation, in opposition to the constantly
repeated admonition of every breeder. And in this case, it is not at
all surprising that the inherent sterility in the hybrids should have
gone on increasing. If we were to act thus, and pair brothers and
sisters in the case of any pure animal, which from any cause had the
least tendency to sterility, the breed would assuredly be lost in a
very few generations.

Although I do not know of any thoroughly well-authenticated cases of
perfectly fertile hybrid animals, I have some reason to believe that
the hybrids from Cervulus vaginalis and Reevesii, and from Phasianus
colchicus with P. torquatus and with P. versicolor are perfectly
fertile. The hybrids from the common and Chinese geese (A. cygnoides),
species which are so different that they are generally ranked in
distinct genera, have often bred in this country with either pure
parent, and in one single instance they have bred inter se. This was
effected by Mr. Eyton, who raised two hybrids from the same parents
but from different hatches; and from these two birds he raised no less
than eight hybrids (grandchildren of the pure geese) from one nest. In
India, however, these cross-bred geese must be far more fertile; for I
am assured by two eminently capable judges, namely Mr. Blyth and Capt.
Hutton, that whole flocks of these crossed geese are kept in various
parts of the country; and as they are kept for profit, where neither
pure parent-species exists, they must certainly be highly fertile.

A doctrine which originated with Pallas, has been largely accepted by
modern naturalists; namely, that most of our domestic animals have
descended from two or more aboriginal species, since commingled by
intercrossing. On this view, the aboriginal species must either at
first have produced quite fertile hybrids, or the hybrids must have
become in subsequent generations quite fertile under domestication.
This latter alternative seems to me the most probable, and I am
inclined to believe in its truth, although it rests on no direct
evidence. I believe, for instance, that our dogs have descended from
several wild stocks; yet, with perhaps the exception of certain
indigenous domestic dogs of South America, all are quite fertile
together; and analogy makes me greatly doubt, whether the several
aboriginal species would at first have freely bred together and have
produced quite fertile hybrids. So again there is reason to believe
that our European and the humped Indian cattle are quite fertile
together; but from facts communicated to me by Mr. Blyth, I think they
must be considered as distinct species. On this view of the origin of
many of our domestic animals, we must either give up the belief of the
almost universal sterility of distinct species of animals when
crossed; or we must look at sterility, not as an indelible
characteristic, but as one capable of being removed by domestication.

Finally, looking to all the ascertained facts on the intercrossing of
plants and animals, it may be concluded that some degree of sterility,
both in first crosses and in hybrids, is an extremely general result;
but that it cannot, under our present state of knowledge, be
considered as absolutely universal.


We will now consider a little more in detail the circumstances and
rules governing the sterility of first crosses and of hybrids. Our
chief object will be to see whether or not the rules indicate that
species have specially been endowed with this quality, in order to
prevent their crossing and blending together in utter confusion. The
following rules and conclusions are chiefly drawn up from Gartner's
admirable work on the hybridisation of plants. I have taken much pains
to ascertain how far the rules apply to animals, and considering how
scanty our knowledge is in regard to hybrid animals, I have been
surprised to find how generally the same rules apply to both kingdoms.

It has been already remarked, that the degree of fertility, both of
first crosses and of hybrids, graduates from zero to perfect
fertility. It is surprising in how many curious ways this gradation
can be shown to exist; but only the barest outline of the facts can
here be given. When pollen from a plant of one family is placed on the
stigma of a plant of a distinct family, it exerts no more influence
than so much inorganic dust. From this absolute zero of fertility, the
pollen of different species of the same genus applied to the stigma of
some one species, yields a perfect gradation in the number of seeds
produced, up to nearly complete or even quite complete fertility; and,
as we have seen, in certain abnormal cases, even to an excess of
fertility, beyond that which the plant's own pollen will produce. So
in hybrids themselves, there are some which never have produced, and
probably never would produce, even with the pollen of either pure
parent, a single fertile seed: but in some of these cases a first
trace of fertility may be detected, by the pollen of one of the pure
parent-species causing the flower of the hybrid to wither earlier than
it otherwise would have done; and the early withering of the flower is
well known to be a sign of incipient fertilisation. From this extreme
degree of sterility we have self-fertilised hybrids producing a
greater and greater number of seeds up to perfect fertility.

Hybrids from two species which are very difficult to cross, and which
rarely produce any offspring, are generally very sterile; but the
parallelism between the difficulty of making a first cross, and the
sterility of the hybrids thus produced--two classes of facts which are
generally confounded together--is by no means strict. There are many
cases, in which two pure species can be united with unusual facility,
and produce numerous hybrid-offspring, yet these hybrids are
remarkably sterile. On the other hand, there are species which can be
crossed very rarely, or with extreme difficulty, but the hybrids, when
at last produced, are very fertile. Even within the limits of the same
genus, for instance in Dianthus, these two opposite cases occur.

The fertility, both of first crosses and of hybrids, is more easily
affected by unfavourable conditions, than is the fertility of pure
species. But the degree of fertility is likewise innately variable;
for it is not always the same when the same two species are crossed
under the same circumstances, but depends in part upon the
constitution of the individuals which happen to have been chosen for
the experiment. So it is with hybrids, for their degree of fertility
is often found to differ greatly in the several individuals raised
from seed out of the same capsule and exposed to exactly the same

By the term systematic affinity is meant, the resemblance between
species in structure and in constitution, more especially in the
structure of parts which are of high physiological importance and
which differ little in the allied species. Now the fertility of first
crosses between species, and of the hybrids produced from them, is
largely governed by their systematic affinity. This is clearly shown
by hybrids never having been raised between species ranked by
systematists in distinct families; and on the other hand, by very
closely allied species generally uniting with facility. But the
correspondence between systematic affinity and the facility of
crossing is by no means strict. A multitude of cases could be given of
very closely allied species which will not unite, or only with extreme
difficulty; and on the other hand of very distinct species which unite
with the utmost facility. In the same family there may be a genus, as
Dianthus, in which very many species can most readily be crossed; and
another genus, as Silene, in which the most persevering efforts have
failed to produce between extremely close species a single hybrid.
Even within the limits of the same genus, we meet with this same
difference; for instance, the many species of Nicotiana have been more
largely crossed than the species of almost any other genus; but
Gartner found that N. acuminata, which is not a particularly distinct
species, obstinately failed to fertilise, or to be fertilised by, no
less than eight other species of Nicotiana. Very many analogous facts
could be given.

No one has been able to point out what kind, or what amount, of
difference in any recognisable character is sufficient to prevent two
species crossing. It can be shown that plants most widely different in
habit and general appearance, and having strongly marked differences
in every part of the flower, even in the pollen, in the fruit, and in
the cotyledons, can be crossed. Annual and perennial plants, deciduous
and evergreen trees, plants inhabiting different stations and fitted
for extremely different climates, can often be crossed with ease.

By a reciprocal cross between two species, I mean the case, for
instance, of a stallion-horse being first crossed with a female-ass,
and then a male-ass with a mare: these two species may then be said to
have been reciprocally crossed. There is often the widest possible
difference in the facility of making reciprocal crosses. Such cases
are highly important, for they prove that the capacity in any two
species to cross is often completely independent of their systematic
affinity, or of any recognisable difference in their whole
organisation. On the other hand, these cases clearly show that the
capacity for crossing is connected with constitutional differences
imperceptible by us, and confined to the reproductive system. This
difference in the result of reciprocal crosses between the same two
species was long ago observed by Kolreuter. To give an instance:
Mirabilis jalappa can easily be fertilised by the pollen of M.
longiflora, and the hybrids thus produced are sufficiently fertile;
but Kolreuter tried more than two hundred times, during eight
following years, to fertilise reciprocally M. longiflora with the
pollen of M. jalappa, and utterly failed. Several other equally
striking cases could be given. Thuret has observed the same fact with
certain sea-weeds or Fuci. Gartner, moreover, found that this
difference of facility in making reciprocal crosses is extremely
common in a lesser degree. He has observed it even between forms so
closely related (as Matthiola annua and glabra) that many botanists
rank them only as varieties. It is also a remarkable fact, that
hybrids raised from reciprocal crosses, though of course compounded of
the very same two species, the one species having first been used as
the father and then as the mother, generally differ in fertility in a
small, and occasionally in a high degree.

Several other singular rules could be given from Gartner: for
instance, some species have a remarkable power of crossing with other
species; other species of the same genus have a remarkable power of
impressing their likeness on their hybrid offspring; but these two
powers do not at all necessarily go together. There are certain
hybrids which instead of having, as is usual, an intermediate
character between their two parents, always closely resemble one of
them; and such hybrids, though externally so like one of their pure
parent-species, are with rare exceptions extremely sterile. So again
amongst hybrids which are usually intermediate in structure between
their parents, exceptional and abnormal individuals sometimes are
born, which closely resemble one of their pure parents; and these
hybrids are almost always utterly sterile, even when the other hybrids
raised from seed from the same capsule have a considerable degree of
fertility. These facts show how completely fertility in the hybrid is
independent of its external resemblance to either pure parent.

Considering the several rules now given, which govern the fertility of
first crosses and of hybrids, we see that when forms, which must be
considered as good and distinct species, are united, their fertility
graduates from zero to perfect fertility, or even to fertility under
certain conditions in excess. That their fertility, besides being
eminently susceptible to favourable and unfavourable conditions, is
innately variable. That it is by no means always the same in degree in
the first cross and in the hybrids produced from this cross. That the
fertility of hybrids is not related to the degree in which they
resemble in external appearance either parent. And lastly, that the
facility of making a first cross between any two species is not always
governed by their systematic affinity or degree of resemblance to each
other. This latter statement is clearly proved by reciprocal crosses
between the same two species, for according as the one species or the
other is used as the father or the mother, there is generally some
difference, and occasionally the widest possible difference, in the
facility of effecting an union. The hybrids, moreover, produced from
reciprocal crosses often differ in fertility.

Now do these complex and singular rules indicate that species have
been endowed with sterility simply to prevent their becoming
confounded in nature? I think not. For why should the sterility be so
extremely different in degree, when various species are crossed, all
of which we must suppose it would be equally important to keep from
blending together? Why should the degree of sterility be innately
variable in the individuals of the same species? Why should some
species cross with facility, and yet produce very sterile hybrids; and
other species cross with extreme difficulty, and yet produce fairly
fertile hybrids? Why should there often be so great a difference in
the result of a reciprocal cross between the same two species? Why, it
may even be asked, has the production of hybrids been permitted? to
grant to species the special power of producing hybrids, and then to
stop their further propagation by different degrees of sterility, not
strictly related to the facility of the first union between their
parents, seems to be a strange arrangement.

The foregoing rules and facts, on the other hand, appear to me clearly
to indicate that the sterility both of first crosses and of hybrids is
simply incidental or dependent on unknown differences, chiefly in the
reproductive systems, of the species which are crossed. The
differences being of so peculiar and limited a nature, that, in
reciprocal crosses between two species the male sexual element of the
one will often freely act on the female sexual element of the other,
but not in a reversed direction. It will be advisable to explain a
little more fully by an example what I mean by sterility being
incidental on other differences, and not a specially endowed quality.
As the capacity of one plant to be grafted or budded on another is so
entirely unimportant for its welfare in a state of nature, I presume
that no one will suppose that this capacity is a SPECIALLY endowed
quality, but will admit that it is incidental on differences in the
laws of growth of the two plants. We can sometimes see the reason why
one tree will not take on another, from differences in their rate of
growth, in the hardness of their wood, in the period of the flow or
nature of their sap, etc.; but in a multitude of cases we can assign
no reason whatever. Great diversity in the size of two plants, one
being woody and the other herbaceous, one being evergreen and the
other deciduous, and adaptation to widely different climates, does not
always prevent the two grafting together. As in hybridisation, so with
grafting, the capacity is limited by systematic affinity, for no one
has been able to graft trees together belonging to quite distinct
families; and, on the other hand, closely allied species, and
varieties of the same species, can usually, but not invariably, be
grafted with ease. But this capacity, as in hybridisation, is by no
means absolutely governed by systematic affinity. Although many
distinct genera within the same family have been grafted together, in
other cases species of the same genus will not take on each other. The
pear can be grafted far more readily on the quince, which is ranked as
a distinct genus, than on the apple, which is a member of the same
genus. Even different varieties of the pear take with different
degrees of facility on the quince; so do different varieties of the
apricot and peach on certain varieties of the plum.

As Gartner found that there was sometimes an innate difference in
different INDIVIDUALS of the same two species in crossing; so Sagaret
believes this to be the case with different individuals of the same
two species in being grafted together. As in reciprocal crosses, the
facility of effecting an union is often very far from equal, so it
sometimes is in grafting; the common gooseberry, for instance, cannot
be grafted on the currant, whereas the currant will take, though with
difficulty, on the gooseberry.

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