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Books: On the Origin of Species

C >> Charles Darwin >> On the Origin of Species

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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
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
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
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 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.

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