species of grain-bearing grass mentioned above is
intentionally planted. In farmer-like manner the
ground upon which it stands is carefully divested of
all other grasses and weeds during the time it is
growing. When it is ripe the grain is taken care of,
the dry stubble cut away and carried off, the paved
area being left unencumbered until the ensuing autumn,
when the same 'ant-rice' reappears within the same
circle, and receives the same agricultural attention
as was bestowed upon the previous crop--and so on year
after year, as I _know_ to be the case, in all
situations when the ant's settlements are protected
from graminivorous animals.
In a second letter Dr. Lincecum, in reply to an inquiry from Mr. Darwin
whether he supposed that the ants plant seeds for the ensuing crop,
says:--
I have not the slightest doubt of it. And my
conclusions have not been arrived at from hasty or
careless observation, nor from seeing the ants do
something that looked a little like it, and then
guessing at the results. I have at all seasons watched
the same ant-cities during the last twelve years, and
I know that what I stated in my former letter is true.
I visited the same cities yesterday, and found the
crop of ant-rice growing finely, and exhibiting also
the signs of high cultivation, and not a blade of any
other kind of grass or weed was to be seen within
twelve inches of the circular row of
ant-rice.--(_Journ. Linn. Soc._, vol. vi. p. 30-1.)
Now, MacCook found the ant-rice growing as described, but only on some
nests. Why it does not grow upon all the nests he does not understand.
So far, then, as his observations go, they confirm those of Dr.
Lincecum; but he does 'not believe that the ants deliberately sow a crop
as Lincecum asserts;' he thinks 'that they have for some reason found it
to their advantage to permit the aristida to grow upon their disks,
while they clear off all other herbage;' but finally concludes 'that
there is nothing unreasonable, nor beyond the probable capacity of the
emmet intellect, in the supposition that the crop is actually sown.
Simply, it is the Scotch verdict--"Not proven."'
The following facts with regard to 'modes of mining' are worth quoting
from MacCook:--
In sinking the galleries the difficulty of carrying is
not great in a moist or tough soil, which permits the
ant to obtain goodly-sized pellets for portage. But
when the soil is light and dry, so that it crumbles
into dust as it is bitten off, the difficulty is
greatly increased. It would be a very tedious task
indeed to take out the diggings grain by grain. This
difficulty the worker overcomes by balling the small
particles against the surface of the gallery, the
under side of the head, or within and against the
mandibles. The fore-feet are used for this purpose,
being pressed against the side face, turned under, and
pushed upward with a motion similar to that of a man
putting his hand upon his mouth. The abdomen is then
swung underneath the body and the apex pressed against
the little heap of grains of dirt massed against the
under side of the mandibles, or between that and the
smooth under surface of the head. Thus the dust is
compressed into a ball which is of sufficient size to
justify deportation.
The same operation is observed in the side-galleries,
where the ants work very frequently upon their sides
or backs, precisely as I have seen colliers do in
Pennsylvania coal-mines.
The following is likewise worth quoting from the same author:--
Seeds are evidently not the only food of our
agriculturals. When the ants at disk No. 2 had broken
through the slight mud-sediment that sealed up their
gate, as described above, they exhibited a peculiar
behaviour. Instead of heading for the roads and
pressing along them, they distributed themselves at
once over the entire disk, radiating from the gate to
all points in the circumference, from which they
penetrated the jungle of grass beyond. In a moment a
large number were returning across the roads, out of
the grass, over the pavement toward the entrance. They
bore in their mandibles objects which I presently
found to be the males and females of white ants
(_Termes flavipes_), which were filling the air,
during and after the rain, in marriage flight. They
had probably swarmed just before the shower. The
agriculturals were under great excitement, and hurried
forth and back at the top of their speed. The number
of ants bearing termites was soon so great that the
vestibule became choked, and a mass of struggling
anthood was piled up around the gate. A stream of
eager insects continually poured out of the door,
pushing their way through the crowd that vainly but
persistently endeavoured to get in with their burdens.
The outcoming ants had the advantage, and succeeded in
jostling through the quivering rosette of antennæ,
legs, heads, and abdomens. Occasionally a worker
gained an entrance by dint of sheer physical force and
perseverance. Again and again would the crowd rush
from all sides upon the gate, only to be pushed back
by the issuing throng. In the meanwhile quite a heap
of termites, a good handful at least, had been
accumulated at one side of the gate, the ants having
evidently dropped them, in despair of entrance, and
hurried off to garner more.
In due time the pressure upon the vestibule
diminished, the laden workers entered more freely, and
in the end this heap was transferred to the interior.
The rapidity with which the ants were distributed to
all parts of their roads, after the first opening of
the gates, was truly surprising. I was greatly
puzzled, at the first, to know what the cause of such
a rush might be. The whole behaviour was such as to
carry the conviction that they knew accurately what
effect the rain would have, had calculated upon it,
and were acting in accordance with previous
experience. I had no doubt at the time, and have none
now, that the capturing of insects beaten down by the
rain is one of the well-established customs of these
ants. I saw a few other insects taken in, and one
milliped, but chiefly the white ants.
That very afternoon I found in a formicary which I
then opened several large colonies, or parts of one
colony of termites, nested within the limits of the
disk and quite at home. The next day numbers of the
winged white ants were found stored within the
granaries of a large formicary. There is no reason to
doubt that these insects were intended for food, in
accordance with the quite universal habit of the
_Formicariæ_.
A curious habit has been noticed by most observers to occur in many
species of ant, and it is one on which Mr. MacCook has a good deal to
say. The habit in question consists in the ants transporting one another
from place to place. The carrying ant seizes her comrade by the middle,
and hurries along with it held aloft--the ant which is carried remaining
quite motionless with all her legs drawn together. Huber supposed the
process to be one enjoyable to both the insects concerned, and to be
performed by mutual understanding and consent; but MacCook, in common
with most other observers, supposes that it is merely a rough and
primitive way of communicating to fellow-workers the locality where
their services are required. He says:--
Keeping these facts in mind, we have a key to the
solution of the press-gang operations which Lincecum
observed among the agriculturals, and which have been
fully described in other species. In the absence of
any common head or directory, and of all executive
officers, a change of location or any other concerted
movement must be carried forward by the willing
co-operation of individuals. At first sight, the act
of seizing and carrying off workers does not appear
like an appeal to free-will. It is indeed coercive, so
far as the first act goes. But, in point of fact, the
coercion ceases the moment the captive is set down
within the precincts of the new movement. The
carrier-ant has depended upon securing her consent and
co-operation by thus bringing her within the circle of
activity for which her service is sought. As a rule,
no doubt, the deported ant at once yields to the
influence around her, and drops into the current of
fresh enterprise, in which she moves with as entire
freedom and as independently as any other worker. But
she is apparently under no restraint, and if she so
please, may return to her former haunts.
_Certain Ants of Africa._--Livingstone says of certain ants of Africa:--
They have established themselves on the plain where
water stands so long annually as to allow the lotus
and other aqueous plants to come to maturity. When all
the ant-horizon is submerged a foot deep, they manage
to exist by ascending to little houses built of black
tenaceous loam on stalks of grass, and placed higher
than the line of inundation. This must have been the
result of experience, for, if they had waited till the
water actually invaded their terrestrial habitations,
they would not have been able to procure materials for
their aërial quarters, unless they dived down to the
bottom for every mouthful of clay.[42]
_The Tree Ant of India and New South Wales._--These ants are remarkable
from their habit of forming nests only in trees. According to Col.
Sykes' account, the shape of the nest is more or less globular, and
about ten inches in diameter. It is formed entirely of cow-dung, which
the insects collect from the ground beneath, and work into the form of
thin scales. These are then built together in an imbricated manner, like
tiles or slates upon the roof of a house, the upper or outer scale,
however, being one unbroken sheet, which covers the whole nest like a
skull-cap. Below this the scales are placed one upon another in a wavy
or scalloped manner, so that numerous little arched entrances are left,
and yet, owing to the imbricated manner in which the scales are
arranged, the interior of the nest is perfectly protected from rain.
This interior consists of a number of irregular cells, the walls of
which are formed by the same process as the exterior.
In New South Wales there is another species of ant which also frequents
trees, but builds within the stem and branches. In the report of Captain
Cook's expedition its habits are thus described:--'Their habitations are
the insides of the branches of a tree, which they contrive to excavate,
by working out the pith almost to the extremity of the slenderest twig;
the tree at the same time flourishing as if it had no such inmate.' On
breaking one of the branches the ants swarm out in legions. Some of our
native species also have the habit of excavating the interior of trees,
though not on so extensive a scale.
_Honey-making Ant_ (Myrmecocystus mexicanus).--This ant is found in
Texas and New Mexico. Capt. W. B. Fleeson has observed its habits, and
his observations have been communicated to the Californian Academy of
Sciences, and also, by Mr. Henry Edwards, to Mr. Darwin. The following
are the chief points of interest in Capt. Fleeson's results:--
The community appears to consist of three distinct
kinds of ants, probably of two separate genera, whose
offices in the general order of the nest would seem to
be entirely apart from each other, and who perform the
labour allotted to them without the least encroachment
upon the duties of their fellows. These three kinds
are--
I. Yellow workers; nurses and feeders of II.
II. Yellow honey-makers; sole function to secrete a
kind of honey in their large globose abdomens, on
which the other ants are supposed to feed. They never
quit the nest, and are fed and tended by I.
III. Black workers, guards, and purveyors; surround
the nest as guards or sentinels, in a manner presently
to be described, and also forage for the food required
for I. They are much larger and stronger insects than
either I. or II., and are provided with very
formidable mandibles.
The nest is placed in sandy soil in the neighbourhood of shrubs and
flowers, is a perfect square, and occupies about four or five square
feet of ground, the surface of which is kept almost unbroken. But the
boundaries of the nest are rendered conspicuous by the guard of black
workers (III.), which continuously parade round three of its sides in a
close double line of defence, moving in opposite directions. In the
accompanying diagram this sentry path is represented by the thick black
lines. These always face the same points of the compass, and the
direction in which the sentries march is one column from south-west to
south-east, and the other column from south-east to south-west--each
column, however, moving in regular order round three sides of a square.
The southern side of the encampment is left unguarded; but if any enemy
approaches on this or any other side, a number of the guards leave their
stations, and sally forth to face the foe--raising themselves on their
hind tarsi on meeting the enemy, and moving their large mandibles in
defiance. Spiders, wasps, beetles, and other insects, if they venture
too near the nest, are torn to pieces by the guard in a most merciless
manner, and the dead body of the vanquished is speedily removed from the
neighbourhood of the nest--the guard then marching back to resume their
places in the line of defence, their object in destroying other insects
being the defence of their encampment, and not the obtaining of food.
The object of leaving the southern side of the square encampment open is
as follows. While some of the black workers are engaged on duty as
guard, another and larger division are engaged on duty as purveyors.
These enter and leave the quadrangle by its open or southern side along
the dotted line marked _a_ to the central point _c_. The incoming line
is composed of individuals each bearing a burden of fragments of
flowers or aromatic leaves. These are all deposited in the centre of the
quadrangle _c_. Along the other diagonal _e_ there is a no less
incessantly moving double line of yellow workers (I.), whose office it
is to convey the supplies deposited by the black workers at _c_ to _b_,
which is the gateway of the fortress. It is remarkable that no black ant
is ever seen upon the line _e_, and no yellow one upon the line _a_;
each keeps his own separate station, and follows his own particular duty
with a steadfastness and apparent adherence to discipline that are most
astonishing. The hole at _d_ seems to be a ventilating shaft; it is
never used as a gateway.
[Illustration: Fig. 7.]
Section of the nest reveals, besides galleries, a small chamber about
three feet below the surface, across which is spread, like a spider's
web, a network of squares spun by the insects, the squares being about
1/4 inch across, and the ends of the whole net being fastened to the
earthen walls of the chamber. In each one of the squares, supported by
the web, sits one of the honey-making ants (II.). Here these honey
makers live in perpetual confinement, and receive a constant supply of
flowers, pollen, &c., which is continually being brought them by (I.),
and which, by a process analogous to that performed by the bee, they
convert into honey.
Such is an epitome of the only account that the world has yet received
of the habits and economy of this wonderful insect, whose instincts of
military organization seem to be not less wonderful than those of the
Ecitons, though in this case they are developed with reference to
defence, and not to aggression. It is especially noteworthy that the
black and yellow workers are believed to belong to 'two separate
genera;' for if this is the case, it is the only one I can recall of two
distinct species co-operating for a common end; for even the nearest
parallel which we find supplied in other species of ants maintaining
aphides, is not quite the same thing, seeing that the aphides are merely
passive agents, like Class II., of the honey-making ant, and not
actively co-operating members of the community, like Class I.
_Ecitons._--We have next to consider the habits of the wonderful
'foraging,' or, as it might be more appropriately called, the military
ant of the Amazon. These insects, which belong to several species of the
same genus, have been carefully watched by Belt, Bates, and other
naturalists. The following facts must therefore be accepted as fully
established.
_Eciton legionis_ moves in enormous armies, and everything that these
insects do is done with the most perfect instinct of military
organization. The army marches in the form of a rather broad and regular
column, hundreds of yards in length. The object of the march is the
capture and plunder of other insects, &c., for food, and as the
well-organised host advances, its devastating legions set all other
terrestrial life at defiance. From the main column there are sent out
smaller lateral columns, the composing individuals of which play the
part of scouts, branching off in various directions, and searching
about with the utmost activity for insects, grubs, &c., over every log,
under every fallen leaf, and in every nook and cranny where there is any
chance of finding prey. When their errand is completed, they return into
the main column. If the prey found is sufficiently small for the scouts
themselves to manage, it is immediately seized, and carried back to the
main column; but if the amount is too large for the scouts to deal with
alone, messengers are sent back to the main column, whence there is
immediately dispatched a detachment large enough to cope with the
requirements. Insects which when killed are too large for single ants to
carry, are torn in pieces, and the pieces conveyed back to the main army
by different individuals. Many insects in trying to escape run up bushes
and shrubs, where they are pursued from branch to branch and twig to
twig by their remorseless enemies, until on arriving at some terminal
ramification they must either submit to immediate capture by their
pursuers, or drop down amid the murderous hosts beneath. As already
stated, all the spoils that are taken by the scouts or by the
detachments sent out in answer to their demands for assistance, are
immediately taken back to the main column. When they arrive there, they
are taken to the rear of that column by two smaller columns of carriers,
which are constantly running, one on either side of the main column,
with the supplies that are constantly pouring in from both sides. Each
of these outside columns is a double line, the ants composing one of the
two lines all running in the same direction as the main army, and the
ants composing the other line all running in the opposite direction. The
former are empty-handed carriers, which having deposited their burdens
in the rear, are again advancing to the van for fresh burdens. Those
composing the other line are all laden with the mangled remains of
insects, pupæ of other ants, &c. On either side of the main column there
are also constantly running up and down a few individuals of smaller
size and lighter colour than the other ants, which seem to play the part
of officers; for they never leave their stations, and while running up
and down the outsides of the column, they every now and again stop to
touch antennæ with some member of the rank and file, as if to give
instructions. When the scouts discover a wasp's nest in a tree, a strong
force is sent out from the main army, the nest is pulled to pieces, and
all the larvæ carried to the rear of the army, while the wasps fly
around defenceless against the invading multitude. Or, if the nest of
any other species of ant is found, a similarly strong force, or perhaps
the whole army is deflected towards it, and with the utmost energy the
innumerable insects set to work to sink shafts and dig mines till the
whole nest is rifled of its contents. In these mining operations the
ants work with an extraordinary display of organized co-operation; for
those low down in the shafts do not lose time by carrying up the earth
which they excavate, but pass on the pellets to those above; and the
ants on the surface, when they receive the pellets, carry them, 'with an
appearance of forethought that quite staggered' Mr. Bates, only just far
enough to ensure that they shall not roll back again into the shaft,
and, after depositing them, immediately hurry back for more. But there
is not a rigid division of labour, although the work 'seems to be
performed by intelligent co-operation amongst a host of eager little
creatures;' for some of them act 'sometimes as carriers of pellets, and
at another as miners, and all shortly afterwards assume the office of
conveyors of the spoil.' Again, as showing the instincts of
co-operation, the following may also be quoted from Bates's account:--
On the following morning no trace of ants could be
found near the place where I had seen them the
preceding day, nor were there signs of insects of any
description in the thicket; but at the distance of
eighty or one hundred yards, I came upon the same
army, engaged evidently on a razzia of a similar kind
to that of the previous evening; but requiring other
resources of their instinct, owing to the nature of
the ground. They were eagerly occupied on the face of
an inclined bank of light earth in excavating mines,
whence, from a depth of eight or ten inches, they were
extracting the bodies of a bulky species of ant of the
genus Formica. It was curious to see them crowding
round the orifices of the mines, some assisting their
comrades to lift out the bodies of the Formicæ, and
others tearing them in pieces, on account of their
weight being too great for a single Eciton; a number
of carriers seizing each a fragment, and carrying it
off down the slope.
These Ecitons have no fixed nest themselves, but live, as it were, on a
perpetual campaign. At night, however, they call a halt and pitch a
camp. For this purpose they usually select a piece of broken ground, in
the interstices of which they temporarily store their plunder. In the
morning the army is again on the march, and before an hour or two has
passed not a single ant is to be seen where the countless multitudes had
previously covered the ground.
Another and larger species of Eciton (_E. humata_) hunts sometimes in
dense armies, and sometimes in columns, according to the kind of prey of
which they are in search. When in columns they are seeking for the nests
of a certain species of ant which have their young in holes of rotten
logs. These Ecitons when seeking for these nests hunt about, like those
just described, in columns, which branch off in various directions. When
a fallen log is reached, the column spreads over it, searching through
all the holes and cracks. Mr. Belt says of them:--
The workers are of various sizes, and the smallest are
here of use, for they squeeze themselves into the
narrowest holes, and search out their prey in the
furthest ramifications of the nests. When a nest of
the _Hypoclinea_ is attacked, the ants rush out,
carrying the larvæ and pupæ in their jaws, but are
immediately despoiled of them by the Ecitons, which
are running about in every direction with great
swiftness. Whenever they come across a _Hypoclinea_
carrying a larva or pupa, they take it from it so
quickly, that I could never ascertain exactly how it
was done.
As soon as an Eciton gets hold of its prey, it rushes
off back along the advancing column, which is composed
of two sets, one hurrying forward, the other returning
laden with their booty, but all and always in the
greatest haste and apparent hurry. About the nest
which they are harrying, all appears in confusion,
Ecitons running here and there and everywhere in the
greatest haste and disorder; but the result of all
this apparent confusion is that scarcely a single
_Hypoclinea_ gets away with a pupa or larva. I never
saw the Ecitons injure the Hypoclineas themselves,
they were always contented with despoiling them of
their young.
The columns of this species 'are composed almost entirely of workers of
different sizes;' but, as in the species previously mentioned, 'at
intervals of two or three yards there are larger and lighter coloured
individuals that often stop, and sometimes run a little backward,
stopping and touching some of the ants with their antennæ,' and looking
'like officers giving orders and directing the march of the column.'
Concerning the other habits of this species, the same author writes:--
The eyes in the Ecitons are very small, in some of the
species imperfect, and in others entirely absent; in
this they differ greatly from the _Pseudomyrma_ ants,
which hunt singly and which have the eyes greatly
developed. The imperfection of eyesight in the Ecitons
is an advantage to the community, and to their
particular mode of hunting. It keeps them together,
and prevents individual ants from starting off alone
after objects that, if their eyesight was better, they
might discover at a distance; the Ecitons and most
other ants follow each other by scent, and, I believe,
they can communicate the presence of danger, of booty,
or other intelligence, to a distance by the different
intensity or qualities of the odours given off. I one
day saw a column of _Eciton hamata_ running along the
foot of a nearly perpendicular tramway cutting, the
side of which was about six feet high. At one point I
noticed a sort of assembly of about a dozen
individuals that appeared in consultation. Suddenly
one ant left the conclave, and ran with great speed up
the perpendicular face of the cutting without
stopping. It was followed by others, which, however,
did not keep straight on like the first, but ran a
short way, then returned, then again followed a little
further than the first time. They were evidently
scenting the trail of the pioneer, and making it
permanently recognisable. These ants followed the
exact line taken by the first one, although it was far
out of sight. Wherever it had made a slight _détour_
they did so likewise. I scraped with my knife a small
portion of the clay on the trail, and the ants were
completely at fault for a time which way to go. Those
ascending and those descending stopped at the scraped
portion, and made short circuits until they hit the
scented trail again, when all their hesitation
vanished, and they ran up and down it with the
greatest confidence. On gaining the top of the
cutting, the ants entered some brushwood suitable for
hunting. In a very short space of time the information
was communicated to the ants below, and a dense column
rushed up to search for their prey. The Ecitons are
singular amongst the ants in this respect, that they
have no fixed habitations, but move on from one place
to another, as they exhaust the hunting grounds around
them. I think _Eciton hamata_ does not stay more than
four or five days in one place. I have sometimes come
across the migratory columns; they may easily be
known. Here and there one of the light-coloured
officers moves backwards and forwards directing the
columns. Such a column is of enormous length, and
contains many thousands if not millions of
individuals. I have sometimes followed them up for two
or three hundred yards without getting to the end.
They make their temporary habitations in hollow trees,
and sometimes underneath large fallen trunks that
offer suitable hollows. A nest that I came across in
the latter situation was open at one side. The ants
were clustered together in a dense mass, like a great
swarm of bees, hanging from the roof but reaching to
the ground below. Their innumerable long legs looked
like brown threads binding together the mass, which
must have been at least a cubic yard in bulk, and
contained hundreds of thousands of individuals,
although many columns were outside, some bringing in
the pupæ of ants, others the legs and dissected bodies
of various insects. I was surprised to see in this
living nest tubular passages leading down to the
centre of the mass, kept open just as if it had been
formed of inorganic materials. Down these holes the
ants who were bringing in booty passed with their
prey. I thrust a long stick down to the centre of the
cluster, and brought out clinging to it many ants
holding larvæ and pupæ, which probably were kept warm
by the crowding together of the ants. Besides the
common dark-coloured workers and light-coloured
officers, I saw here many still larger individuals
with enormous jaws. These they go about holding wide
open in a threatening manner.
It was this ant which, as previously stated, showed sympathy and
fellow-feeling with companions in difficulties.
The habits of _E. drepanophora_ are closely similar to those of the
species already described; and, indeed, except in matters of detail,
all the species of Ecitons have much the same habits. Mr. Bates records
an interesting observation which he made on one of the moving columns of
this species. He says: 'When I interfered with the column or abstracted
an individual from it, news of the disturbance was quickly communicated
to a distance of several yards to the rear, and the column at that point
commenced retreating.' The main column is in this species narrower,
viz., 'from four to six deep,' but extends to a great length, viz., half
a mile or more. It was this species of Eciton that the same naturalist
describes as enjoying periods of leisure and recreation in the 'sunny
nooks of the forest.'
Next we have to consider _E. prædator_, of which the same observer
writes:--
This is a small dark reddish species, very similar to
the common red stinging ant of England. It differs
from all other Ecitons in its habit of hunting, not in
columns, but in dense phalanxes consisting of myriads
of individuals, and was first met with at Ega, where
it is very common. Nothing in insect movements is more
striking than the rapid march of these large and
compact bodies. Wherever they pass, all the rest of
the animal world is thrown into a state of alarm. They
stream along the ground and climb to the summits of
all the lower trees, searching every leaf to its apex,
and whenever they encounter a mass of decaying
vegetable matter, where booty is plentiful, they
concentrate, like other Ecitons, all their forces upon
it, the dense phalanx of shining and quickly-moving
bodies, as it spreads over the surface, looking like a
flood of dark red liquid. They soon penetrate every
part of the confused heap, and then, gathering
together again in marching order, onward they move.
All soft-bodied and inactive insects fall an easy prey
to them, and, like other Ecitons, they tear their
victims in pieces for facility of carriage. A phalanx
of this species, when passing over a tract of smooth
ground, occupies a space of from four to six square
yards; on examining the ants closely they are seen to
move, not all together in one straightforward
direction, but in variously spreading contiguous
columns, now separating a little from the general
mass, now reuniting with it. The margins of the
phalanx spread out at times like a cloud of
skirmishers from the flanks of an army. I was never
able to find the hive of this species.
Lastly, there are two species of Eciton totally blind, and their habits
differ from those of the species which we have hitherto considered.
Bates writes of them:--
The armies of _E. vastator_ and _E. erratica_ move, as
far as I could learn, wholly under covered roads, the
ants constructing them gradually but rapidly as they
advance. The column of foragers pushes forward step by
step, under the protection of these covered passages,
through the thickets, and on reaching a rotting log,
or other promising hunting-ground, pour into the
crevices in search of booty. I have traced their
arcades, occasionally, for a distance of one or two
hundred yards; the grains of earth are taken from the
soil over which the column is passing, and are fitted
together without cement. It is this last-mentioned
feature that distinguishes them from the similar
covered roads made by termites, who use their
glutinous saliva to cement the grains together. The
blind Ecitons, working in numbers, build up
simultaneously the sides of their convex arcades, and
contrive, in a surprising manner, to approximate them
and fit in the key-stones without letting the loose
uncemented structure fall to pieces. There was a very
clear division of labour between the two classes of
neuters in these blind species. The large-headed
class, although not possessing monstrously lengthened
jaws like the worker-majors in _E. hamata_ and _E.
drepanophora_, are rigidly defined in structure from
the small-headed class, and act as soldiers, defending
the working community (like soldier termites) against
all comers. Whenever I made a breach in one of their
covered ways, all the ants underneath were set in
commotion, but the worker-minors remained behind to
repair the damage, whilst the large-heads issued forth
in a most menacing manner, rearing their heads and
snapping their jaws with an expression of the fiercest
rage and defiance.
_Annornia arcens._--This is the so-called 'driver' or 'marching' ant of
West Africa, which in habits and intelligence closely resembles the
military ants of the other hemisphere. I shall therefore not wait again
to describe these habits in detail. Like the Ecitons, the marching ants
of Africa have no fixed nest, but make temporary halts in the shade of
hollow trees, overhanging rocks, &c. They march in large armies, and,
like the Ecitons, always in the form of a long close column; but in this
case the relative position of the carriers of spoil and larvæ is
reversed, for while these occupy the middle place the soldiers and
officers march on either side. These have large heads armed with
powerful jaws, and never take part in carrying; their function is to
maintain order, act as scouts, and attack prey. The habits of these ants
resemble most closely those of the blind Ecitons in that they very
frequently, and indeed generally, build covered ways; they do so
apparently in order to protect themselves from the heat of the African
sun. Their line of march is therefore marked by a continuous arch or
tunnel, which is always being constructed by the van of the column. The
structure is made of earth moulded together by saliva, and is very
quickly built. But it is only built in places where the line of march is
exposed to the sunlight; at night, or in the shadow of trees or long
grass, it is not made. If their camp is flooded by a tropical rainstorm,
the ants congregate in a close mass, with the younger ants in the
centre; they thus form a floating island.
It is remarkable that ants of different hemispheres should manifest so
close a similarity with respect to all these wonderful habits. The
Chasseur ants of Trinidad, and, according to Madame Merian, the ants of
visitation of Cayenne, also display habits of the same kind.
_General Intelligence of Various Species._
Many of the foregoing facts display an astonishing degree of
intelligence as obtaining among ants; for I think that however much
latitude we may be inclined to allow to 'blind instinct' in the way of
imitating actions elsewhere due to conscious purpose, some at least of
these foregoing facts can only be fairly reconciled with the view that
the insects know what they are doing and why they are doing it. But as I
am myself well aware of the difficulty that arises in all such cases of
drawing the line between purposeless instinct and purposive
intelligence, I have thought it desirable to reserve for this concluding
division of the present chapter several isolated facts which have been
observed among sundry species of ants, and which do not seem to admit
of being reasonably comprised under the category of instinctive action,
if by the latter we mean action pursued without knowledge of the
relation between the means adopted and the ends attained.
It will be remembered that our test of instinctive as distinguished from
truly intelligent action is simply whether all individuals of a species
perform similar adaptive movements under the stimulus supplied by
similar and habitual circumstances, or whether they manifest individual
and peculiar adaptive movements to meet the exigencies of novel and
peculiar circumstances. The importance of this distinction may be
rendered manifest by the following illustrations.
[Illustration: Fig. 8.]
We have already seen that the ants which Sir John Lubbock observed
display many and complex instincts, which together might seem to justify
us in anticipating that animals which present such wonderful instincts
must also present sufficient general intelligence to meet simple though
novel exigencies by such simple adaptations as the unfamiliar
circumstances require. Yet experiments which he made in this connection
seem to show that such is not the case, but that these ants, with all
their wealth of instinctive endowments, are utterly destitute of
intelligent resources; they have abundance of common and detailed
knowledge (supposing the adaptations to be made consciously) how to act
under certain complex though familiar circumstances, but appear quite
unable to originate any adaptive action to obviate even the simplest
conceivable difficulty, if this is of a kind which they have not been
previously accustomed to meet. Thus, on a horizontal rod B supported in
a saucer of water S, and therefore inaccessible to the ants from
beneath, he placed some larvæ A. On the nest N he then placed a block of
wood C D, constructed so that the portion D should touch the larvæ at
A. When the ants had made a number of journeys over C D A and back
again, he raised the block C D so that there was an interval 3/10 of an
inch between the end of the block D and the larvæ at A.
The ants kept on coming, and tried hard to reach down
from D to A, which was only just out of their
reach. . . . After a while they all gave up their
efforts and went away, losing their prize in spite of
most earnest efforts, because it did not occur to them
to drop 3/10 of an inch. At the moment when the
separation was made there were fifteen ants on the
larvæ. These could, of course, have returned if one had
stood still and allowed the others to get on its back.
This, however, did not occur to them; nor did they
think of letting themselves drop from the bottom of the
paper (P) on to the nest. Two or three, indeed, fell
down, I have no doubt by accident; but the remainder
wandered about, until at length most of them got into
the water.
In another experiment he interposed a light straw bridge on the way
between the nest and the larvæ, and when the ants had well learnt the
way, he drew the bridge a short distance towards the nest, so that a
small chasm was made in the road. The ants tried hard and ineffectually
to reach across it, but it did not occur to them to _push_ the straw
into its original position.
The following experiment is still more illustrative of the absence of
intelligence, because the adjustive action required would not demand the
exercise of such high powers of imagination and abstraction as would
have been required for the moving forwards of the paper drawbridge.
To test their intelligence I made the following
experiments: I suspended some honey over a nest of
_Lasius flavus_ at a height of about 1/2 an inch, and
accessible only by a paper bridge more than 10 feet
long. Under the glass I then placed a small heap of
earth. The ants soon swarmed over the earth on to the
glass, and began feeding on the honey. I then removed
a little of the earth, so that there was an interval
of about 1/3 of an inch between the glass and the
earth; but though the distance was so small, they
would not jump down, but preferred to go round by the
long bridge. They tried in vain to stretch up from the
earth to the glass, which, however, was just out of
their reach, though they could touch it with their
antennæ; but it did not occur to them to heap the
earth up a little, though if they had moved only half
a dozen particles of earth they would have secured for
themselves direct access to the food. This, however,
never occurred to them. At length they gave up all
attempts to reach up to the glass, and went round by
the paper bridge. I left the arrangement for several
weeks, but they continued to go round by the long
paper bridge.
Another and somewhat similar experiment consisted in placing an upright
stick A, supporting at an angle another stick B, which nearly but not
quite touched the ground at C. At the end of the stick B there were
placed some larvæ in a horizontal glass cell at D. Into this cell were
also placed a number of ants along with the larvæ. The drop from D to C
was only 1/2 an inch; 'still, though the ants reached over and showed a
great anxiety to take this short cut home, they none of them faced the
leap, but all went round by the sticks, a distance of nearly 7 feet.'
Sir John then reduced the interruption to 2/5 of an inch, so that the
ants could even touch the glass cell with their antennæ; yet all day
long the ants continued to go the long way round rather than face the
drop. Next, therefore, he took still longer sticks and tapes, and
arranged them as before, only horizontally instead of vertically. He
also placed some fine earth under the glass cell containing the larvæ.
The ants as before continued to go the long way round (16 feet), though
the drop could not have hurt either themselves or the larvæ, and though
even this drop might have been obviated by heaping up the fine earth
into a little mound 1/8 of an inch high, so as to touch the glass cell.
It is desirable, however, here to state that all species of ants do not
show this aversion to allowing themselves to drop through short
distances; for Moggridge describes the harvesting ants of Europe as
seeming rather to enjoy acrobatic performances of this kind; and the
same fact is recorded by Belt of the leaf-cutting ants of the Amazons.
Dr. Bastian, in his work on 'Brain as an Organ of Mind,' suggests that
the 'seeming lack of intelligence betrayed by our English ants, from
their disinclination to take a small leap, may be due simply to their
defective sight' (pp. 241-2). But even this consideration does not
extenuate the stupidity of the ants which failed to heap up the fine
earth to reach the glass cell which they were able to touch with their
antennæ.
That the species of ants on which Sir John Lubbock experimented were
not, however, quite destitute of intelligence is proved by the result of
the following experiment:--
I put some provisions in a shallow box with a glass
top and a single hole in one side; I then put some
specimens of _Lasius niger_ to the food, and soon a
stream of ants was at work busily carrying supplies
off to the nest. When they had got to know their way
thoroughly, and from thirty to forty were so occupied,
I poured some fine mould in front of the hole, so as
to cover it to a depth of about 1/2 an inch. I then
took out the ants which were actually in the box. As
soon as the ants had recovered from the shock of this
unexpected proceeding on my part, they began to run
all round and about the box, looking for some other
place of entrance. Finding none, however, they began
digging down into the earth just over the hole,
carrying off the grains of earth one by one and
depositing them without any order all round at a
distance of from 1/2 to 6 inches, until they had
excavated down to the doorway, when they again began
carrying off the food as before.
This experiment was several times repeated on _L. niger_ and on _L.
flavus_, always with the same result.
Thus, then, we may conclude that the reasoning power of these ants,
although shown by the first experiments to be almost _nil_, is shown by
this experiment to be not quite _nil_; for the attempt to meet the
exigencies of the case by first going round the box to seek another
entrance, before taking the labour to remove the earth from the known
entrance, implies a certain rudimentary degree of adaptive capacity
which belongs to the category of the rational.
Another point of considerable interest, as bearing on the general
intelligence of ants, is one that was brought out as the result of a
laborious series of hourly observations, extending without intermission
from 6.30 A.M. to 10 P.M. for a period of three months. The object of
these observations was to ascertain whether the principle of the
division of labour is practised by the ants. The result of these
observations was to show that during the winter-time, when the ants are
not active, certain individuals are told off to forage for supplies, and
that when any casualty overtakes these individuals, others are told off
to supply their places. Thus, in the words of Sir John Lubbock's
analysis of his lengthy tables,--
The feeders at the beginning of the experiment were
those known to us as Nos. 5, 6, and 7. On the 22nd of
November a friend, registered as No. 8, came to the
honey, and again on the 11th December; but with these
two exceptions the whole of the supplies were carried
in by Nos. 5 and 6, with a little help from No. 7.
Thinking now it might be alleged that possibly these
were merely unusually active or greedy individuals, I
imprisoned No. 6 when she came out to feed on the 5th.
As will be seen from the table, no other ant had been
out to the honey for some days; and it could therefore
hardly be accidental that on that very evening another
ant (then registered as No. 9) came out for food. This
ant, as will be seen from the table, then took the
place of No. 6 (No. 5 being imprisoned). On the 11th
January No. 9 took in all the supplies, again with a
little help from No. 7. So matters continued until the
17th, when I imprisoned No. 9, and then again, _i.e._
on the 19th, another ant (No. 10) came out for the
food, aided, on and after the 22nd, by another (No.
11). This seems to me very curious. From the 1st
November to the 5th January, with two or three casual
exceptions, the whole of the supplies were carried in
by three ants, one of whom, however, did comparatively
little. The other two are imprisoned, and then, but
not till then, a fresh ant appears on the scene. She
carries in the food for a week, and then she being
imprisoned, two others undertake the task. On the
other hand, in nest 1, when the first foragers were
not imprisoned, they continued during the whole time
to carry in the necessary supplies.
The facts, therefore, certainly seem to indicate that certain ants are
told off as foragers, and that during winter, when but little food is
required, two or three such foragers are sufficient to provide it.
Although Sir John Lubbock's ants showed such meagre resources of
intelligent adjustment, other species of ants, which we have already had
occasion to consider, appear to be as remarkable in this respect as they
are in respect of their instinctive adjustments. Unfortunately
observations on this subject are very sparse, but such as they are they
hold out a strong inducement for any one who has the opportunity to
experiment with the view of testing the intelligence of those species in
connection with which the following observations have been made.
Réaumur states that ants will make no attempt to enter
an inhabited beehive to get at the contained honey,
knowing that the bees will slaughter them if they do
so. But if the hive is uninhabited, or the bees all
dead, the ants will swarm into the hive as long as any
honey is to be found there.
P. Huber records that a wall which had been partly erected by ants was
observed by him--
As though it were intended to support the still
unfinished arched roof of a large room, which was
being built from the opposite side. But the workers
which had begun the arch had given it too low an
elevation for the wall on which it was to rest, and if
it had been continued on the same lines it would have
met the partition wall halfway up, and this was to be
avoided. I had just made this criticism to myself,
when a new arrival, after looking at the work, came to
the same conclusion. For it began at once to destroy
what had been done, and to heighten the wall on which
it was supported, and to make a new arch with the
materials of the old one under my very eyes. When the
ants begin an undertaking it seems exactly as if an
idea slowly ripened into execution in their minds.
Thus if one of them finds two stalks lying crosswise
on the nest, which make possible the formation of a
room, or some little rafters which suggest the walls
and the corners, it first observes the various parts
accurately, and then quickly and neatly heaps little
pellets of earth in the interspaces and alongside the
stalks. It brings from every side materials that seem
appropriate, and sometimes takes such from the
uncompleted works of its companions, so much is it
urged on by the idea which it has once conceived, and
by the desire to execute it. It goes and comes and
turns back again, until its plan is recognisable by
the others.
Ebrard, in his 'Etudes de Moeurs' (p. 3), gives the following
remarkable instance of the display of intelligence of _F. fusca_:--
The earth was damp and the workers were in full
swing. It was a constant coming and going of ants,
coming forth from their underground dwelling, and
carrying back little pellets of earth for building. In
order to concentrate my attention I fixed my gaze on
the largest of the rooms which were being built,
wherein several ants were busy. The work had made
considerable progress; but although a projection could
be plainly seen along the upper edge of the wall,
there remained an interspace of about twelve or
fifteen millimetres to fill in. Here would have been
the place, in order to support the earth still to be
brought in, to have had recourse to those pillars,
buttresses, or fragments of dried leaves, which many
ants are wont to use in building. But the use of this
expedient is not customary with the ants I was
observing (_F. fusca_). Our ants, however, were
sufficient for the occasion. For a moment they seemed
inclined to leave their work, but soon turned instead
to a grass-plant growing near, the long narrow leaves
of which ran close together. They chose the nearest,
and weighted its distal end with damp earth, until its
apex just bent down to the space to be covered.
Unfortunately the bend was too close to the extremity,
and it threatened to break. To prevent this
misfortune, the ants gnawed at the base of the leaf
until it bent along its whole length and covered the
space required. But as this did not seem to be quite
enough, they heaped damp earth between the base of the
plant and that of the leaf, until the latter was
sufficiently bent. After they had thus attained their
object, they heaped on the buttressing leaf the
materials required for building the arched roof.
The characteristic _trait_ of the building of ants,
says Forel, is the almost complete absence of an
unchangeable model, peculiar to each species, such as
is found in wasps, bees, and others. The ants know how
to suit their indeed little perfect work to
circumstances, and to take advantage of each
situation. Besides, each works for itself and on a
given plan, and is only occasionally aided by others
when these understand its plan. Naturally many
collisions occur, and some destroy that which others
have made. This also gives the key to understanding
the labyrinth of the dwelling. For the rest, it is
always those workers which have discovered the most
advantageous method, or which have shown the most
patience, which win over to their plan the majority of
their comrades and at last the whole colony, although
not without many fights for supremacy. But if one
succeeds in obtaining a second to follow it, and this
second draws the others after it, the first is soon
lost again in the crowd.
Espinas also observed ('Thierischen Gesellschaften,' German translation,
1879, p. 371) that each single ant made its own plan and followed it
until a comrade, which had caught the idea, joined it, and then they
worked together in the execution of the same plan.
Moggridge says of the harvesters of Europe,--
I have observed on more than one occasion that when in
digging into an ants' nest I have thrown out an
_elater_ larva, the ants would cluster round it and
direct it towards some small opening in the soil,
which it would quickly enlarge and disappear down. At
other times, however, the ants would take no notice of
the _elater_, and it is my belief that the attentions
paid to it on former occasions were purely selfish,
and that they intended to avail themselves of the
tunnel thus made down into the soil, with the view of
reopening communications with the galleries and
granaries concealed below, the approaches to which had
been covered up. I have frequently watched the ants
make use of these passages mined by the _elater_ on
these occasions.
And again, as showing apparently intelligent adaptation of their usual
habits to altered circumstances, he gives an account of the behaviour of
these ants when a great crowd of them were confined by him in a glass
jar containing earth. He says:--
On the following morning the openings were ten in
number, and the greatly increased heaps of excavated
earth showed that they must probably have been at work
all night. The amount of work done in this short time
was truly surprising, for it must be remembered that,
eighteen hours before, the earth presented a perfectly
level surface, and the larvæ and ants, now housed
below, found themselves prisoners in a strange place,
bounded by glass walls, and with no exit possible.
It seems to me that the ants displayed extraordinary
intelligence in having thus at a moment's notice
devised a plan by which the superabundant number of
workers could be employed at one time without coming
in one another's way. The soil contained in the jar
was of course less than a tenth part of that comprised
within the limits of an ordinary nest, while the
number of workers was probably more than a third of
the total number belonging to the colony. If therefore
but one or two entrances had been pierced in the soil,
the workers would have been for ever running against
one another, and a great number could never have got
below to help in the all-important task of preparing
passages and chambers for the accommodation of the
larvæ. These numerous and funnel-shaped entrances
admitted of the simultaneous descent and ascent of
large numbers of ants, and the work progressed with
proportionate rapidity. After a few days only three
entrances, and eventually only one, remained open.
Concerning the harvesting ant of Texas, the following quotation may be
made, under the present head, from MacCook. After remarking that these
ants always select sunny places wherein to build their nests, or disks,
he goes on to say that within a few paces of his tent--
A nest was made which was partly shaded by a small
mesquite tree that stood just beyond the margin of the
clearing. The sapling had probably grown up after the
location of the community, and for some reason had
been permitted to remain until too old to kill off.
The shadow thrown upon the pavement was very slight;
nevertheless, fifteen feet distant a new formicary was
being established. The path from the ranch to the
spring ran between this new hill and the old one, and
ants were in communication between the two. An opening
had been made in the ground, and the beginnings of a
new formicary were quite apparent. This is the only
instance observed of what seemed an attempt at
colonising or removing, and I associated it with the
presence of the small but growing shadow of the young
tree.
He also gives us a still more remarkable observation, which indeed, I
must candidly say, does not appear to me credible. I am, therefore, glad
to add that it does not appear very distinctly from the account whether
the author himself made the observation, or had it narrated to him by
his guide. But here is the observation in his own words:--
While studying the habits of the cutting ant I was
tempted to make a night visit to a farm some distance
from camp, by the farmer's story of depredations made
by these insects upon certain plants and vegetables. A
long, dark tramp, a blind and vain search among the
fields, compelled us at last to call out the
countryman from his bed. He led us directly to one of
the cutting ants' nests, which was overshadowed by a
young peach tree. 'There they be, sir,' cried he
triumphantly. They were agriculturals! So also were
the other nests shown. The reason for this confounding
of the two ants on the part of the people hereabouts,
and the reason for the 'cutting' operations of our
harvesters, will be explained farther on. It is only
in point here to say that the farmer affirmed that the
ants under the peach tree had stripped off the first
tender leaves last spring, so that scarcely one had
been left upon the limbs. I am convinced that the
reason for this onslaught was the desire to be rid of
the obnoxious shade, and open the formicary to the
full light of the sun.
From this account it is not very clear whether the writer himself saw
evidence of the former denudation of the tree, and if so whether there
was any indication, other than the word of the farmer, that the
denudation had been effected by the ants. To make this conclusion
credible the best conceivable evidence would be required, and this,
unfortunately, is just what we find wanting. Somewhat the same remarks
may be made on the following quotation from the same writer, though in
this case his view is to some extent supported by an observation of
Moggridge, as well as by that of Ebrard already quoted:--
Here I observed what appeared to be a new mode of
operation. The workers, in several cases, left the
point at which they had begun a cutting, ascended the
blade, and passed as far out toward the point as
possible. The blade was thus borne downward, and as
the ant swayed up and down it really seemed that she
was taking advantage of the leverage thus gained, and
was bringing the augmented force to bear upon the
fracture. In two or three cases there appeared to be a
division of labour; that is to say, while the cutter
at the roots kept on with her work, another ant
climbed the grass blade and applied the power at the
opposite end of the lever. This position may have been
quite accidental, but it certainly had the appearance
of a voluntary co-operation. I was sorry not to be
able to establish this last inference by a series of
observations, as the facts were only observed in this
one nest.
The observation of Moggridge, to which I have alluded as in some measure
rendering support to the foregoing, is as follows. Speaking of European
harvesters which he kept in an artificial nest for the purposes of close
observation, he says:--
I was also in this way able to see for myself much
that I otherwise could not have seen. Thus I was able
to watch the operation of removing roots which had
pierced through their galleries, belonging to seedling
plants growing on the surface, and which was performed
by two ants, one pulling at the free end of the root,
and the other gnawing at its fibres where the strain
was greatest, until at length it gave way.
And again,--
Two ants sometimes combine their efforts, when one
stations itself near the base of the peduncle, and
gnaws it at the point of greatest tension, while the
other hauls upon and twists it. . . . I have occasionally
seen ants engaged in cutting the capsules of certain
plants, drop them, and allow their companions below to
carry them away.
Lastly, the statements of these three observers taken together serve to
render credible the following quotation from Bingley,[43] who says that
in Captain Cook's expedition in New South Wales ants were seen by Sir
Joseph Banks and others--
As green as a leaf, which live upon trees and build
their nests of various sizes, between that of a man's
head and his fist. These nests are of a very curious
structure: they are formed by bending down several of
the leaves, each of which is as broad as a man's hand,
and gluing the points of them together so as to form a
purse. The viscous matter used for this purpose is an
animal juice. . . . Their method of bending down leaves
we had no opportunity to observe; but we saw thousands
uniting all their strength to hold them in this
position, while other busy multitudes were employed
within, in applying this gluten, that was to prevent
their returning back. To satisfy ourselves that the
leaves were bent and held down by the efforts of these
diminutive artificers, we disturbed them in their
work; and as soon as they were driven from their
station, the leaves on which they were employed sprang
up with a force much greater than we could have
thought them able to conquer by any combination of
their strength.
This remarkable fact also seems to be corroborated by the following
independent observation of Sir E. Tennent:--
The most formidable of all is the great red ant, or
Dimiya. It is particularly abundant in gardens and on
fruit-trees; it constructs its dwellings by gluing the
leaves of such species as are suitable from their
shape and pliancy into hollow balls, and these it
lines with a kind of transparent paper, like that
manufactured by the wasp. I have watched them at the
interesting operation of forming these dwellings;--a
line of ants standing on the edge of one leaf bring
another into contact with it, and hold both together
with their mandibles till their companions within
attach them firmly by means of their adhesive paper,
the assistants outside moving along as the work
proceeds. If it be necessary to draw closer a leaf too
distant to be laid hold of by the immediate workers,
they form a chain by depending one from the other till
the object is reached, when it is at length brought
into contact, and made fast by cement.
I shall now pass on to the remarkable observation communicated to Kirby
by Colonel Sykes, F.R.S., and which is thus narrated by Kirby in his
'History, Habits, and Instincts of Animals:'--
When resident at Poona, the dessert, consisting of
fruits, cakes, and various preserves, always remained
upon a small side table, in a verandah of the
dining-room. To guard against inroads, the legs of the
table were immersed in four basins filled with water;
it was removed an inch from the wall, and, to keep off
dust from open windows, was covered with a tablecloth.
At first the ants did not attempt to cross the water,
but as the strait was very narrow, from an inch to an
inch and a half, and the sweets very tempting, they
appear, at length, to have braved all risks, to have
committed themselves to the deep, to have scrambled
across the channel, and to have reached the objects of
their desires, for hundreds were found every morning
revelling in enjoyment: daily vengeance was executed
upon them without lessening their numbers; at last the
legs of the table were painted, just above the water,
with a circle of turpentine. This at first seemed to
prove an effectual barrier, and for some days the
sweets were unmolested, after which they were again
attacked by these resolute plunderers; but how they
got at them seemed totally unaccountable, till Colonel
Sykes, who often passed the table, was surprised to
see an ant drop from the wall, about a foot above the
table, upon the cloth that covered it; another and
another succeeded. So that though the turpentine and
the distance from the wall appeared effectual
barriers, still the resources of the animal, when
determined to carry its point, were not exhausted, and
by ascending the wall to a certain height, with a
slight effort against it, in falling it managed to
land in safety upon the table.
Colonel Sykes was a good observer, so that this statement, standing upon
his authority, ought not, perhaps, to be questioned. But in all cases of
remarkable intelligence displayed by animals, we naturally and properly
desire corroboration, however good the authority may be on which the
statement of such cases may rest. I will, therefore, add the following
instances of the ingenious and determined manner in which ants overcome
obstacles, and which so far lend confirmation to the above account.
Professor Leuckart placed round the trunk of a tree, which was visited
by ants as a pasture for aphides, a broad cloth soaked in tobacco-water.
When the ants returning home down the trunk of the tree arrived at the
soaked cloth, they turned round, went up the tree again to some of the
overhanging branches, and allowed themselves to drop clear of the
obnoxious barrier. On the other hand, the ants which desired to mount
the tree first examined the nature of the barrier, then turned back and
procured from a distance little pellets of earth, which they carried in
their jaws and deposited one after another upon the tobacco-cloth till a
road of earth was made across it, over which the ants passed to and fro
with impunity.
This interesting, and indeed surprising observation of Leuckart's is, in
turn, a corroboration of an almost identical one made more than a
century ago by Cardinal Fleury, and communicated by him to Réaumur, who
published it in his 'l'Histoire des Insectes' (1734). The Cardinal
smeared the trunk of a tree with birdlime in order to prevent the ants
from ascending it; but the insects overcame the obstacle by making a
road of earth, small stones, &c., as in the case just mentioned. In
another instance the Cardinal saw a number of ants make a bridge across
a vessel of water surrounding the bottom of an orange-tree tub. They did
so by conveying a number of little pieces of _wood_, the choice of which
material instead of earth or stones, as in the previous case, seems to
betoken no small knowledge of practical engineering.
Büchner, after quoting these cases, proceeds to say (_loc. cit._, p.
120),--
The ants behaved in yet more ingenious fashion under
the following very similar circumstances. Herr G.
Theuerkauf, the painter (Wasserthorstr. 49, Berlin),
writes to the author, November 18, 1875: 'A maple tree
standing on the ground of the manufacturer, Vollbaum,
of Elbing (now of Dantzic), swarmed with aphides and
ants. In order to check the mischief, the proprietor
smeared about a foot width of the ground round the
tree with tar. The first ants who wanted to cross
naturally stuck fast. But what did the next? They
turned back to the tree and carried down aphides,
which they stuck down on the tar one after another
until they had made a bridge over which they could
cross the tarring without danger. The above-named
merchant, Vollbaum, is the guarantor of this story,
which I received from his own mouth on the very spot
whereat it occurred.'
Büchner also gives the following case on the authority of Karl Vogt
(_loc. cit._, p. 128). An apiary of a friend was invaded by ants:--
To make this impossible for the future, the four legs
of the beehive-stand were put into small, shallow
bowls filled with water, as is often done with food in
ant-infested places. The ants soon found a way out of
this, or rather a way into their beloved honey, and
that over an iron staple with which the stand was
attached to a neighbouring wall. The staple was
removed, but the ants did not allow themselves to be
defeated. They climbed into some linden trees standing
near, the branches of which hung over the stand, and
then dropped upon it from the branches, doing just the
same as their comrades do with respect to food
surrounded by water, when they drop upon it from the
ceiling of the room. In order to make this impossible,
the boughs were cut away. But once more the ants were
found in the stand, and closer investigation showed
that one of the bowls was dried up, and that a crowd
of ants had gathered in it. But they found themselves
puzzled how to go on with their robbery, for the leg
did not, by chance, rest on the bottom of the bowl,
but was about half an inch from it. The ants were seen
rapidly touching each other with their antennæ, or
carrying on a consultation, until at last a rather
larger ant came forward and put an end to the
difficulty. It rose to its full height on its hind
legs, and struggled until at last it seized a rather
projecting splinter of the wooden leg, and managed to
take hold of it. As soon as this was done other ants
ran on to it, strengthened the hold by clinging, and
so made a little living bridge, over which the others
could easily pass.
The same author publishes the following very remarkable observation,
quoted from a letter to him by Dr. Ellendorf:--
It is a hard matter to protect any eatables from these
creatures, let the custody be ever so close. The legs
of cupboards and tables in or on which eatables are
kept are placed in vessels of water. I myself did
this, but I none the less found thousands of ants in
the cupboard next morning. It was a puzzle to me how
they crossed the water, but the puzzle was soon
solved; for I found a straw in one of the saucers,
which lay obliquely across the edge of the pan and
touched the leg of the press: this they had used for a
bridge. Hundreds were drowned in the water, apparently
because disorder had reigned at first, those coming
down with booty meeting those going up. But now there
was perfect order; the descending stream used one side
of the straw, the ascending the other. I now pushed
the straw about an inch away from the cupboard leg; a
terrible confusion arose. In a moment the leg
immediately over the water was covered with hundreds
of ants, feeling for the bridge in every direction
with their antennæ, running back again and coming in
ever larger swarms, as though they had communicated to
their comrades within the cupboard the fearful
misfortune that had taken place. Meanwhile the
new-comers continued to run along the straw, and not
finding the leg of the cupboard the greatest
perplexity arose. They hurried round the edge of the
pan, and soon found out where the fault lay. With
united forces they quickly pulled and pushed at the
straw, until it again came into contact with the wood,
and the communication was again restored.
This observation is strikingly, though unconsciously, confirmed by a
recent writer in the _Leisure Hour_ (1880, pp. 718-19), who having been
much troubled by small red ants in the tropics swarming over his
provisions, placed the latter in a meat-safe detached from the wall and
standing on four legs, each of which was placed in a little tin vessel
containing water. Eight or ten days afterwards he found his provisions
in the safe swarming with ants as before, and on investigating their
mode of access to them found--
Proceeding along the whitewashed wall a string of ants
going and coming from the outer door to a height of
four feet on my wall, and corresponding with that of
the safe; and looking between it and the wall, I
discovered the secret--the bridge which these
persevering little insects had made. It consisted of a
broken bit of straw, which rested with one end on a
mud buttress fixed to the wall, and the other on the
overhanging or projecting top of the safe, which came
within an inch and a half of the wall. So they must
have carried the straw up from the floor, and resting
their end of it on the support they had prepared, let
it fall until its other end reached the safe, and then
crossed and completed the structure, for it was
fastened at both ends with the mortar composed of
their saliva and fine earth. Ruthlessly I destroyed
the bridge, and moving the safe farther from the wall,
managed to prevent their inroads for that season at
least. Since then I have frequently seen short
bridges, composed entirely of the concrete or mortar
which the white ants use to cover up their workings,
extending from a damp earthen wall to anything not
more than three-quarters of an inch from it.
Of the Ecitons Mr. Belt says:--
I shall relate two more instances of the use of a
reasoning faculty in these ants. I once saw a wide
column trying to pass along a crumbling, nearly
perpendicular slope. They would have got very slowly
over it, and many of them would have fallen, but a
number having secured their hold, and reaching to each
other, remained stationary, and over them the main
column passed. Another time they were crossing a
watercourse along a small branch, not thicker than a
goose-quill. They widened this natural bridge to three
times its width by a number of ants clinging to it and
to each other on each side, over which the column
passed three or four deep; whereas excepting for this
expedient they would have had to pass over in single
file, and treble the time would have been consumed.
Can it be contended that such insects are not able to
determine by reasoning powers which is the best way of
doing a thing?
Another observer, writing from the same part of the world to Büchner,
gives a still more wonderful account of the ingenuity of Ecitons in
crossing water. This observer is Herr H. Kreplin, of Heidemühl (Station
Ducherom), 'who lived for nearly twenty years in South America as an
engineer, and had often the opportunity of seeing the driver ants in the
forests there.' He writes to Büchner, under date May 10, 1876, as
follows:--
On both sides of the train, at about 10 mm. distance
from each other, stronger ants are to be seen,
distinguishable from the others by their foxy colour
and very thick heads with gigantic mandibles. These
'thickheads' play the same _rôle_ in the ant-state for
which they are cast in cultured communities. They look
after the order of the march, and allow none to turn
either to the right or left. The least confusion in
the regularity of the march makes them turn round and
put things straight again. While the procession of the
brown workers streams on unceasingly with a swarming
motion, the 'officers,' as the natives call these
thickheads, run constantly backwards and forwards,
ready to take the command on meeting any difficulty.
The crossing of streams by these creatures is the most
interesting point. If the watercourse be narrow, the
thickheads soon find trees, the branches of which meet
on the bank on either side, and after a short halt the
column set themselves in motion over these bridges,
rearranging themselves in the narrow train with
marvellous quickness on reaching the further side. But
if no natural bridge be available for the passage,
they travel along the bank of the river until they
arrive at a flat sandy shore. Each ant now seizes a
bit of dry wood, pulls it into the water, and mounts
thereupon. The hinder rows push the front ones even
further out, holding on to the wood with their feet
and to their comrades with their jaws. In a short time
the water is covered with ants, and when the raft has
grown too large to be held together by the small
creatures' strength, a part breaks itself off and
begins the journey across, while the ants left on the
bank busily pull their bits of wood into the water,
and work at enlarging the ferry-boat until it again
breaks. This is repeated as long as an ant remains on
shore. I had often heard described this method of
crossing rivers, but in the year 1859 I had the
opportunity of seeing it for myself.
It is remarkable that the military or driving ants of Africa exhibit
precisely similar devices for the bridging of streams, namely, by
forming a chain of individuals over which the others pass. By means of
similar chains they also let themselves down from trees. It must be
observed, however, that these and all the above observations, being
independently made and separately recorded, serve to corroborate one
another so strongly that we can entertain no reasonable doubt concerning
the wonderful facts which they convey.
I shall now bring these numerous instances to a close with a quotation
from Mr. Belt, which reveals in the most unequivocal manner surprising
powers of observation and rational action on the part of the
leaf-cutting ants of South America, whose general habits we have already
considered:--
A nest was made near one of our tramways, and to get
to the trees the ants had to cross the rails, over
which the waggons were continually passing and
repassing. Every time they came along a number of ants
were crushed to death. They persevered in crossing for
some time, but at last set to work and tunnelled
underneath each rail. One day, when the waggons were
not running, I stopped up the tunnels with stones; but
although great numbers carrying leaves were thus cut
off from the nest, they would not cross the rails, but
set to work making fresh tunnels underneath them.
_Anatomy and Physiology of Nerve-centres and Sense-organs._
The foregoing facts concerning the intelligence of ants fully justifies
Mr. Darwin's observation that 'the brain of an ant is one of the most
marvellous atoms of matter in the world, perhaps more so than the brain
of a man.' It may therefore be interesting in this particular case to
depart from the lines otherwise laid down throughout the present work,
and to devote a short section to the anatomy and physiology of this
nerve-centre with its appended organs of sense.
The brain of an ant, then, is proportionally larger than that of any
other insect. (See Titus Graber, 'Insects,' vol. i. p. 255.) In
structure, also, the brain of an ant is in advance of that of other
insects, its nearest analogue being the brain of a bee. The superiority
of development is particularly remarkable with reference to the 'stalked
bodies' of Dujardin; and these are largest in neuter workers, which are
the most intelligent members of the community.
Injury of the brain causes, as in higher animals, tetanic spasms and
involuntary reflex movements, followed by stupefaction.
An ant, whose brain has been perforated by the pointed
mandibles of an amazon, remains as though nailed to
its place; a shudder runs from time to time through
its body, and one of its legs is lifted at regular
intervals. It occasionally makes a short and quick
step, as though driven by an unseen spring, but, like
that of an automaton, aimless and objectless. If it is
pulled, it makes a movement of avoidance, but falls
back into its stupefied condition as soon as it is
released. It is no longer capable of action
consciously directed to a given object; it neither
tries to escape, nor to attack, nor to go back to its
home, nor to rejoin its companions, nor to walk away;
it feels neither heat nor cold, it knows neither fear
nor desire for food. It is merely an automatic and
reflex machine, and is exactly similar to one of those
pigeons from which Flourens removed the hemispheres of
the cerebrum. Just in the same way behaves the body of
an ant from which the head has been taken away. In the
numerous fights between amazons and other ants,
countless cases have been observed of slight injury to
the brain, which have caused the most remarkable
phenomena. Many of the wounded were seized with a mad
rage, and flung themselves at every one that came in
their way, whether friend or foe. Others assumed an
appearance of indifference, and walked serenely about
in the midst of the fighting. Others exhibited a
sudden failure of strength; but they still recognised
their enemies, approached them, and tried to bite them
in cold blood, in a way quite foreign to the behaviour
of healthy ants. They were also often observed to run
round and round in a circle, the motion resembling the
_manège_, or riding-school action of mammals, when one
of the crura cerebri has been removed.
If an ant is cut in half through the thorax, so that
the great nerve ganglia of the pro-thorax remain
untouched, the behaviour of the head shows that
intelligence also remains untouched. Ants mutilated in
this way try to go forwards with their two remaining
legs, and beg with their antennæ for their companions'
aid. If one of these latter lets itself be stopped,
then we observe a lively interchange of thanks and
sympathy expressed by the actively moving antennæ.
Forel placed near to each other two such mutilated
bodies of the _F. rufibarbis_. They conversed with
each other in the above-described way, and appeared
each to beg for help. But when he put in some
similarly mutilated ants of a hostile species, _F.
sanguinea_, the picture was changed; war broke out
between these cripples just in the same way and with
the same fury as between perfect ants.[44]
The antennæ appear to be the most important of the sense-organs, as
their removal produces an extraordinary disturbance in the intelligence
of the animal. An ant so mutilated can no longer find its way or
recognise companions, and therefore is unable to distinguish between
friends and foes. It is also unable to find food, ceases to engage in
any labour, and loses all its regard for larvæ, remaining permanently
quiet and almost motionless. A somewhat similar disturbance, or rather
destruction, of the mental faculties is observable as a result of the
same mutilation in the case of bees.[45]
FOOTNOTES:
[19] While this MS. is passing through the press Sir John Lubbock has
read another paper before the Linnæan Society, which contains some
important additional matter concerning the sense of direction in ants.
It seems that in the experiment above described, the hat-box was not
provided with a cover or lid, i.e. was not a 'closed chamber,' and that
Sir John now finds the ants to take their bearings from the direction in
which they observe the light to fall upon them. For in the experiment
with the uncovered hat-box, if the source of light (candle) is moved
round together with the rotating table which supports the box, the ants
continue their way without making compensating changes in their
direction of advance. The same thing happens if the hat-box is covered,
so as to make of it a dark chamber. Direction of light being the source
of their information that their ground is being moved, we can understand
why they do not know that it is being moved when it is moved in the
direction of their advance, as in the experiment with the paper slip.
[20] It is to be noted that although ants will attack stranger ants
introduced from other nests, they will carefully tend stranger larvæ
similarly introduced.
[21] _The Naturalist in Nicaragua_, 1874, p. 26.
[22] See _Leisure Hour_, 1880, p. 390.
[23] _Introduction to Entomology_, vol. ii. p. 524.
[24] Vol. vii. pp. 443-4.
[25] Büchner, _Geistesleben der Thiere_, pp. 66-7.
[26] _Origin of Species_, 6th ed. pp. 207-8.
[27] _Loc. cit._ p. 121.
[28] _Loc. cit._ p. 123.
[29] _Origin of Species_, 6th ed. p. 218.
[30] _Geistesleben der Thiere_, pp. 145-9.
[31] _Loc. cit._
[32] _Loc. cit._ p. 337.
[33] _Loc. cit._ p. 97.
[34] _Harvesting Ants and Trap-door Spiders_, London, 1873 and
Supplement, 1874.
[35] _Journal Linn. Soc._, vol. vi. p. 29, 1862.
[36] _Agricultural Ant of Texas_, Philadelphia, 1880.
[37] _Trans. Ent. Soc. Lond._, i. 103, 1836.
[38] _Madras Journ. Lit. Sc._, 1851.
[39] For this see Moggridge, _loc. cit._ pp. 6-10, where, besides Prov.
iv. 6-8, and xxx. 25, quotations are given from Horace, Virgil, Plautus,
and others.
[40] Proc. Phil. Acad. Nat. Sci., xii. p. 445.
[41] Agricultural Ant of Texas (Lippincott & Co., Philadelphia, 1880).
[42] _Missionary Travels_, p. 328.
[43] _Animal Biography_, 'Ants.'
[44] Büchner, _Geistesleben der Thiere_, English translation, p. 49.
[45] While this work is passing through the press, an interesting Essay
has been published by Mr. MacCook on the Honey-making Ant. I am not here
able to refer to this Essay at greater length, but have done so in a
review in _Nature_ (March 2, 1882.)--G. J. R.