ready to come forth, her instinctive jealousy would lead her to
attack them all as successively produced; and being so much older and
stronger, the probability is that she would destroy them; in which case
there could be no swarms, and the race would perish. But this is wisely
prevented by a circumstance which invariably takes place--that the
first swarm is conducted by this queen, and not by a newly disclosed
one, as Reaumur and others have supposed. Previously to her departure,
after her great laying of male eggs in the month of May, she oviposits
in the royal cells when about three or four lines in length, which the
workers have in the mean time constructed. These however are not all
furnished in one day,--a most essential provision, in consequence of
which the queens come forth successively, in order to lead successive
swarms. There is something singular in the manner in which the workers
treat the young queens that are to lead the swarms. After the cells are
covered in, one of their first employments is to remove here and there
a portion of the wax from their surface, so as to render it unequal;
and immediately before the last metamorphosis takes place, the walls
are so thin that all the motions of the inclosed pupa are perceptible
through them. On the seventh day the part covering the head and trunk
of the young female, if I may so speak, is almost entirely unwaxed.
This operation of the bees facilitates her exit, and probably renders
the evaporation of the superabundant fluids of the body of the pupa
more easy.
You will conclude, perhaps, when all things are thus prepared for the
coming forth of the inclosed female, that she will quit her cell at the
regular period, which is seven days:--but you would be mistaken. Were
she indeed permitted to pursue her own inclinations, this would be the
case: but here the bees show how much they are guided in their instinct
by circumstances and the wants of their society; for did the new queen
leave her cell, she would immediately attack and destroy those in the
other cells; a proceeding which they permit, as I have before stated,
when they only want a successor to a defunct or a lost sovereign. As
soon therefore as the workers perceive--which the transparency of
the cell permits them to do--that the young queen has cut circularly
through her cocoon, they immediately solder the cleft up with some
particles of wax, and so keep her a prisoner against her will. Upon
this, as if to complain of such treatment, she emits a distinct sound,
which excites no pity in the breasts of her subjects, who detain her a
prisoner two days longer than nature has assigned for her confinement.
In the interim, she sometimes thrusts her tongue through the cleft she
has made, drawing it in and out till she is noticed by the workers,
to make them understand that she is in want of food. Upon perceiving
this they give her honey, till her hunger being satisfied she draws her
tongue back--upon which they stop the orifice with wax[168].
You may think it perhaps extraordinary that the workers should thus
endeavour to retard the appearance of their young females beyond its
natural limit; but when I explain to you the reason for this seeming
incongruity of instinct, you will adore the wisdom that implanted it.
Were a queen permitted to leave her cell as soon as the natural term
for it arrived, it would require some time to fit her for flight,
and to lead forth a swarm; during which interval a troublesome task
would be imposed upon the workers, who must constantly detain her
a prisoner to prevent her from destroying her rivals, which would
require the labours and attention of a much larger number than are
necessary to keep her confined to her cell. On this account they
never suffer her to come forth till she is perfectly fit to take her
flight. When at length she is permitted to do this, if she approaches
the other royal cells, the workers on guard seem greatly irritated
against her, and pull and bite and chase her away; and she enjoys
tranquillity only while she keeps at a distance from them. As her
instinct is constantly urging her to attack them, this proceeding
is frequently repeated. Sometimes standing in a particular and
commanding attitude, she utters that authoritative sound which so
much affects the bees; they then all hang down their heads and remain
motionless; but as soon as it ceases, they resume their opposition.
At last she becomes violently agitated, and communicating her
agitation to others, the confusion more and more increases, till
a swarm leaves the hive, which she either precedes or follows. In
the same manner the other young queens are treated while there are
swarms to go forth; but when the hive is sufficiently thinned, and it
becomes troublesome to guard them in the manner here described, they
come forth unnoticed, and fight unimpeded till one alone remains to
fill the deserted throne of the parent hive.--You see here the reason
why the eggs that produce these queens are not laid at the same time,
but after some interval, that they may come forth successively. For
did they all make their appearance together, it would be a much more
laborious and difficult task to keep them from destroying each other.
When the bees thus delay the entrance of the young queens into their
world, they invariably let out the oldest first; and they probably
know their progress to maturity by the emission of the sound lately
mentioned. The accurate Huber took the trouble to mark all the royal
cells in a hive as soon as the workers had covered them in, and he
found that they were all liberated according to seniority. Those
first covered first emit the sound, and so on successively; whence
he conjectures that this is the sign by which the workers discover
their age. As their captivity, however, is sometimes prolonged to
eight or ten days, this circumstance in that time may be forgotten.
In this case he supposes that their tones grow stronger as they grow
older, by which the workers may be enabled to distinguish them. It
is remarkable that no guard is placed round the mute queens bred
according to the Lusatian method, which, when the time for their
appearance is come, are not detained in captivity a single moment;
but, as you have heard, are left to fight, conquer, or die[169].
You must not think, however, from what I have been saying, that the old
queen never destroys the young ones previously to her leading forth
the earliest swarm. She is allowed the most uncontrolled liberty of
action; and if she chooses to approach and destroy the royal cells, her
subjects do not oppose her. It sometimes happens, when unfavourable
weather retards the first swarm, that all the royal progeny perishes
by the sting of their mother, and then no swarm takes place. It is to
be observed that she never attacks a royal cell till its inhabitant is
ready to assume the pupa, therefore much will depend upon their age.
When they arrive at this state, her horror of these cells, and aversion
to them, are extreme: she attacks, perhaps, and destroys several; but
finding it too laborious, for they are often numerous, to destroy the
whole, the same agitation is caused in her as if she were forcibly
prevented, and she becomes disposed to depart, rather than remain in
the midst of her rivals, though her own offspring.
But though the bees, in one of these cases, appear such unconcerned
spectators of the destruction of royal personages, or rather, the
applauders and inciters of the bloody fact; and in the other show
little respect to them, put such a restraint upon their persons,
and manifest such disregard to their wishes; yet when they are once
acknowledged as governors of the hive, and leaders of the colony, their
instinct assumes a new and wonderful direction. From this moment they
become the "_publica cura_," the objects of constant and universal
attention; and wherever they go, are greeted by a homage which evinces
the entire devotion of their subjects. You seemed amused and interested
in no slight degree by what I related in a former letter of the marked
respect paid by the ants to their females[170]: but this will bear no
comparison with that shown by the inhabitants of the hive to their
queen. She appears to be the very soul of all their actions, and the
centre of their instincts. When they are deprived of her, or of the
means of replacing her, they lose all their activity, and pursue no
longer their daily labours. In vain the flowers tempt them with their
nectar and ambrosial dust: they collect neither; they elaborate no wax,
and build no cells; they scarcely seem to exist; and, indeed, would
soon perish, were not the means of restoring their monarch put within
their reach. But, if a small piece of comb containing the brood grubs
of workers be given to them, all seem endued with new life: their
instincts revive; they immediately set about building royal cells; they
feed with their appropriate food the grubs they have selected, and
every thing proceeds in the usual routine. Virgil has described this
attachment of the bees to their sovereign with great truth and spirit
in the following lines:
"Lydian nor Mede so much his king adores,
Nor those on Nilus' or Hydaspes' shores:
The state united stands while he remains,
But should he fall, what dire confusion reigns!
Their waxen combs and honey, late their joy,
With grief and rage distracted, they destroy:
He guards the works, with awe they him surround,
And crowd about him with triumphant sound;
Him frequent on their duteous shoulders bear,
Bleed, fall, and die for him in glorious war."
M. Huber thus describes the consequences of the loss of a
queen.--When the queen is removed from a hive, at first the bees
seem not to perceive it, their order and tranquillity not being
disturbed, and their labours proceeding as usual. About an hour after
her departure, inquietude begins to manifest itself amongst them;
the care of the young brood no longer engages their attention, and
they run here and there, as if in great agitation. This agitation,
however, is at first confined to a small portion of the community.
The bees that are first sensible of their loss meet with others, they
mutually cross their antennæ, and strike them lightly. By this action
they appear to communicate the sad intelligence to those who receive
the blow, who in their turn impart it in the same way to others.
Disorder and confusion increase rapidly, till the whole population is
in a tumult. Then the workers may be seen running over the combs,
and against each other; impetuously rushing to the entrance and
quitting the hive; from thence they spread themselves all around,
they re-enter, and go out again and again. The hum in the hive
becomes very loud, and increases the tumult, which lasts two or three
hours, rarely four or five: they then return and resume their wonted
care of the young; and if the hive be visited twenty-four hours after
the departure of the queen, it will be seen that they have taken
steps to repair their loss by filling some of the cells with a larger
quantity of jelly than is the usual portion of common larvæ; which
however is intended, it seems, not for the food of the inhabitant,
but for a cushion to elevate it, since it is found unconsumed in
the cell when the grub has descended into the pyramidal habitation
afterwards prepared for it[171].
If, after being removed, their old queen is restored to the hive, they
instantly recognise her, and pay her the usual attentions; but if a
strange one be introduced within the first twelve hours after the
old one is lost, she is kept a close prisoner till she perishes: if
twenty-four hours, as I have before hinted, have expired since they
lost their queen, and you introduce a new one, at the moment you set
this stranger upon a comb, the workers that are near her first touch
her with their antennæ, and then pass their proboscis over all parts
of her body: place is next given to others, who salute her in the
same manner:--all then beat their wings at the same time, and range
themselves in a circle round their new sovereign. A kind of agitation
is now communicated to the whole surface of the comb, which brings all
the bees upon it to see what is going forward. This may be called the
first shout of the applauding multitude to welcome the arrival of their
new sovereign. The circle of courtiers increases, they vibrate their
wings and bodies, but without tumult, as if their sensations were very
agreeable. When she begins to move, the circle opens to let her pass,
and all follow her steps. She is received with similar demonstrations
of loyalty in the other parts of the hive, is soon acknowledged queen
by all, and begins to lay eggs.--Reaumur put some bees into a hive
without their queen, and then introduced to them one that he had taken
when half perished with cold, and kept in a box, in which she had
covered herself with powder. The bees immediately owned her for their
queen, employed themselves very anxiously in cleaning her and warming
her, sometimes turning her upon her back for this purpose--and then
began to construct cells in their new habitation[172]. Even when the
bees have got young brood, have built or are building royal cells, and
are engaged in feeding these hopes of their hive, knowing that their
great aim is already accomplished, they cease all these employments
when this intruder comes amongst them.
With regard to the ordinary attention and homage that they pay
to their sovereigns--the bees do more than respect their queen,
says Reaumur, they are constantly on the watch to make themselves
useful to her, and to render her every kind office; they are for
ever offering her honey; they lick her with their proboscis, and
whereever she goes she has a court to attend upon her[173]. It may
here be observed, that the stimulant which excites the bees to
these acts of homage is the pregnant state of their queen, and her
fitness to maintain the population of the hive; all they do being
with a view to the public good: for while she remains a virgin she is
treated with the utmost indifference, which is exchanged, as soon as
impregnation has taken place, for the above marks of attachment[174].
The instinct of the bees, however, does not always enable them to
distinguish a partially fertile queen from one that is universally so.
What I mean is this--A queen, whose impregnation is retarded beyond the
twenty-eighth day of her whole existence, lays only male eggs, which
are of no use whatever to the community, unless they are at the same
time provided with a sufficient supply of workers. Yet even a queen
of this description, and sometimes one that is entirely sterile, is
treated by them with the same respect and homage as a fertile one.
This seems to evince an amiable feeling in these creatures, attachment
to the person as well as to the functions of the sovereign; which is
further manifested by their unwillingness at first to receive a new
sovereign upon the loss or death of their old one. Nay, this respect is
sometimes shown to the carcase of a defunct queen, which Huber assures
us he has seen bees treat with the same attention that they had shown
her when alive; for a long time preferring her inanimate corpse to
the fertile queens that he offered to them[175]. He attributes this
to some agreeable sensation which they experience from their queens,
independent of their fecundity. But since virgin queens, as we have
seen, do not excite it, more probably it is a remnant of their former
attachment, first excited by her fecundity, and afterwards strengthened
and continued by habit.
I may here introduce an interesting anecdote related by Reaumur, which
strongly marks the attachment of bees to their queen when apparently
lifeless. He took one out of the water quite motionless, and seemingly
dead, which had lost part of one of its legs. Bringing it home, he
placed it amongst some workers that he had found in the same situation,
most of which he had revived by means of warmth; some however still
being in as bad a state as the poor queen. No sooner did these revived
workers perceive the latter in this wretched condition, than they
appeared to compassionate her case, and did not cease to lick her with
their tongues till she showed signs of returning animation; which the
bees no sooner perceived, than they set up a general hum, as if for joy
at the happy event. All this time they paid no attention to the workers
who were in the same miserable state[176].
On a former occasion I have mentioned the laying of the eggs by the
queen[177]; but as I did not then at all enlarge upon it, I shall now
explain the process more in detail. In a subsequent letter I shall
notice, what has so much puzzled learned apiarists--her fecundation:
which is now ascertained beyond contradiction, from the observations
of M. Huber, to take place in the open air, and to be followed by the
death of the unfortunate male[178]. It is to be recollected that,
from September to April, generally speaking, there are no males in
the hives; yet during this period the queen often oviposits: a former
fecundation, therefore, must fertilize all the eggs laid in this
interval. The impregnation, in order to ensure complete fertility,
must not be too long retarded: for, as I before observed, if this be
delayed beyond the twenty-eighth day of her existence, her ovaries
become so vitiated, that she can no longer lay eggs that will produce
workers, but can only furnish the hive with a male population; which,
however high a privilege it may be accounted amongst men, is the
reverse of it amongst the bees. When this is the case, the abdomen of
the queen becomes so enlarged that she is no longer able to fly[179];
and, what is remarkable, she loses that instinctive animosity which
stimulates the fertile ones to attack their rivals[180]. Thus she
seems to own that she is not equal to the duties of her station, and
can tolerate another to discharge them in her room. When we consider
how much virgin queens are slighted by their subjects, we may suppose
that nature urges them to take the opportunity of the first warm day,
when the males fly forth, to pair with one of them.
When fecundation has not been retarded, forty-six hours after it has
taken place, the queen begins to lay eggs that will produce workers,
and continues for the subsequent eleven months, more or less, to lay
them solely; and it is only after this period that an uninterrupted
laying of male eggs commences.--But when it has been retarded, after
the same number of hours she begins laying male eggs, and continues to
produce these alone during her whole life. From hence it should seem
to follow, that the former kind of eggs are first in the oviducts,
and, if impregnation be not effected within a given time, that all
the worker embryos perish. Yet how this can take place with respect
to those that in a fertile queen should succeed the laying of male
eggs, or be produced in the second year of her life, seems difficult
to conceive;--or how the male embryos escape this fate, which destroys
all the female, both those that are to precede them and those that are
to follow them. Is it impossible that the sex of the embryo may be
determined by the period at which the _aura seminalis_ vivifies it, and
by the state of the ovary at that time? In one state of the ovary this
principle may cause the embryos to become workers, in another males.
And something of this kind perhaps may be the cause of hermaphrodites
in other animals. But this I give merely as conjecture[181]: the truth
seems enveloped in mystery that we cannot yet penetrate. Huber is of
opinion that a single impregnation fertilizes all the eggs that a queen
will produce during her whole life, which is sometimes more than two
years[182]. But of this enough.
I said that forty-six hours after impregnation the queen begins
laying worker eggs;--this is not, however, invariable. When her
impregnation takes place late in the year, she does not begin laying
till the following spring. Schirach asserts, that in one season a
single female will lay from 70,000 to 100,000 eggs[183]. Reaumur
says, that upon an average she lays about two hundred in a day, a
moderate swarm consisting of 12,000, which are laid in two months;
and Huber, that she lays above a hundred. All these statements, the
observations being made in different climates, and perhaps under
different circumstances, may be true. The laying of worker eggs
begins in February, sometimes so early as January[184]. After this,
in the spring, the great laying of male eggs commences, lasting
thirty days; in which time about 2000 of these eggs are laid. Another
laying of them, but less considerable, takes place in autumn. In
the season of oviposition, the queen may be discerned traversing
the combs in all directions with a slow step, and seeking for cells
proper to receive her eggs. As she walks she keeps her head inclined,
and seems to examine, one by one, all the cells she meets with. When
she finds one to her purpose, she immediately gives to her abdomen
the curve necessary to enable it to reach the orifice of the cell,
and to introduce it within it. The eggs are set in the angle of the
pyramidal bottom of the cell, or in one of the hollows formed by the
conflux of the sides of the rhombs, and, being besmeared with a kind
of gluten, stand upright. If, however, it be a female that lays only
male eggs, they are deposited upon the lowest of the sides of the
cell, as she is unable to reach the bottom[185].
While our prolific lady is engaged in this employment, her court
consists of from four to twelve attendants, which are disposed
nearly in a circle, with their heads turned towards her. After
laying from two to six eggs, she remains still, reposing for eight
or nine minutes. During this interval the bees in her train redouble
their attentions, licking her fondly with their tongues. Generally
speaking, she lays only one egg in a cell; but when she is pressed,
and there are not cells enough, from two to four have been found in
one. In this case, as if they were aware of the consequences, the
provident workers remove all but one. From an experiment of Huber's,
it appears that the instinct of the queen invariably directs her to
deposit worker eggs in worker cells; for when he confined one, during
her course of laying worker eggs, where she could only come at male
cells, she refused to oviposit in them; and trying in vain to make
her escape, they at length dropped from her; upon which the workers
devoured them. Retarded queens, however, lose this instinct, and
often, though they lay only male eggs, oviposit in worker cells and
even in royal ones. In this latter case the workers themselves act as
if they suffered in their instinct from the imperfect state of their
queen; for they feed these male larvæ with royal jelly, and treat
them as they would a real queen. Though male eggs deposited in worker
cells produce small males, their education in a royal cell with
"royal dainties" adds nothing to their ordinary dimensions[186].
The _swarming_ of bees is a very curious and interesting subject, to
which, since a female is the _sine quâ non_ on this occasion, I may
very properly call your attention here. You will recollect that I
said something upon the principle of emigrations, when I was amusing
you with the history of ants[187]; but the object with them seems to
be merely a change of station for one more convenient or less exposed
to injury, and not to diminish a superabundant population. Whereas
in the societies of the hive-bee, the latter is the general cause of
emigrations, which invariably take place every year, if their numbers
require it; if not, when the male eggs are laid, no royal cells are
constructed, and no swarm is led forth. What might be the case with
ants, were they confined to hives, we cannot say. Formicaries in
general are capable of indefinite enlargement, therefore want of
room does not cause emigration;--but bees being confined to a given
space, which they possess not the means of enlarging,--to avoid
the ill effects resulting from being too much crowded, when their
population exceeds a certain limit, they must necessarily emigrate.
Sometimes--for instance, when wasps have got into a hive--the bees
will leave it, in order to fly from an inconvenience or enemy which
they cannot otherwise avoid; but it does not very often happen that
they wholly desert a hive.