BEES AND WASPS.
ARRANGING this chapter under the same general headings as the one on
ants, we shall consider first--
_Powers of Special Sense._
Bees and wasps have much greater powers of sight than ants. They not
only perceive objects at a greater distance, but are also able to
distinguish their colours. This was proved by Sir John Lubbock, who
placed honey on slips of paper similarly formed, but of different
colours; when a bee had repeatedly visited a slip of one colour (A), he
transposed the slips during the absence of the bee; on its return the
insect did not fly to slip B, although this now occupied the _position_
which had been previously occupied by slip A, but again visited slip A,
although this now occupied the position which had been previously
occupied by slip B. Therefore, as these experiments were again and again
repeated both on bees and wasps with uniform results, there can be no
question that the insects by their first visits to slip A established an
association between the colour of A and the honey upon it, such that,
when they again returned and found B in the place of A, they were guided
by their memory of the colour rather than by their memory of the
position. It was thus shown that the insects could distinguish green,
red, yellow, and blue. These experiments also brought out the further
fact that both bees and wasps exhibit a marked preference for some
colours over others. Thus, in a series of black, white, yellow, orange,
green, blue, and red slips, two or three bees paid twenty-one visits to
the orange and yellow, and only four to all the other slips. The slips
were then moved, after which, out of thirty-two visits, twenty-two were
to the orange and yellow. Another colour to which a similar preference
is shown is blue.
As regards scent, Sir John found that on putting a few drops of eau de
Cologne at the entrance of a beehive, 'immediately a number (about 15)
came out to see what was the matter.' Other scents had a similar effect;
but on repetition several times the bees became accustomed to the scent,
and no longer came out.
As in ants, so in bees, Sir John's experiments failed to yield any
evidence of a sense of hearing. But in this connection we must not
forget the well-known fact, first observed by Huber, that the queen bee
will answer by a certain sound the peculiar piping of a pupa queen; and
again, by making a certain cry or humming noise, will strike
consternation suddenly on all the bees in the hive--these remaining for
a long time motionless as if stupefied.
_Sense of Direction._
The following are Sir John Lubbock's observations upon this subject in
the case of bees and wasps:--
Every one has heard of a 'bee-line.' It would be no
less correct to speak of a wasp-line. On August 6 I
marked a wasp, the nest of which was round the corner
of the house, so that her direct way home was not out
at the window by which she had entered, but in the
opposite direction, across the room to a window which
was closed. I watched her for some hours, during which
time she constantly went to the wrong window, and lost
much time in buzzing about at it. For ten consecutive
days this wasp paid numerous visits, coming in at the
open window, and always trying, though always
unsuccessfully, to return to her nest in the
'wasp-line' of the closed window--buzzing about that
window for hours at a time, though eventually on
finding it closed she returned and went round through
the open window by which she always entered.
This observation shows how strong must be the instinct in a wasp to take
the shortest way home, and how much the insect depends upon its sense of
direction in so doing. It also shows how long a time it requires to
learn by individual experience the properties of a previously unknown
substance such as glass. But to this latter point we shall presently
have occasion to return.
Next we must adduce evidence to show that in way-finding the 'sense of
direction' in bees appears to be largely supplemented by observation of
particular objects.
Sir John Lubbock observes: 'I never found bees to return if brought any
considerable distance at once. By taking them, however, some twenty
yards each time they came to the honey, I at length _trained_ them to
come to my room;' that is to say, bees require to _learn_ their way
little by little before they can return to a store of honey which they
may have been fortunate enough to find; their general sense of direction
is not in itself a sufficient guide. This, at least, is the case where,
as in the experiments in question, the bees are _carried_ from the hive
to the store of honey (here a distance of less than 200 yards): possibly
if they had found the honey by themselves flying towards it, and so
probably taking note of objects by the way, one journey might have
proved sufficient to teach them the way. But, whether or not this would
have been the case, the fact that when carried they required also to be
taught the way piece by piece, is conclusive proof that their sense of
direction _alone_ is not sufficient to enable them to traverse a route
of 200 yards a second time.
The same result is brought out by other experiments conducted on a
different plan, though not apparently with this object. 'My room is
square, with two windows on the south-west side, where the hive was
placed, and one on the south-east.' Besides the ordinary entrance from
outside, the hive had a small postern door opening into the room.
At 6.50 a bee came out through the little postern
door. After she had fed, she evidently did not know
her way home; so I put her back.
At 7.10 she came out again. I again fed her and put
her back.
At 10.15 she came out a third time; and again I had to
put her back.
At 10.55 she came out again, and still did not
remember the door. Though I was satisfied that she
really wished to return, and was not voluntarily
remaining outside; still, to make the matter clear, I
turned her out of a side window into the garden, when
she at once returned to the hive.
At 11.15 she came out again; and again I had to show
her the way back.
At 11.20 she came out again; and again I had to show
her the way back (this makes five times); when,
however,--
At 11.30 she came out again after feeding, she
returned straight to the hive.
At 11.40 she came out, fed, and returned straight to
the hive.
At 11.50 she came out, fed, and returned straight to
the hive; she then stayed in for some time.
At 12.30 she came out again, but seemed to have
forgotten the way back; after some time, however, she
found the door and went in.
Again:--August 24 at 7.20 a bee came through the
postern: I fed her; and though she was not frightened
or disturbed, when she had finished her meal she flew
to the window and had evidently lost her way; so at 8
o'clock I in pity put her back myself.
August 29.--A bee came out to the honey at 10.10; at
10.12 she flew to the window, and remained buzzing
about till 11.12, when, being satisfied that she could
not find her way, I put her in.
Nay, even those who seemed to know the postern, if
taken near the other window, flew to it, and seemed to
have lost themselves.
This cost me a great many bees. Those which got into
my room by accident continually died on the floor near
the window.
These observations show that even when a bee is not _carried_ from the
hive to the honey, but herself _flies_ to it, her sense of direction is
not alone sufficient to enable her to find the way back to the hive--or,
rather, to the unaccustomed entrance to the hive from which she had come
out. Probably if the side window had been open, the bee would have
returned to the hive round the corner of the house, and through the
entrance to which she was most accustomed. But as it was she had to
_learn_, by five or six journeys, the way between the postern entrance
and the food.
But the following observation on a wasp is in this connection the most
conclusive.
A marked wasp visited honey exposed in the room before mentioned. 'The
next morning she came--
At 7.25, and fed till 7.28, when she began flying
about the room and even into the next; so I thought it
well to put her out of the window, when she flew
straight away to her nest. My room, as already
mentioned, had windows on two sides; and the nest was
in the direction of a closed window, so that the wasp
had to go out of her way in going out through the open
one.
At 7.45 she came back. I had moved the glass
containing the honey about two yards; and though it
stood conspicuously, the wasp seemed to have much
difficulty in finding it. Again she flew to the window
in the direction of her nest, and I had to put her
out, which I did at 8.2.
At 8.15 she returned to the honey almost straight.
8.21, she flew again to the closed window, and
apparently could not find her way; so at 8.35 I put
her out again. It seems obvious from this that wasps
have a sense of direction, and do not find their way
merely by sight.
At 8.50 back to honey, and 8.54 again to wrong window;
but finding it closed, she took two or three turns
round the room, and then flew out through the open
window.
At 9.24 back to the honey; and 9.27 away, first,
however, paying a visit to the wrong window, but
without alighting.
At 9.36 back to the honey, and 9.39 away, but, as before, going
first to wrong window.
She was away therefore 9 minutes.
9.50 " 9.53 away, this time straight. " 11 "
10 " 10. 7 " " 11 "
10.19 " 10.22 " " 12 "
10.35 " 10.39 " " 13 "
10.47 " 10.50 " " 9 "
11. 4 " 11. 7 " " 14 "
11.21 " 11.24 " " 14 "
11.34 " 11.37 " " 10 "
11.49 " 11.52 " " 1 "
12. 3 " 12. 5 " Away therefore 11 minutes.
12.13 " 12.15½ " " 8 "
12.25 " 12.28 " " 10 "
12.39 " 12.43 " " 11 "
12.54 " 12.57 " " 11 "
1.15 " 1.19 " " 18 "
1.27 " 1.30 " " 8 minutes,'
&c., &c., the way being now clearly well learnt.
But that the sense of direction is of much service to bees in finding
the locality of their hives seems to be indicated by the following
observation thus narrated, on the authority of the authors themselves,
by Messrs. Kirby and Spence:--
In vain, during my stay at St. Nicholas, I sallied out
at every outlet to try to gain some idea of the extent
and form of the town. Trees, trees, trees, still met
me, and intercepted the view in every direction; and I
defy any inhabitant bee of this rural metropolis,
after once quitting its hive, ever to gain a glimpse
of it again until nearly perpendicularly over it. The
bees, therefore, . . . must be led to their abodes by
instinct, &c.
The observation, however, is not so conclusive as its authors suppose;
for there is nothing to show that the bees did not take note of
particular objects on their accustomed routes, and so learn these routes
by stages. It would be worth while in this connection to try the effect
of hooding the eyes of a bee, or, if this were deemed too disturbing an
experiment, removing the hive bodily to a distance from its accustomed
site, and observing whether the bees start away boldly as before for
long flights, or learn their new routes by stages.
In this connection I may quote the following.
Mr. John Topham, of Marlborough House, Torquay, writing to 'Nature,'[46]
says:--
On October 29, 1873, I removed a hive of bees in my
garden, after it was quite dark, for a distance of 12
yards from the place in which it had stood for several
months; and between its original situation and the new
one there was a bushy evergreen tree, so that all
sight of its former place was obstructed to a person
looking from the new situation of the hive.
Notwithstanding this change, the bees every day flew
to the locality where they formerly lived, and
continued flying around the site of what had been
their home until, as night came on, they many of them
sank upon the grass exhausted and chilled by the cold.
Numbers, however, returned alive to their new
position, after having looked in vain for their hive
in its old place. At night I picked the exhausted bees
up, and having restored warmth to them (by leaving
them for a time on my coat-sleeve), I returned them to
their companions.
Here was an illustration that the faculty of memory
was superior to that of observation; but that was not
all. Nearly every bee which I picked up during the 23
days through which this effort of memory lasted was an
_old one_, as was easily deduced from observing the
worn edges of the wings; showing that whilst the young
insects were quick in receiving new impressions and in
correcting errors, the nervous system of the old bees
continued _acting in the direction which early habit
had effected_. So true it is that 'one touch of nature
makes the whole world kin.'
A closely similar observation has been told me by a friend, Mr. George
Turner. He found that when he removed a beehive only a yard or two from
its accustomed site, the bees, on returning home, flew in swarms around
the latter, and for a long time were unable to find the hive. And
several other similar cases might be adduced. Lastly, Thompson says:--
It is highly remarkable that they [bees] know their
hive more from its locality than from its appearance,
for if it be removed during their absence and a
similar one be substituted, they enter the strange
one. If the position of a hive be changed, the bees
for the first day take no distant flight till they
have thoroughly scrutinised every object in its
neighbourhood.[47]
On the other hand, the writer of the article on 'Bees' in the
'Encyclopædia Britannica' says that in certain parts of France it is the
habit of bee-keepers to place a number of hives upon a boat, which, in
charge of a man, floats slowly down a river. The bees are thus
continuously changing their pasture-ground, and yet do not lose their
locomotive hives.
It may be here worth while to add, parenthetically, as the only
authentic observation with which I am acquainted concerning the distance
that bees are accustomed to forage, the following statement of Prof.
Hugh Blackburn. Writing from Glasgow University to 'Nature,'[48] he says
that bees are found in a certain peach-house every spring at the time of
blossom, although, so far as he can ascertain, the beehives nearest to
the peach-house in question are his own, and these are at a distance of
ten miles.
On the whole, then, and in the absence of further experiments, we must
conclude it to be probable that the sense of direction with which
hymenopterous insects are, as shown by some of Sir John Lubbock's
experiments, unquestionably endowed, is of no small use to them in
finding their way from home to food and _vice versâ_; although it
appears certain, from other of his experiments, that this sense of
direction is not in all cases a sufficient guide, and therefore requires
to be supplemented by the definite observation of landmarks.
But the most conclusive evidence on this latter point is afforded by a
highly interesting observation of Mr. Bates on the sand-wasps at
Santurem, which may here be suitably introduced, as the insects are not
distantly allied. He describes these animals as always taking a few
turns in the air round the hole they had made in the sand before leaving
to seek for flies in the forest, apparently in order to mark well the
position of the burrow, so that on their return they might find it
without difficulty. This observation has been since confirmed in a
striking manner by Mr. Belt, who found that the sand-wasp takes the most
precise bearings of an object the position of which she desires to
remember. This observation is so interesting that it deserves to be
rendered _in extenso_:--
A specimen of _Polistes carnifex_ (_i.e._ the
sand-wasp noticed by Mr. Bates) was hunting about for
caterpillars in my garden. I found one about an inch
long, and held it out towards it on the point of a
stick. It seized it immediately, and commenced biting
it from head to tail, soon reducing the soft body to a
mass of pulp. It rolled up about one-half of it into a
ball, and prepared to carry it off. Being at the time
amidst a thick mass of a fine-leaved climbing plant,
it proceeded, before flying away, to take note of the
place where it was leaving the other half. To do this,
it hovered in front of it for a few seconds, then took
small circles in front of it, then larger ones round
the whole plant. I thought it had gone, but it
returned again, and had another look at the opening in
the dense foliage down which the other half of the
caterpillar lay. It then flew away, but must have left
its burden for distribution with its comrades at the
nest, for it returned in less than two minutes, and
making one circle around the bush, descended to the
opening, alighted on a leaf, and ran inside. The green
remnant of the caterpillar was lying on another leaf
inside, but not connected with the one on which the
wasp alighted, so that in running in it missed it, and
soon got hopelessly lost in the thick foliage. Coming
out again, it took another circle, and pounced down on
the same spot again, as soon as it came opposite to
it. Three small seed-pods, which here grew close
together, formed the marks that I had myself taken to
note the place, and these the wasp seemed also to have
taken as its guide, for it flew directly down to them,
and ran inside; but the small leaf on which the
fragment of caterpillar lay not being directly
connected with any on the outside, it again missed it,
and again got far away from the object of its search.
It then flew out again, and the same process was
repeated again and again. Always when in circling
round it came in sight of the seed-pods down it
pounced, alighted near them, and recommenced its quest
on foot. I was surprised at its perseverance, and
thought it would have given up the search; but not so,
it returned at least half-a-dozen times, and seemed to
get angry, hurrying about with buzzing wings. At last
it stumbled across its prey, seized it eagerly, and as
there was nothing more to come back for, flew straight
off to its nest, without taking any further note of
the locality. Such an action is not the result of
blind instinct, but of a thinking mind; and it is
wonderful to see an insect so differently constructed
using a mental process similar to that of man.
_Memory._
We may here first allude to an observation of Sir John Lubbock already
quoted in another connexion (see p. 147). It is here evident that the
wasp, after finding the store of honey in the room, and after finding
the window closed in the 'wasp-line' direction to its nest, required
three repeated _lessons_ from Sir John before she _learnt_ that the
window on the other side of the room, and away from the direction of her
nest, afforded no obstacle to her exit. Having learnt this, the fourth
time she came she again flew to the closed window as before, and then,
as if but dimly remembering that there was another opening somewhere
that offered no such mysterious resistance to her passage, 'she took two
or three turns round the room, and then flew out through the open
window.' Having now taken the bearings of all the room upon her own
wings, and having again found the difference between the two windows in
respect of resistance, although in all other respects so much alike, the
next time she came she made in the first instance as it were an
experimental flight towards the closed window, but clearly had the
alternative of going to the open one in her memory; for on finding the
window closed as before, she did not alight, but flew straight from the
closed to the open window. The same thing happened once again, but now,
with the distinction between the two windows thus fully learnt, and with
it the perception that in this case 'the shortest cut was the longest
way round,' she never again flew to the closed window; in the forty
successive visits which she paid through the remainder of that day, and
the hundred visits or so which she made during the two following days,
she seems to have uniformly flown to the open window.
As evidence of _forgetfulness_, it will be enough to refer to the case
of another wasp which, under precisely similar circumstances to those
just detailed, learnt her way out of the open window one day, having
made fifty passages through it in five hours. Yet Sir John remarks,--
It struck me as curious that on the following day this
wasp seemed by no means so sure of her way, but over
and over again went to the closed window.
It is further of interest to note, as showing the similarity of the
memory displayed by these insects with that of the higher animals, that
there are considerable individual differences to be found in the degree
of its manifestation.
In this respect they certainly differ considerably.
Some of the bees which came out of the little postern
door (already described) were able to find their way
back after it had been shown to them a few times.
Others were much more stupid; thus one bee came out on
the 9th, 10th, 11th, 12th, 14th, 15th, 16th, 17th,
18th, and 19th, and came to the honey; but though I
repeatedly put her back through the postern, she was
never able to find her way for herself.
I often found that if bees which were brought to honey
did not return at once, still they would do so a day
or two afterwards. For instance, on July 11, 1874, a
hot thundery day, and when the bees were much out of
humour, I brought twelve bees to some honey; only one
came back, and that one only twice; but on the
following day several of them returned.
This latter observation is important, as proving that bees can remember
for at least a whole day the locality where they have found honey only
once before, and that they so far think about their past experiences as
to return to that locality when foraging.
As the association of ideas by contiguity is the principle which forms
the basis of all psychology, it is desirable to consider still more
attentively this the earliest manifestation that we have of it in the
memory of the Hymenoptera. That it is not exercised with exclusive
reference to _locality_ is proved by the following observation of Sir
John Lubbock:--
I kept a specimen of _Polistes Gallica_ for no less
than nine months.[49] . . . I had no difficulty in
inducing her to feed on my hand; but at first she was
shy and nervous. She kept her sting in constant
readiness. . . . Gradually she became quite used to me,
and when I took her on my hand apparently expected to
be fed. She even allowed me to stroke her without any
appearance of fear, and for some months I never saw
her sting.
One other observation which goes to prove that other things besides
locality are noted and remembered by bees may here be quoted. Sir John
placed a bee in a bell jar, the closed end of which he held towards a
window. The bee buzzed about at that end trying to make for the open
air. He then showed her the way out of the open end of the jar, and
after having thus learnt it, she was able to find the way out herself.
This seems to show that the bee, like the wasp on the closed
window-pane, was able to appreciate and to remember the difference
between the quality of glass as resisting and air as permeable, although
to her sense of vision the difference must have been very slight. In
other words, the bee must have remembered that by first flying _away_
from the window, round the edge of the jar, and then _towards_ the
window, she could surmount the transparent obstacle; and this implies a
somewhat different act of memory from that of associating a particular
object--such as honey--with a particular locality. It is noteworthy that
a fly under similar circumstances did not require to be taught to find
its way out of the jar, but spontaneously found its own way out. This,
however, may be explained by the fact that flies do not always direct
their flight towards windows, and therefore the escape of this one was
probably not due to any act of intelligence.
While upon the subject of memory in the Hymenoptera, it is indispensable
that we should again refer to the observation of Messrs. Belt and Bates
already alluded to on pages 150-51. For it is from that observation
rendered evident that these sand-wasps took definite pains, as it were,
to _teach themselves_ the localities to which they desired to return.
Mr. Bates further observed that after thus taking a careful mental note
of the place, they would return to it without a moment's hesitation
after an absence of an hour. The observation of Mr. Belt, already quoted
_in extenso_, proves that these mental notes may be taken with the
utmost minuteness, so that even in the most intricate places the insect,
on its return, is perfectly confident that it has not made a mistake.
With regard to the duration of memory, Stickney relates a case in which
some bees took possession of a hollow place beneath a roof, and having
been then removed into a hive, continued for several years to return and
occupy the same hole with their successive swarms.[50]
Similarly Huber relates an observation of his own showing the duration
of memory in bees. One autumn he put some honey in a window, which the
bees visited in large numbers. During the winter the honey was taken
away and the shutters shut. When they were again opened in the spring
the bees returned, although there was no honey in the window.
These two cases amply prove that the memory of bees is comparable with
that of ants, which, as we have seen from analogous facts, also extends
at least over a period of many months.
_Emotions._
Sir John Lubbock's experiments on this head go to show that the social
sympathies of bees are even less developed than he found them to be in
certain species of ants. Thus he says:--
I have already mentioned with reference to the
attachment which bees have been said to show for one
another, that though I have repeatedly seen them lick
a bee which had smeared herself in honey, I never
observed them show the slightest attention to any of
their comrades who had been drowned in water. Far,
indeed, from having been able to discover any evidence
of affection among them, they appear to be thoroughly
callous and utterly indifferent to one another. As
already mentioned, it was necessary for me
occasionally to kill a bee; but I never found that the
others took the slightest notice. Thus on the 11th of
October I crushed a bee close to one which was
feeding--in fact, so close that their wings touched;
yet the survivor took no notice whatever of the death
of her sister, but went on feeding with every
appearance of composure and enjoyment, just as if
nothing had happened. When the pressure was removed,
she remained by the side of the corpse without the
slightest appearance of apprehension, sorrow, or
recognition. It was, of course, impossible for her to
understand my reason for killing her companion; yet
neither did she feel the slightest emotion at her
sister's death, nor did she show any alarm lest the
same fate should befall her also. In a second case
exactly the same occurred. Again, I have several
times, while a bee has been feeding, held a second bee
by the leg close to her; the prisoner, of course,
struggled to escape, and buzzed as loudly as she
could; yet the selfish eater took no notice whatever.
So far, therefore, from being at all affectionate, I
doubt whether bees are in the least fond of one
another.
Réaumur, however ('Insects,' vol. v., p. 265), narrates a case in which
a hive-bee was partly drowned and so rendered insensible; the others in
the hive carefully licked and otherwise tended her till she recovered.
This seems to show that bees, like ants, are more apt to have their
sympathies aroused by the sight of ailing or injured companions than by
that of healthy companions in distress; but Sir John Lubbock's
observations above quoted go to prove that even in this case display of
sympathy is certainly not the rule.
_Powers of Communication._
Huber says that when one wasp finds a store of honey 'it returns to its
nest, and brings off in a short time a hundred other wasps;' and this
statement is confirmed by Dujardin, who witnessed a somewhat similar
performance in the case of bees--the individual which first found a
concealed store informing other individuals of the fact, and so on till
numberless individuals had found it.
Although the systematic experiments of Sir John Lubbock have not tended
to confirm these observations with regard to bees and wasps, we must not
too readily allow his negative results to discredit these positive
observations--more especially as we have seen that his _later_
experiments have fully confirmed the opinion of these previous authors
with respect to ants. His experiments on bees and wasps consisted in
exposing honey in a hidden situation, marking a bee or wasp that came to
it, and observing whether it afterwards brought any companions to share
the booty. He found that although the same insect would return over and
over again, strangers came so rarely that their visits could only be
attributed to accidental and independent discovery. Only if the honey
were in an exposed situation, where the insects could _see_ one another
feeding, would one follow the other to the food.
But we have the more reason not to accept unreservedly the conclusion
to which these experiments in themselves might lead, because the very
able observer F. Müller states an observation of his own which must be
considered as alone sufficient to prove that bees are able to
communicate information to one another:--
Once (he says[51]) I assisted at a curious contest,
which took place between the queen and the other bees
in one of my hives, which throws some light on the
intellectual faculties of these animals. A set of
forty-seven cells have been filled, eight on a newly
completed comb, thirty-five on the following, and four
around the first cell of a new comb. When the queen
had laid eggs in all the cells of the two older combs
she went several times round their circumference (as
she always does, in order to ascertain whether she has
not forgotten any cell), and then prepared to retreat
into the lower part of the breeding-room. But as she
had overlooked the four cells of the new comb, the
workers ran impatiently from this part to the queen,
pushing her, in an odd manner, with their heads, as
they did also other workers they met with. In
consequence the queen began again to go around on the
two older combs; but as she did not find any cell
wanting an egg she tried to descend, but everywhere
she was pushed back by the workers. This contest
lasted for a rather long while, till the queen escaped
without having completed her work. Thus the workers
knew how to advise the queen that something was as yet
to be done, but they knew not how to show her _where_
it had to be done.
Again, Mr. Josiah Emery, writing to 'Nature,'[52] with reference to Sir
John Lubbock's experiments, says that the faculty of communication which
bees possess is so well and generally known to the 'bee-hunters' of
America, that the recognised method of finding a bees' nest is to act
upon the faculty in question:--
Going to a field or wood at a distance from tame bees,
with their box of honey they gather up from the
flowers and imprison one or more bees, and after they
have become sufficiently gorged, let them out to
return to their home with their easily gotten load.
Waiting patiently a longer or shorter time, according
to the distance of the bee-tree, the hunter scarcely
ever fails to see the bee or bees return accompanied
with other bees, which are in like manner imprisoned
till they in turn are filled, when one or more are
let out at places distant from each other, and the
direction in each case in which the bee flies noted,
and thus, by a kind of triangulation, the position of
the bee-tree proximately ascertained.
Those who have stored honey in their houses understand
very well how important it is to prevent a single bee
from discovering its location. Such discovery is sure
to be followed by a general onslaught from the hive
unless all means of access is prevented. It is
possible that our American are more intelligent than
European bees, but hardly probable; and I certainly
shall not ask an Englishman to admit it. Those in
America who are in the habit of playing first, second,
and third fiddle to instinct will probably attribute
this seeming intelligence to that principle.
According to De Fravière, bees have a number of different notes or tones
which they emit from the stigmata of the thorax and abdomen, and by
which they communicate information. He says:--
As soon as a bee arrives with important news, it is at
once surrounded, emits two or three shrill notes, and
taps a comrade with its long, flexible, and very
slender feelers, or antennæ. The friend passes on the
news in similar fashion, and the intelligence soon
traverses the whole hive. If it is of an agreeable
kind--if, for instance, it concerns the discovery of a
store of sugar or of honey, or of a flowering
meadow--all remains orderly. But, on the other hand,
great excitement arises if the news presages some
threatened danger, or if strange animals are
threatening invasion of the hive. It seems that such
intelligence is conveyed first to the queen, as the
most important person in the state.
This account, which is quoted from Büchner, no doubt bears indications
of imaginative colouring; but if the observation as to the emission of
sounds is correct--and, as we shall see, this point is well confirmed by
other observers--it is most likely concerned in communicating by tone a
general idea of good or harm: probably in the former case it acts as a
sign, 'follow me;' and in the latter as a signal of danger. Büchner
further says that, according to Landois, if a saucer of honey is placed
before a hive, a few bees come out, which emit a cry of tut, tut, tut.
This note is rather shrill, and resembles the cry of an attacked bee.
Hereupon a large number of bees come out of the hive to collect the
offered honey.
Again,--
The best way to observe the power of communication
possessed by bees by means of their interchange of
touches, is to take away the queen from a hive. In a
little time, about an hour afterwards, the sad event
will be noticed by a small part of the community, and
these will stop working and run hastily about over the
comb. But this only concerns part of the hive, and the
side of a single comb. The excited bees, however, soon
leave the little circle in which they at first
revolved, and when they meet their comrades they cross
their antennæ and lightly touch the others with them.
The bees which have received some impression from this
touch now become uneasy in their turn, and convey
their uneasiness and distress in the same way to the
other parts of the dwelling. The disorder increases
rapidly, spreads to the other side of the comb, and at
last to all the people. Then arises the general
confusion before described.
Huber tested this communication by the antennæ by a
striking experiment. He divided a hive into two quite
separate parts by a partition wall, whereupon great
excitement arose in the division in which there was no
queen, and this was only quieted when some workers
began to build royal cells.
He then divided a hive in similar fashion by a
trellis, through which the bees could pass their
feelers. In this case all remained quiet, and no
attempt was made to build royal cells: the queen could
also be clearly seen crossing her antennæ with the
workers on the other side of the trellis.
Apparently the feelers are also connected with the
exceedingly fine scent of the bees, which enables
them, wonderful as it may seem, to distinguish friend
and foe, and to recognise the members of their own
hive among the thousands and thousands of bees
swarming around, and to drive back from the entrance
stranger or robber bees. The bee-masters, therefore,
when they want two separate colonies or the members of
them to unite in one hive, sprinkle water over the
bees, or stupefy them with some fumigating substance,
so as to make them to a certain extent insensible to
smell, in order to attain their object. It is always
possible to unite colonies by making the bees smell of
some strong-smelling stuff, such as musk.[53]
Lastly, under the present heading I shall quote one other observation,
for which I am also indebted to Büchner's very admirable collection of
facts relating to the psychology of Hymenoptera:--
Herr L. Brofft relates, in 'der Zoologische Garten'
(XVIII. Year, No. 1, p. 67), that a poor and a rich
hive stood next each other on his father's bee stand,
and the latter suddenly lost its queen. Before the
owner had come to a decision thereupon the bees of the
two hives came to a mutual understanding as to the
condition of their two states. The dwellers in the
queenless hive, with their stores of provisions, went
over into the less populous or poorer hive, after they
had assured themselves, by many influential
deputations, as to the state of the interior of the
poor hive, and, as appeared, especially as to the
presence of an egg-laying queen!
_General Habits._
The active life of bees is divided between collecting food and rearing
young. We shall therefore consider these two functions separately.
The food collected consists of two kinds, honey (which, although stored
in the 'crop' for the purpose of carriage from the flowers to the cells,
appears to be but the condensed nectar of flowers) and so-called
'bee-bread.' This consists of the pollen of flowers, which is worked
into a kind of paste by the bees and stored in their cells till it is
required to serve as food for their larvæ. It is then partly digested by
the nurses with honey, so that a sort of chyle is formed. It is
observable that in each flight the 'carrier bees' collect only one kind
of pollen, so that it is possible for the 'house bees' (which, by the
way, are the younger bees left at home to discharge domestic duties with
only a small proportion of older ones, left probably to direct the more
inexperienced young) to sort it for storage in different cells. In the
result there are several different kinds of bee-bread, some being more
stimulating or nutritious than others. The most nutritious has the
effect, when given to any female larva, of developing that larva into a
queen or fertile female. This fact is well known to the bees, who only
feed a small number of larvæ in this manner, and the larvæ which they
select so to feed they place in larger or 'royal' cells, with an obvious
foreknowledge of the increased dimensions to which the animal will grow
under the influence of this food. Only one queen is required for a
single hive; but the bees always raise several, so that if any mishap
should occur to one, other larvæ may be ready to fall back upon.
Besides honey and bee-bread two other substances are found in beehives.
These are propolis and beeswax. The former is a kind of sticky resin
collected for the most part from coniferous trees. This is used as
mortar in building, &c. It adheres so strongly to the legs of the bee
which has gathered it, that it can only be detached by the help of
comrades. For this purpose the loaded bee presents her legs to her
fellow-workers, who clean it off with their jaws, and while it is still
ductile, apply it round the inside of the hive. According to Huber, who
made this observation, the propolis is applied also to the insides of
the cells. The workers first planed the surfaces with their mandibles,
and one of them then pulled out a thread of propolis from the heap
deposited by the carrier bees, severed it by a sudden throwing back of
the head, and returned with it to the cell which it had previously been
planing. It then laid the thread between the two walls which it had
planed; but, proving too long, a portion of the thread was bitten off.
The properly measured portion was then forced into the angle of the cell
by the fore-feet and mandibles. The thread, now converted into a narrow
ribbon, was next found to be too broad. It was therefore gnawed down to
the proper width. Other bees then completed the work which this one had
begun, till all the walls of the cells were framed with bands of
propolis. The object of the propolis here seems to be that of giving
strength to the cells.
The wax is a secretion which proceeds from between the segments of the
abdomen. Having ingested a large meal of honey, the bees hang in a thick
cluster from the top of their hive in order to secrete the wax. When it
begins to exude, the bees, assisted by their companions, rub it off into
heaps, and when a sufficient quantity of the material has been thus
collected, the work begins of building the cells. As the cells are used
both for storing food and rearing young, I shall consider them later
on. Now we have to pass to the labours incidental to propagation.
All the eggs are laid by one queen, who requires during this season a
large amount of nourishment, so much, indeed, that ten or twelve working
bees (_i.e._ sterile females) are set apart as her feeders. Leaving the
'royal cell,' she walks over the nursery-combs attended by a retinue of
workers, and drops a single egg into each open cell. It is a highly
remarkable fact that the queen is able to control the sex of the eggs
which she lays, and only deposits drone or male eggs in the drone cells,
and worker or female eggs in the worker cells--the cells prepared for
the reception of drone larvæ being larger than those required for the
worker larvæ. Young queens lay more worker eggs than old queens, and
when a queen, from increasing age or any other cause, lays too large a
proportion of drone eggs, she is expelled from the community or put to
death. It is remarkable, also, under these circumstances, that the queen
herself seems to know that she has become useless, for she loses her
propensity to attack other queens, and so does not run the risk of
making the hive virtually queenless. There is now no doubt at all that
the determining cause of an egg turning out male or female is that
which Dzierzon has shown, namely, the absence or presence of
fertilisation--unfertilised eggs always developing into males, and
fertilised ones into females. The manner, therefore, in which a queen
controls the sex of her eggs must depend on some power that she has of
controlling their fertilisation.
The eggs hatch out into larvæ, which require constant attention from the
workers, who feed them with the chyle or bee-bread already mentioned. In
three weeks from the time that the egg is deposited, the white worm-like
larva has passed through its last metamorphosis. When it has emancipated
itself its nurses assemble round it to wash and caress it, as well as to
supply it with food. They then clean out the cell which it has left.
When so large a number of the larvæ hatch out as to overcrowd the hive,
it is the function of the queen to lead forth a swarm. Meanwhile several
larval queens have been in course of development, and matters are so
arranged by the foresight of the bees, that one or more young queens are
ready to emerge at a time when otherwise the hive would be left
queenless. But the young queen or queens, although perfectly formed,
must not escape from their royal prison-houses until the swarm has
fairly taken place; the worker bees will even strengthen the coverings
of these prison-houses if, owing to bad weather or other causes,
swarming is delayed. The prisoner queens, which are fed through a small
hole in the roof of their cells, now continually give vent to a
plaintive cry, called by the bee-keepers 'piping,' and this is answered
by the mother queen. The tones of the piping vary. The reason why the
young queens are kept such close prisoners till after the departure of
the mother queen with her swarm, is simply that the mother queen would
destroy all the younger ones, could she get the chance, by stinging
them. The workers, therefore, never allow the old queen to approach the
prisons of the younger ones. They establish a guard all round these
prisons or royal cells, and beat off the old queen whenever she
endeavours to approach. But if the swarming season is over, or anything
should prevent a further swarm from being sent out, the worker bees
offer no further resistance to the jealousy of the mother queen, but
allow her in cold blood to sting to death all the young queens in their
nursery prisons. As soon as the old queen leaves with a swarm, the young
queens are liberated in succession, but at intervals of a few days; for
if they were all liberated at once they would fall upon and destroy one
another. Each young queen as it is liberated goes off with another
swarm, and those which remain unliberated are as carefully guarded from
the liberated sister queen as they were previously guarded from the
mother queen. When the season is too late for swarming the remaining
young queens are liberated simultaneously, and are then allowed to fight
to the death, the survivor being received as sovereign.
The bees, far from seeking to prevent these battles,
appear to excite the combatants against each other,
surrounding and bringing them back to the charge when
they are disposed to recede from each other; and when
either of the queens shows a disposition to approach
her antagonist, all the bees forming the cluster
instantly give way to allow her full liberty of
attack. The first use which the conquering queen makes
of her victory is to secure herself against fresh
dangers by destroying all her future rivals in the
royal cells; while the other bees, which are
spectators of the carnage, share in the spoil,
greedily devouring any food which may be found at the
bottom of the cells, and even sucking the fluid from
the abdomen of the pupæ before they toss out the
carcasses.[54]
Similarly, when a strange queen is put into a hive already provided with
a queen--
A circle of bees instinctively crowd around the
invader, not, however, to attack her--for a worker
never assaults a queen--but to respectfully prevent
her escape, in order that a combat may take place
between her and their reigning monarch. The lawful
possessor then advances towards the part of the comb
where the invader has established herself, the
attendant workers clear a space for the encounter,
and, without interfering, wait the result. A fearful
encounter then ensues, in which one is stung to death,
the survivor mounting the throne. Although the workers
of a _de facto_ monarch will not fight for her
defence, yet, if they perceive a strange queen
_attempting_ to enter the hive, they will surround
her, and hold her until she is starved to death; but
such is their respect for royalty that they never
attempt to sting her.[55]
All these facts display a wonderful amount of apparently sagacious
purpose on the part of the workers, although they may not seem to
reflect much credit on the intelligence of the queens. But in this
connection we must remember the observation of F. Huber, who saw two
queens, which were the only ones left in the hive, engaged in mortal
combat; and when an opportunity arose for each to sting the other
simultaneously, they simultaneously released each other's grasp, as if
in horror of a situation that might have ended in leaving the hive
queenless. This, then, is the calamity to avert which all the instincts
both of workers and queens are directed. And that these instincts are
controlled by intelligence is suggested, if not proved, by the
adaptations which they show to special circumstances. Thus, for
instance, F. Huber smoked a hive so that the queen and older bees
effected their escape, and took up their quarters a short distance away.
The bees which remained behind set about constructing three royal cells
for the purpose of rearing a new queen. Huber now carried back the old
queen and ensconced her in the hive. Immediately the bees set about
carrying away all the food from the royal cells, in order to prevent the
larvæ contained therein from developing into queens. Again, if a strange
queen is presented to a hive already provided with one, the workers do
not wait for their own queen to destroy the pretender, but themselves
sting or smother her to death. When, on the other hand, a queen is
presented to a hive which is without one, the bees adopt her, although
it is often necessary for the bee-master to protect her for a day or two
in a trellis cage, until her subjects have become acquainted with her.
When a hive is queenless, the bees stop all work, become restless, and
make a dull complaining noise. This, however, is only the case if there
is likewise a total absence of royal pupæ, and of ordinary pupæ under
three days of age--_i.e._ the age during which it is possible to rear an
ordinary larva into a queen.
As soon as the queen has been fertilised, and the services of the drones
therefore no longer required, the worker bees fall upon their
unfortunate and defenceless brothers to kill them, either by direct
stinging or by throwing them out of the hive to perish in the cold. The
drones' cells are then torn down, and any remaining drone eggs or pupæ
destroyed. Generally all the drones--which may number more than a
thousand--are slaughtered in the course of a single day. Evidently the
object of this massacre is that of getting rid of useless mouths; but
there is a more difficult question as to why these useless mouths ever
came into existence. It has been suggested that the enormous
disproportion between the present number of males and the single fertile
female refers to a time before the social instincts became so complex
or consolidated, and when, therefore, bees lived in lesser communities.
Probably this is the explanation, although I think we might still have
expected that before this period in their evolution had arrived bees
might have developed a compensating instinct, either not to allow the
queen to lay so many drone eggs, or else to massacre the drones while
still in the larval state. But here we must remember that among the
wasps the males do work (chiefly domestic work, for which they are fed
by their foraging sisters); so it is possible that in the hive-bee the
drones were originally useful members of the community, and that they
have lost their primitively useful instincts. But whatever the
explanation, it is very curious that here, among the animals which are
justly regarded as exhibiting the highest perfection of instinct, we
meet with perhaps the most flagrant instance in the animal kingdom of
instinct unperfected. It is the more remarkable that the drone-killing
instinct should not have been better developed in the direction of
killing the drones at the most profitable time--namely, in their larval
or oval state--from the fact that in many respects it seems to have been
advanced to a high degree of discriminative refinement. Thus, to quote
Büchner,--
That the massacre of the drones is not performed
entirely from an instinctive impulse, but in full
consciousness of the object to be gained, is proved by
the circumstance that it is carried out the more
completely and mercilessly the more fertile the queen
shows herself to be. But in cases where this fertility
is subject to serious doubt, or when the queen has
been fertilised too late or not at all, and therefore
only lays drones' eggs, or when the queen is barren,
and new queens, to be fertilised later, have to be
brought up from working-bee larvæ, then all or some of
the drones are left alive, in the clear prevision that
their services will be required later. . . . This wise
calculation of consequences is further exemplified in
that sometimes the massacre of the drones takes place
before the time for swarming, as, for instance, when
long-continued unfavourable weather succeeds a
favourable beginning of spring, and makes the bees
anxious for their own welfare. If, however, the
weather breaks, and work again becomes possible, so
that the bees take courage anew, they then bring up
new drones, and prepare them in time for the swarming.
This killing of drones is distinguished from the
regular drone massacre by the fact that the bees then
only kill the developed drones, and leave the drone
larvæ, save when absolute hunger compels their
destruction. Not less can it be regarded as a prudent
calculation of circumstances when the bees of a hive,
brought from our temperate climate to a more southern
country, where the time of collecting lasts longer, do
not kill the drones in August, as usual, but at a
later period, suitable to the new conditions.
But the philosophy of drone-killing is, I think, even more difficult in
the case of the wasps than in that of the bees. For, unlike the bees,
whose communities live from year to year, the wasps all perish at the
end of autumn, with the exception of a very few fertilised females. As
this season of universal calamity approaches, the workers destroy all
the larval grubs--a proceeding which, in the opinion of some writers,
strikingly exemplifies the beneficence of the Deity! Now, it does not
appear to me easy to understand how the presence of such an instinct in
this case is to be explained. For, on the one hand, the individual
females which are destined to live through the winter cannot be
conspicuously benefited by this slaughter of grubs; and, on the other
hand, the rest of the community is so soon about to perish, that one
fails to see of what advantage it can be to it to get rid of the grubs.
If the whole human race, with the exception of a few women, were to
perish periodically once in a thousand years, the race would profit
nothing by destroying, a few months before the end of each millennium,
all sick persons, lunatics, and other 'useless mouths.' I have not seen
this difficulty with regard to the massacring instinct in wasps
mentioned before, and I only mention it now in order to draw attention
to the fact that there seems to be a more puzzling problem presented
here than in the case of the analogous instinct as exhibited by bees.
The only solution which has suggested itself to my mind is the
possibility that in earlier times, or in other climates, wasps may have
resembled bees in living through the winter, and that the grub-slaying
instinct is in them a survival of one which was then, as in the case of
the bees now, a clearly beneficial instinct.
For some days before swarming begins, there is a great excitement and
buzzing in the hive, the temperature of which rises from 92° to 104°.
Scouts having been previously sent out to explore for suitable quarters
wherein to plant the new colony, these now act as guides. The swarm
leaves the hive with their queen. The bees which remain behind busy
themselves in rearing out the pupæ, which soon arriving at maturity,
also quit the hive in successive swarms. According to Büchner,
'secondary swarms with young queens send out no scouts, but fly at
random through the air. They clearly lack the experience and prudence of
the older bees.' And, regarding the behaviour of the scouts sent out by
primary swarms, this author says:--
M. de Fravière had the opportunity of observing the
manner in which such an examination is carried on, and
with what prudence and accuracy. He placed an empty
beehive, made in a new style, in front of his house,
so that he could exactly watch from his own window
what went on inside and out without disturbance to
himself or to the bees. A single bee came and examined
the building, flying all round it and touching it. It
then let itself down on the board, and walked
carefully and thoroughly over the interior, touching
it continually with its antennæ so as to subject it on
all sides to a thorough investigation. The result of
its examination must have been satisfactory, for after
it had gone away it returned accompanied by a crowd of
some fifty friends, which now together went through
the same process as their guide. This new trial must
also have had a good result, for soon a whole swarm
came, evidently from a distant spot, and took
possession. Still more remarkable is the behaviour of
the scouts when they take possession of a satisfactory
hive or box for an imminent or approaching swarm.
Although it is not yet inhabited they regard it as
their property, watch it and guard it against stranger
bees or other assailants, and busy themselves
earnestly in the most careful cleansing of it, so far
as this cleansing is impossible to the setter up of
the hive. Such a taking possession sometimes occurs
eight days before the entrance of the swarm.
_Wars._--As with ants, so with bees, the great cause of war is plunder;
and facts now well substantiated by numberless observers concerning
'robber-bees' indicate a large measure of intelligence. These aim at
lessening their labour in collecting honey by plundering the store of
other hives. The robberies may be conducted singly or in concert. When
the thieving propensity is developed only in individual cases, the
thieves cannot rely on force in plundering a foreign state, and so
resort to cautious stealth. 'They show by their whole behaviour--creeping
into the hive with careful vigilance--that they are perfectly conscious
of their bad conduct; whereas the workers belonging to the hive fly in
quickly and openly, and in full consciousness of their right.' If such
solitary burglars are successful in obtaining plunder, their bad example
leads other members of their own community to imitate them; thus it is
that the whole bee-nation may develop marauding habits, and when they do
this they act in concert to rob by force. In this case an army of bees
precipitates itself upon the foreign hive, a battle ensues, and if
successful in overcoming resistance, the invaders first of all search
out the queen-bee and put her to death, whereby they disorganise their
enemies and plunder the hive with ease. It is observed that when this
policy is once successful, the spirit of aggrandisement is encouraged,
so that the robber-bees 'find more pleasure in robbery than in their own
work, and become at last formidable robber-states.' When an invaded hive
is fairly overcome by the invaders killing the queen, the owners of the
hive, finding that all is lost, not only abandon further resistance, but
very often reverse their policy and join the ranks of their conquerors.
They assist in the tearing down of their cells, and in the conveyance of
the honey to the hive of their invaders. 'When the assailed hive is
emptied, the next ones are attacked, and if no effective resistance is
offered, are robbed in similar fashion, so that in this way a whole
bee-stand may be gradually destroyed.' Siebold observed the same facts
in the case of wasps (_Polistes gallica_). If, however, the battle turns
in favour of the defenders, they pursue the flying legions of their
enemies to a distance from their home. It sometimes happens that the
plundered hive offers no resistance at all, owing to the robbers having
visited the same flowers as the robbed, and so probably (having much the
same smell) not being recognised as belonging to a different community.
The thieves, when they find such to be the case, may become so bold as
to stop the bees that are returning to the hive with their loads, of
which they deprive them at the entrance of the hive. This is done by a
process which one observer, Weygandt,[56] calls 'milking,' and it seems
that the milking bee attains the double advantage of securing the honey
from the milked one and disarming suspicion of the other bees by
contracting its smell and entering the hive loaded, into which it is
admitted without opposition to continue its plunder.
Sometimes robber-bees attack their victims in the fields at a distance
from the hives. This sort of highway robbery is generally conducted by a
gang of four or five robber-bees which set upon a single honest bee,
'hold him by the legs, and pinch him until he unfolds his tongue, which
is sucked in succession by his assailants, who then suffer him to depart
in peace.'
It is strange that hive-bees of dishonest temperaments seem able to coax
or wheedle humble-bees into the voluntary yielding of honey.
'Humble-bees have been known to permit hive-bees to take the whole honey
that they have collected, and to go on gathering more, and handing it
over, for three weeks, although they refuse to part with it, or seek
refuge in flight, when wasps make similar overtures.'[57]
Besides theft and plunder, there are other causes of warfare among bees,
which, however, are only apparent in their effects. Thus, for some
undiscernible reason, duels are not infrequent, which generally end in
the death of one or both combatants. At other times, equally without
apparent reason, civil war breaks out in a hive, which is sometimes
attended with much slaughter.
_Architecture._--Coming now to the construction of the cells and combs,
there is no doubt that here we meet with the most astonishing products
of instinct that are presented in the animal kingdom. A great deal has
been written on the practical exhibition of high mathematical principles
which bees display in constructing their combs in the form that secures
the utmost capacity for storage of honey with the smallest expenditure
of building material. The shortest and clearest statement of the subject
that I have met with is the following, which has been given by Dr.
Reid:--
There are only three possible figures of the cells
which can make them all equal and similar, without any
useless interstices. These are the equilateral
triangle, the square, and the regular hexagon.
Mathematicians know that there is not a fourth way
possible in which a plane may be cut into little
spaces that shall be equal, similar, and regular,
without useless spaces. Of the three figures, the
hexagon is the most proper for convenience and
strength. Bees, as if they knew this, make their cells
regular hexagons.
Again, it has been demonstrated that, by making the
bottoms of the cells to consist of three planes
meeting in a point, there is a saving of material and
labour in no way inconsiderable. The bees, as if
acquainted with these principles of solid geometry,
follow them most accurately. It is a curious
mathematical problem, at what precise angle the three
planes which compose the bottom of a cell ought to
meet, in order to make the greatest possible saving,
or the least expense of material and labour. This is
one of the problems which belong to the higher parts
of mathematics. It has accordingly been resolved by
some mathematicians, particularly by the ingenious
Maclaurin, by a fluctionary calculation, which is to
be found in the Transactions of the Royal Society of
London. He has determined precisely the angle
required, and he found, by the most exact mensuration
the subject would admit, that it is the very angle in
which the three planes in the bottom of the cell of a
honeycomb do actually meet.[58]