Anyway, no drug, not even alcohol, causes the fundamental ills of society. If we're looking for the source of our troubles, we shouldn't test people for drugs, we should test them for stupidity, ignorance, greed and love of power. P. J. O'Rourke
Chapter 8: Instinct
By: Charles Darwin, 1859
MANY instincts are so wonderful that their development will probably
appear to the reader a difficulty sufficient to overthrow my whole
theory. I may here premise that I have nothing to do with the origin
of the 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 faculties in animals of 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 require experience
to enable us to perform, when performed by an animal, more
especially by a very young one, without 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 are universal. A little
dose of judgment or reason, as Pierre Huber expresses it, often
comes into play, even with animals low in the scale of nature.
Frederic Cuvier and several of the older metaphysicians have
compared instinct with habit. This comparison gives, I think, an
accurate notion of the frame of mind under which an instinctive action
is performed, but not necessarily 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, 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 reperformed 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 deriving any benefit
from 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 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 a 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 acquired by habit.
It will be universally admitted that instincts are as important as
corporeal structures 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 was 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 in many
cases of subordinate importance to the effects of the natural
selection of what may be called spontaneous 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. 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, &c; in which case either the 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 to
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, as was first
observed by Huber, 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, 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. It is certain, from
the observations of Huber, that the aphides show no dislike to the
ants: if the latter be not present they are at last compelled to eject
their excretion. But as the excretion is extremely viscid, it is no
doubt a convenience to the aphides to have it removed; therefore
probably they do not excrete solely for the good of the ants. Although
there is no evidence that any animal performs an action for the
exclusive good of another species, yet each tries to take advantage of
the instincts of others, as each takes advantage of the weaker
bodily structure of other species. So again 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 be
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 the nests of the same species in
the northern and southern United States. Why, it has been asked, if
instinct be variable, has it not granted to the bee "the ability to
use some other material when wax was deficient"? But what other
natural material could bees use? They will work, as I have seen,
with wax hardened with vermilion or softened with lard. Andrew
Knight observed that his bees, instead of laboriously collecting
propolis, used a cement of wax and turpentine, with which he had
covered decorticated trees. It has lately been shown that bees,
instead of searching for pollen, will gladly use a very different
substance, namely oatmeal. 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. The fear of man is slowly acquired, as I have
elsewhere shown, by the various animals which inhabit desert
islands; and we see an instance of this even in England, in the
greater wildness of all our large birds in comparison with 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 mental qualities of animals of the same kind, born in a
state of nature, vary much, could be shown by many facts. Several
cases could also be adduced of occasional and strange habits in wild
animals, which, if advantageous to the species, might have given rise,
through natural selection, to new instincts. But I am well aware
that these general statements, without the facts in detail, will
produce but a feeble effect on the reader's mind. I can only repeat my
assurance, that I do not speak without good evidence.
Inherited Changes of Habit or Instinct in Domesticated Animals
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 be
enabled to see the part which habit and the selection of so-called
spontaneous variations have played in modifying the mental qualities
of our domestic animals. It is notorious how much domestic animals
vary in their mental qualities. With cats, for instance, one naturally
takes to catching rats, and another mice, and these tendencies are
known to be inherited. One cat, according to Mr. St. John, always
brought home gamebirds, another hares or rabbits, and another hunted
on marshy ground and almost nightly caught woodcocks or snipes. A
number of curious and authentic instances could be given of various
shades of disposition and of taste, and likewise of the oddest tricks,
associated with certain frames of mind or periods of time, being
inherited. But let us look to the familiar case of the breeds of the
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 behold 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 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 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: the act of
pointing is probably, as many have thought, only the exaggerated pause
of an animal preparing to spring on its prey. When the first
tendency to point 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 in
progress, as each man tries to procure, without intending to improve
the breed, dogs which stand and hunt best. On the other hand, habit
alone in some cases has sufficed; hardly any 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 can hardly
suppose that domestic rabbits have often been selected for tameness
alone; so that we must attribute at least the greater part of the
inherited change from extreme wildness to extreme tameness, to habit
and long-continued close confinement.
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 largely and how
permanently the minds of our domestic animals have been modified. 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 and some
degree of selection have 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 with them; for I am informed by Captain
Hutton that the young chickens of the parent-stock, the Gallus
bankiva, when reared in India under a hen, are at first excessively
wild. So it is with young pheasants reared in England 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 under domestication 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 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 habit and selection have probably
concurred.
Special Instincts
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,- namely, the instinct which leads the cuckoo
to lay her eggs in other birds' nests; the slave-making instinct of
certain ants; and the cell-making power of the hive-bee. These two
latter instincts have generally and justly been ranked by
naturalists as the most wonderful of all known instincts.
Instincts of the Cuckoo.- It is supposed by some naturalists that
the more immediate cause of the instinct of the cuckoo 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 migrates 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 both asserted and denied that the
American cuckoo occasionally lays her eggs in other birds' nests;
but I have lately heard from Dr. Merrell, of Iowa, that he once
found in Illinois a young cuckoo together with a young jay in the nest
of a blue jay (Garrulus cristatus); and as both were nearly full
feathered, there could be no mistake in their identification. I
could also 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, and that she occasionally laid an egg
in another bird's nest. If the old bird profited by this occasional
habit through being enabled to migrate earlier or through any other
cause; or if the young were made more vigorous by advantage being
taken of the mistaken instinct of another species than when reared
by their own mother, encumbered as she could 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 us 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 more successful in rearing their young. By a
continued process of this nature, I believe that the strange
instinct of our cuckoo has been generated. It has, also, recently been
ascertained on sufficient evidence, by Adolf Muller, that the cuckoo
occasionally lays her eggs on the bare ground, sits on them, and feeds
her young. This rare event is probably a case of reversion to the
long-lost, aboriginal instinct of nidification.
It has been objected that I have not noticed other related instincts
and adaptations of structure in the cuckoo, which are spoken of as
necessarily co-ordinated. But in all cases, speculation on an instinct
known to us only in a single species, is useless, for we have hitherto
had no facts to guide us. Until recently the instincts of the European
and of the nonparasitic American cuckoo alone were known. now, owing
to Mr. Ramsay's observations, we have learnt something about three
Australian species, which lay their eggs in other birds' nests. The
chief points to be referred to are three: first, that the common
cuckoo, with rare exceptions, lays only one egg in a nest, so that the
large and voracious young bird receives ample food. Secondly, that the
eggs are remarkably small, not exceeding those of the skylark,- a bird
about one-fourth as large as the cuckoo. That the small size of the
egg is a real cause of adaptation we may infer from the fact of the
non-parasitic American cuckoo laying full-sized eggs. Thirdly, that
the young cuckoo, soon after birth, has the instinct, the strength,
and a properly shaped back for ejecting its foster-brothers, which
then perish from cold and hunger. This has been boldly called a
beneficent arrangement, in order that the young cuckoo may get
sufficient food, and that its foster-brothers may perish before they
had acquired much feeling!
Turning now to the Australian species; though these birds
generally lay only one egg in a nest, it is not rare to find two or
even three eggs in the same nest. In the bronze cuckoo the eggs vary
greatly in size, from eight to ten times in length. Now if it had been
of an advantage to this species to have laid eggs even smaller than
those now laid, so as to have deceived certain foster-parents, or,
as is more probable, to have been hatched within a shorter period (for
it is asserted that there is a relation between the size of eggs and
the period of their incubation), then there is no difficulty in
believing that a race or species might have been formed which would
have laid smaller and smaller eggs; for these would have been more
safely hatched and reared. Mr. Ramsay remarks that two of the
Australian cuckoos, when they lay their eggs in an open nest, manifest
a decided preference for nests containing eggs similar in colour to
their own. The European species apparently manifests some tendency
towards a similar instinct, but not rarely departs from it, as is
shown by her laying her dull and pale-coloured eggs in the nest of the
Hedge-warbler with bright greenish-blue eggs. Had our cuckoo
invariably displayed the above instinct, it would assuredly have
been added to those which it is assumed must all have been acquired
together. The eggs of the Australian bronze cuckoo vary, according
to Mr. Ramsay, to an extraordinary degree in colour; so that in this
respect, as well as in size, natural selection might have secured
and fixed any advantageous variation.
In the case of the European cuckoo, the offspring of the
foster-parents are commonly ejected from the nest within three days
after the cuckoo is hatched; and as the latter at this age is in a
most helpless condition, Mr. Gould was formerly inclined to believe
that the act of ejection was performed by the foster-parents
themselves. But he has now received a trustworthy account of a young
cuckoo which was actually seen, whilst still blind and not able even
to hold up its own head, in the act of ejecting its foster-brothers.
One of these was replaced in the nest by the observer, and was again
thrown out. With respect to the means by which this strange and odious
instinct was acquired, if it were of great importance for the young
cuckoo, as is probably the case, to receive as much food as possible
soon after birth, I can see no special difficulty in its having
gradually acquired, during successive generations, the blind desire,
the strength, and structure necessary for the work of ejection; for
those young cuckoos which had such habits and structure best developed
would be the most securely reared. The first step towards the
acquisition of the proper instinct might have been more
unintentional restlessness on the part of the young bird, when
somewhat advanced in age and strength; the habit having been
afterwards improved, and transmitted to an earlier age. I can see no
more difficulty in this, than in the unhatched young of other birds
acquiring the instinct to break through their own shells;- or than
in young snakes acquiring in their upper jaws, as Owen has remarked, a
transitory sharp tooth for cutting through the tough egg-shell. For if
each part is liable to individual variations at all ages, and the
variations tend to be inherited at a corresponding or earlier age,-
propositions which cannot be disputed,- then the instincts and
structure of the young could be slowly modified as surely as those
of the adult; and both cases must stand or fall together with the
whole theory of natural selection.
Some species of Molothrus, a widely distinct genus of American
birds, allied to our starlings, have parasitic habits like those of
the cuckoo; and the species present an interesting gradation in the
perfection of their instincts. The sexes of Molothrus badius are
stated by an excellent observer, Mr. Hudson, sometimes to live
promiscuously together in flocks, and sometimes to pair. They either
build a nest of their own, or seize on one belonging to some other
bird, occasionally throwing out the nestlings of the stranger. They
either lay their eggs in the nest thus appropriated, or oddly enough
build one for themselves on the top of it. They usually sit on their
own eggs and rear their own young; but Mr. Hudson says it is
probable that they are occasionally parasitic, for he has seen the
young of this species following old birds of a distinct kind and
clamouring to be fed by them. The parasitic habits of another
species of Molothrus, the M. bonariensis, are much more highly
developed than those of the last, but are still far from perfect. This
bird, as far as it is known, invariably lays its eggs in the nests
of strangers; but it is remarkable that several together sometimes
commence to build an irregular untidy nest of their own, placed in
singularly ill-adapted situations, as on the leaves of a large
thistle. They never, however, as far as Mr. Hudson has ascertained,
complete a nest for themselves. They often lay so many eggs- from
fifteen to twenty- in the same foster-nest, that few or none can
possibly be hatched. They have, moreover, the extraordinary habit of
pecking holes in the eggs, whether of their own species or of their
foster-parents, which they find in the appropriated nests. They drop
also many eggs on the bare ground, which are thus wasted. A third
species, the M. pecoris of North America, has acquired instincts as
perfect as those of the cuckoo, for it never lays more than one egg in
a foster-nest, so that the young bird is securely reared. Mr. Hudson
is a strong disbeliever in evolution, but he appears to have been so
much struck by the imperfect instincts of the Molothrus bonariensis
that he quotes my words, and asks, "Must we consider these habits, not
as especially endowed or created instincts, but as small
consequences of one general law, namely, transition?"
Various birds, as has already been remarked, occasionally lay
their eggs in the nest of other birds. This habit is not very uncommon
with the Gallinaceae, and throws some light on the singular instinct
of the ostrich. In this family several hen-birds 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 with the cuckoo, at
intervals of two or three days. The instinct, however, of the American
ostrich, as in the case of the Molothrus bonariensis, 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 regularly lay their eggs in the nests
of other kinds of bees. This case is more remarkable than that of
the cuckoo; for these bees have not only had their instincts but their
structure modified in accordance with their parasitic habits; for they
do not possess the pollen-collecting apparatus which would have been
indispensable if they had stored up food for their own young. Some
species of Sphegidea (wasp-like insects) are likewise parasitic; 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, yet that, when this insect finds
a burrow already made and stored by another species, it takes
advantage of the prize and becomes for the occasion parasitic. In this
case, as with that of the Molothrus or 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 feloniously appropriated, be not thus exterminated.
Slave-making instinct.- 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
female do no work of any kind, and 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 own 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
speculate how so wonderful an instinct could have been perfected.
Another species, 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 existence of so extraordinary an instinct as that of
making slaves. Hence, I will give the observations which I 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
(F. fusca) 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 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 together 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 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 year 1860, 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, the slaves in
Switzerland 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 witnessed a migration of F. sanguinea from one
nest to another, and it was a most interesting spectacle to behold the
masters carefully carrying their slaves in their jaws instead of being
carried by them, as in the case of F. rufescens. 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 slavemaking 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 their 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 slavemaking 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 returning home and entering their nests, carrying
the dead bodies of F. fusca (showing that it was not a migration)
and numerous pupae. I traced a long file of ants burthened with booty,
for about forty yards back, 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, an image of despair 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 continental 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 the pupae of other species, if scattered near
their nests, it is possible that such pupae originally stored as
food might become developed; and the foreign 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, natural selection might increase and modify 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 rufescens.
Cell-making instinct of the Hive-Bee.- 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
effected by a crowd of bees working in a dark hive. Granting
whatever instincts you please, 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 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 this 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 an inverted pyramid, 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 flat surfaces, according as the cell
adjoins two, three, or more other cells. When one cell rests on
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
base 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, and what is more important, labour, 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 cells.
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 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 squares 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. But I hear from Prof. Wyman, who has made
numerous careful measurements, that the accuracy of the workmanship of
the bee has been greatly exaggerated; so much so, that whatever the
typical form of the cell may be, it is rarely, if ever, realised.
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 that Melipona to have the
power of forming 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
many insects make in wood, 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 to a certain extent; and then she unites the points of
intersection by perfectly flat surfaces. By such modifications of
instincts which in themselves are 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, rectangular 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 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
scalloped 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, rectangular 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 bases; and these flat bases, formed by
thin little plates of the vermilion wax 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 some parts, only small portions, in other parts,
large portions of a rhombic plate were thus 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 in circularly gnawing away and deepening the basins on both sides
of the ridge of vermilion wax, in order to have thus succeeded in
leaving flat plates between the basins, by stopping work at the 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
plate, that they could have effected this by gnawing away the convex
side; and I suspect that the bees in such cases stand on opposite
sides 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 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 this
away 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 that 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 would 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 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. These walls, as Professor
Miller has kindly ascertained for me, vary greatly in thickness;
being, on an average of twelve measurements made near the border of
the comb, 1/352nd of an inch in thickness; whereas the basal
rhomboidal plates are thicker, nearly in the proportion of three to
two, having a mean thickness, from twenty-one measurements, of 1/229th
of an inch. By the above 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 it 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 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 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 were to 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.
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: cells constructed like those of the
bee or the wasp gain in strength, and save much in labour and space,
and in the materials of which they are constructed. With respect to
the formation of wax, 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 proved that from twelve to fifteen pounds
of dry sugar are consumed by a hive of bees for the secretion of a
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 and the time
consumed in collecting the honey must be an important element of
success to any family of bees. Of course the success of the species
may be dependent on the number of its enemies, or parasites, or on
quite distinct causes, and so be altogether independent of the
quantity of honey which the bees can collect. But let us suppose
that this latter circumstance determined, as it probably often has
determined, whether a bee allied to our humble-bees could exist in
large numbers in any country; and let us further suppose that the
community lived through 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 imaginary humble-bee if a slight modification in
her instincts 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 labour and wax. Hence it would
continually be more and more advantageous to our humble-bees, if
they were to make their 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 the adjoining cells, and much labour
and 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
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
labour and 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 the construction of
cells of due strength and of the proper size and shape for the larvae,
this being effected with the greatest possible economy of labour and
wax; that individual swarm which thus made the best cells with least
labour, and least waste of honey in the secretion of wax, having
succeeded best, and having transmitted their newly-acquired economical
instincts to new swarms, which in their turn will have had the best
chance of succeeding in the struggle for existence.
Objections to the Theory of Natural Selection as applied to
Instincts: Neuter and Sterile Insects
It has been objected to the foregoing view of the origin of
instincts that "the variations of structure and of instinct must
have been simultaneous and accurately adjusted to each other, as a
modification in the one without an immediate corresponding change in
the other would have been fatal." The force of this objection rests
entirely on the assumption that the changes in the instincts and
structure are abrupt. To take as an illustration the case of the
larger titmouse (Parus major) alluded to in a previous chapter; this
bird often holds the seeds of the yew between its feet on a branch,
and hammers with its beak till it gets at the kernel. Now what special
difficulty would there be in natural selection preserving all the
slight individual variations in the shape of the beak, which were
better and better adapted to break open the seeds, until a beak was
formed, as well constructed for this purpose as that of the
nuthatch, at the same time that habit, or compulsion, or spontaneous
variations of taste, led the bird to become more and more of a
seed-eater? In this case the beak is supposed to be slowly modified by
natural selection, subsequently to, but in accordance with, slowly
changing habits or taste; but let the feet of the titmouse vary and
grow larger from correlation with the beak, or from any other
unknown cause, and it is not improbable that such larger feet would
lead the bird to climb more and more until it acquired the
remarkable climbing instinct and power of the nuthatch. In this case a
gradual change of structure is supposed to lead to changed instinctive
habits. To take one more case: few instincts are more remarkable
than that which leads the swift of the Eastern Islands to make its
nest wholly of inspissated saliva. Some birds build their nests of
mud, believed to be moistened with saliva; and one of the swifts of
North America makes its nest (as I have seen) of sticks agglutinated
with saliva, and even with flakes of this substance. Is it then very
improbable that the natural selection of individual swifts, which
secreted more and more saliva, should at last produce a species with
instincts leading it to neglect other materials, and to make its
nest exclusively of inspissated saliva? And so in other cases. It
must, however, be admitted that in many instances we cannot conjecture
whether it was instinct or structure which first varied.
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 have originated; cases, in which no
intermediate gradations are known to exist; cases of instincts of 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 progenitor, and
consequently must believe that they were independently acquired
through 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 the 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 especial difficulty
in this having been effected through 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 wonderful difference in this
respect between the workers and the perfect females, would have been
better exemplified by the hive-bee. If a working ant or other neuter
insect had been an ordinary animal, I should have unhesitatingly
assumed that all its characters had been slowly acquired through
natural selection; namely, by individuals having been born with slight
profitable modifications, which were inherited by the offspring; and
that these again varied and again were 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 inherited structure which are correlated
with certain ages, and with either sex. We have differences correlated
not only with one sex, but with that short period 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 the oxen of other
breeds, relatively to the length of the horns in both the bulls and
cows of these same breeds. Hence I can see no great difficulty in
any character becoming 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. Breeders of cattle wish the flesh and fat to be
well marbled together: an animal thus characterised has been
slaughtered, but the breeder has gone with confidence to the same
stock and has succeeded. Such faith may be placed in the power of
selection, that a breed of cattle, always yielding oxen with
extraordinarily long horns, could, it is probable, be formed by
carefully watching which individual bulls and cows, when matched,
produced oxen with the longest horns; and yet no ox would ever have
propagated its kind. Here is a better and real illustration: according
to M. Verlot, some varieties of the double annual Stock from having
been long and carefully selected to the right degree, always produce a
large proportion of seedlings bearing double and quite sterile
flowers; but they likewise yield some single and fertile plants. These
latter, by which alone the variety can be propagated, may be
compared with the fertile male and female ants, is ants, and the
double sterile plants with the neuters of the same community. As
with the varieties of the stock, so with social insects, selection has
been applied to the family, and not to the individual, for the sake of
gaining a serviceable end. Hence we may conclude that slight
modifications of structure or of instinct, correlated with the sterile
condition of certain members of the community, have proved
advantageous: consequently the fertile males and females have
flourished, and transmitted to their fertile offspring a tendency to
produce sterile members with the same modifications. This process must
have been repeated many times, 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 acme 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 commonly 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 Myrmecoeystus, 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
and 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 the theory. In
the simpler case of neuter insects all of one caste, which, as I
believe, have been rendered different from the fertile males and
females through natural selection, we may conclude from the analogy of
ordinary variations, that the successive, slight, profitable
modifications did not first arise in all the neuters in the same nest,
but in some few alone; and that by the survival of the communities
with females which produced most INSTINCT is neuters having the
advantageous modifications, all the neuters ultimately came to be thus
characterised. According to this view we ought occasionally to find in
the same nest neuter insects, presenting gradations of structure;
and this we do find, even not rarely, considering how few neuter
insects out of Europe have been carefully examined. Mr. F. Smith has
shown that the neuters of several British ants differ surprisingly
from each other in size and sometimes in colour; and that the
extreme forms can be linked together by individuals taken out of the
same nest: I have myself compared perfect gradations of this kind.
It sometimes happens that the larger or the smaller sized workers
are the most numerous; or that both large and small are numerous,
whilst those of an intermediate size are scanty in numbers. Formica
lava has larger and smaller workers, with some few 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 here we 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
were in this condition; we should then have had a species of ant
with neuters in nearly the same condition as 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
occasionally to find gradations of important structures 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 in addition 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 size, yet they graduate insensibly into each
other, as does the widely-different structure of their jaws. I speak
confidently on this latter point, as Sir J. Lubbock made drawings
for me, with the camera lucida, of the jaws which I dissected from the
workers of the several sizes. Mr. Bates, in his interesting Naturalist
on the Amazons, has described analogous cases.
With these facts before me, I believe that natural selection, by
acting on the fertile ants or parents, could form a species which
should regularly produce neuters, all of large size with one form of
jaw, or all of small size with widely different jaws; or lastly, and
this is the greatest 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 first been formed, as
in the case of the driver ant, and then the extreme forms having
been produced in greater and greater numbers, through the survival
of the parents which generated them, until none with an intermediate
structure were produced.
An analogous explanation has been given by Mr. Wallace, of the
equally complex case, of certain Malayan butterflies regularly
appearing under two or even three distinct female forms; and by
Fritz Muller, of certain Brazilian crustaceans likewise appearing
under two widely distinct male forms. But this subject need not here
be discussed.
I have now explained how, 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 ants, on the same principle that the division
of labour is useful to civilised man. Ants, however, work by inherited
instincts and by inherited organs or tools, whilst man works by
acquired knowledge and manufactured instruments. But I must confess,
that, with all my faith in natural selection, I should never have
anticipated that this principle could have been efficient in so high a
degree, had not the case of these neuter insects led me to this
conclusion. 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 may be effected by the accumulation of
numerous, slight, spontaneous variations, which are in any way
profitable, without exercise or habit having been brought into play.
For peculiar habits confined to the workers or sterile females,
however long they might be followed, could not possibly affect the
males and fertile females, which alone leave descendants. I am
surprised that no one has hitherto advanced this demonstrative case of
neuter insects, against the well-known doctrine of inherited habit, as
advanced by Lamarck.
Summary
I have endeavoured in this chapter briefly 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
there is no real difficulty, under changing conditions of life, in
natural selection accumulating to any extent slight modifications of
instinct which are in any way useful. In many 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 can be shown to have been produced for the good of other
animals, though animals take 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 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 tropical
South America lines its nest with mud, in the same peculiar manner
as does our British thrush; how it is that the hornbills of Africa and
India have the same extraordinary instinct of plastering up and
imprisoning the females in a hole in a tree, with only a small hole
left in the plaster through which the males feed them and their
young when hatched; how it is that the male wrens (Troglodytes) of
North America build "cocknests," to roost in, like the males of our
kittywrens,- 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
ichneumonidea 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.
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