Chapter IV: Bees and Wasps (1)

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.