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NEW OBSERVATIONS

ON

BEES.

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NEW OBSERVATIONS ,

ON THE

NATURAL HISTORY

BEES.

BY

FRANCIS FIUBER.

THIRD EDITION.

ILLUSTRATED BY FIVE ELATES.

EDINBURGH:

PRINTED FOR W. & C. TAIT, AND
LONGMAN, HURST, REES, OR31B, AND RROW'N,
LONDON.

1821.

Edinburgh :
Printed hj J, cj C. Muirhead.

THE EIGHT HONOUEABLE

SiE JOSEPH BANKS, Baet.

KNIGHT OF THE MOST HONOURABLE ORDER OF
THE BATH,

A PRIVY COUNCILLOR,

PRESIDENT OF THE UOYAL SOCIETY OF LONDON,

THIS translation:

IS INSCRIBED.

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TRANSLATOR’S PREFACE.

The facts contained in this volume are
deeply interesting to the naturalist. They
not only elucidate the history of those in-
dustrious creatures whose properties are the
peculiar subject of investigation, but they
present some new and singular features in
animal physiology, hitherto unknown, and
even unsuspected.

In all countries, and in every age, the
labour of bees has proved a fertile source of
admiration, and mankind have endeavoured
unremittingly to convert it to gratification
or emolument. What can be more wonder-
ful, indeed, than to witness an insect of such
apparent insignificance rendering each dif-
ferent flower tributary to itself or the neces-
sities of its young, and fabricating structures
which no human art can approach or imitate!

PREFACE.

Vlll'

But it is from the impulse of its propensi-
ties, and from the united efforts of myriads,
that we are enabled to gain those valuable
products, which otherwise would be utter-
ly unattainable..

Curiosity and avidity being equally a-
wahened by the industry of these diminutive
beings, innumerable theories, observations,
and experiments have followed regarding
them, and uncommon patience has attended
the inquiry. Yet the real discoveries, com-
pared with the multitude of investigators,
have been surprisingly few. Vague specu-
lations have been substituted for rational
researches into the nature of bees, and super-
ficial inspection deemed' satisfactory analysis
of their works. Hence the properties ac-
tually ascertained are so interwoven with
errors, that no subject has been the parent
of greater absurdities. Unfortunately, also,
some of those treatises, the most accessible,
and, at the present day, the most popular,
only contribute- to- their wider dispersion,
and enhance the difficulties of the philoso-
phic naturalist attempting their eradication.
A considerable portion of the subsequent
work is devoted to this purpose. It belongs
to the reader to judge of the deductions

PREFACE.

IX

whereby the author concludes that he has
established facts from experiment.

Perhaps no treatise of equal compass, per-
haps no treatise whatever contains as^many
novelties in the history of bees. Their na-
ture, organization, senses, instinct, and mode
of perpetuation, are all illustrated. The
origin of wax, the faculty of obtaining it
from honey or sugar, its application to use
in the structure of cells, and the formation
of combs, are perspicuously discussed, while
several points are established which had been
previously the alternate theme of conjecture
and controversy. But the general approba-
tion given here, as well as on the Continent,
to a modest and unobtrusive work, wherein
both instruction and amusement are com-
bined, constitutes the best testimony of its
merits. Thus, to use the words of Sue, a
foreign author, “ the observations are so
consistent, and the deductions so conclusive,
that, when occupied with this treatise, we
feel as if we had assisted the author in each
experiment, and pursued it with correspon-
ding zeal and interest. Let us invite the
admirers of nature to its perusal : few are
of such quality, or so faithfully describe the
properties and habits of bees.”

X

PKEFACE.

It is a circumstance too remarkable to be
overlooked, that the author’s defective sight
induced him to employ an assistant in con-
ducting his experiments. Some of these
have been since coufirined by various observ-
ers, and among others by his son Peter
Huber, whose elegant taste for natural histo-
ry is testified in his Researches concerning
the Habits of Ants. The experiments them-
selves seem to have been so judiciously
adapted to the object in view, and the con-
clusions so strictly logical, as evidently to
preclude important errors. It is not uncom-
mon, indeed, to hear authority challenged.
Even persons the least qualified venture to
oppose their opinions to the truly learned,
with a specious air of confidence, very apt
to delude the unthinking part of mankind.
But those whose minds are the most copi-
ously stored, are not invariably the most
urgent to disclose the full scope of their in-
formation. Therefore, although certain facts
be not specified, it is far from a necessary
consequence that they have been unseen :
Nor although we fail to procure the same
result under what we deem the same circum-
stances, are we> hastily to denounce the ac-
curacy of our preciusors.

PREFACE.

XI

After the talents of the author’s philoso-
phic assistant, Francis Burnens, had been
assiduously devoted to his service for a num-
ber of successive years, he was called on to
reside in a different quarter, where he now
discharges some respectable public functions;
and it appears that of late he had not for-
saken their common pursuits.

A long interval elapsed between the pub-
lication of the different memoirs, forming
the first and second part of this volume.
Here the whole are presented somewhat in
an abbreviated form, particularly the portion
relative to anatomical details, the theory of
respiration, and the architecture of bees. All
the substance, however, is preserved: and
amidst the perplexity and intricacy in which
it is partly involved, the narrative is render-
ed a§ explicit and concise as the subject ad-
mits. Nevertheless, a few passages have
proved of doubtful interpretation, and a few
unintelligible. Naturalists are frequently
betrayed into an unprofitable prolixity, les-
sening the interest of their works, and even
clouding them with obscurity.

The cultivation of bees forms a branch of
rural economy, which may be carried to very
great extent. Its utility is obvious ; and

XU

PREFACE.

where so much benefit might accrue, especi-
ally to the more dependent classes of the
community, it is to be regretted that it
does not receive greater attention. Proba-
bly the cultivator will always find his cares
sufficiently rewarded by success.

CONTENTS.

PART I.

Page.

Letter I. — On the impregnation of the queen

bee^ 1

II. — Sequel of observations on the im-
pregnation of the queen bee, 33

III. — The same subject continued; obser-

vations on retarding the fecun-
dation of queens, 36

IV. — On Schirach’s discovery,.. 63

V,— Experiments proving that there are
sometimes common bees which
lay fertile eggs,..,.,,. 74

VI. — On the combats of queens ; the mas-
sacre of the males ; and what suc-
ceeds in a hive where a stran-
ger queen is substituted for the
natural one,.,, go

lx

XIV

CONTENTS.

Page.

Letter VII.— -Sequel of experiments on the re-
ception of a stranger queen ; M.
de Reaumur’s observations on the
subject, 114.

VIII. — Is the queen oviparous What
influence has the size of the cells,
where the eggs are deposited, on
the bees produced ? Researches
on the mode of spinning the coc-

coons, 121

IX.— On the formation of swarms, 143

X. — The same subject continued, 16S

XI. — The same subject continued, 186

XII. — Additional observations on queens
laying only the eggs of drones,
and on those deprived of the an-
tennae, 197

XIII.— Economical considerations on bees, 211
PART II.

Chapter I. — Schirach’s discovery confirmed....... 229

II. — On the senses of bees, and particu-
larly that of smelling, 253

III. — Researches on the use of the an-

tennae in some complicated opera-
tions of bees, 272

IV. — On the respiration of bees, 287

V. — On a new enemy of bees, 305

CONTENTS.

XV

Page.

Chapter VI. — New opinions on wax, 320

VII. — On the origin of wax, , 329

VIII, On the architecture of bees, 349

IX. — Modifications of the architecture

of bees, 377

X. — Completion of the cells, 401

Appendix 417

Explanation of the Plates 424

Analytical Index.. 429

Page 68, line 1, page 61, line 4. Perhaps the passages
should describe the bees as brooding on the cells alluded to.

Page 226. The distance specified for the excursions of
bees seems to be corroborated by a communication to IM,
Lombard from the author, in April 1810.

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II

PART I.

LETTER I.

ON THE I31PREGNATION OF THE QUEEN BEE.

Sir, as you desirctl me to transmit a writ-
ten detail of my principal experiments on
bees, when I gave you an account of them
at Genthod, I hasten to extract the follow-
ing observations from my journal. No-
thing can be more flattering than the in-
terest you take in my researches : therefore,
permit me to remind you of your promise to
suggest new subjects for investigation.* '

All the letters In the first portion of the work are adilress-
C(1 to the celebrated naturalist M. Bonnet, wliosc labours
in this department of science :u'e well known and justly ap-
preciated. Ilis decease in lT9d, during the prosecution of
the author’s experiments, explains the reason for dividing
the volume into two parts. — 'I'.

B

IMPREGNATION OE

Glass hives, constructed after !M. dc
Reaumur’s principles, are of a form unfa-
vourable to the observer; because their width
allowing the bees to build two combs par-
allel, whatever passes between them is con-
cealed from his view. Long experience of
this has induced you to recommend hives
much flatter or thinner ; the panes of which
should be separated by so small an interval,
that only a single row of combs could be
erected between them. From having felt
the same inconvenience, I have profited by
your counsel in providing hives reduced to
an inch and a half in width, wherein swarms
have been established without any difiicul-
ty. Here, however, the charge of construct-
ing a single comb must not be committed
to the bees : they are taught by nature to
make more than one, and all parallel to each
other — a law from which they never dero-
gate, unless when constrained by some par-
ticular arrangement. Therefore, if left to
themselves in these flat hives, which cannot
admit of two combs parallel to the jflane
of the sides, they will form several small
ones perpendicular to it ; and in that case
all will be equally lost to the observer.
Thus previous dispositions become essential

THE QUEEN BEE.

for the direction of the comhs. I so con-
trived that, while tliey WTre built perpendi-
cular to the horizon, the lateral surfaces
should be three or four lines from the ])anes
constituting the sides of the hive.* This
distance in allowing sufficient latitude for
the motions of the bees, prevented them
from collecting in too large clusters on the
surface of the comb. By such precautions,
they were easily established in very thin
hives, where they pursued their labours with
the same assiduity and regularity; and every
cell being exposed to view, none of their
proceedings could be concealed.

It is true, that by compelling these in-
sects to live in a habitation w’here they could
construct only a single row of combs, I had,
in a certain measure, changed their natural
condition, and this circumstance possibly
might have affected their instinct.f There-
fore, to obviate every objection, I invented
a kind of hives, which, without losing the
auvantages of those very thin, at the same

* The different measurements are expressed in lines, of
which le are in an inch. — T.

t A single comb of very large dimensions, attached un-
der the bouglis of a tree, is said to be consUuctcd bv a
species of bees in India.— T.

4

impregnation of

time approached the figure of common hives
where bees form several rows of combs.

I procured several small fir frames, a foot
square and an inch and a quarter broad;
and connecting them together by hinges,
the whole, like so many divisions, could be
opened and shut as the leaves of a book :
Plate 1, fig. 1, 2. When using a hive of
this description, we previously fixed a comb
ill each frame, and then introduced all the
bees which were required for the particular
experiment. Opening the difiTerent divisions
ill succession, we daily inspected both sur-
faces of every comb : there was not a single
cell where we could not see distinctly what-
ever passed at all times, nor a single bee, I
may almost say, with which we were not
particularly acquainted. Indeed this ap-
paratus is nothing more than the union of
several very flat hives capable of separation.
But bees must not be visited, in such a ha-
bitation, before their combs are fixed se-
curely in the frames, otherwise they may
kill or maim them by falling out, or excite
that degree of irritation as will expose the
observer to being stung, which is always
painful, and sometimes dangerous : but they
soon become accustomed to their situation,

THE QUEEN EEE.

o

and are in some respect tamed by it. In
three days we may begin to operate on the
hive, to open it, remove part of the combs,
and substitute others, without the bees ex-
hibiting too formidable symptoms of dis-
pleasure. You will remember. Sir, that on
visiting my retreat, I showed you a hive of
this kind that had been a long time in
experiment, and how much you were sur-
prised that the bees so quietly allowed us to
open it.

In these hives, I have repeated all my ob-
servations, and obtained exactly the same
results as in the thinnest. Thus, I think,
already to have obviated any objections that
may arise concerning the supposed incon-
venience of flat hives. Besides, I cannot
regret the repetition of my labours; by
going over the same course several times, i
am much more certain of having avoided er-«
ror ; and it also appears, that some advan-
tages are found in these which may be called
Tiooh, or Lecif-ltivcs, as they prove extremely
useful in the economical treatment of bees,
which shall be afterwards detailed.

Having now come to the particular object
of this letter, the fecundation of tlie queen
bee, I shall, in a few words, examine the

6

IMPREGNATION OF

different opinions of naturalists; next, I shall
state the most remarkable observations
which their conjectures have induced me to
make, and then describe the new experi-
ments by which I think I have solved the
problem.*

Swammerdam, who studied bees with un-
remitting attention, and who could never see
the real union of a drone and a queen, was
satisfied of its being unnecessary for fecun-
dation of the eggs; but having remarked
that, at certain times, the drones exhaled a
very strong odour, he thought it an emana-
tion which operated fecundation by penetrat-
ing the body of the female. His conjec-
ture was confirmed on dissecting the male
sexual organs ; for he was so much struck
with the disproportion between them and
those of the female, that he did not believe
their union possible. Besides, his opinion
concerning the influence of the odour was
plausible, from affording a good reason for

* I ciinnot insist that my readers, tlie better to compre-
hend what is here said, shall peruse the Memoirs of M. de
Reauirmr on Bees, and those of the Lusatian Society ; hnt
1 must request them to examine M. Bonnet’s works, tom. 5,
4to edit, and tom. 10, 8vo. where they will find a short
and distinct abstract of all that naturedLsts have hitherto
discovered on the subject.

THE QUEEN BEE.

7

the prodigious number of males. There are
frequently fxfteen hundred or two thousand
in a hive ; and, according to Swammerdam,
it is necessary they should be numerous, that
the emanation proceediug from them may
have an intensity or energy sufficient to
effect impregnation.

Though ]\I. de lleaiimur has refuted this
hypothesis by just and conclusive reasoning,
he has failed to make the sole experiment
that could support or overturn it : which was
confining all the drones of a hive in a tin
case, perforated with minute apertures, that
might allow the emanation of the odour to
escape, but prevent transmission of their or-
gans. The case should have been then pla-
ced in a hive well peopled, but completely de-
prived of males, both of large and small size,
and the consequences observed. It is evi-
dent, had the queen laid eggs after matters
were thus disposed, that Swammerdam’s
hypothesis would have acquired probability;
and, on the contrary, it would have been con-
futed had slie produced no eggs, or only
sterile ones. However, the experiment has
been made by us, and the female remained
barren; therefore, it is undoubted, that the

8

IMPREGNATION OF

emanation of the odour of the males does
not impregnate queens.

hi. de Reaumur was of a different opi-
nion. He thought that the queen’s fecunda-
tion followed actual union. Having confin-
ed several drones in a glass vessel along with
a virgin queen he saw the females make
many advances to the males ; but, unable to
observe any thing so intimate that it could
be denominated their union, he leaves the
question undecided. W e have repeated this
experiment also ; we have frequently confin-
ed virgin queens with drones of all ages : we
have done so at every season, and witnessed
all their advances and solicitations towards
the males : we have even believed that we
saw a kind of union between them, but so
short and imperfect that it was unlikely to
effect impregnation. Yet, to neglect no-
thing, we confined to her hive a virgin
queen, that had suffered the ap]>roaches of
the male. During a month that her im-
prisonment continued, she did not lay a sin-
gle egg ; therefore, these momentary junc-
tions do not accomplish fecundation.

In the Contemplation de In Nature, you
have cited the observations of the English
naturalist IMr. Debraw^ whidi, from their

THE QUEEN BEE.

9

apparent accuracy, seemed at last to eluci-
date the mystery. Favoured by chance,
that observer one day perceived at the bot-
tom of cells containing eggs, a matter, ap-
parently prolific, at least very diflPerent from
the substance or jelly which bees commonly
collect around their new hatched worms.
Solicitous to learn its origin, and conjectur-
ing that it might be masculine matter, he be-
gan to watch the motions of every drone in the
hive, on purpose to seize the moment when it
should be received by the eggs. He assures
us, that he saw several drones insinuate the
posterior part of the body into the cells for
that purpose. After frequent repetition of
the first, he entered on a long series of other
experiments. He confined a number of
workers in glass bells along with a queen
and several males. They were supplied with
pieces of comb containing honey, but want-
ing brood. He saw the queen lay eggs,
which were bedewed by the males, and from
which larvae were hatched consequently,

Ihe larva is the 'worm or caterpillar wliieh comes
from the egg of an insect : it then changes into another
state caUed the pupa, chrysalis, or nymph, and, lastly, in-
to the perfect animal. Eater naturalists incline to substi-
tute this name in all cases where the worm is not .seen un-
der its final aspect. — 2\

10

niPREGNATION OF

he could not hesitate in advancing as a fact
demonstrated, that male bees fecundate the
queen’s eggs in the manner of frogs and
tislies, that is, after they are produced.*

There was something very specious in this
explanation : the experiments on which it
was founded seemed correct ; and it afforded
a satisfactory reason for the prodigious num-
ber of males in a hive. At the same time,
the author had neglected to obviate one
strong objection. Larvae appear when there
are no drones. From the month of Septem-
ber until April, hives are generally destitute
of males, yet, notwithstanding their absence,
the queen then lays fertile eggs. Thus, the
prolific matter cannot be required for their
impregnation, unless we shall suppose that
it is necessary at a certain time of the year,
while at every otlier season it is useless.

To discover the truth, amidst these facts
apparently so contradictory, I determined to
repeat IMr. Debraw’s experiments, and to
observe more precaution than he himself had
done. First, I sought for that matter, which
he supposes the prolific, in cells containing
eggs. Several were actually found with such

* Philosophical Transactions, Vol. IxviL

THE QUEEN BEE.

11

an appearance, and, during the first days of
observation, neither iny assistant nor myself
doubted the reality of the discovery. But
^ve afterwards found it an illusion arising
from the reflection of the light, for nothing
like a fluid was visible, except when the so-
lar rays reached the bottom of the cells. This
part is commonly covered by shining frag-
ments of the cocoons of worms successively
hatched, and the reflection of the light from
these, when much illuminated, produces an
illusory effect. W e proved it by the strict-
est examination, for no vestiges of a fluid
w’ere perceptible when the cells were detached
and cut asunder.

Though the first observation inspired us
with some distrust of ]\Ir. Debraw’s disco-
very, we repeated his other experiments with
the utmost care. On the 6th of August
17B7j we immersed a hive,’^ and, with scru-
pulous attention, examined all the bees while
in the bath. We ascertained that there was
no male, either large or small ; and having

Bees may be long immersed in water without destroy-
ing life. Reaumur found them recover after nine hours
immersion ; other animals of analogous sjiecies exhibit still
more wonderful resurrections. — T.

12

IMPREGNATION OF

examined every comb, we found neither male
nymph, nor worm. When the bees were dry,
we replaced the whole, along with the queen,
in their habitation, and transported them
into my cabinet. They were allowed full
liberty ; therefore, they flew about, and made
their usual collections ; but, it being neces-
sary that no male should enter the hive dur-
ing the experiment, a glass tube was adapt-
ed to the entrance, of such dimensions that
two bees only could pass at once; and we
watched the tube attentively during the four
or five days that the experiment continued.
We should have instantly observed and re-
moved any male appearing, that the result
of the experiment might be uudistm-bed,
and I can positively affirm that not one was
seen. However, from the first day, which
was the 6tli of August, the queen deposited
fourteen eggs in the workers’ cells ; and all
these were hatched on the tenth of the same
month.

This experiment is decisive ; since the
eggs laid by the queen of a hive where there
were no males, and rvhere it was impossible
one could be introduced, since these eggs, I
say, were fertile, it becomes indubitable that

THE QUEEN BEE.

13

aspevsiou with the masculine matter is not
required to effect their exclusion.

Though it did not appear that any rea-
sonable objection could be started against
such an inference, yet, as I had been accus-
tomed in all my experiments to investigate
the most trifling difficulties which could oc-
cur, I conceived that Mr. Debraw’s partisans
might maintain, that the bees, deprived of
drones, perhaps would search for those in
other hives, and carry the fecundative mat-
ter to their own habitations for the purpose
of depositing it on the eggs.

It was easy to appreciate the force of this
objection ; for the only thing necessary was
repetition of the former experiments, and
confinement of the bees so closely to their
hives that none could possibly escape. You
know very well. Sir, that these animals can
live three or four months confined in a hive
w'ell stored with honey and wax, if apertures
are left for circulation of the air. Tliis experi-
ment was made on the tenth of August ; and
I ascertained, by means of immersion, that
no male was present. The bees were confined
four days in the closest manner, and then I
found forty young larvae, recently hatched.

14

IMPREGNATION OF

I extended my precautions so far as to
immerse the same hive a second time, to be
assured that no male had escaped my re-
searches. Each of the bees was separately
examined, and none was there that did not
display its sting. The coincidence of this
experiment with the other, proved that the
eggs were not externally fecundated.

In terminating the confutation of Mr.
Debraw’s opinion, I have only to explain
what led him into error. He employed
queens in his experiments, with whose his-
tory he was not acquainted from their ori-
gin. When he observed that the eggs pro-
duced by a queen confined along with males
were fertile, he thence determined that they
had been bedewed by the prolific matter in
the cells. But, to have rendered his. con-
clusion just, he should have first ascertained
that the female was in a virgin state, and
this he neglected. The truth is, that, with-
out knowing it, he had used a queen after
her commerce with the male. Had he taken
a virgin queen, the moment she came from
the royal cell, and confined her in his vessels
along with drones, the result would have
been opposite; for, even amidst a seraglio of

THE QUEEN BEE.

15

males, this young queen never would have
laid, as I shall afterwards prove.

The Lusatian observers, and Hattorf in
particular, thought the queen was fecundated
of herself, without concourse with the males.
I shall here give an abstract of the experi-
ment on which this opinion is founded.^

Hattorf took a queen whose virginity he
could not doubt. He excluded all the males
of the large and likewise of the small spe-
cies, and, in several days, found both eggs
and worms. He asserts that there were no
drones in the hive during the course of the
experiment ; but, although they were ab-
sent, the queen laid eggs, from which worms
proceeded : whence he considers that she is
impregnated by herself.

Reflecting on this experiment, I did not
find it sufficiently accurate. Males pass with
great facility from hive to hive ; and Hattorf
took no precaution against any being intro-
duced into his. He says, indeed, there was
no male, but is silent respecting the means
adopted to prove the fact : And althougli
he might be satisfied of no large drone

Vide in Scliirach’s History of Bees, a memoir by Hat-
torf, entitled, Physical Researches whether the Qtiee?i lies
f<-’fj>iires fecundation by Drones ?

16

IMPREGNATION OF

being present, still a small one might have
escaped his vigilance, and fecundated the
queen. With a view to clear up the doubt,
I resolved to repeat his experiment, in the
manner described by him, and without great-
er care or precaution.

I put a virgin queen into a hive, from
which all the males were excluded, but the
bees left at perfect liberty. Several days
afterwards I visited the hive, and found new
hatched worms in it. Here then was the
same result as Hattorf obtained ! But be-
fore deducing the same consequence, we had
to ascertain beyond dispute that no male had
entered the hive. Thus, it was necessary
to immerge the bees, and examine each se-
parately : By which operation we actually
found four small males. Therefore, to render
the experiment decisive, not only w^as it re-
quisite to remove all the drones, but also, by
some infallible method, to prevent any from
being introduced, which the German natur-.
alist had neglected.

I prepared to repair this omission, by
putting a virgin queen into a hive, from
which the whole males were carefully re-
^noved; and to be physically certain that
none could obtain access, a glass tube was

THE QUEEN BEE.

17

adapted at the entrance of such dimensions,
that the working bees could freely pass and
repass, but too narrow for the smallest male,
blatters continued thus for thirty days ; the
workers departing and returning, pertormed
their usual labours ; but the queen remain-
ed sterile. At the expiration of that time,
her belly was equally slender as at the mo-
ment of her origin. I repeated the experi-
ment several times, and always with the same
result.

Therefore, as a queen, rigorously separated
from all commerce with the male, remains
sterile, it is evident she cannot impregnate
herself, and that Hattorf’s opinion is ill-
founded.

Hitherto, by endeavouring to confute or
verify the conjectures of all the authors who
had preceded me, by new experiments, I had
acquired the knowledge of new facts, but
these were apparently so contradictory as to
render the solution of the problem still more
difficult. bVhile examining Mr. Hebraw’s
hypothesis, I confined a queen in a hive,
from which all the drones were removed ;
yet she was fertile. AA^hcn considering the
opinion of Hattorf, on the contrary, I put
one of whose virginity I was perfectly satiS'*

18

IMPREG NATION OE

fiecl ill the same situation, she remained
sterile.

Embarrassed by so many difficulties, I was
on the point of abandoning the subject of
iny researches, when at length, on more at-
tentive reflection, I thought these contradic-
tions might arise from experiments made
indifferently on virgin queens, with whose
history I was not acquainted from the origin,
and which perhaps had been fecundated un-
knowm to me. Impressed with this idea, I
undertook a new^ method of observation, not
on queens fortuitously taken from the hive,
but oil females decidedly in a virgin state,
and whose history was known to me from the
instant of their leaving the cell.

From a very great number of hives, I re-
moved all the reigning females, and substi-
tuted for each a queen taken at the moment
of her birth. The hives were then divided
into two classes. All the males, both large
and small, were taken from the first, and I
adapted a glass tube at the entrance, so nar-
row that no drone could pass, but large
enough for the free passage of the common
bees. In the hives of the second class, I
left the wliole drones belonging to them, and
%ven introduced more ; and to prevent them

THE QUEEN BEE.

19

from escaping, a glass tube, also too narrow
for the males, was adapted to their entrance.

For more than a month, I carefully watch-
ed this experiment, made on a large scale ;
but, much to my surprise, every queen re-
mained sterile. Thus it was proved that
queens confined in a hive would continue
barren though amidst a seraglio of males.

This result induced me to suspect that
the females could not be fecundated in the
interior of the hive, and that it was neces-
sary for them to leave it for receiving the
approaches of the male. To ascertain the
fact was easy, by a direct experiment ; and
as the point is important, I shall relate in
detail what was done by my secretary and
myself on the 29th of June 1788.

Aware that in summer the males usually
leave the hive in the warmest part of the day,
it was natural for me to conclude that if the
queens were obliged to go out for fecunda-
I tion, instinct would induce them to do so at
the same time as the others.

A t eleven in the forenoon, we placed our-
selves opposite to a hive containing an unim-
prcgnatcd queen five days old. The sun had
shone from his rising; the air was very warm ;
and the males began to leave the hives.

20

impregnation or

then enlarged the entrance of that selected
for observation, and paid great attention to
the bees entering and departing. The males
appeared, and immediately took flight. Soon
afterwards, the young queen came to the en-
trance ; at first she did not fly, but during
a little time traversed the board, brushing
her belly with her hind legs ; neither work-
ers nor males bestowing any notice on her.
At last .she took flight. When several feet
from the hive, she returned and approached
it as if to examine the place of her depar-
ture, perhaps judging this precaution neces-
sary to recognise it; she then flew away, de-
scribing horizontal circles twelve or fifteen
feet above the earth. We contracted the en-
trance of the hive that she might not return
unobserved, and placed ourselves in the centre
of the circles described in her flight, the more
easily to follow her and witness all her mo-
tions, But she did not remain long in a situa-
tion favourable for our observations, and ra-
pidly rose out of sight. We resumed our
place before the liivc ; and in seven minutes
the young queen returned to the entrance of
a habitation whicli she had left for the first
time. Having found no external evidence
of fecundation, v,e allowed her to enter.

THE QUEEN BEE.

X

In a quarter of an hour she re-appeared ;
and, after brushing herself as before, took
flight. Then returning to examine the hive,
she rose so high that we soon lost sight of
her. This second absence was much longer
than the first ; it occupied twenty-seven mi-
nutes. We now found her in a state very
different from that in which she was after her
former excursion : the organs distended by a
substance, thick and hard, very much re-
sembling the matter in the vessels of the
male ; completely similar to it indeed in
colour and consistence.*

But more evidence than mere resemblance
being requisite to establish that the female
had returned with the prolific matter of the
males, we allowed this queen to enter the

* It will afterwards appear, that what we took for the
generative matter, was the male organs left in the body of
the female ; a discovery which we owe to a circumstance

I that shall be immediately related. Perhaps I should avoid
prolixity, by suppressing all m*y first observations on the
impregnation of the queen, and passing directly to the ex-
periments that prove she carries away the genital organs ;
but in such observations, which are both new and delicate,
and where it is so easy to be deceived, I consider that a
candid avowal of my errors is doing the reader service.
This is an additional proof to so many others, of the abso-
lute necessity that an observer should repeat all his experi-
ments a thousand times to obtain the certainty of seeing
facts as they really exist.

IMPREGNATION OF

hive, and confined her there. In two days,
we found her belly swoln ; and she had al-
ready laid nearly an hundred eggs in the
workers’ cells.

To confirm our discovery, we made several
other experiments, and with the same suc-
cess. I shall continue to transcribe my jour-
nal.

On the second of July, the weather being
very fine, numbers of males left the hives ;
and we set at liberty a young virgin queen,
eleven days old, whose hive had been always
deprived of them. Having quickly left the
hive, she returned to examine it, and then
rose out of sight. She came back in a few
minutes without any external marks of im-
pregnation, and departed again in a quarter
of an hour, with so rapid a flight that we
could scarcely follow her a moment. This
absence continued thirty minutes ; but on
her return, the last ring of the body was
open, and the organs full of the whitish sub-
stance already mentioned. Slie was then re-
placed in the hive from which all the males
were excluded. In two days we found her
impregnated.

These observations at length demonstrate
why Hattorf obtained results so different

THE QUEEN BEE.

23

from ours. His queens, though in hives de-
prived of males, had been fecundated ; and
he thence concludes that sexual intercourse
is not requisite for their impregnation. But
not having confined the queens to their
hives, they had profited by their liberty to
unite with the males. We, on the contrary,
have surrounded our queens with a number
of males, yet they continued sterile ; because
the precautions for confining the males to
their hives, had also prevented the queens
from departing to seek that fecundation
without, which they could not obtain within.

The same experiments were repeated on
queens twenty, twenty-five, and thirty days
old. All became fertile after a single im-
pregnation. However, w'e have remarked
some essential peculiarities in the fecundity
of those remainiirg in the virgin state until
the twentieth day of their existence; but w^e
shall defer speaking of the fact, until being
able to present naturalists with observations
sufficiently correct and numerous to merit
their attention : Yet let me add a few words
: to what I have already said. Though neither
my assistant nor myself have witnessed the
commerce of a queen and a drone, we think
that, after the detail which has been just

24

IMPIIEGNATION OF

commenced, no doubt of the fact can remain,
nor can its necessity to effect impregnation
be disputed. The sequel of our experb
inents, made with every possible precaution,
appears demonstrative. The uniform sterility
of queens in hives wanting males, and in
those where they are confined along with
them ; the departure of these queens from
the hives; and the very conspicuous evi-
dence of impregnation with which they re-
turn, are proofs against which no objections
can stand. But we do not despair of being
able next spring to obtain the complement
of this proof, by seizing the female at the
very moment of her union.

Naturalists always have been extremely
embarrassed to account for the number of
males in most hives, and which seem only a
burden on the community, since they fulfil
no function. But we now begin to discern
the object of nature in multiplying them to
such an extent. As fecundation cannot be
accomplished within, and as the queen is
obliged to traverse the expanse of the at-
mosphere, it is requisite the males should
be numerous, that she may have the chance
of meeting some one of them. Were only
two or three in each hive, there would be i

THE QUEEN BEE.

25

little probability of their departure at tlie
same instant with the queen, or that they
^vould meet in their excursions; and most
of the females thus would remain sterile.*^

But why has nature prohibited sexual
union within the hives ? This is a secret still
unknown to us. It is possible, however, that
some favoumhle circumstance may enable
us to penetrate it in the course of our obser-
vations. V arious conjectures may be formed ;
but at this day we require facts,, and reject
gratuitous suppositions. It should be remem-
bered, that bees do not form the sole re-
public among insects presenting a similar
phenomenon ; female ants are also obliged to
leave the ant-hills previous to fecundation.f

* Remarkable in-egularity subsists in the number of males,
compared with the other inhabitants of a hive. Swammer-
dam found 693 along with 84'94 workers. Previous to the
swarming of a large hive, Reaumur counted 700 among
26,426 common bees, and one queen. In another, contain-
ing only 2900 w'orkers, he found 693 males. He computed
50,000 cells in the former, of which 20,000 were full of
brood. About 2520 cells w'ere appropriated for breeding
males, and above lialf of them were occupied by larvic and
nymphs. Thus, including the 700 in the perfect state, he
observes, that this hive tvould be provided with above 2000
males. Tom. v. p. 561. — T.

■|' The males and females of ants are winged insects ; the
former perish some time after their amours, and the females
lose their tvings a certain period after impregnation.— 7'.

C

26

IMPREGNATION OE

I cannot, request, Sir, tliat you will com-
municate those reflections whicli your genius
will excite concerning the facts I have relat-
ed- This is a favour to which I am not yet
entitled^ But as new experiments unques-
tionably will occur to you, w^hether on the
inpregnation of the queen or on other points,
may I solicit you to suggest them ? They
shall be executed with all possible care ; and
I shall esteem this mark of friendship and
interest as the most flattering encourage-
ment that the continuance of my labours
can receive. — Pregny, IMh August 1789.

I

letter from M. bonnet to M. HUBER,

You have surprised me most agreeably*
Sir, with your interesting discovery of the
impregnation of the queen bee. It was a for-
tunate conjecture, that she left the hive to be
fecundated ; and your method of ascertain-
ing the fact was extremely judicious, and
well adapted to the object.

Let me remind you, that male and fe-
male ants unite in the air; and that after I
impregnation the females return to the ant- j

THE QUEEN BEE. 27

Mils to deposit tlieir eggs. Contemplation
rlc la Natuir, Fart. II, chap. 22, note 1.
It would be necessary to seize the instant
when the drone unites with the female. But
how remote from the power of the observer
are the means of ascertaining their com-
merce in the air ! If you have satisfactory
evidence that the matter seen on the last
rings of the female is truly inascuHne, it is
more than mere presumption in favour of
the truth. Perhaps it may be necessary
that the male should seize the female under
the belly, which cannot be easily done but
in the air. The large opening at the ex-
tremity, which you have observed in so par-
ticular a condition, seems to correspond with
the singular size of the organs of the male.

You wish that I should suggest some
new experiments on these industrious repub-
licans. In doing so, I shall take the greater
pleasure and interest, as I know to what ex-
tent you possess the valuable art of com-
bining ideas, and of deducing from tins com-
bination results adapted to the discovery of

new facts. A few at this moment occur to
me.

It may be proper to attempt the artificial
fecundation of a virgin (piceii with a pciici].

28

IMPREGNATION OP

at the same time observing every precau-
tion to avoid error. This experiment, you
are aware, has already succeeded with more
than one animal.

To ascertain that the queen, which has
left the hive for impregnation, is the same
that returns to deposit her eggs, you will
find it necessary to colour the thorax with
some varnish resisting humidity. It will
be proper also to paint the thorax of a con-
siderable number of workers, in order to dis-
cover the duration of their life, which is a
more secure method than slight mutilations.

That the worm may be hatched, the egg
must be fixed almost vertically by one end
near the bottom of the cell. Is it true that
it is unproductive, unless when in this po-
sition? Unable to determine the fact, I
leave it to the decision of experiment.

I formerly mentioned to you that I had
long doubted the real nature of the small
ovular substances deposited by the queens
in the cells, and my inclination to suppose
them minute worms not yet begun to ex-
pand. Their elongated figure seems to fii-
vour my conjecture. It would be expedient,
therefore, to watch them with the utmost
assiduity, from the instant of production

THE QUEEN EEE.

29

until the period of exclusion. If the in-
tegument bursts, there can be no doubt that
these minute substances are real eggs.

I return to the mode of the union taking
place. The height that the queen and the
males rise to in the air prevents us from
seeing what passes between them : on which
account, the hive should be put into an
apartment with a very lofty ceiling. M. de
Reaumur’s experiment confining a queen
with several males in a glass vessel, merits
lepetition ; and if, instead of a vessel, a
glass tube some inches in diameter and seve-
ral feet long, were used, perhaps something
satisfactory might be discovered.

\ on have Iqul the fortune to observe the
small queens mentioned by the Abbe Need-
ham, but which he never saw. It will be of
gieat importance to dissect them, for the
purpose offinding their ovaries. When Mr
Ricm inforraedme that he had confined three
hundred workers, along with a comb contain-
ing no eggs, and afterwards found hundreds
m It, I strongly recommended that he should
dissect the workers. He did so; and in-
formed me that eggs were discovered in
lec. robably without being aware of it,
e las dissected small queens. As small
Clones exist, it is not surprising if small

30

IMPREGNATION OF

queens are produced also, and undoubtedly
by the same external causes.

It is of much consequence to be intimate-
ly acquainted with this species of queens, for
they may have great influence on different
experiments, and may embarrass the observ-
er: we should ascertain whether they in-
habit pyramidal cells smaller than the cam-
mon or hexagonal ones,

M. Schirach’s famous experiment on the
supposed conversion of a common worm into
a royal one cannot 1x3 too often repeated,
though the Lusatian observers already have
done so frequently. I am anxious to learn
whether, as the discoverer maintains, the ex-
periment will succeed only with worms three
or four days old, and never with simjile eggs.

The Lusatian observers, and the naturalist
of the Palatinate, affirm, that when common
bees arc confined with combs absolutely void of
eggs, they then lay none but those of drones.
Thus, there may be small queens producing
the eggs of males only, for it is evident they
must have produced those supposed to come
from workers. But how is it possible to
conceive that their o\ aries contain male eggs
alone ?

According to ]\J . de Ileaumur, the life of
chrysalids may be prolonged by keeping

THE QUEEN BEE.

31

them in a low temperature, such as that of
an ice-house. The same experiment should
be made on the eggs of a queen ; and on the
nymphs of drones and workers.

Another interesting experiment would be
removing all the combs composed by the
common cells, and leaving none but those
destined for the larvae of males. By this
means we should learn whether the eggs of
common worms, laid by the queen in the
large cells, will produce large workers. It
is very probable, however, that deprivation
of the common cells might discourage the
bees, because they require them for their
honey and wax. A^everthelcss, it is likely,
that by taking away only part of them, the
workers may be forced to lay common eggs
in the cells of drones.

I should also wish to have the young lar-
va gently removed from the royal cell, and
deposited at the bottom of a common one,
along with some of the royal food.

As the figure of hives has much influ-
ence on the respective disposition of the
combs, it would be a satisfactory experiment
greatly to diversify their shape and internal
dimensions. Nothing could be better adapt-
ed to instiuct us how bees can regulate and
apply their labours to existing circiunstan-

IMPREGNATION OF

ces. This may enable us to discover parti-
cular facts which we cannot foresee.

The royal eggs, and those producing
drones, have not yet been carefully compar-
ed with the eggs from which workers pro-
ceed. But it ought to be done, that we
may ascertain whether these different eggs
have secret distinctive characteristics.

The food supplied by the workers to the
royal worm, is not the same with that given
to the common worm. Could we not en-
deavour, with the point of a pencil, to re-
move a little of the royal food, and give it
to a common worm deposited in a cell of the
largest dimensions ? I have seen common
cells hanging almost vertically, where the
queen had laid ; and these I should prefer
for such an experiment.

Various facts, demanding corroboration,
were collected in my INIemoirs on Bees;
of which number are my own observations.
You can select what is proper. You have
already enriched the history of bees so much,
that every thing may be expected from your
understanding and perseverance. You
know the sentiments with which you have
inspired the Contemplator of Nature.
Gentkod, I8//4 August 1789-

THE QUEEN BEE.

33

LETTER II.

SEQUEL OF OBSERVATIONS ON THE IM-
PREGNATION OF THE QUEEN BEE.

\ All the experiments, related in niy pre-

i ceding letter, were made in the years 1787
and 1788. They seem to establish two
facts, which had been previously the subject
of vague conjecture: 1. That the queen bee
is not impregnated of herself, hut is feeuu-
I dated by union with the male. 2. That
I this is accomplished without the hive, and
|i in the air.

* The latter appeared so extraordinary, that
(j notwithstanding all the evidence obtained,
I we eagerly desired to take the queen in the
I fact; but, as she constantly rose to a great
ft height, we never could see what passed. On

34 IMPREGNATION OF

that account you advised us to cut some
part off the wings of virgin queens. We
endeavoured, in every possible manner, to
benefit by your advice, but, to our great
regret, when the wings lost much, the bees
could no longer fly ; and, by cutting oflP only
an inconsiderable portion, w'e did not di-
minish the rapidity of their flight. Proba-
bly there is a medium, though w^e w^ere un-
able to attain it. On your suggestion, also,
we tried to render their vision less acute, by
covering the eyes with an opaque varnish,
an experiment equally fruitless.

We likewise attempted artificial fecun-
dation, and took every possible precaution
to insure success ; yet the result w'as alw^ays
unsatisfactory. Several queens became the
victims of our curiosity ; and those surviving
remained sterile. Notwithstanding these
different experiments w'ere unsuccessful, it
was proved that queens leave their hives to
seek the males, and that they return with
undoubted evidence of fecundation. Satis-
fied with this, we could only trust to time
or accident for decisive proof of an actual
union. We were far from suspecting a
most singular discovery, made by us in July
this year, which affords complete demon-

THE QUEEN BEE.

35

stration of the supposed event, namely, that
the sexual organs of the male remain with
the female.'*

I conjecture that the males perish after
losing their organs ; but why so great a sacri-
fice is exacted by nature involves a mystery
which I cannot pretend to unveil. I am un-
acquainted with any analogous fact in na-
tural history ; hut as there are two kinds of
insects, namely, ephemeras and ants, whose
union seems to take place only in the air,
it would be extremely interesting to ascer-
tain, whether their males also lose the same
parts in similar circumstances, and whether,
as with drones, enjoyment in their flight is
the prelude of death .f

* The remainder of this chapter chiefly consists of ana-
tomical details, which may be rather considered an in-
terruption of the narrative. The Translator has judged
it expedient to transfer an abstract of them to an Ap-
pendix,

t 29th May 1S13. — I have not witnessed the sexual
commerce ofephemerce ; but was it attended witli the muti-
lation of tlie males, this would not have escaped M. de
Geer, who has done so. In regard to ants, their males ai-e
so little liable to such a penalty, that, as I have assured my-
self by repeated observations, the successive fecundation
of several females can follow.

36

EFFECTS OF

LETTER III.

^-vvxvw%-vxwv%

THE SAME SUBJECT CONTINUED. — OBSER-
VATIONS ON RETARDING THE FECUN- ■
DATION OF QUEENS.

In my first letter, I remarked, that when I
queens were prevented from receiving the 1
approaches of the male until the twenty-fifth
or thirtieth day of their existence, the result J
presented very interesting peculiarities. My ^
experiments at that time were not sufficient- I
ly numerous; but they have been since so j
often repeated, and the result so uniform, '
that I no longer hesitate to announce, as a
certain discovery, the singularities which re-
tarded fecundation produces on the ovaries
of the queen. If she receives the male dur-
ing the first fifteen days of her life, she re-

Iio

EETAEDED IMPREGNATION. 3T

mains capable of laying the eggs both of
workers and of drones ; but should impreg-
nation be retarded until the twenty-second
day, her ovaries are vitiated in such a man-
ner that she becomes unfit for laying the
eggs of workers, and will produce only those
of drones.

In June 1787, being occupied in re-
searches relative to the formation of swarms,
I had occasion, for the first time, to observe
a queen that laid none but the eggs of males.
AVhen a hive is ready to swarm, I had pre-
viously remarked, that the moment of
’ departure is always preceded by a very lively
f agitation, which first affects the queen, is
; then communicated to the workers, and ex-
f cites such a tumult among them, that they
i abandon their labours, and rush in disorder
) to the outlets. I then knew very well the
B cause of the queen’s agitation, and it is de-
D scribed in the history of swarms, but I was
ignorant how the delirium was communi-
B cated to the workers ; and this difficulty in-

0 terruptcd my researches. Therefore I thought

1 of investigating, by direct experiments, whe-
el ther at all times, when the queen was great-
y ly agitated, even not in the time of the hive
V swarming, her agitation would be communi-

38

EFFECTS OF

cated in like manner to the workers. The
moment one was hatched, I confined her to
the hive by contracting the entrances. When
assailed by the imperious desire of union
with the males, I could not doubt that she
would make great exertions to escape, and
that the impossibility of accomplishing it
would produce a kind of delirium. We had
the patience to observe this queen thirtv-
five days. Every morning about eleven
o’clock, when the weather was fine, and the
sunshine invited the males to leave their
hives, we saw her impetuously traverse every
corner of her habitation, seeking to escape.
Her fruitless efforts threw her into an extra-
ordinary agitation, the symptoms of which I
shall describe elsewhere, and all the com-
mon bees were affected by it. As she ne-
ver was out during this time, she could not
be impregnated. At length, on the thirty-
sixth day, 1 set her at liberty. She soon
took advantage of it; and was not long of
returning with the most evident marks of
fecundation.

Satisfied with the particular object of the
experiment, I was far from any hopes that
it would lead to the knowledge of another
very remarkable fact. How great was my as-

retaeded impregnation. 39

toiiishment, therefore, on finding that this
female, which, as usual, began to lay forty-
six hours after being fructified, produced the
eggs of drones, hut none of workers, and that
she continued ever after to lay thos€ of drones
only.

At first, I exhausted myself with conjec-
tures regarding so singular a fact ; the more
I reflected on it, the more did it seem inex-
plicable. At length, by attentively meditat-
ing on the circumstances of the experiment,
it appeared that there were two principles,
the influence of which I should first of all
endeavour to appreciate separately. On the
I one hand, the queen had suffered long con-
i finement ; on the other, her fecundation had
I been extremely retarded. You know. Sir,
! that the commerce of these females general-
' ly takes place about the fifth or sixth day ;
; but here it had been postponed until the thir-
; ty-sixth. That the peculiarity now demon-
i strated had resulted from confinement, was
' exceedingly improbable ; for queens, in the
I natural state, leave their hives only once to
i< seek the males ; all the rest of their life
I they remain voluntary prisoners. Thus it
« was unlikely that captivity would produce
i: the consequences which I wished to cluci-

40

EFFECTS OF

date. At the same time, as it was essential
to neglect nothing in a subject so new, I was
of desirous ascertaining whether it was ow-
ing to the length of confinement, or to re-
tarded fecundation..

Investigating these points was no easy
matter. To discover whether captivity, and
not retarded fecundation would vitiate the
ovaries, I found it necessary to allow a fe-
male to receive the approaches of a male,
and also to keep her imprisoned. Now this
could not be, for the union of bees never
takes place in hives. For the same reason,
it was impossible to retard this event, with-
out keeping the queen in confinement. I
was long embarrassed by tlie difficulty. At
length, I contrived an apparatus, which,
though imperfect, nearly fulfilled my pur-
pose.

I put a queen, at the moment of her last
metamorphosis, into a hive well stored, and
sufficiently provided with workers and males;
the entrance was contracted so as to prevent
her exit, but allowing free p^^ssage to the
workers. I also made another opening for
the queen, and adapted a glass tube to it,
communicating with a cubical glass box eight
feet high. Hither she could at all times

RETARDED IMPREGNATION. 41

1-epair aud fly about, enjoying a purer air than
was found within the hive : but she could
not be fecundated; for though the males
flew about within the same bounds, the space
was too limited to admit of any union be-
i tween them. By the experiments related
I in my first letter, their connection takes
I place high in the air only : therefore, in this
apparatus, I found the advantage of retard-
ing fecundation, while the liberty which the
queen now had, did not render her situation
too remote from the natural state. I watch-
ed the experiment fifteen days. Every fine
morning, the young captive left her hive ;

I she traversed her glass prison, and flew much
about, and with great facility. She laid
) none during this interval, for she had not
[ united with a male. On the sixteenth day
I set her at liberty : she left the hive, rose
1 aloft in the air, and soon returned with full
I evidence of impregnation. In two days, she
I laid the eggs of workers first, and afterwards
B as many as the most fertile queens.

It thence followed, 1. That captivity had
i not altered the organs of the queen : 2. Fe-
5 cundation having taken place within the first
i sixteen days, she produced both species of
! <?ggs.

42

EFFECTS OF

This was an important experiment. It
rendered my labours much more simple, by
clearly pointing out the method to be pur-
sued : it absolutely precluded the supposed
influence of captivity; and left nothing for
investigation, but the consequences of retard-
ed fecundation.

With that view, I repeated the experi-
ment ; but, instead of giving the virgin
queen liberty on the sixteenth day, I detain-
ed her until the twenty-first. She departed,
rose high in the air, was fecundated, and re-
turned. Thirty-six hours afterwards, she be-
gan to lay: but it was the eggs of males only,
and, although very fruitful afterwards, she
laid no other kind.

During the remainder of 1787, and the
two subsequent years, I occupied myself with
experiments on retarded fecundation, and
had constantly the same results. It is un-
doubted, therefore, that when the sexual
union of a queen is retarded beyond the twen-
tieth day, only an imperfect impregnation
is operated : instead of laying the eggs of
workers and males equally, she will lay none
hut those of males.

I do not aspire to the honour of explain-
ing this remarkable circumstance. AVhen

EETARDED IMPREGNATION.

43

the course of my experiments led me to ob-
serve that some queens laid only the eggs
of drones, it was natural to investigate the
proximate cause of such a singularity ; and
I ascertained that it arose from retarded fe-
cundation. My evidence is demonstrative,
for I can always prevent queens from lay-
ing the eggs of workers, by retarding their fe-
cundation until the twenty-second or twenty-
third day of their existence. But what is
the remote cause of such a peculiarity ? or,
in other words, why does the delay of im-
pregnation render queens incapable of lay-
ing the eggs of workers ? This is a problem
on which analogy throws no light : nor in
all physiology am I acquainted with any
fact that bears the smallest similarity.

The problem becomes still more difficult
by reflecting on the natural state of things,
that is, when fecundation has not been post-
poned. The queen then lays the eggs of
workers forty-six hours after her union, and
continues for the subsequent eleven months to
produce these alone ; and it is only after this
period that a considerable and uninterrupted
laying of the eggs of drones commences.^

* It appears that this period is liable to modification,
and that the great epoch of' laying the eggs of males may
be accelerated or retarded by the state of the atmospherg^
promoting or impeding tlie collection, of the bees,

44

EFFECTS OF

When, on the contrary, impregnation is re-
tarded after the twentieth day, the queen be-
gins, from the forty-sixth hour, to lay the eggs
of males, and produces no other kind during
her whole life. As, in the natural state, she
lays the eggs of workers only, during tlie
first eleven months, it is clear that these, and
the male eggs, are not indiscriminately mix-
ed in the oviducts. Undoubtedly they oc-
cupy a situation corresponding to the princi-
ples that regulate her laying : the eggs of
workers are first, and those of drones behind
them. Farther, it appears that the queen
can lay no male eggs until those of workers,
occupying the first place in the oviducts, are
discharged. Why, then, is this order in-
verted by retarded impregnation ? How does
it happen that the whole workers’^ eggs which
the queen ought to lay, if her commerce had
been in due time, now wither and disappear,
yet do not obstruct the passage of the eggs
of drones, which occupy only the second place
in the ovaries ? Nor is this all. I have sa-
tisfied myself that a single union is sufficient
to impregnate the whole eggs that a queen
will lay in the course of at least two years.
I have even reason to think that it impreg-
nates all the eggs that she will lay during
her entire life : but I want absolute proof

HETAllDED IMPEEGNATION.

45

for more than two years. This, wliich is
truly a very admirable fact in itself, renders
the influence of retarded fecundation still
more difficult to be accounted for. Since a
single junction suffices, it is clear that the
masculine matter acts from the first moment
on all the eggs that the queen will lay in
two years. According to your principles,
it gives them that degree of animation which
afterwards effects their successive expansion.

I Having received the first impressions of life,
they grow, they mature, so to speak, until
the day they are produced ; and as the laws
I of laying are invariable, because the eggs
[ of the first eleven months are always those
I of workers, it is evident that those which
j appear first are also the eggs that come soon-
s est to maturity. Thus, in the natural state,
[ the space of eleven months is necessary for
! the male eggs to acquire that degree of in-
I crement they must have attained when laid.
I This consequence, which to me seems im-
I mediate, renders the problem insoluble. How
) can the eggs, which should grow slowly for
( eleven months, suddenly acquire their full
I expansion in forty-eight hours, when fccun-
^ dation has been retarded twenty-one days,
\ and by the effect of retardation alone ? 0)>

i

a

46

EFFECTS OF

serve, I Leseech you, that the hypothesis o
successive expansion is not gratuitous ; it
rests on the principles of sound philosophy.
Besides, to be convinced that it is well found-
ed, we have only to look at the figures given
by Swammerdam of the ovaries of the queen
bee. There we see the eggs in that part of the
oviducts contiguous to the extremity, much
farther advanced and larger than those con-
tained in the parts more remote. There-
fore the difficulty remains in full force : it
is an abyss wherein I am lost.

The state of certain vegetable seeds, which,
although in good preservation, lose the fa-
culty of germination from age, is the only
known fact bearing any relation to that now
described. The eggs of workers may also
retain, but for a very short time, the proper-
ty of receiving the impressions of the pro-
lific matter; and after this period, which is
about fifteen or eighteen days, elapses, they
may become so disorganized as to be no long-
er susceptible of its stimulus. I am sensible
that the comparison is very imperfect; be-
sides, it explains nothing, nor does it even
put us on the way of making any new ex-
periments. Let me add but one reflection
more.

Iietarded impregnation. 4*7

Hitherto no other effect has been observ-
ed from the retarded impregnation of ani-
mals, but their becoming absolutely sterile.
The first instance of a female still preserv-
ing the faculty of engendering males, is pre-
sented by the queen bee. But as no fact in
nature is unique, it is most probable that
the same peculiarity will be also found in
other creatures. An extremely curious oh-
1 ject of research would be to consider insects
in this new point of view ; I say insects, for
I do not conceive that any thing similar
I will be found in other species of animals,
i The experiments now suggested will neces-
I sarily begin with insects the most analo-
j gous to bees; as wasps, humble bees, mason
I bees, all kinds of flies, and the like. Some
experiments might be also made on but-
terflies; and, perhaps, an animal may be
found whose retarded fecundation will be
attended with the same effects as that
of queen bees. Should the animal be lar-
I ger, dissection will be more easily accom-
j plished; and we may be able to discover
I what happens to • the eggs when retarded
j impregnation prevents their expansion. At
least, we may hope that some fortunate

■1

48

Effects of

circumstance will lead to solution of the
.problem.*

Let us now return to my experiments. In
]\Iay 1789, 1 took two queens just after
having undergone their last metamorphosis :
one w^as put into a leaf hive well provided
with honey and wax, and sufficiently peo-
pled by workers and males, and the other
into a hive exactly similar, from which all
the drones were removed. The entrances of
these hives were too confined for the passage
of the females and drones, hut left per-
fect freedom to the common bees. After
thirty days’ imprisonment, the queens being
released, they departed and returned impreg-
nated. On visiting the hives in the begin-
ning of July, I found much brood, but
wholly consisting of the worms and nymphs
of males. Actually there was not a single
workers worm or nymph. Both queens laid
without interruption until autumn, and uni-

* The experiments suggested in this paragraph recal a
singular reflection of M. de Reaumur. Where treating of
oviparous flies, he says, it woilld not be impossible for a
hen to produce a living chicken, if after fecundation, die
■eggs she should lay first could be by any means rctaineil
twenty-one days in the oviducts. Mem. sxtr. Ics Insect,
tom. ‘t. 7nem. 10.

RETARDED IMPREGNATION. 49

formly the eggs of drones, and they ceased
as any other queens in the first week of No-
vemher.

I was much interested to learn what
would become of them in the subsequent
spring, whether they would resume laying,
or if they required new fecundation ; and if
they did lay, of what species the eggs would
be. However, the hives being very weak,
I was apprehensive of their perishing during
the winter. Fortunately, we were able to
preserve them ; and from April 1 790, the
queens recommenced laying, while the pre-
cautions we had taken prevented tlrem from
receiving any new approaches of the male.
Their eggs were still those of drones.

It would have been extremely gratifying
to have followed the history of these two fe-
males still farther, but, to my s-reat regret,
the workers abandoned their hives on the
fourth of INIay, and that same day I found
iboth queens dead. No weevils were in the
Alive, which could disturb the bees; and the
■ honey was still very plentiful; hut as, in the
I course of the preceding year, no workers
had been produced, and winter had destroy-
lied many, their numbers were too few in
spring for engaging in their wonted labours,

I D

50

EFFECTS OF

and, from discouragement, they deserted
their habitation to occupy the neighbour-
ing hives.

In my Journal, I find a detail of many
experiments on retarded impregnation — so
many, that it would be tedious to transcribe
the whole. I may repeat, however, that
there was not the least variation in the
jn'inciple : and that whenever the union of
queens was postponed beyond the twenty-
first day, the eggs of males only were pro-
duced. Therefore, I shall limit my narra-
tive to those experiments that have taught
me some remarkable facts.

A queen being hatched on the fourth of
October 1789, 've put her into a leaf-hive.
Though the season was well advanced, con-
siderable numbers of males were still in it ; :■

and here it became important to learn, whe- -
ther at this period of the year they could I'
equally effect fecundation ; also, in case it |;
succeeded, whether the queen’s laying, be- I
o-un in the middle of autumn, would be in- •
terrupted or continued during winter. Thus,
we allowed her to leave the hive. She de- •
parted, indeed, but made four-and-twenty j
fi'uitless attempts before returning with tlie ii
evidence of fecundation. Finally, on the > ‘

RETARDED IMPREGNATION. 51

tliirfcy-first of October, she was more fortu-
nate. Departing, she brought back uii-
doubtecl proof. of the success of her amours :
and now being twenty-seven days old, fe-
cundation had been retarded. She ought
to have begun laying wuthin forty-six hours,
but the weather was cold, and she laid
none; which proves, as we may cursorily re-
mark, that refrigeration of the atmosphere
is the principal agent that suspends the
laying of queens during winter. I vwas ex-
cessively impatient to learn whether, on the
return of spring, she would prove fertile,
without a new union. The means of ascer-
taing the fact was easy ; for the entrances of
the hives only required contraction, so as to
. prevent her from escaping. She was con-
fined from the end of October until May.
In the middle of March, visiting the combs,
we found a considerable number of eggs,
but none being yet hatched, it could not be
known whether they would produce work-
^ ers or males. On tlie fourth of April,
t however, having again examined the state
of the hive, we found a prodigious quan-
5 tity of nymphs and worms, all of drones ;
iiior had this queen laid a single worker’s
■egg.

EFFECTS OF

%J/0l

Here, as well as in the preceding cxperi-*
inent, retardation had rendered the queens
incapable of laying the eggs of workers*

But this result is the more remarkable, as
the queen did not commence laying until
four months and a half after fecundation.

It is not rigorously true, therefore, that the
term of forty-six hours elapses between the
sexual union of the female and her laying ;
the interval may be much longer, if the
weather grows cold. Lastly, it follows, that
although cold will retard the laying of a
queen impregnated in autumn, she will be-
gin to lay in spring without requiring com-
merce anew.

It may be added, that the fecundity of
the queen, whose history is given here, was ;
astonishing. On the first of May, we found
in her hive, besides six hundred males, al- '
ready in the winged state, tw'o thousand four
hundred nnd thirty-eight cells, containing
either eggs or nymphs of drones. Thus,
she had laid more than three thousand male
eggs during March and ^pril, w'hich is
above fifty each day. Her death occurring
soon afterwards, unfortunately interrupted
my observations. I intended to calculate i
the total number of male eggs that she would I

EETAEDED IMPREGNATION. 52

\

j have laid throughout the year, and compare it
with the laying of queens whose fecundation
had not been retarded. You know, Sir, that
the latter lay about two thousand male eggs
in spring ; and another laying, but less con-
siderable, is resumed in August ; also that
in the interval, they produce the eggs of
workers almost exclusively. But it is other-
wise with the females whose union has been
retarded; they produce no workers’ eggs.
For four, five, or six months following, they
[ lay the eggs of males witliout interruption,

! and in such numbers, that, in this short time,

. I suppose one queen gives birth to more
I drones tlian are produced in tlic course of
two years by a female whose fecundation has
L not been retarded. It gives me much re-
j gret that I have not been able to verify this
If conjecture.

I should also describe the very remarkable
ii manner in which queens, laying only the
3 eggs of drones, sometimes deposit them in
the cells. Instead of being jdaced in the
)l lozenges forming the bottom, they are fre-
J quently fixed on the lower side of the cells,
i two lines from the mouth; which results
from the body of such queens being shorter
i than that of those whose fecundation has

54

EFFECTS OF

not been retarded. The extremity remains
slender, while the first two rings next the
thorax are uncommonly enlarged. Thus, in
disposing themselves for laying, the extremi-
ty cannot attain the bottom of the cells on
account of the swolu rings, consequently the
egg must remain attached to the part reached
by it. The worms proceeding from the eggs
pass their vermicular state in the same place
where they are deposited, which proves that
bees are not charged with the care of trans-
porting the eggs, as has been supposed. But
here another plan is followed: they advance
those cells wherein they observe eggs two
lines from the orifice, beyond the surface of
the comb. It is not essential, however, that
they shall be fixed by one end at the bottom
of the cell.

Permit me, Siivto digress a moment from
the subject, and give the I'esult of an expe-
riment which seems interesting. Bees, I
say, are not charged with the care of trans-
porting into cells the eggs misplaced by the
queen : and, judging by the single instance
I have related, you will think me well enti-
tled to deny this feature of their industry.
Yet, as several authors have maintained the
reverse, and even demanded oui admiration

retarded impregnation. 55

of them in conveying the eggs, I should ex-
plain clearly that they are deceived.

I had a glass hive constructed of two stages;
the higher was filled with combs composed
of large cells, and the lower with those of
common ones. A kind of division or dia-
phragm, separated these two stages from
each other, having an opening at each side
for the passage of the workers from one stage
to the other, but too narrow for the queen.
I put a considerable number of bees into
this hive; and, in the upper part, confined
a very fertile queen that had just finished
her great laying of male eggs ; therefore she

Ihad only those of workers to lay, and she
was obliged to deposit them in the surround-
ing large cells from the want of others. INIy
object in this arrangement will be already
anticipated. The reasoning was simple. If
the queen laid workers’ eggs in the large

i cells, and the bees were charged with trans-
porting them if misplaced, they would in-
fallibly take advantage of the liberty allowed
i them to pass from either stage ; they would
i seek the eggs deposited in the large cells,
i and carry them down to the lower stage con-
i taiuing the cells adapted for that species.
I If, on the contrary, they left the common
I eggs in the large cells, I should obtain cor-

56

EFFECTS OF

tain proof that they were not entrusted with
transporting them.

The issue of this experiment excited my
curiosity extremely ; and we observed the
queen several days without intermission.
During the first twenty-four hours, she ab-
stained from laying a single egg in the sur-
rounding cells ; she examined them one after
another, but passed on without insinuating
her belly into any. She restlessly traversed
the combs in all directions : her eggs ap-
peared an oppressive burden, but she per-
sisted in retaining them rather than they
should be deposited in cells of unsuitable
diameter. The bees, however, did not cease
to pay her homage, and treat her as a mo-
ther. It was amusing to observe, when
reaching the edges of the division separating
the two stages, how she gnawed at them to
enlarge the passage : the workers approached
her and also, labouring with their teeth,
made every exertion to augment the en-
trance to her prison, but ineffectually. On
the second day, the queen could no longer
retain her eggs ; they escaped in spite of
her, and fell at random : yet eight or ten
were seen in the cells which next day had
disappeared. Then we conceived that
the bees had conveyed them into the

RETARDED IMPREGNATION.

I

i

' 1

•?

B

[(

r

C

0

c

I

0

57

' small cells of tlie lower stage, where we
sought them with the utmost assiduity ;
hut I can safely affirm not one was there.
The eggs that the queen still laid on the
third day, disappeared as the first. We
again sought them iu the small cells, but
none were found. The fact is, they were ate
by the workers ; and this is what has de-
ceived the naturalists, who supposed them
carried away. They have remarked the
disappearance of the misplaced eggs, and,
without farther investigation, have asserted
that the bees convey them elsewhere : they
take them, indeed, not to be carried off, but
to be devoured.'*^

Thus nature has not entrusted bees with
the care of placing the eggs in the cells ap-
propriated for them, but she has ins])ired fe-
males themselves with sufficient instinct to
know the species of eggs which they are
about to lay, and to deposit them in those
suitable. Phis has been already observed
by M. de Reaumur, and here my observa-

* This is not peculiar to the workers of these animals.
Among humble bees, which also live in society, the work-
ers endeavour to destroy the eggs : and after the females
have deposited them in the cells, they have to contend vi-
gorously with the common bees for their preservation.— 7',

58

EFFECTS OF

tions correspond with his. Therefore it is
certain, that in the natural state, when fe-
cundation -ensues at the proper time, and
the queen has suffered from nothing, she is
never mistaken in the choice of the cells
where her eggs are to be deposited ; she ne-
ver fails to lay those of workers in small
cells, and those of males in large ones. The
distinction is important, for the same uner-
ring instinct is no longer conspicuous in the
conduct of females whose impregnation has
been deferred. I was oftener than once de-
ceived respecting the eggs laid by such
queens, for they were deposited indiscrimi-
nately in small cells and in those of drones;
and, not aware of their instinct being af-
fected, I conceived that the eggs in small
cells would produce workers ; whence I was
very much surprised to see the bees close up
the cells at the moment they should have
been hatched, and demonstrate, by anticipa-
tion, that the included worms would change
into drones. They actually became males ;
those produced in small cells were small,
those in large cells large. Thus I must
warn observers, who would repeat my expe-
riments on queens laying only the eggs of

RETAKDED IMPREGNATIOX.

59

males, not to be misled by these circiiin-
stances, and expect that eggs of males will
be deposited in the workers’ cells.

It is a singular fact also, that females,
if whose fecundation has been retarded, some-
I times lay the eggs of males in royal cells.
[ I shall prove, in the history of swarms, that
i immediately when queens, in the natural
? state, begin their great laying of male eggs,
numerous royal cells are constructed. Un-
1 doubtedly there is some secret relation
' between the appearance of these eggs and
I their formation; for bees never derogate
I from this law of nature. It is not surprising,
therefoie, that such cells are constructed in
: hives governed by queens laying the eggs
1 of males only : it is no longer extraordinary
' that these queens deposit, in the royal cells,

! eggs of the only species they can lay, for in
! general their instinct seems affected. But
what I cannot comprehend is, why the bees
I take exacBy the same care of the male eggs
deposited in royal cells, as of those that shoidd
I become queens. They provide them more
I plentifully wUh food, they build up the cells
■ as if containing a royal worm ; in a word,
t^hey labour with such regularity that we
I have been frequently deceived. More than

60

EFFECTS OF

once, in the firm persuasion of finding royal
nymphs, we have opened the cells after they
were sealed, yet the nymph of a drone al-
ways appeared. Here the instinct of the
workers seems defective. In the natural
state, they can accurately distinguish the
male worms from those of common bees, as
they never fail to give a particular covering
to the cells containing the former. Why
then can they no longer distinguish the
worms of drones, when deposited in the royal
cells ? The fact deserves much attention. I
am convinced that in investigating the in-
stinct of animals, we must carefully observe
where it appears to err.^

Perhaps I should have begun this letter
with an abstract of the observations of prior
naturalists, on queens laying none but the
eggs of males ; however, I shall here repair
the omission.

In a work, Histoire dc la Reine des
Aheillcs, translated from the German by
Rlassiere, there is printed a letter from
Schirach to you, dated April 15, 1771, where
he speaks of certain hives the whole brood

* It is proved by the author’s later observations, that the
instinct of the workers is not bo defective as he is inclined
licrc to conclude. — T.

RETARDED IMPREGNATION. 61

of which changed into drones. You will re-
member that he ascribes this circumstance
to some unknown vice in the ovaries of the
queen ; but he was far from suspecting that
it resulted from retarded fecundation. He
justly felicitated himself on discovering a
method of preventing the destruction of
i hives in this condition, which was simple,

I for it consisted in removing the queen that
I laid the eggs of males only, and substituting
' one, whose ovaries were not impaired, for
' her. But to render the substitution effec-
j tual, it w'as necessary to procure queens at
( pleasure ; a secret reserved for Schirach,
[. and of which I shall speak in the following
3 letter. You observe that the whole expe-
i riments of the German naturalist tended to
i the preservation of the hives whose queens
fi laid none except male eggs ; and that he
i! did not attempt to discover the cause of the
7 evident vitiation of their ovaries.

M. de Reaumur also says a few words
i( somewhere of a hive containing many more
>1 drones than workers, but advances no con-
5 jectures on the cause. However, he adds,
I as a remarkable circumstance, that the males
! 0 were tolerated in tliis hive until tlic subsc-
^ J quent spring. It is true, that bees governed

62 RETARDED IMPREGNATION.

by a queen laying only male eggs, or by a
virgin queen, preserve their drones several
months after they have been massacred in
other hives, for which I can ascribe no rea-
son ; but it is a fact that I have witnessed
several times during my long course of ob-
servations on retarded impregnation. In
general it has appeared, that while the queen
lays male eggs, bees do not destroy the
males already perfect in the hive. Pregny»
August, 1791.

LARV^ CONVERTED TO QUEENS. 63

LETTER IV.

ON sciiirach’s discovery.

When you found it necessary, Sir, in the
new edition of your works, to give an ac-
count of Scliiracli’s beautiful experiments
on the conversion of common worms into
royal ones, you invited naturalists to repeat
them. Indeed such an important discovery
required the confirmation of several testi-
monies, and for this reason, I hasten to in-
form you that all my researches establish its
: truth. During ten years that I have studi-
ed bees, I have repeated Schirach’s ex-
periment so often, and with such uniform
• success, that I can have no longer the least
doubt on the subject. Therefore, I consider

64

LAUViE OF WORKERS

it an established fact, that when bees lose
their queen, while several workers’ worms
are preserved in the hive, they enlarge some
of their cells, and supply them not only with
a different kind of food, hut with a greater
quantity of it, and that the worms reared in
this manner, instead of changing to common
bees, become real queens. I request my
readers to reflect on the explanation you
have given of so uncommon an incident, and
the philosophical consequences deduced from
it, Contemplation de la Nature, part 11,
chap. 27.

Here I shall content myself with some ac-
count of the figure of the royal cells con-
structed by bees around the worms destin-
ed for the royal state, and terminate with
discussing some points wherein my ob-
servations are at variance with those of
Schirach.

Bees soon become sensible of having lost
their queen, and in a few' hours commeuce
the labour necessary to repair their misfor-
tune. First, they select the young common
worms, w'hich, by a certain treatment, are to
be converted into queens, and immediately
begin enlarging the cells where they are de-
posited. Their mode of proceeding is cnri-

CONVERTED TO QUEENS.

65

ous; and tlie better to illustrate it, I shall
describe the labour bestowed on a single cell,
which will apply to all the rest containing
worms destined for queens.

The bees having selected a worm, three of
the contiguous cells are sacrificed: they sup-
ply it with food, and surround it by a cylindri-
cal enclosure, rendering the cell a perfect tube,
with a rhomboidal bottom, for this part re-
j mains untouched.^ Were it injured, three
i corresponding cells on the opposite surface
' of the comb would be laid open, and the
[I worms would perish ; therefore, their destruc-
W tion being unnecessary, nature has opposed it.
i Leaving the bottom of the cell rhomhoidal,
the bees are content with raising a cylindri-
u cal tube around its worm which, like the
i other cells of the comb, is horizontal. But
i this habitation is adapted to the larva called
to the royal state only for the first three
^ days of its existence : it requires another
^ position for the other two that it is still a
f worm. During these which constitute so
^ small a portion of its life, it must occupy a
f cell nearly of a pyramidal figure, hanging in

* The bottom of a cell consists of three lozenges or
' S rhomboids, as is amply explained in the second part of
,1 the work. — T-

66

LARViE OF WORKERS

a perpendicular direction. We may affirm
that the workers are aware of this ne-
cessity; for, after tlie worm has completed
the third day, they prepare the place to be
occupied by its new lodging. Gnawing
away the cells environing the cylindrical
tube, they mercilessly sacrifice their worms,
and employ the wax in constructing a
ncAv pyramidal tube, which they solder at
right angles to the first, and work it down-
wards. The diameter of this pyramid de-
creases insensibly from the base, which is
very wide, down to the point. During the i
two days that it is inhabited by the worm,
a bee constantly keeps its head more or less,
inserted into the cell, and, on quitting it,
another comes to occupy its place. In pro-
portion as the worm grows, the bees labour
in extending the cell, and bring food, which
they place before its mouth, and around its
body. The worm, which can move only in
a spiral direction, turns incessantly to take
the food before its head ; it insensibly de-
scends, and at length arrives at the orifice
of the cell. Now is the time of transfor- !
ination to a nymph. As any fartlier care |
would be unnecessary, the bees close the cell |
with a peculiar substance appropriated for I

CONVERTED TO QUEENS^ 67

it, and there the worm undergoes both its
metamorphoses.

Though Schirach supposes that none but
I worms three days old are selected for the
I royal treatment, I am certain of the contra-
ry ; and that the process succeeds equally
well with those of two days only. I must
1 be permitted to relate at length the evi-
I dence I have of the fact, which will demon-
I strate both the reality of common worms
I being converted into queens, and the little
I influence which their age has on the effect
f of the treatment.

I I put some pieces of comb containing

[workers’ eggs, in the cells, of the same kind
as those already hatched, into a hive depriv-
ed of the queen. The same day several
cells were enlarged by the bees, and convert-
ed into royal cells, and the worms supplied
with a thick bed of jelly. Five were then
removed by us from these cells, and five
common worms, which had come from the
egg forty-eight hours previously before us,
were substituted for them. The bees did
not seem aware of the change ; they watch-
ed over the new worms as over those chosen
by themselves : they continued enlarging
the cells, and closed them at the usual time.

^8

larvae of workers

When they had hatched on them seven
days,* we took away the cells to preserve
the queens that would be produced. Two
were excluded, almost at the same moment,
of the largest size, and well formed in every
respect. 'The term of the other cells hav-
ing elapsed without any queen appearing,
we opened them, and found in one a dead
queen, but still a nymph ; the other two
were empty. After spinning their silk coc-
coons, the worms had died before passing into
their nymphine state, and presented only a
dry skin. I can conceive nothing more
conclusive than this experiment. It demon-
strates that bees have the power of convert-
ing the worms of workers into queens; since
they succeeded in procuring them by oper-
ating on the worms which we ourselves had
selected. It is equally demonstrated, that
the success of the process does not depend
on the worms being three days old, as those :
entrusted to the bees were only two. Nor i
is this all; bees can convert worms still:
younger into queens, for the following expe- •
rimcnt shows, that when the queen is lost,,
they destine worms only a few hours old to )
replace her.

* The author’s meaning here is obscure. — T,

CONVERTED TO QUEENS.

69

A hive in my possession having been long
deprived of the femalej had neither egg nor
worm. I provided a queen of the greatest
fertility; and she immediately began lay-
ing in the cells of workers; but I removed
I her before being quite three days in the hive,
and before any of her eggs were hatched.

' The following morning, that is on the fourth
day, we counted fifty minute worms, the
oldest scarcely hatched twenty-four hours.
However, several were already destined for
. queens, which was proved by the bees depo-
I siting around them a much more abundant
S provision of food than is supplied to com-

tmon worms. Next day, the worms being
nearly forty hours old, the bees had enlarged
and converted their hexagonal cells into cy-
■a lindrical ones of the greatest capacity. Dur-
I ing the subsequent days, they still laboured
\:k at them, and closed them on the fifth from
A the origin of the worms. Seven days after
J sealing of the first of these royal cells, a
|A queen of the largest size issued from it, and
W immediately rushing tow'ards the other roy-
al cells, she endeavoured to destroy their
ofl nymphs and worms. In another letter, I
j » shall relate the effects of her fury.

70

LARV^ OF WORKERS

From these details, you will observe, Sir,
that Schirach’s experiments had not been
sufficiently diversified when he affirmed that
it was essential for the conversion of com-
mon worms into queens, they should be
three days old. It is undoubted, that equal
success attends the experiment, not only
with worms two days old, but also when
they have been only a few hours in exist-
ence.

After my researches to corroborate
Schirach’s discovery, I was desirous of learn-
ing whether, as this observer conceives, the
only means which the bees have of procur-
ing a queen, is giving the common w’orms a
certain kind of aliment, and rearing them in
the largest cells. You will remember, that
]\I. de Reaumur’s sentiments are very differ-
ent : “ The mother 'should lay, and she ac-
tually lays, eggs from which bees fit for be-
ing mothers must in their turn proceed.
She docs so ; and it is evident the workers
know what she is to do. Bees, to which the
mother is so precious, seem to take a pecu-
liar interest in the egg that is to produce a
mother, and to consider it of the greatest
value. They construct particular cells where
such are to be deposited. The figure of a royal

CONVERTED TO QUEENS. 71

cell only begun, very much resembles a cup,
or, more correctly. speaking, the cup that has
lost its acorn.”

M. cle Reaumur, though not suspecting
the possibility of a common worm being
; converted into a queen, conceived that the
queen bee laid a particular species of eggs
in the royal cells, from which worms should

■ come that would be queens. According to
Schirach, on the other hand, bees always
having the power of procuring a queen, by
bringing up worms three days old in a par-

I ticular manner, it would be needless for
: nature to grant females the faculty of lay-
! ing royal eggs. Such prodigality is, in his

■ eyes, inconsistent with her ordinary laws.
Therefore he maintains, in direct terms,
that the queen does not lay royal eggs in
cells purposely prepared to receive them.
He considers the royal cells only as common
ones, enlarged by the bees at the moment
when the included worm is destined for a

‘ queen; and adds, that the royal cell is al-

' ways too long for the belly of the mother to
: reach its bottom.

I admit that M. de Reaumur nowhere
says he has seen the queen lay in the royal
owe\ei, he did not doubt the fact;

72

LARV^ OF TTORKER3

and, after all my observations, I must es^
teem his opinion just. It is quite certain
that, at particular periods of the year, bees
prepare royal cells ; that the females deposit
their eggs in them ; and that worms, which
will become queens, issue from these eggs.

Schirach’s objection, concerning the length
of the cells, proves nothing ; for the queen
does not postpone depositing her eggs till
they are finished. While only sketched and
shaped like the cup of an acorn, she lays in
them. This naturalist, dazzled by the bril-
liancy of his discovery, saw but part of the
truth. He was the first to find out the re-
source granted to bees by nature for repair-
ing the loss of their queen ; and too soon per-
suaded himself that she had provided none
other for the production of females. His
error arose from not observing bees in very
flat hives : had he used such as mine, he
would have found a confirmation of ^I. de
Keaumur’s opinion on opening them in
spring. Tlien, which is the season of swarm-
ing, hives in good condition are governed by
a very fruitful queen. There are royal cells
of a figure widely different from those con-
structed around the worms destined by the i
bees for queens : large, attached to the comb i

CONVERTED TO QUEENS. 73

by a stalk, and hanging vertically like sta-
lactites ; such, in short, as described by M.
de Reaumur. The females lay in them be-
fore completion, and we have surprised a
queen depositing the egg when the cell was
] only as the cup of an acorn. The workers
never lengthen them until it has been laid.
1 In proportion as the worm grows, they are
t enlarged, and closed by the bees when the
i first transformation approaches. Thus, it is
I true, that, in spring, the queen deposits eggs
t in royal cells previously prepared, from which
? winged insects of her own species are to come!
\ Nature has, tlierefore, provided a twofold
t means for the multiplication and conserva-
i tion of their race. Pregny, ^4>t/i August
\ 1791.

1

P.

74

SOME WORKERS

LETTER V.

EXPERIMENTS PROVING THAT THERE ARE
SOMETIMES COMMON BEES WHICH LAY
FERTILE EGGS.

The singular discovery of Riem, cou-
cerniiig the existence of fertile workers, has
appeared very doubtful to you, Sir. You have
suspected that the eggs ascribed to work-
ers by this naturalist were actually produced,
by small queens, which, on account of their
size, had been confounded with common
bees. But you do not positively insist that
Riem is deceived ; and, in the letter which
you did me the honour to address to me,
you requested me to investigate, by new ex-
periments, whether there are truly working
bees capable of laying fertile eggs. I have

LAV FEPvTILE EGGS. 75

made such experiments with great care : and
it is for you to judge of the confidence which
they merit.

On the fifth of August 1788, we found
the eggs and worms of large drones in two
hives, hoth of which had been some time de-
prived of queens. We also observed the
I rudiments of several royal cells appended
like stalactites to the edges of the combs,
and containing the eggs of males. Being
perfectly certain that there was no queen of
: large size among the bees of these two hives,
it was evident to us that the eggs, daily
. becoming more numerous, were laid, either
by queens of small size, or by fertile workers.
I had reason to believe that it was actually
by common bees, for we had frequently ob-
[: served them inserting the posterior part of
the body into the cells, and assuming the
; same attitude as the queen when laying.
.But, notwithstanding every exertion, we
never had been able to seize one in this posi-
tion to examine it more narrowly : and we
were unwilling to assert any thing positively,
without having the bees in our hands that
had actually laid. Therefore our observa-
tions were continued with equal assiduity,
in hopes that, by some fortunate chance, or

76

SOME WOEKEIIS

ill a moment of address, we could secure one
of them. JNIore than a month all oiu- endea-
vours were abortive.

My assistant then offered to perform a
task requiring both courage and patience,
and which I could not resolve to suggest,
though the same expedient had occm-red
to myself. He proposed to examine each
bee in the hive separately, in order to dis-
cover whether some small queen had not
insinuated herself among them, and had
escaped our first researches. This was an
important experiment ; for, shoidd no small
queen be found, it would be demonstrative
evidence that the eggs had been laid by
simple workers.

To accomplish this investigation with all
possible exactness, immersing the bees was
not enough. You know. Sir, that tlie con-
tact of watei^ stiffens their organs, that it
produces a certain alteration of their exter-
nal figure ; and, from the resemblance of
small queens to workers, the slightest altcr-
'ation of shape would have prevented us
from distinguishing with sufficient accuracy
to which species those that were immersed
might belong. Therefore it was necessary
to seize the whole bees of both hives, not-

LAY FERTILE EGGS.

77

withstanding their irascible nature, and ex-
amine their specific character with the ut-
most care. This my assistant undertook,
and executed with great address. Eleven
days were employed in it ; and, during all
that time, he scarcely allowed himself any
relaxation, hut what the relief of his eyes
required. He took every bee in his hand ;
he attentively examined the trunk, the
I hind limbs, and the sting. Not one ap-
I peared without the characteristics of the
; common bee, that is, the little basket on the
j hind legs, the long trunk, and the straight
j sting. Having previously prepared glass
i cases containing combs, he put each bee in-
j to them after examination. It is superflu-
(< ous to say they were confined, which was a
^ precaution indispensible, until termination
I of the experiment. Neither was it enough
to establish that the whole were workers;
^ we had also to protract the experiment, and

i observe whether any of tliem would pro-
duce eggs. Thus we inspected the cells for
I scveial days, wherein we soon discovered
t new laid eggs, from which the worms of
I drones came at the proper time. My as-
9 sistant held in his hands the bees that pro-
I) t need them ; and as he w'as perfectly cer-

78

SOME WORKERS

tain they were common ones, it is proved
that there are sometimes fertile workers in
hives.

Plaving corroborated Schirach’s discovery
by so decisive an experiment, we replaced
all the bees examined in flat hives, only an
inch and a half wide, which being capable
of admitting but a single row of combs,
were very favourable to the observer. We
expected, by persisting in strictly watching
them, that a fertile one might be sur-
prised in the act of laying, which we were
desirous of doing, for the purpose of ascer-
tainino: the difference of its ovaries from
those of queens by dissection.

At length, on the eighth of September,
we had the good fortune to succeed. A
])ee appeared in the position of a female lay-
ing : before having time to leave the cell,
we suddenly opened the hive and seized it.
This insect presented all the external charac- i
teristics of common bees ; the only difference
we could recognize, and that a very slight
one, consisted in the belly seeming less, and
more slender than that of workers. On
dissection, the ovaries were found to be
double, like those of (luecns, but more fra-
o-ilc, smaller, and composed of fewer ovi-

LAY FERTILE EGGS.

79

ducts. The filaments containing the eggs
were extremely fine, and exhibited swell-
ings at equal distances. We counted ele-
ven eggs of sensible size, some of which ap-
peared to have come to matin ity.

On the ninth of September, we seized
another fertile worker the instant it laid,
and dissected it. The ovary was still less
expanded than that of the preceding bee,
and only four eggs had attained maturity.
INly assistant extracting one from the ovi-
ducts, succeeded in fixing it by an end
on a glass slider. We may take this op-
portunity of remarking, that it is in Ixie ovi-
ducts themselves the eggs are imbued with
the viscous liquid with which they are pro-
duced, and not in passing from the spherical
sac, as Swammertlam believed.*' During

* It is not easy to understand this passage without re-
sorting to delineations of’ the ovaries, which the reader will
find in .Swammerdam’s Bihlia Naturie, pi. 19 : and pi. 32,
tom. 5, of Reaumur’s IMemoirs. A duct leads from each
ovary into a canal, through which the eggs are transmitted
from the queen. From this common canal, a spherical sac
rises near the union of the two ducts, where the secretion
of some particular fluid was su))posed. The eggs of most
insects have a kind of viscosity, by means of which, per-
haps, they are attached to the places where deposited. Thus,
we observe many rows lying horizontally, or fixed by one
end, on the leaves of plants, and adhering there until the
worm is hatched. — T.

80

SOME WORKERS

the remainder of this month, we found ten
fertile workers in the same hives, and dis-
sected them all. The ovaries of the greater
part were easily distinguished, but in some
we could not discern the faintest traces of
them. In the latter, the oviducts to all ap-
pearance were hut imperfectly developed,
and more address than we had acquired in
dissection was necessary for their discovery.

Fertile workers never lay the eggs of com-
mon bees ; they produce none but those of
males. Riem had already ascertained
this singular fact ; and here all my observa-
tions correspond with his. I shall only add
to what he says, that fertile workers are not
absolutely indifferent in the choice of the
cells for depositing their eggs. They always
prefer large ones, and resort to small cells
hut when unable to find those of greater dia-
meter. Flowever, they so far correspond
with queens whose impregnation has been ij
retarded, that they sometimes lay in royal |
cells.

When speaking of females laying male
eggs alone, I have already expressed my sur- ;
prise that bees bestow such care and atten- I
tion on those deposited in royal cells, as to
feed their worms, and, at the period of trans- f

t

LAY FERTILE EGGS.

81

formation, to close tliem up. But I know
i not, Sir, why I omitted to remark, that
■ after sealing the royal cells, the workers
: build them up, and sit on them until the
, last metamorphosis of the included male.'*
I The treatment of the royal cells where fer-
tile workers lay the eggs of drones is very
i different. They begin indeed with bestow-
’ ing every care on their eggs and worms ;
i they close the cells at a suitable time, but
1 never fail to destroy them three days after-
! wards.

; Having finished these first experiments
i with success, I had still to discover the cause
i of the evolution of the sexual organs of fer-
. tile workers. Biem had not engaged in this
D interesting problem ; and at first I dreaded
] that I should have no other guide towards
p its solution than mere conjecture. Yet, on
t serious reflection, it appeared that some
I light to direct my progress in this new re-
i search might be obtained, from connecting
( the preceding facts.

I reasoned thus: From Schirach’s ele-
i gant discoveries, it is beyond all doubt that

* It IS difficult to discover wliether the author tliiuks, as

1 some naturalists do, that bees are instrumental in hatchiiic
3 the eggs. — T.

82

SOME WORKEllS

common bees are originally of the female
sex.^' They have derived from nature the
germs of an ovary, but she has admitted of
its expansion only in the particular case of
their receiving a certain aliment while a
worm. Thus it must be the peculiar object
of inquiry, whether the fertile workers ob-
tain that aliment while worms.

All my experiments convince me that
bees, capable of laying, are produced in hives
that have lost the queen. A great quantity
of royal jelly is then prepared for feeding
the larvae destined to replace her. There-
fore, if fertile workers are produced under
these circumstances alone, it is evident
their origin is only in those hives where
bees’ prepare the royal jelly, and towards
this 1 must bend all my attention. It in-
duces me to suspect, that when bees give
the royal treatment to certain worms, they
either by accident or by a particular instinct,
the principle of which is unknown to me,
drop some particles of royal jelly into cells

* It is said by Warder, a practical observer in tlie
earlier part of last century, “ After great ]iains taken, and
curious observations made, with the use of the best glasses,
I must come to this certain conclusion, that all the working
bees arc females.” The True Amazons, or Monarchy of
the Bees, chap. 1. — T,

LAY FERTILE EGGS.

83

J contiguous to tliose containing the worms
i destined for queens. The larvae of workers
I that liave accidentally received portions of
i so active an aliment, must be more or less
^ affected by it'; and their ovaries should ac-
) quire a degree of expansion. But this ex-
] pansion will be imperfect; why? because
3 the royal food has been administered only
1 in small portions, and, besides, the larvas
( having lived in cells of the smallest dimeii-
i sions, their parts cannot enlarge beyond the
I ordinary proportions. Thus, the bees pro-
f duced from them will resemble common
t workers in size, and in all the external cha-
t racteristics. Added to that, they will have
f. the faculty of laying some eggs, solely from
I. the effect of the trifling portion of royal jel-
[ ly mixed with their aliment.

' That we may judge of the truth of this
I theory, it is necessary to consider fertile
t workers from their origin ; to investigate
whether the cells, where they are brought
\ up, are constantly in the vicinity of the
loyal cells, and if their food is mixed with
I pai tides of the royal jelly. Unfortunately
I the execution of these experiments is very
} difficult. \V hen pure, the royal jelly is re-
} cognised by its sharp and pungent taste ;

84

SOME WOllKEllS

but when mixed with other substances, its
peculiar savour is imperfectly distinguished.

Thus I conceived that my investigation
should be limited to the situation of the cells;
and, as the subject is important, permit me
to enter a little into detail.*

In June 1790, I observed that one of my
thinnest hives had wanted the queen for seve-
ral days, and that the bees had no means of
replacing her, there being no workers’ worms.
I then provided them with a small portion
of comb, each cell containing a young worm
of the working species. Next day, the bees
prolonged several cells around the w'orms
destined for queens, into the form of royal
ones. They also bestowed some care on the
worms in the adjoining cells. Four days
afterwards, all the royal cells were shut,
and we counted nineteen small cells also
perfected, and closed by a covering almost
flat, In these were worms that had not
received the royal treatment ; but as they
had lived in the vicinity of the worms des-
tined for replacing the queens, it was very
interesting to follow their history, and cs-
.sential to watch the moment of their last

Tlic original is extremely confused in tlie preceding
passages. — T.

LAY FERTILE EGGS.

85

transformation. I removed the nineteen
cells into a grated box, which was intro-
duced among the bees. I also removed the
royal cells, for it was of great importance
that the queens they would produce should
not disturb or derange the result of the ex-
periment. But here another precaution was
also requisite. It was to he dreaded that
the bees, being deprived of the produce of
1 their labour and the object of their hope,

I might be totally discouraged ; therefore, I
I supplied them with another piece of comb,

( containing the brood of workers, having it
1 in my power to destroy the young when

> necessary. This plan succeeded admirably.
The bees, in bestowing all their attention

i on these latter worms, forgot those that had

> been removed.

When the moment of transformation of
l3 the nymphs in the nineteen cells arrived, I
i examined the grated box frequently every
B day, and at length found six bees exactly si-
ri milar to common hees. The worms of the
r remaining thirteen had perislicd without
r changing.

The portion of brood comb that had been
•1 brought into the hive to prevent the discou-
A ragement of the bees was then removed. I

86

S03IE WORKERS

jnit aside the queens produced in the royal
cells ; and having painted the thorax of the six
bees red, and amputated the right antenna,
I transferred the whole into the hive, where
they were well received.

You easily conceive my object. Sir, in
this course of observations. I knew there
was neither a large nor small queen in the
hive: therefore, if, in the sequel, I should
find new laid eggs in the combs, how very
probable must it be that they had been pro-
duced by some of the six bees? But, to
attain absolute certainty, it w'as necessary
to take them in the act of laying. Some
ineffaceable mark was also required for dis-
tinguishing them in particular.

These proceedings were attended with the
most ample success. ^Ve soon found eggs
in the hive; their number increased daily;
and their worms were all drones. But a
long interval elapsed before we could take
the bees that laid them. At length, by
means of assiduity and perseverance, we per-
ceived one introducing the posterior part in-
to a cell; we opened the hive, and caught
the bee : We saw the egg it had deposited,
and by the colour of the thorax, and priva-
tion of the right antenna, instantly recog-

LAY TERTILE EGGS.

87

iiiscd it as one of the six that had passed
the vermicular state iii tlie vicinity of the

royal cells.

I could no longer doubt the certainty of
my conjecture ; at the same time, I know
not whether the truth will appear as rigor-
ous to you, Sir, as it does to myself. But
I reason in the following manner : If it be
certain that fertile workers are always pro-
duced in the vicinity of royal cells, it is no less
true, that in itself that vicinity is indiffer-
ent ; for the size and figure of these cells
can have no effect on the worms in those
surrounding them ; there must be something
more. W^e know that a particular aliment
is conveyed to the royal cells ; we also know,
that this aliment has a very powerful effect
on the ovaries; that it alone can unfold the
germ. Thus we must necessarily suppose,
that the worms in the adjacent cells have
had a portion of the same food : — and this

is what they gain by vicinity to the royal
cell. Numbers of bees, in their course
thither, will pass over them, stop and drop
some portions of the jelly destined for the
3 royal larvae. Such reasoning, I presume,
iis consistent with the principles of sound

88

SOME WORKERS

I have repeated the experiment now de
scribed so often, and weighed all the con-
comitant circumstances with so much care,
that whenever I please I can obtain fertile
workers in my hives. The method is sim-
ple. I remove the queen from a hive ; and
very soon the bees labour to replace her, by
enlarging several cells containing the brood
of workers, and by supplying the included
worms with the royal jelly. Portions of the
same aliment also fall on the young larvse
deposited in the adjacent cells, and occasion
a certain evolution of the ovaries. Fertile
workers are constantly produced in hives
where the bees labour to replace their queen;
but we very rarely find them, because they
are attacked and destroyed by the young
queens reared in the royal cells. There-
fore, all their enemies must be removed for
their preservation, and the larvaj of the roy-
al cells taken away before undergoing their
last metamorphosis. Then the fertile work-
ers, being free of rivals at the time of their
origin, will be well received; and, by taking
the precaution of marking them, it will be
seen, in a few days, that they produce the
eggs of males. Thus, the whole secret of
this process consists in removing the royal

LAY FERTILE EGGS.

89

cells at tlie proper time ; that is, after being
sealed, and previous to the young queens
leaving them.*

I shall add hut a few words to this long
letter. There is nothing so very surprising
in the production of fertile workers, when
we attend to the consequences of Schirach’s
beautiful discovery But why do they lay
male eggs only? I can conceive, indeed,

I that the reason of their laying few is from
the imperfect evolution of their ovaries, but
) I can form no conclusions why all the eggs
I should be those of males, neither can I ac-
f count any better for their use in hives ; and
j hitherto I have not made any experiments
L on their mode of fecundation. Pregny,
I 9,5th August 1791.

I* I have frequently seen queens, at the moment they
are produced, begin first by attacking the royal cells, and
then the common ones beside them. Being unacquainted
with fertile workers when I first observed this fact, I could
^ not conceive from what motive the fury of the queen was
n thus directed towards the common cells. But now I know
V that they can distinguish the species included, and have the
g same instinctive jealousy or aversion at them as against the
H nymphs of queens properly so denominated,

1

!*

90

COMBATS OF QUEENS.

LETTER VL

ON THE COMBATS OF QUEENS: THE MAS-
SACRE OF THE MALES: AND WHAT SUC-
CEEDS IN A HIVE M'HEEE A STRANGER
QUEEN IS SUBSTITUTED FOR THE NA-
TURAL ONE.

M. DE Reaumur had not witnessed every
tiling relative to bees when he composed his
liistory of these industrious animals. Seve-
ral observers, and tliose of Lusatia in par-
ticular, have discovered many important
facts that escaped him ; and I, in my turn,
have made various observations on some, of
which he liad no suspicion : at the same
time, and this is a very remarkable circum-
stance, not only all that he expressly de- I
dares he saw has been verified by succeed- '
ing naturalists, but all his conjectures are

COMBATS OF QUEENS.

91

foiiiicl just. Tlic Ggittisii iicitiircilisthj Sclii-
rad), Hattorf, and Riem, sometimes contra >
diet him,, indeed, in their memoirs; but I
can maintain that, while combating the opi-
nions of M. de Reaumur, it is they who are
almost always wrong; of which several in-
stances might be adduced.

IVhat 1 shall now proceed to say, will give
me an opportunity of detailing some inter-
esting facts.

IM. de Reaumur observed, that when any
supernumerary queen was either produced
in a hive, or came into it, one of the two
soon perished. He has not actually wit-
nessed the combat in which she falls, but
he conjectures there is a mutual attack, and
that the empire remains with the strongest
or the most fortunate. Schirach, on the other
ihand, and after him Riem, thinks that the
working bees assail the stranger, and sting
[her to death. I cannot comprehend by
[what means they were able to make this ob-
fservation. Having used none except very
thick hives, containing several rows of combs,
they could at most see nothing but the com-
mencement of hostilities. AVhile the coin-
mat lasts, the bees move with great rapidity;
they fly on all sides; and, gliding between

92

COMBATS OF QUEENS.

the combs, conceal their motions from the
observer. For my part, though using the
most favourable hives, I have never seen a :
combat between the queen and the workers, .
but I have very often beheld one between
the queens themselves.

Ill one of my hives in particular, there I
were five or six royal cells, each including a ?
nymph. The eldest first underwent its trans- '
formation. Scarcely did ten minutes elapse
from the time of her leaving her cradle,
vdien she visited the other royal cells still 1
close. Furiously attacking the nearest, by
dint of labour she succeeded in opening the I
top, and we saw her tearing the silk of the ^
coccoon with her teeth ; but probably her «
efforts were inadequate to the object, for 1
she abandoned this end of the cell, and be- I
gan at the other, where she effected a larger 1
aperture. When it was of sufficient size, 1
she endeavoured to introduce her belly, and m
made many exertions, until she succeeded in 9
giving her rival a deadly wound with her 9
sting. Then quitting the cell, all the bees 9
that hitherto had been spectators of her la- 9
hour began to enlarge the opening, ‘“inda
drew out the dead body of a queen scarce- J
ly come from its envelope of a nymph.

COMBATS OF QUEENS. 93

^leauwhile;, the victorious young queen
attacked another royal cell, but did not en-
deavour to introduce her extremity into it.
Here there was only a royal nymph, and no
queen, come to maturity, as in the first cell.
In all probability, the nymphs of queens in-
spire their rivals with less animosity. Still
! they do not escape destruction : because,
! whenever a royal cell has been opened be-
I fore the proper time, the bees extract the
contents in whatever form they may be,
whether worm, nymph, or queen : and
wdien this victorious female had abandoned
the second cell, the workers, enlarging the
opening which she had effected, extracted
the included nymph. Lastly, the young

I queen attacked a third cell, but could
not succeed in penetrating it. She labour-
I ed languidly, appearing as if exhausted by
9 her previous exertions. As we now required
I queens for some particular experiments, w^e
^ resolved to remove the other royal cells, yet in
^ safety, to secure them from her resentment.
1 After these incidents, we wished to see
I what ensued on two queens leaving their
tj I cells at the same time, and in what manner
bone of them perished. I find an observation
I on this head in my Journal, 15th May 1790.

94

COMBATS OF QUEENS.

Two queens quitted their cells in one of
our thinnest hives almost at the same mo-
ment, and, immediately on perceiving each
other, rushed together, apparently with
great fury. Their position was such, that
the antennae of each were seized by the
other’s teeth, while the head, breast, and
belly of both were mutually opposed. The
extremity of their bodies had only to be
curved, that they might be reciprocally
pierced with the stings, and fall dead together.
However it seems as if nature has not or-
dained that both combatants shall perish in
the duel ; but rather that, when finding
themselves in the situation now described,
namely, opposite, and belly to belly, they
shall recede at that instant with the utmost
precipitation. Thus, when these two rivals
felt the extremities about to meet, they dis-
engaged themselves, and each fled away.
You will observe, Sir, that I have repeated
this observation very often, so that it leaves
no room for doubt: and I tliink that here
we may penetrate the intention of nature.

There ought to be none but one queen iu
a hive : therefore it is necessary, if by chance
a second be either produced or comes into
it, that one of the two must be destroyed. !

Combats of queens.

Combats of queens. 95

This cannot be committed to the working
bees, because, in a republic Composed of so
many individuals, a simultaneous consent
cannot be supposed always to exist ; it might
liappen frequently that one group of bees
destroyed one of the females, while a se-
cond would massacre the other, and the hive
thus be deprived of queens. Therefore it
is necessary that they themselves should
be entrusted with the destruction of their
rivals : but as, in these combats, nature de-
mands only a single victim, she has wisely
arranged that, at the moment when, from
their position, the two combatants might
lose their lives, both feel so great an alarm

that they think only of flight, and not of
using their stings.

I am well aware of the hazard of error in

minute researches into the causes of the most

tiifling facts. But here the object and the

means seem so iilain, that I have ventured

to advance my conjectures. You will judge

setter than I can, whether they are well

ounded.— Let me now return from this di-
gression.

A few minutes after tlic two queens se-

Cu t I T™*'

to seek l,er rival. I„u„eaiately on

96

COMBATS OF QUEENSr

coming in sight, they rushed together, seized
one another, and resumed exactly their for-
mer position. The result of this rencontie
was the same as before. When their bellies
approached, they hastily disengaged them-
selves, and fled with precipitation. During
all this time, the workers seemed in great
agitation; and the tumult appeared to in-
crease when the adversaries separated. Two
different times we observed them stop the
flight of the queens, seize their limhs, and
retain them prisoners above a minute. At
last, that queen, which was either the strong-
est or the most enraged, darted on her rival
at a moment when unperceived, and with
her teeth caught the origin of the wing ,
then rising above her, she brought the ex-
tremitv of her own body under the belly ot
the other ; and, by this means, easily pierced
her with the sting. Quitting hold of the
wing, her sting was withdrawn ; the van-
quished queen fell down, dragged herself
languidly along, and, her strength failing,

she soon expired. ^

This observation proved that virgin queens
engage in single combats : but we wished to
discover whether those that were fecundated,
and mothers, entertained the same animosity.

COMBATS OF QUEENS.

97

On tlie 22d of July we selected a flat
hive, containing a very fertile queen : and
f being curious to learn whether, as virgin
» queens, she would destroy the royal cells,
j three were introduced into the middle of
j the comb. Whenever she became sensible of
I it, she sprung forward on the whole, and
r pierced them towards the bottom ; nor did
1 she desist until the included nymphs were
? exposed. The workers, which hitherto had
} been spectators of this destruction, now came
« to carry the nymphs away ; they greedily de-
^ voured the food remaining at the bottom of
f the cells, and also sucked the fluid from
i the abdomen of the nymphs: and termi-
nated^with destroying the ceils from which
they had been extracted.

In the next place, we introduced a very
fertde queen into the same hive; after paint
>ng the thorax to distinguish her from the
re.gnmg queen. A circle of bees quickly
formed around the stranger, but their in
ten mn was not to caress and receive her
[well for they insensibly accumulated so
touch, and surrounded her so closely, that
n scarcely a mittute she lost her liberi; and
Name a prisoner. It is a remarkable efr
kmstance, that other workers at the salj

98

COMBATS OF QUEEXS.

time collected around the reigning queen and
restrained all her motions ; we instantly saw
her confined like the stranger. Perhaps it
may be said, that the bees anticipated the
combat in which these queens were about
to engage, and were impatient to behold the
issue of it, for they retained their prisoners
only when they appeared to withdraw from
each other ; and if one less restrained seemed
desirous of approaching her rival, all the bees
forming the clusters gave w^ay to allow her
full liberty for the attack; then if the queens
testified a disposition to fly, they returned
to inclose them.

We have repeatedly witnessed this fact,
but it presents so new and singular a cha-
racteristic in the policy of bees, that it must
be seen again a thousand times before any
positive assertion can be made on the sub-
ject. I would therefore recommend that i
naturalists should attentively examine the i
combats of queens, and particularly ascertain |
what part is taken by the workers. Is their ■
object to accelerate the combat? Do they i
by any secret means excite the fury of the I ^
combatants ? AAniencc does it happen that, i ^
accustomed to bestow every care on their
queen in certain circumstances, they op- '

COMBATS OF QUEENS.

99

pose her preparations to avoid impending
danger?

A long series of observations is necessary

!

I

to solve these problems. It is an immense
field for experiment, which will afford infi-
nitely curious results. I intreat you to par-
don my frequent digressions. The subject
is deeply philosophical : genius such as your’s
is requisite to treat it properly ; and I shall
now be satisfied with proceeding in the de-
scription of the combat.

The cluster of bees that surrounded the
reigning qileen having allowed her some free^-
dom, she seemed to advance towards that
part of the comb where her rival stood ; then,
all receded before her, the multitude of
workers separating the two adversaries, gra-
dually dispersed, until only two remained ;
these also removed, and allowed the queens
to come in sight. At this moment the
reigning queen rushed on the stranger, with
her teeth seized her near the origin of the
wing, and succeeded in fixing her against
the comb without any possibility of motion
or resistance. Next curving her body, she
pierced this unhappy victim of our curiosity
with a mortal wound.

' In the. last place, to exhaust every com-
1 bination, we had still to examine whether

100

COMBATS OP QUEEXS.

a combat would ensue between two queens,
one impregnated, and the other a virgin ;
and what circumstances attended it.

On the 18th of September, we introdu-
ced a very fruitful queen into a glass hive,
already containing a virgin queen, putting
her on the opposite side of the comb, that
we might have time to see how the work-
ers would receive her. She was immedi-
ately surrounded, but confined only a mo-
ment. Being oppressed with the necessity
of laying, she dropped some eggs ; how-
ever, we could not discover what became of
them; certainly the bees did not convey
them to the cells, for, on inspection, we
found none there. The group surrounding !
this queen having dispersed a little, she ad-
vanced towards the edge of the comb, and '
soon approached very near the virgin queen.
When in sight, they rushed together ; the i
virgin queen got on the back of the other,
and gave her several stings in the belly, but,
being aimed at the scaly part, they did her
no injury, and the combatants separated.

In a few minutes, they returned to the
charge ; and this time the impregnated queen
mounted on Jier rival ; however, she sought
in vain to pierce her, for the sting did not

COMBATS OF QUEENS. 101

enter; the virgin queen then disengaged
lierself and fled ; and she also succeeded in
escaping another attack, where her adver-
sary had the advantage of position. These
rivals appeared nearly of equal strength;
and it was difficult to foresee to which side
victory would incline, until at last, by a suc-
cessful exertion, the virgin queen mortally
wounded the stranger, and she expired in a
moment. The sting had penetrated so far
that the victor was unable to retract it, and
she was overthrown by the fall of her ene-
my. She made great exertions to disen-
gage the sting : but could succeed by no
other means than timiiug on the extremi-
ty of her belly, as on a pivot. Probably its
barbs fell by this motion, and, closing spi-
rally around the stem, came more easily
from the wound.

These observations. Sir, I think will satis-
fy you respecting the conjecture of our ce-
lebrated Reaumur. It is certain that if
several queens are introduced into a hive,
one alone will preserve the empire ; that the
others will perish from her attacks ; and that
the workers at no time will attempt to em-
ploy their stings against a stranger queen.
I can conceive what has misled Riem and

102 COMBATS OF QUEENS.

Scliiracli ; but it is necessary, in explaining
it, that I should relate a new feature in the
policy of bees at considerable length.

In the natural state of hives, several
queens, issuing from different royal cells,
may sometimes exist at the same moment,
and they will remain either until formation
of a swarm or a combat among them decides
to which the throne shall appertain. But
excepting this case, there never can be su-
jiernumerary queens ; and if an observer
wislies to introduce one, he can accomplish
it only by force, that is, by opening the hive.
In a word, no queen can insinuate herself
into a hive in the natural state, for the fol-
lowing reasons.

Bees preserve a sufficient guard, day and
night, at the entrance of their habitation.
These vigilant centinels examine whatever
is presented : and, as if distrusting their
eyes, they touch with the antenme every in-
dividual endeavouring to penetrate the hive,
and also the various suhstances put within
their reach ; which affords us an opportuni-
ty of observing that the antenme are cer-
tainly the organs of feeling. If a stranger
queen appears, her entry is prevented by the
bees on guard instantly laying hold of her

COMBATS OF QUEENS.

103

C legs or wings with their teeth, and crowding
i so closely around her, that she cannot move.
Ij Other bees, from the interior of the hive,
gradually come to their assistance, and con-
fine her still more narrowly, all keeping their
heads towards the centre where the queen
is inclosed ; and they remain with such evi-
dent anxiety, eagerness, and attention, that
the cluster formed by them may be carried
about for some time without their being sen-
sible of it. A stranger queen, so closely
i! confined and hemmed in, cannot possibly
( penetrate the hive. If the bees retain her

i

I

)

>

[

too long imprisoned, she perishes, her deatli
probably ensuing from hunger, or the pri-
vation of air. It is undoubted, at least,
that she never is stung. Except in a single
instance, we never have seen the bees direct
their stings against her, and then it was
owing to ourselves. Compassionating a
queen s situation, we endeavoured to remove
her from the centre of a cluster ; the bees
became enraged; and, in darting out their
stings, some struck the queen, and killed
her. It is so certain that the stings were
not purposely directed against her that seve-
ral of the workers were tlieraselves killed ;
and surely they could not intend destroyiirg

104

SUBSTITUTION OF

one another. Had not we interfered, they
would have been content with confining
the queen, and she would not have perished.

It was in similar circumstances that
Riem saw the workers anxiously pursue a
queen. Believing that they designed to
sting her, he concluded that the office of
the common bees is to kill supernumerary
queens. You have quoted his observations
in the Contemplation de la Nature, part
11, chap. 27, note 7. But you are sen-
sible, Sir, from these details, that he has
been mistaken. He did not know the at-
tention paid by bees to what passes at the
entrance of their hive, and he was entirely
ignorant of the means they take to prevent
the access of supernumerary queens.

After ascertaining that workers in no si-
tuation sting such supernumerary females, we
were curious to learn how a stranger queen
would be received in a hive deprived of the
reigning one. To elucidate this matter, we
made numerous experiments, the detail of
which would protract this letter too much ;
therefore I shall relate only the principal

results. I

Bees are not immediately aware of the j
removal of their queen ; their labours ar^

A STEANGER QUEEN. 105

I

1 uninterrupted ; they watch over the young,

1 and perform the whole of their ordinary oc-
I cupations. But, in a few hours, agitation
ensues ; all appears a scene of tumult in the
hive. A singular humming is heard ; the
bees desert their young and rush over the
■ surface of the combs with delirious impetuo-
sity. Then they discover that their queen
is no longer among them. But how do they
ascertain it? How do the bees on the sur-
i| face of one comb discover that the queen is
|l or is not on the next comb ? It treating of
) another characteristic of these insects, you
have yourself. Sir, proposed the same ques-
tion. I am incapable of answering it in-
deed ; but I have collected some facts, that
may perhaps facilitate elucidation of the
mystery.

I cannot doubt that the agitation arises
from the workers having lost their queen ;
for on restoring her, tranquillity is instantly
regained among them ; and, what is very
singular, they recognise her : you must in-
terpret this expression strictly. Substitution
of another queen is not attended with the
same effect, if she is introduced into the
I hive within the first twelve liours after re-
I moval of the reigning one. Here the agi-

106

S^UBSTITUTIOX OF

tation continues; and the bees treat the
stranger just as they do when the presence of
their own leaves them nothing to desire.
They surround, seize, and keep her a very
long time captive, in an impenetrable clus-
ter ; and she commonly dies either from hun-
ger or privation of air.

If eighteen hours elapse before substitu-
tion of a stranger queen for the native one
removed, she is treated at first in the same
manner, but the bees leave her sooner, nor
is the surrounding cluster so close; they
gradually disperse ; and the queen is at last
liberated. She m6ves languidly; and some-
times expires in a few minutes. However,
some queens have escaped in good health
from an imprisonment of seventeen hours ;
and ended with reigning in the hives w’here
they had been originally ill received.

If, before substituting the stranger queen,
twenty-four hours elapse, slie will be well
received, and reign from the moment of her
introduction into the hive. Here I speak
of the good reception given to a queen after
an interregnum of twenty-four hours. But
as this w'ord reception is very indefinite, it
is proper to enter into some detail to ex->
plain the precise sense in which I use it.

A STRANGEll QUEEN.

107

Oil the 15th of August I introduced a
fertile queen, eleven months old, into a glass
hive. The bees had been twenty-four hours
deprived of their queen, and had alieady
begun the construction of twelve royal cells,
such as described in the preceding chaptei.
Immediately on placing this female stranger
on the comb, the workers near her touched
her with their antennae, and, passing their
trunks over every part of her body, they
' gave her honey. Then these gave place to
I others, tliat treated her exactly in the same
I manner. All vibrated their wings at once,

I and ranged themselves in a circle around
j their sovereign. Hence resulted a kind of
I agitation, which gradually communicated to
[ the workers situated on the same surface of
j the comb, and induced them to come and
I reconnoitre, in their turn, what was going

!on. They soon arrived ; and, having broke
through the circle formed by the first, ap-
proached the queen, touclied her with the
antennal, and gave her honey. After this
little ceremony they retired ; and, placing
ij themselves behind the others, enlarged the
i| circle, where they vibrated their wings,
*u and buzzed without tumult or disorder, and
■ 8) as if experiencing some very agreeable sen-

108

SUBSTITUTION OF

sation. The queen had not yet left the
spot where I had put her, but in a quarter
of an hour she began to move; when the
bees, far from offering any opposition, opened
the circle at that part to which she turned,
followed her, and surrounded her with a
guard. She was oppressed with the neces-
sity of laying, and dropped her eggs. Fi-
nally, after an abode of four hours, she
began to deposit male eggs in the cells she
met with.

While these events passed on the surface
of tlie comb where the queen stood, all w'as
quiet on the other side. There the work-
ers apparently were unconscious of a queen’s
arrival in the hive. They laboured with
great activity at the royal cells, as if igno-
rant that they no longer stood in need of
them : they watched over the royal w'orms,
supplied them with jelly and the like. But
the queen having at length come hither,
she was receiveil with the same respect that
slie had experienced from their companions
on the other side of the comb. They en-
compassed lier, gave her honey, and touch-
ed her with their antennas; and what
proved better that they treated her as a mo-
tlier, was their immediately desisting from

A STRANGER QUEEN.

109

work at the royal cells ; they removed the
worms, and devoured the food collected
; around them. From this moment the
queen was recognised by all her people, and
conducted herself in this new habitation as
if it had been her native hive.

These particulars will give a just idea of
the manner in which bees receive a stranger
queen. When they have had time to forget
their own, she is treated exactly as if she
> were their natural one, except that there is
* perhaps at first greater interest testified in
1 her, or more conspicuous demonstrations of
I it. I am conscious of the impropriety of
these expressions, but JM. de Reaumur in
some degree authorises them. He does
not scruple to say, that bees pay attention,
homage, and respect, to their queen, and
from his example similar words have escaped
most authors who treat on bees.

Twenty-four or thirty hours’ absence is
sufficient to make them forget their first

queen ; but I can hazard no conjecture on
the cause.

Before terminating this letter, which is
I full of combats arid disastrous scenes, per-
i haps I should give you an account of some

110

MASSACRE OF DRONES.

more pleasing and interesting facts relative
to their industry. However, to avoid re-
turning to duels and massacres, I shall here
subjoin my observations on the destruction
of the males.

You will remember. Sir, it is agreed by
all observers, that at a certain period of the
year, the workers expel and kill the drones.

M. de Reaumur speaks of these executions
as a horrible massacre. He does not ex-
pressly affirm, indeed, that he has himself
witnessed it, but what we have seen corres-
ponds so well with his account, that there
can be no doubt he has beheld its peculiari-
ties.

It is usually in the months of July and
August, that bees free themselves of the I
males. Then they are drove away and pur- '
sued to the inmost parts of the hive, where
they collect in numbers ; and as at the same
time we have found many dead onjthe ground
before the hives, it appeared indubitable
that after being expelled, the bees stung
them to death. Yet on the surface of the
comb, we do not see the sting used against ;
them ; for the bees are content to ])ursue
and drive them away. You remark this.

Sir, yourself in the new notes added to the.

MASSACRE OF DRONES.

Ill

Contemplation de la Nature; and you
j seem disposed to think, that the drones,

’ forced to retire to the extremity of the hive,

■ perish from hunger. Your conjectures were
extremely probable. Still it was possible
that the carnage might have taken place in
the bottom of the hive, and had been unob-
served, because that part is dark, and escapes
1 the observer’s eye.

To appreciate the justice of this suspi-
I cion, we thought of making the support of
\ the hive of glass, and of placing ourselves
j below to see what passed in the scene of ac-
j tion. Therefore, a glass table was construct-
i ed, on which were put six hives with swarms

!i of the same year ; and lying under it, we
I endeavoured to discover how the drones were
i destroyed. This contrivance succeeded to
I admiration. On the 4th of July, we saw
the workers actually massacre the males, in
1 the whole six swarms, at the same hour, and
I with the same peculiarities. The glass
I table was covered with bees full of anima-
tion, rushing upon the drones as they came
f from the bottom of the hive ; they seized
! them by the antennae, the limbs, and the
' wings, and after having dragged them about,
It or, so to speak, after quartering them, they

112

MASSACRE OF DRONES.

killed them by repeated stings directed be-
tween the rings of the belly. The moment
that this formidable weapon reached them,
was the last of their existence; they stretch-
ed their wings, and expired. At the same
time, as if the workers did not consider
them as dead as they appeared to us, they
still darted it so deep, that it could hardly be
withdrawn, and they were obliged to turn
round upon themselves before the stings
could be disengaged.

Next day, having resumed our former po-
sition, we witnessed new scenes of caniage.
During three hours, the bees furiously de-
stroyed the males. They had massacred all
their own on the preceding evening, but
now they attacked those which, driven from
the neighbouring hives, had taken refuge
amongst them. We saw them also tear some
remaining nymphs of this species from the
€ombs ; they greedily sucked all the fluid
from the abdomen, and then carried them
away. The following day no drones re-
mained in the hives.

These two observations seem to me deci-
sive. It is incontestible that nature has
charged the workers witli destruction of
the males at certain seasons of the year.

massacre of drones.

113

But what means does she use to excite their
fury against them ? This also is one of those
questions that I cannot pretend to answer,
j However, an observation I have made, may
i one day lead to solution of the problem.

I The males are never destroyed in hives de-
t prived of queens; on the contrary, while a
ij savage massacre prevails in other places, they
here find an asylum. They are tolerated
and fed, and many are seen even in the
I middle of January. They are also preserved

iin hives which, without a queen properly so
called, have some individuals of that species
that lay the eggs of males, and in those
whose half-fecundated queens, if I may use
the expression, propagate only drones.
Therefore, the massacre takes place in none
hut hives where the queens are completely
fertile, and it never begins until the season
of swarming is past.* — PiiEGNY, 28/A Au-
igust 1791.

i* Several interesting; observations, by an intelligent and
impartial Scotisli naturalist, the Rev. Mr. Dunbar, have
recently appeared, relative to the eombats of queens, and
some other topics in the history of bees. — T-

114

RECEPTION OF

LETTER VII.

SEOUEL OF EXPERIMENTS ON THE RE-
CEPTION OF A STRANGER QUEEN. M.
DE REAUMUR’S OBSERVATIONS ON THE
SUBJECT.

I HAVE frequently testified my admiration
of M. de Reaumur’s observations on bees.
I feel sensible pleasure in acknowledging,
that if I have truly made any progress in
the art of observation, I am indebted for it
to profound study of the works of this excel-
lent naturalist. In general his authority
has such weight, that I can scarcely trust
my own experiments when the results are
different from those obtained by him. Like-
wise, on finding myself in opposition to the
historian of bees, I repeat my experiments ;

A STRANGER QUEEN.

115

I vary the mode of conducting them ; I ex-
amine with the utmost caution all the cir-
cumstances that might mislead me ; and ne-
ver are my labours interrupted before acquir-
ing the moral certainty of avoiding error.
AVith the aid of these precautions, I have dis-
covered the justice of M. de Reaumur’s sug-
gestions, and I have a thousand times seen,
that if certain experiments seemed to com-
bat them, it was from incorrectness of execu-
tion. Yet I n'lust except some cases, where
my results have been constantly different
from his. Those respecting the reception
of a stranger queen substituted for the na-
tural one are of the number.

If, after removing the natural queen, a
stranger is immediately substituted, the
usurper is ill received. I never could suc-
ceed in making the bees adopt her, but by
allowing an interval of twenty or twenty-
four hours to elapse. Then they appeared to
have forgot their own queen ; and respect-
fully received any female put in her place.
M. de Reaumur, on the contrary, affirms,
that should the original queen be removed,
and another introduced, this new one will
he perfectly well received from the begin-
ning. As evidence of his assertion, he gives

116

RECEPTION OF

the detail of an experiment which ought to
be read in his work, for here I shall quote
only an extract of it.* He induced four or
five hundred bees to leave their native hive
and enter a glass box, containing a small
piece of comb towards the top. At first
they were in great agitation ; and, to pacify
or console them, he presented a new queen.
From that moment, the tumult ceased, and
the stranger queen was received with all
respect.

I do not dispute the truth of this experi-
ment ; but, in my opinion, it does not war-
rant the conclusion deduced by INI. de Reau-
mur. His apparatus removed the bees too
much from their natural condition, to al-
low him to judge of their instinct and dis-
positions. Ill other situations, he has him-
self observed these animals reduced to
small numbers lose their industry and ac-
tivity, and feebly continue their ordinary
labours. Thus their instinct is affected by
every operation that diminishes their num-
ber too much.f To render such an experi-
ment truly conclusive, it must be made in a

* Edit. 4to, tom. v. p. 258.

t The author elsewhere observes, tliat they even cease
to keep a guard at the entrance of the hives in such situa-
tions.— T.

A STRANGER QUEEN.

117

populous hive ; and on removing the native
qu6en, the stranger must be immediately sub*
stituted in her place. Had this been done,
I am quite persuaded, that M. de Reaumur
would have seen the bees imprison the
usurper, confine her at least twelve or fif-
teen hours among them, and frequently suf-
focate her : nor would he have witnessed any
favourable reception before an interval of
twenty-four hours after removal of the ori-
ginal queen. No variation has occurred in
my experiments regarding this fact. Their
number, and the attention bestowed on
them, make me presume that they merit
I your confidence.

M. de Reaumur, in another passage of
1 the same Memoir, affirms, that bees, hav-
i ing a queen they are satisfied with, are
j nevertheless disposed to give the best pos-^
^ sible reception to any female seekmg re^-
' fuge among them. In the preceding letter,
1 1 have related my experiments on this head :

: their issue has been very different from that
I of M. de Reaumur. I have proved that
i the workers never employ their stings against
i the queen ; but this cannot be called wel-
come reception of a stranger. Tliey retain
I her within their ranks, and seem to allow

118

RECEPTION OE

her liberty only when she prepares to coin*
bat the reigning queen. This observation
cannot be made except in the thinnest hives.
Those used by M. de Reaumur always had
two parallel combs at leasts which must have ;
prevented him from, remarking some very
important circumstances that influence the
conduct of w'orkers when supplied with se-
veral females. The first circles formed
around a stranger queen he has taken for
caresses ; and, from the little that his queen
•could advance in view between the combs,
it must have been impossible for him to ob-
serve that the circles, which were always
gradually contracting, ended in restraint of’
the females there inclosed. Had he used
thinner hives, he would have discovered that
what he supposed an indication of favourable
reception w^as the prelude of actual impri-
sonment.

I am reluctant to assert that IM. de Reau-
mur was deceived. Yet I cannot admit
that, on certain occasions, bees tolerate a
plurality of females in their hives. Neither
will the experiment on which this affirma-
tion rests be considered decisive. In the
month of December, he introduced a "stran-
ger queen into a glass hive in his cabinet, and

A STllA^’GEIl QUEEN.

119

confined her there, while the bees had not an
opportunity of going out. The stranger was
well received ; her presence awakened the
workers from their lethargic state, into which
they did not relapse ; she excited no car-
nage ; the number of dead bees on the board
of the hive did not sensibly increase ; and no
dead queens were found.

Before concluding any thing favourable to
the plurality of queens, it was necessary to
ascertain whether the native female was liv-
ing when the new one was introduced into
the hive: however the author neglected it ;
and it is very probable that the hive had
lost its queen, since the bees were languid,
and the presence of a stranger restored their
activity.

I trust. Sir, that you will pardon this
slight criticism. Far from industriously
seeking faults in our celebrated Reaumur,
I derive the greatest pleasure when my ob-
servations coincide with his, and more espe-
cially when my experiments justify his con-
jectures. But I think it proper to point
out those cases where the imperfections of
his hives have led him into error, and to ex-
plain from what causes I have not seen cer-
tain facts under the same aspect he did. I

120 RECEPTION OF A QUEEN.

feel particular anxiety to merit your confi-
dence ; and I am aware that the greatest ex-
ertions are necessary, when I have to com-
bat the historian of bees. I confide in your
judgment, and pray you to be assured of my
respect. — Pregny, 30/A August 1791.

THE QUEEN BEE IS OVIPAROUS. 121

LETTER VIII.

Is THE QUEEN OVIPAROUS? WHAT IN-
FLUENCE HAS THE SIZE OF THE CELLS,
WHEREIN THE EGGS ARE DEPOSITED,
ON THE BEES PRODUCED?— RESEARCH-
ES ON THE MODE OF SPINNING THE
COCCOONS.

IXn this letter I shall collect together some
isolatetl observations relative to various
points in the history of bees, concerning
^ which you wished me to engage.

, You were desirous that I should invest!.
I gate wliether the queen be really ovina-
t rous ; a question left undecided by M de
" Rf^iuinur, who says he never saw the
’'‘“<=’’0''. ™<1 is content with remark-

G

i

122

THE QUEEN BEE

ing, that worms are found in cells where
eggs have been deposited three days previ-
ously. If we resolve to attempt to seize the
moment of the worm quitting the egg, it is
necessary that our inquiries be carried be-
yond the interior of the hive : for there
the continual motion of the bees obscures
what passes at the bottom of the cells. The
egg must be taken out, presented to the
microscope, and every change attentively
watched. One other precaution is essential.
As a certain degree of heat is requisite for
hatching the worms, should the eggs be
too soon deprived of it they wither and pe-
rish. Therefore, the sole method to succeed
in observing the worm come out, consists in
watching the queen while she lays, in mark-
ing the egg so as to he recognised, and in re-
moving it from the hive to the microscope
only an hour or two before the three da)'s
elapse. The worm certainly will be hatched
provided it has been exposed as long as pos-
sible to the full degree of heat. Such is the
. course I have pursued; and the following
arc the results obtained.

In the month of August, we removed
several cells containing eggs that had been
three days deposited : we cut off the top of

IS OVIPAROUS.

123

the cells, and put the pyramidal bottom,
wliere the egg was fixed, on a glass slider.
Slight motions were soon perceptible in one of
the eggs. At first, we could not discern any
external organization : the worm was entire-
ly concealed from us by its pellicle, and we
then prepared to examine the egg with a
powerful magnifier. However, during the
intern al, it burst its surrounding membrane,
and cast off part of the envelope, w'hich was
torn and ragged on different parts of the
body, and more evidently so towards the last
rings. It curved and stretched itself alter-
nately with very lively action, and occupied
twenty minutes in casting off the spoil; when,
ceasing its exertions, it lay down, bent the
body, and seemed to take that rest which it
now required. An egg laid in a worker’s cell
produced this insect, which would have be-
come a worker itself.

We next directed our attention to the
moment when a male worm should be
hatched. All egg was exposed to the sun
on glass slider; and, with a good magnifier

ithm the transparent pellicle. This mem
brane was still entire, and the worm per-
fectly motionless. The two loiigitndiiinl

M

124

THE QUEEN BEE

lines of tracliso were visible on the surface,
with many ramifications proceeding from
them. We never lost siglit of the egg a
single instant, and now succeeded in ob-
serving the first motions of the worm. The
thick end alternately straightened and curv-
ed, and extended almost to the part where
the sharp extremity was fixed. These ex-
ertions burst the membrane, first on the up-
per part towards the head, then on the back,
and afterwards on all the rest of the body
successively : the ragged pellicle remained
in folds on different places of it, and then
fell off. Thus, it is beyond dispute, that
the queen is oviparous.

Some observers affirm, that the workers
watch on the eggs before the worms are
hatched ; and it is certain that, at whatever
time the hive is examined, we always see
them with the head and thorax inserted into
cells containing eggs, and remaining motion-
less several minutes in that position. It is im-
possible to discover what they do, for the inte-
rior of the cell is concealed by tbeir bodies ;
but it is very easily ascertained that, in this
attitude, they are doing nothing to the eggs.

If, at the moment the queen lays, her
eggs are put into a grated box, and deposited

IS OVIPAROUS.

12o

in a strange hive, where there is the neces-
sary degree of heat, the worms come out at
the usual time, just as if they had been left
in tlie cells. Thus no extraordinary aid or
attention is required for promoting their ex-
clusion.

When the workers penetrate the cells,
and remain fifteen or twenty minutes mo-
tionless, I have reason to believe that it is
only to repose from their labours. ]\Iy ob-
servations on this subject seem correct. You
know, Sir, that some irregular shaped cells
are frequently constructed on the panes of
the hive. Tliese, being glass on one side,
are exceedingly convenient to the observer,
since all that passes within is exposed. I
have often seen bees enter such cells when
nothing could attract them. They contained
neither eggs nor honey, nor did they need
farthei completion. Therefore the workers
repaired thither only to enjoy some repose.
Indeed, they were fifteen or twenty minutes
so perfectly motionless, that had not the di-
latation of the rings showed their respiration,
we might have concluded them dead. The
queen also sometimes penetrates the large
cells of the males, and continues very
long motionless. Her position prevents

126

MODE OF SPINNING

the bees from paying their full homage to
her, yet even then they do not fail to sur-
round her in a circle, and brush the part of
her belly that remains uncovered.

Drones do not enter the cells while re-
posing, but cluster together on the combs ;
and sometimes remain in this position eigh-
teen or twenty hours without the slightest
motion.

As it is important, in many experiments,
to know the exact time that the three spe-
cies of bees exist before assuming their ulti-
mate form, I shall here subjoin my own ob-
servations on the point.

The worm of workers passes three days
in the egg, five in the vermicular state, and
then the bees close up its cell with a wax
covering. It now begins spinning its coc-
coon, in which operation thirty-six hours are
consumed. In three days, it changes to a
nymph, and passes six in this form. It is
only on the twentieth day of its existence,
counting from the moment the egg is laid,
that the winged state is attained.

The royal worm also passes three days in
the egg, and is five a worm, when the bees
close its cell; and it immediately begins
spinning the coccoon, which occupies twenty-

THE COCCOON.

127

four hours. The tenth and eleventh day it
remains in complete repose, and even six-
teen hours of the twelfth. Then the trans-
formation to a nymph takes place ; in which
state four days and the third part of a fifth
are passed. Thus it is not before the six-
teenth day that the perfect state of queen is
attained.

The male passes three days in the egg,
six and a half as a worm, and metamor-
phoses into a winged animal on the twenty-
fourth day after the egg is laid.

Though the larvas of bees are apodal, they
are not condemned to remain absolutely mo-
tionless in their cells; for they can move in
a spiral direction. During the first three
days, this motion is so slow as scarcely to
be peiceptible, but it afterwards becomes

tmore evident. I have then observed them
perform two complete revolutions in an hour
I and three quarters. When the period of
f transformation arrives, they are only two
|! lines from the orifice of the cells. As their
« position is constantly the same, bent in
I an arc, it follows that the larvse in the

8 cells of workers and drones are perpcndicu-

I lar to the horizon, while those in the royal
‘ cells he horizontally. It might be thought.

i

128

MODE OF SPINNING

that the difference of position has much
influence on the increment of the various
larvae ; yet it has none. By reversing combs
containing common cells full of brood, I have
brought the worms into a horizontal posi-
tion ; but they were not injured. I have
also turned the royal cells, so that the worms
came into a vertical direction ; however their
increment was neither slower nor less perfect.

The mode of bees spinning their coccoons
has given me great surprise, and there I
have witnessed many new and interesting
facts. The worms both of workers and males
fabricate complete coccoons in their cells;
that is, close at both ends, and surrounding
the whole body. On the other hand, the
royal larvse spin imperfect coccoons, open
behind, and enveloping only the head, thorax,
and first ring of the abdomen. The dis-
covery of this difference, which at first may
seem trifling, has been the source of extreme
pleasure to me, for it evidently demonstrates
the admirable art with which nature con-
nects the various characteristics of the iu-
dustry of bees.

You recollect, 'Sir, the evidence that I
I gave you of the mutual aversion of I'

THE COCCOON.

129

queens, of the combats in which they en-
gage, and the animosity that leads them to
destroy one another. Of several royal nymphs
in a hive, the first transformed attacks the
rest, and stings them to death. But were
these nymphs enveloped in complete coc-
coons, slie could not accomplish it. Why ?
because the silk is of so close a texture, the
sting could not penetrate, or if it did, the
barbs would be retained by the meshes of
the coccoon, and the queen, unable to re-
tract it, would become the victim of her
own fury. Thus, that she may destroy
her rivals, it is necessary the last rings of
the body shall remain uncovered ; and the
royal nymphs must therefore form only im-
perfect coccoons. You will observe that the
last rings alone should be exposed, for the
sting can penetrate no other part : the head
and thorax are protected by connected shelly
plates, which it cannot pierce.

Hitherto philosophers have claimed our
admiration of nature for her care in pre-
serving and multiplying the species. But
from the facts I relate, we must now ad-
mire her precautions in exposing certain in-
dividuals to a mortal hazard.

130

MODE OF SPINNING

The detail on which I have just entered
clearly indicates the final cause of the open-
ing left by the royal worms in their coc-
coons; but it does not show w^hether they
do so in consequence of a particular instinct,
or whether the wideness of their cells pre-
vents them from stretching the thread up
to the top. This question interested me
very much, and the only method of deciding
it was to observe the worms while spinning,
which could not be done from their cells
being opaque. It then occurred to me to
dislodge them from their own habitations,
and introduce them into glass tubes, blown
in exact imitation of the different kind of
cells. The most difficult part of the ope-
ration consisted in extracting the worms and
introducing them here ; but my assistant
accomplished it with much address. He
opened several sealed royal cells, where we
knew the larvfe were about to be^in their
coccoons, and, taking them gently out, in-
troduced one into each of my glass cells
without the smallest injury.

They soon prepared to work ; and com-
menced by stretching the anterior "part of
the body in a straight line, while the other
was bent in a curve : thus forming an arc of

THE COCCOON.

131

which the longitudinal sides of the cells
f were tangents, and afforded two points oi
I support. The head was next conducted to
the different parts of the cell which it could
reach, and carpeted the surface with a thick
bed of silk. We remarked that the threads
r w'ere not carried from one side to another,
r which would have been impracticable, for
1 the wwms being obliged to support them-
i selves, had to keep the posterior rings curved ;

and the free and moveable part of the body
F. was not long enough to admit of the mouth
i reaching the sides diametrically opposite to
[i fix the threads to them. You will remem-
f her. Sir, that the royal cells are of a pyra-
!l midal form, with a wide base, and a long
b diminished top. They hang perpendicu-
I larly in the hive, the point downwards ; from
If which position the royal w^orm can be siis-

i tained in the cell, only when the curvature
I of the posterior part gains two points of
8 support : and it cannot obtain this support
I without resting on the lower part, or to-
» wards the extremity. Therefore, if attempt-

ii ing to stretch out and spin towards the wide
r end of the cell, it could not reach both sides
i; from being too distant. One' part would be

touched by its extremity, and the othc^by

132

MODE OF SPINNING

its back, consequently it would tumble
down. I have particularly ascertained the
fact in glass cells that were too large, and
of which the diameter was greater towards
the point than is usual in cells ; there they
were unable to sustain themselves.

These first experiments obviated the pro-
bability of any particular instinct iii the
royal worms. They proved, that if spinning
incomplete • coccooiis, it was because they
were forced to do so by the figure of their cells.
H owever, wishing to have evidence still
more direct, I put them into cylindrical glass
cells, or portions of glass tubes resembling
common cells, where I had the satisfaction
of seeing them spin complete coccoons, as
the worms of workers do. Lastly, I put
common worms into very wide cells, and
they left tfie coccoon open. It is demon-
strated, therefore, that the royal worms, and
those of workers, have the same instinct
and the same industry, or in other words,
when situated in similar circumstances, the
course they follow is the same. I may here
add, that the royal worms artificially lodged
in cells, where they can spin complete coc-
coons, undergo all their metamorphoses
equally well. Thus the necessity which na-

THE COCCOOX.

133

ture imposes on them, of leaving the coc-
coon open, is not on account of their incre-
ment; nor has it any other object than that
of exposing them to the certainty of perish-
ing by the wounds of their natural enemy ;
an observation new and truly singular.

I should relate my experiments on the
influence that the size of the cells has on
bees. It is to you, Sir, that I am indebted
for suggesting them.

As we sometimes found males smaller
than they ought to* be, and also queens more
diminutive than usual, it was desirable to
obtain a general explanation, to what degree
the cells, where bees pass the first period of
their existence, have an influence on their
size. With this view, you advised me to
remove all the combs composed of common
cells, and to leave those consisting of large
cells only. It was evident, if the common
eggs which the queen would lay in these
laige cells produced workers of larger size,

I we were bound to conclude that the size of
the cells had a decided influence on the
size of the bees. The first time I made
t le experiment, it did not succeed, be-
cause weevils having lodged in the hive
discouraged the bees. But I repeated it

134

INFLUENCE OF THE

afterwards, and the result was very remark-
able.

I removed the whole comb, consisting of
common cells, from one of my best glass
hives, and left that composed of the cells of
males alone : and, to avoid vacuities, I sup-
plied others of the same kind. This was in
June, the season most favourable to bees.
1 expected that the bees would have quickly
repaired the ravages produced by the oper-
ation in their dwelling; that they would
labour at the breaches, and unite the new
combs to the old. But I was very much
surprised to see that they did not begin to
work. Expecting the resumption of their
activity, I continued observing them several
days ; however, my hopes were disappoint-
ed. Their homage to the queen was not in-
terrupted indeed ; but except in this, their
conduct to her was quite different from what
it is usually; they clustered on the combs
without exciting any sensible heat. A
thermometer among them rose only to 81°,
though standing at 77 ° in the open air. In
a word, they appeared in a state of the
greatest despondency.

The queen herself, though very fertile, and
though she must have been oppressed by her

SIZE OF THE CELLS.

135

I

eggs, hesitated long before depositing them
in the large cells ; she chose rather to drop
them at random, than they should be laid in
those unsuitable. However, on the second
day, we found six that had been dejiosited
there with all regularity. The worms were
hatched three days afterwards, and then we
began to study their history. Notwithstand-
ing the bees provided them with food, they
did not carefully attend to it ^ yet I was in
hopes that they might be reared. I was
again disappointed ; for next morning the
whole worms had disappeared, and their cells
remained empty. Profound silence reigned
in the hive ; few bees left it, and these re-
turned without pellets on their limbs ; all
was cold and inanimate. To promote a lit-
tle motion, I thought of supplying the hive
ynth a comb, composed of small cells, full of
male brood of every age. The bees, after
twelve days obstinately abstaining from
working in wax, did not unite this comb to
their own. However, their industry was
awakened in a way that I had not antici-
pated. They removed the brood from it,
c eancd out the whole cells, and prepared
them for receiving new eggs. I cannot de-
einiiiie w.iether they expected the queen

136

INFLUENCE OF THE

to lay, but it is certain if they did so they
were not deceived. From this moment, she
no longer dropped her eggs ; but laid such a
number in the new comb, that we found five
or six together in the same cell. I then re- '
moved all the combs composed of large cells,
to substitute small cells in their place, an
operation which restored complete activity
among the bees.

The peculiarities of this experiment seem
worthy of attention. It proves that nature
does not allow the queen the choice of the
eggs she is to lay. It is ordained that, at a
certain time of the year, she shall produce
those of males, and at another time the eggs
of workers; an order which cannot be in-
verted. We have seen that another fact
led me to the same consequence; and as
that was extremely important, I am delight-
ed to have it confirmed by a new observa-
tion. Let me repeat, therefore, that the
eggs are not indiscriminately mixed in the
ovaries of the queen, but arranged so, that,
at a particular season, she can lay only a cer-
tain kind. Thus, it would be vain at that
time of the year, when the queen should lay
the eggs of workers, to attempt forcing her
to lay male eggs, by filling the hives with

SIZE OF THE CELLS.

137

large cells ; for, by the experiment just de-
scribed, we learn, that she will rather drop
the workers’ eggs by chance than deposit
them in an unsuitable place ; and that she
will not lay the eggs of males. I cannot
yield to the pleasure of allowing this queen
discernment or foresight, for. I observe a
kind of inconsistency in her conduct. If
she refused to lay the eggs of workers in
large cells, because nature had instructed
her that their size was neither proportioned
to the size nor necessities of common worms,
would not she also have been instructed to
abstain from laying several eggs in one cell ?
It seems much easier to rear a worker’s worm
in a large cell, than to rear several of the
same species in a small one. Therefore, the
supposed discrimination of the queen bee is
not very conspicuous. Here the most pro-
minent feature of industry appears in the
common bees. When supplied with a comb
of small cells, full of male brood, their ac-
tivity was awakened : but instead of bestow-
ing the necessary care on this brood, as they
•' would have done in every other situation,

I they destroyed the whole nymphs and larvae,

I and cleaned out their cells, that the queen!

3 now oppressed with the necessity of laying,

138

INFLUENCE OF THE

might suffer no impediment to depositing
her eggs. Could we allow them either rea-
son or reflection, this would be an interest-
ing proof of their affection for her.

The experiment, now detailed at length,
not having fulfilled my object in determin-
ing the influence of the size of the cells on
that of the worms, I devised another which
proved more successful.

Having selected a comb composed of large
cells, containing the eggs and worms of
males, I removed all the worms from their
farina; and my assistant substituting those
of workers a day old in their place, he in-
troduced it into a hive that had the queen.
The bees did not abandon these substituted
worms ; they covered their cells with a top
almost flat, of a kind quite different from
what is put on the cells of males; which
shows that they were well aware that,
though inhabiting large cells, they were not
males. This comb remained eight days in
the hive, counting from the time that the
cells were sealed. I then removed it to ex-
amine the included nymphs, which proved
those of workers in different stages of ad-
vancement ; but, as to size and figure, they
perfectly resembled what had grown in the

SIZE OF THE CELLS.

139

smallest cells. Thence I concluclecl, that the
larvse of workers do not acquire greater size
in large than in small cells. Although the
experiment was made only once, it seems
decisive. Nature has appropriated cells of
certain dimensions for the worms of workers
while in their vermicular state ; undoubted-
ly she has ordained that their organs shall
be fully expanded, and as there is sufficient
space for that purpose, more would be use-
less. Their evolution ought to be no greater
in the most spacious cells than in those appro-
priated for them. If some cells smaller than
common ones are found in combs, and the
eggs of workers are deposited there, the size
of the bees probably will be less than that
of common workers, because they have been
cramped in the cells ; but it does not ensue,
that a larger cell will admit of them grow-
ing to a greater size.

The effect produced on the size of drones

!by the size of the cells their worms inhabit,
may serve as a rule for what should happen
to the larvae of workers in the same circum-
stances. The large cells of males are suffi-
ciently capacious for the full expansion of
•their organs. Thus, although reared in cells
1 of still greater capacity, they will grow no

I

140

INFLUENCE OF THE

larger than common drones. We have had
evidence of this in those produced by queens
whose fecundation has been retarded. You
will remember, Sir, that they sometimes lay
male eggs in the royal cells. Now, the
males proceeding from these eggs, and rear-
ed in cells much more spacious than nature
has appropriated for them, are no larger than
common males. Therefore it is certain,
that whatever be the size of the cells where
the worms acquire their increment, the bees
will attain no greater size than is peculiar
to their species. But if, in their primary
form, they live in cells smaller than they
should be, as their growth then will be
checked, they will not attain the usual size,
of which there is proof in the following ex-
periment.

I had one comb consisting of the cells of
large drones, and another with those of
workers, which also served for the male
worms. Of these, my assistant took a cer-
tain number from the smallest cells, and de-
posited them on a quantity of food purposely
prepared in the large ones ; and in return he
introduced into the small eells the worms
that had been hatched in the others ; then
committing both to the care of the workers

St^E OF THE CELLS.

141

in a hive where the queen laid the eggs of
males only. The bees were not affected by
this change; they took equal care of the
worms ; and when the period of metamor-
phosis arrived, gave both kinds of cells that
convex covering usually put on those of
males. Eight days afterwards we removed
the combs, and found, as I had expected,
nymphs of large males in the large cells,
and those of small males in the small
ones.

On your suggestion I carefully made a-
nother experiment, but it met with an un-
foreseen obstacle. To appreciate the influ-
ence of the royal food on the evolution of
the worms, you desired me to supply the
worm of a worker in a common cell with it.
Twice I have attempted this operation with-
out success c nor do I think it can ever suc-
ceed. If bees get the charge of worms in
whose cells the royal food is deposited, and
if at the same time they have a queen, they
soon remove the worms, and greedily devour
the food. When, on the contrary, they are
depiivcd of a queen, they change the com-
mon cells containing worms into cells of the
llaigest kind, and then the worms will be in-
fallibly converted to queens.

k

142

INFLUENCE OF CELLS.

But there is another situation where w’e
can judge of the influence of the royal food
administered to worms in common cells. I
have spoke at great length in my letter on
the existence of fertile workers. You can-
not forget. Sir, that the evolution of their
sexual organs is owing to the reception of
some particles of royal jelly, while in the
vermicular form : but, for want of new ob-
servations, I must refer you to what is pre-
viously said on the subject. — ^Pregny, 4//i
September 1791.

FOPtMATION OF SWARMS.

143

LETTER IX.

ON THE FORMATION OF SWARMS.

I CAN add but few facts to the information
which M. de Reaumur has communicated
relative to swarms.

A young queen, according to this celebrat-
ed naturalist, is always or almost always at
the head of a swarm ; but he does not as-
sert the fact positively, and had some doubts
on the subject. ^ “ Is it certain,” says he,
“ as we have hitherto supposed, in coinci-
dence with all who have treated of bees, that
the new colony is uniformly conducted by a
young mother? May not the old mother
be disgusted with her habitation ? or may
she not be influenced by some particular

144 FOUMATION OF SWARMS.

circumstances, to abandon all her possessions
to the young female ? I wish it had been
in my power to solve this question otherwise
than by mere probabilities, and that some
misfortune had not befallen all the bees
whose queen I had marked red on the thorax.”
These expressions seem to indicate, that
M. de Reaumur suspected that the old
queens sometimes conduct the young swarms.
From tlie following details, you will ob-
serve that his conjectures are fully justified.

In the course of spring and summer, the
same hive may throw several swarms. The
old queen is always at the head of the first
colony ; the others are conducted by young
queens. Such is the fact which I shall now
prove; and the peculiarities attending it
shall be also related.

But previous to entering on this subject,
I shall repeat what aLread)'’ has been fre-
quently observed, that the leaf or flat hives
are indispensable iu studying the industry
and instinct of bees. When these insects
are left at liberty to construct several rows
of parallel combs, we are precluded from ob- •
serving what continually passes between
them, or they must be dislodged by w'ater I
or smoke, for the purpose of examining their

FORMATION OF SWARMS. 145

structures ; a Violent proceeding, which
has a material influence on their instinct,
and consequently exposes an observer to the
risk of supposing simple accidents perma-
nent laws.

I now proceed to experiments proving
that an old queen always conducts the first
swarm.

One of my glass hives consisted of three pa-
rallel combs, placed in frames opening like the
leaves of a hook. It was well peopled, and
abundantly provided with honey and w^ax ^
and with brood, of every different age. On
the fifth of J\Iay 1788, I removed its queen,
and, on the sixth, transferred all the bees
from another hive into it, with a fertile
queen at least a year old. They entered
easily and without fighting, and were in ge-
neial well received. The old inhabitants of
the hive, which, since privation of their
queen, had begun twelve royal cells, also
gave the fertile queen a good reception-
they presented her with honey, and sur-
rounded her in regular circles. However
there was a little agitation in the evening’
though confined to the surface of the comb
where we had put the queen, and which she

II

146 FOEMATION OF SWARMS.

had not quitted, for all was perfectly quiet
on the other side.

On the inorniug of the seventh, the bees
had destroyed the twelve royal cells, but,
independent of that, order continued preva-
lent in the hive ; the queen laid the eggs of
males in the large cells, and those of work-
ers in the small ones, respectively.

Towards the twelfth, we found the bees
occupied in constructing twenty-two royal
cells, of the same species described by M.
de Reaumur, that is, the bases not in the
plane of the comb, but appended perpendi-
cularly by pedicles or stalks of different
length, like stalactites, on the edge of the
passage made by the bees through their
combs. They bore considerable resemblance
to the cup of an acorn, the longest being
only about two lines and a half in depth
from the bottom to the orifice.

On the thirteenth, the queen seemed al-
ready more slender than when introduced
into the hive ; however she still laid some
eggs, both in common cells and in those of
males. We also surprised her this day laying
in a royal cell : she first dislodged the work-
er there employed, by pushing it away with

FORMATION OF SWARMS. 147

her head, and then supported herself by the
adjoining cells while depositing the egg.

On the fifteenth, the size of the queen was
yet farther reduced ; the bees continued their
attention to the royal cells, which were all
unequally advanced, some to the height of
three or four lines, while others were already
an inch long ; thus proving that the queen
had not laid in the whole at the same time.

At the moment when least expected, the
hive swarmed on the nineteenth ; we were
warned of it by the noise in the air; and hast-
ened to collect and put the bees into a hive
purposely prepared. Though we had over-
looked the facts attending the departure of
this swarm, the object of the experiment was
fulfilled ; for, on examination of all the beeS,
we were convinced they had been conducted
by the old queen, by her that we introduced
on the sixth of the month, and which had
been deprived of one of the antennae. Ob-
serve, there was no other queen in this colo-
ny. In the hive she had left we found se-
ven royal cells close at the top, but open at
the side, and quite empty. Eleven more
were sealed, and some others newly begun •
no queen remained in the hive ’

148 rORMATION OF SWARMS.

The new swarm next became the object of
our attention : we observed it during the
rest of the year, in winter and the subse-
quent spring ; and, in April, we had the sa-
tisfaction of seeing another depart with the
same queen at its head that had conducted
the former one in May of the preceding year.

You will remark, Sir, that this experiment

is positive. We put an old queen in a glass

hive while laying the eggs of males : the bees

received her well, and at that time began to

construct roval cells ; next she laid in one of
✓

them before us; and in the last place led
forth the swarm.

We have repeated the same experiment
several times with equal success. Thus it
appears incontestible, that the old queen al-
ways conducts the first swarm, but never
quits the hive before depositing eggs in the
royal cells, from which other queens will
proceed after her departure. These cells
are prepared by the bees only while the queen
lays male eggs ; which is attended by a re-
markable fiict, namely, that after this laying
terminates, her belly being considerably di-
minished, she can easily fly, whereas, it is
previously so heavy that she can hardly drag

FORMATION OF SWARMS. 149

it along. Therefore it is necessary she
should lay, in order to be in a state for un-
dertaking her journey, which sometimes may
be very long.

But this single condition is not enough.
It is also requisite that the bees be very nu-
merous ; they should be even superabun-
dant, and it may be said that they are aware
of it: for, if the hive is thin, no royal cells
are constructed when the male eggs are laid,
which is done solely at the period that the
queen is able to conduct a colony. This fact
wvas proved by the following experiment on
a large scale.

On the third of May 1788, we divided
each of eighteen hives, whose queens were
about a year old, into two portions. Thus
each portion of the hivqs had but half the
bees that were originally there. Eighteen
halves wanted queens, but the other eighteen
had very fertile ones. They soon began to
lay the eggs of males; but the bees being few,
they did not construct royal cells, and none
ol the hives threw a swarm. — Therefore, if
the hive containing the old queen is not
veiy populous, she remains in it until the
subsequent spring ; and if the population is
then sulficicnt, royal cells will be construct-

150 FORMATION OF SWARMS.

eel ; she will begin to lay male eggs, and,
after depositing them, will issue forth at the
head of a colony, before the young queens
are produced.

Such is a brief abstract of my observa-
tions on swarms conducted by old queens.
You must excuse the long detail on which
I am about to enter, coneerniug the history
of the royal cells left by the queen in the
hive. Every thing relative to this part of
the history of bees has been hitherto very
obscure. A long course of observations,
protracted even during several years, was
necessary to remove, in some degree, the
veil concealing the mystery. I have been
indemnified for the trouble, indeed, by the
pleasure of seeing my experiments recipro-
cally confirmed ; but, considering the assi-
duity required in these researches, they were
truly very laborious.

Having established in 1788 and 1789,
that queens a year old conducted the first
swarm, and that they left worms or nymphs
in the hive, which were to transform into
queens in their turn, I endeavoured, in
1790, to profit by the goodness of the spring,
to study all that related to these young

FORMATION OF SWARMS. 151

queens ; and I shall now extract the chief
experiments from my Journal.

On the fourteenth of May, we transferred
two portions of bees, from their straw hives,
into a very thin large glass hive ; and allow-
ed them only one queen of the preceding
year, which had already commenced laying
in her native hive. We introduced her on
the fifteenth. She was very fertile : the
bees received her well, and she quickly be-
gan to lay in small and large cells alter-
nately.

On the twentieth, we saw the fabrica-
tion of twelve royal cells, all on the edges
of the communications, or passages through
the combs, and shaped liked stalactites.

Ten were much, though unequally en-
larged on the twenty-seventh ; but none so
long as when the worms are hatched.

On the twenty-eighth, previous to which
the queen had not ceased laying, her belly
was very slender, and she began to exhibit
signs of agitation. Her motion soon be-
came more lively, yet she still continued
examining the cells as when about to lay ;
sometimes introducing half her belly, hut
suddenly withdrawing it, without having
done so. At other times, inserting it no

152 FOEMATION OF SWARMS.

farther, she deposited an egg, which was
not fixed by an end to the bottom of the
cell, hut rested irregularly on one side of
the hexagon. The queen produced no dis-
tinct sound in her course, nor did we hear
any thing different from the ordinal'}’’ hum-
ming of bees. She sometimes stopped while
passing over those in her way ; and the bees
also stopped as if to consider her. Advancing
briskly, they struck her with their antennae,
and mounting on her back, she proceeded
carrying several above her. None gave her
honey, but she took it voluntarily in passing
over the cells. She was no longer surround-
ed and enclosed by bees in regular circles.
The first that were aroused by her motions,
followed her running in the same manner,
and in their passage, excited those still
tranquil on the combs. The track of the
queen was evident after she left it, by the
agitation created, which was never after-
wards quelled.

She had soon visited every part of the
hive, and disseminated a general sensation :
if some places still remained tranquil, the
bees in agitation arrived, and communicated
their disorder. The queen ceased to depo- -
sit her eggs in cells ; she dropped them at

FORMATION OF SM'ARMS. 153

random: nor did the bees continue to watch
over the young ; they ran through the hive
in every different direction ; even those re-
turning from the fields before the confusion
came to its height, no sooner entered their
dwelling than they participated in these tu-
multuous motions. Neglecting to free them-
selves of the waxen pellets on their limbs,
they hurried blindly about. At last the
whole rushed precipitately towards the out-
lets of the hive, and the queen along with
them.

As it was of much consequence to see the
formation of new swarms in this hive, I
wished it still to continue very populous,
and on that account removed the queen at
the moment she came out, that the bees
might not fly too far, and likewise that they
might return. In fact, after losing their
female, they did return to the hive. To
increase the population still more, I added
another swarm, which had come from a
straw hive on the same morning, and re-
moved its queen also.

Although these facts were clear, and ap-
parently not susceptible of any error, I was
particularly earnest to discover whether old
queens always followed a similar course.

154 FORMATION OF SWARMS.

Therefore, on the twenty-ninth, I replaced
in her glass hive the queen, a year old,
hitherto the subject of my experiments, and
which had just begun to lay the eggs of
males. On the same day, finding one of
the royal cells, left by the preceding queen,
larger than the rest, we supposed from its
length that the included worm was two days
old ; that it W'as hatched on the twenty-
seventh, and that the egg had been laid on
the twenty-fourth. On the thirtieth the
queen laid a great deal in the large and small
cells alternately. Then and on the two fol-
lowing days, the bees enlarged several royal
cells, but unequally, which proved that they
included larvae of different ages. One was
closed on the first of June, another on the
second, and some new ones were also com-
menced. All was perfectly quiet at eleven
in the morning; but, at mid-day, the queen,
from the utmost tranquillity, became evi-
dently agitated ; and her disorder insensibly
communicated to the workers in every part
of their dwelling. In a few minutes they
precipitately crowded to the outlets, and,
along with the queen left the hive. After
they had settled on the branch of a neigh-
bouring tree, I sought for the queen ; thinks

FORMATION OF SWARMS. 155

iug that, by removing her, the bees might
return ; which actually ensued. Their first
care seemed devoted to seeking the female;
they were still in great agitation, but gra-
dually calmed ; and in three hours complete
tranquillity was restored.

On the third they had resumed their usual
occupations : they attended to the young,
worked within the open royal cells, and also
watched on those that were shut. They made
a waved work on them, which is not done
by applying wax bands, but by removing
wax from the surface. Towards the top this
waved work is almost imperceptible ; it be-
comes deeper above, and the workers exca-
vate it still more from thence to the base of
the pyramid. The cell, when once shut,
also becomes thinner ; and is so mueli redu-
ced, immediately preceding the queen’s meta-
morphosis from a nymph, that all the motions
within are perceptible through the thin co-
vering of wax on which the waved work is
founded. It is a very remarkable circum-
stance, that in making the cells thinner,
from the moment they are closed, the bees
i know to regulate their labour, so^that it tcr-
D minates only when the nymph is ready to
t undeigo its last metamorphosis.

156 FORMATION OF SWARMS.

On the seventh day the coccoou is almost
completely unwaxed, if I may use the ex-
jiression, at the part next to the head and
thorax of the queen. This operation facili-
tates her exit ; for she has nothing to do but
to cut the silk forming the coccoon. Most
probably the object here is to promote eva-
poration of the superabundant fluids of the
nymph. I have engaged in some direct ex-
periments to ascertain the fact, hut they are
yet unfinished. A third royal cell was
closed by the bees on the same day, the
third of June, twenty-four hours after clos-
ing the second, and the like followed with
respect to other royal cells successively, dur-
ing the subsequent days.

Every moment of the seventh we expected
the queen to leave the royal cel], which was
shut on the thirtieth of May. On the pre-
ceding evening her period of seven days had
elapsed : and the waving of her cell was so
deep, that what passed within was pretty
perceptible. We could discern that the silk
of the coccoon was cut circularly, a line and
a half from the extremity ; but the bees be-
ing unwilling that she should yet quit her
cell, had soldered the covering to it with
some particles of wax. Mliat seemed most

rOUMATION OF SWARMS. 157

singular was, this female emitting a very
distinct sound or clacking, from her prison,
which became still more audible in the even-
ing, and even consisted of several monoto-
nous notes in rapid succession.

The same sound issued from the second
royal cell on the eighth. Several bees kept
guard around each of the royal cells.

The first opened on the ninth. Its
young queen was lively, slender, and of a
brown colour. l\ow, we understood why
j bees retain the females captive in their cells
j after the period for transformation has elaps-
I ed ; it is, that they may be able to fly the
I instant they are liberated. The new queen
( occupied all our attention. When she ap-
i preached the other royal cells, the bees on
I guard pulled, bit her, and chased her away ;
( their irritation seemed to be greatly ex-
I cited against her, and she enjoyed tranquil-
f lity only when at a considerable distance
^ fiom these cells. Ihis proceeding was fre-
I quently repeated throughout the day. She
f- twice emitted the sound; standing while
I doing so with her thorax against a comb, and
! her wings crossed on her back, in motion,
I but without being unfolded or farther open-
) cd. Whatever might be the cause of her

158

FORMATION OF SWARMS.

assuming this attitude, the bees were affect-
ed by it ; all hung down their heads, and
remained motionless.

The hive presented the same appearances
on the following day. Twenty-three royal
cells yet remained, assiduously guarded by
a great many bees. When the queen ap-
proached, the guards became agitated,
crowded her on all sides, bit, and common-
ly drove her away; sometimes when, in
these circumstances, she emitted her sound, j
assuming the position just described, from ;
that moment the bees became motionless.

The queen confined in the second cell,
which she had not yet left, was heard to
pipe several times;* and we accidentally
discovered how the bees fed her. By at-
tentive inspection, a small aperture was dis-
cernible in the end of the coccoon, which
she had cut, in order to escape, and which
her guards had agaiu covered with wax to
retain her still longer. She alternately
thrust her proboscis through the cleft, and |
retracted it unperceived by the bees, until at j

* It is not evident whether the author means that she
emitted a sound similar to that just described ; hut a note>
commonly called piping, is heard from some organ yet uu*
discovered. — T.

FORMATION OF SWARMS. 159

length attracting their attention, one of them
came to apply its own to that of the captive
queen, and then gave way to others also ap-
proaching her with honey. When satisfied,
she retracted her trunk, and the bees once
more closed up the opening with wax.

The queen this day between twelve and
one became extremely agitated. The royal
cells had midtiplied very much; she could
go no where without finding them, and on
her approach she was very roughly treated.
Then she fled, but to obtain no better recep-
i tion. At last these things agitated the
I bees ; they precipitately rushed through the
I outlets of the hive, and settled on a tree in
[ the garden. It singularly happened that
I the queen was unable to follow them and
1, herself head the swarm. She had attempted
3 to pass between two royal cells, before they
i were abandoned by the bees guarding them,
t' but she was so confined and maltreated as to
>• be incapable of moving ; we transferred her
ti into a separate hive prepared for a particu-
( lar experiment. The bees, which had clus-
'1 tered on a branch, soon discovered tlieir
n queen was not present, and returned to the
i hive of their own accord. Such is an ac-
'( count of the second colony of this hive.

160 FORMATION OF SWARMS.

We were extremely solicitous to ascer-
tain what would become of the other royal
cells. Four of the close ones had attained
complete maturity, and the queens woidd
have left them had not the bees prevented
it. They were not open either previous to
the agitation of the colony, or at the mo-
ment of swarming.

None of the queens were at liberty on
the eleventh. The second should have
transformed on the eighth; thus she had
been three days confined, a longer period
than the first which had occasioned the for-
mation of the swarm. We could not dis-
cover what caused the difference in their
captivity.

On the twelfth, the queen was at last li-
berated, as we found her in the hive. She
had been treated exactly as her predecessor;
the bees allowed her to rest in quiet when
distant from the royal cells, but tormented
her cruelly when she approached them.
We watched this queen a long time; but
not aware that she would lead out a colony,
we left the hive for a few hours, and return-
ing at mid-day, were greatly surprised to
find it almost totally deserted. During our
absence, it had thrown a prodigious swarm,

FORMATION OF SWARMS. 161

which still clustered on the branch of a
neighbouring tree. W e also saw with

astonishment the third cell open, and its
top connected to it as by a hinge. In all
probability the captive queen, profiting by the
confusion that preceded the swarming, had
escaped. Thus there was no doubt of both
queens being in the swarm. So it proved ;
and we removed them, that the bees might
return to the hive, which they did very soon
afterwards.

While we were occupied in this oper-
ation, the fourth captive queen left her pri-
son, and the bees found her on returning.
At first they were very much agitated, but
calmed towards the evening, and resumed
their wonted labours. They formed a strict
1 watch around^ eighteen royal cells now re-
maining to be guarded, and took great care
to repulse the queen whenever she attempt-
ed to approach them.

The fifth queen left her cell at ten at
night ; therefore, two queens were now in
the hive. They immediately began fight-
ing, but came to disengage themselves from
each other. However, they fought several
times during the night without any thing de-
cisive. Next day, the thirteenth, we witness-

162 FORMATION OF SWARMS.

eel the death of one, which fell by the wounds
of her enemy. This duel was quite similar
to what is said of the combats of queens.

The victorious queen now presented a
very singular spectacle. She approached a
royal cell, and took this moment to utter
the sound, and assume that posture, which
strike the bees motionless. For some mi-
nutes, we conceived that, profiting by the
dread exhibited by the workers on guard,
she would open it, and destroy the young
female; also she prejiared to mount the cell;
but in doing so she ceased the sound, and
quitted the attitude paralysing the bees.
The guardians of the cell instantly took
courage; and, by means of tormenting and
biting the queen, drove her away.

On the fourteenth, the sixth young queen
appeared, and the hive threw a swarm, with
all the concomitant disorder before describ-
ed. The agitation was so considerable, that
a sufficient number of bees did not remain
to guard the royal cells, whence several of
the imprisoned queens were enabled to make
their escape. Three were in the cluster
formed by the swarm, and other three re-
mained in the hive. We removed those
that had led the colony, to force the bees to

FORMATION OF S\VARMS. 163

return. They entered the hive, resumed
their post around the royal cells, and mal-
treated the queens when approaching them.

A duel took place in the night of the fif-
teenth', in which one queen fell. We found
her dead next morning before the hive ; but
three still remained, as one had been hatch-
ed during night. Next morning also we
saw a duel. Both combatants were extreme-
ly agitated, either with the desire of fight-
ing, or by the treatment of the bees when
they came near the royal cells. Their agi-
tation quickly communicated to the rest of
the colony, whicli at mid-day departed impe-
tuously along with the two females. This
was the fifth swarm that had left the hive
between the thirtieth of May and fifteenth
of .June. On the sixteenth, a sixth swarm
cast, of which I shall give you no account,
as it showed nothing new.

Unfortunately we lost this last, which was
a very strong swarm ; the bees flew out of
sight, and never could be found. The hive
was now very thinly inhabited. Only the
few bees that had not participated in the
general agitation remained, together with
those that returned from the fields after the
swarm had departed. The royal cells, thcre^

164

FORMATION OF SWARMS.

fore, were slenderly guarded ; the queens es-
caped from them, and engaged in several
combats, until the throne pertained to one
proving the most successful.

Notwithstanding the victories of this
queen, she was treated with great indiffer-
ence from the sixteenth to the nineteenth,
that is, during the three days that she pre-
served her virginity. At length, having
gone forth to seek the males, she returned
with all the external signs of fecundation,
and was thenceforth received with every
mark of respect. Her first eggs were laid
forty-six hours after fecundation.

Behold, Sir, a simple and faithful account
of my observations on the formation of
swarms. That the narrative might be the
more connected, I have avoided interrupting
it by the detail of several particular experi-
ments which I made at the same time, with
the view of elucidating various obscure
points of their history. These shall be the
subject of future letters : for although I
have said so much, I hope still to interest
you. — Pregny, Qth Septcinber 1791.

P. S. — In revising this letter, I find I
Lave neglected to anticipate an objection

FORMATION OF SWARMS. l65

that may embarrass my readers, and which
ought to be obviated.

After the first five swarms had thrown,
I always returned the bees to the hive :
therefore, it is not surprising that it was
continually stocked so sufficiently that each
colony was numerous. But matters are
otheiwise in the natural state i the bees com-
posing a swarm do not return to their dwell-
ing ; and it will be asked, AVhat resource
enables a common hive to swarm three or four
times without being too much weakened ?

I cannot lessen the difficulty. I have
observed that the agitation, which precedes
swarming, is often so considerable, that most
of the bees quit the hive ; and in that case
we cannot well comprehend how, in three
or four days afterwards, it can be in a state
to send out another colony equally strong.

But remark, in the first place, that the
queen leaves a prodigious quantity of work-
ers brood, which soon transforms to bees ;
and in this way the population sometimes
becomes almost as great after swarming as
before it.

Thus the hive is perfectly capable of af-
tording a second colony without being too
much impaired. The third and fourth

166 FORMATION OF SWARMS.

swarm weaken it more sensibly ; but the in-
habitants always remain in sufficient num-
ber to preserve the course of their labours
uninterrupted ; and the losses are soon re-
placed by the great fecundity of the queen,
as she lays above an hundred eggs a day.

If, in some cases, the agitation of swarm-
ing is so great, that all the bees participate
in it, and leave the hive, the desertion lasts
but for a moment. The hive throws only
during the finest part of the day, and it is
then that the bees are ranging through the
country. Those that are out, therefore, can-
not participate in the agitation ; having re-
turned to the hive, they quietly resume their
labours ; and their number is not small, for,
when the weather is fine, at least a third are
employed in the fields at once.

Even in the most embarrassing case,
namely, where the whole bees desert the
hive, it does not follow that all those endea-
vouring to depart become members of the
new colony. When this agitation or deli-
rium seizes them, the whole rush forward
and accumulate towards the entrance of the
hive, and are heated in such a manner that
they perspire copiously. Those near the
bottom, and supporting the weight of all

FORMATION OF SWARMS. 167

the rest, seem perfectly drenched; their wings
grow moist ; they are incapable of flight ;
and even when able to escape, they advance
no farthei than the board of the hive, and
soon return.

Those that have lately left their cells re-
main behind the swarm, still feeble, for they
could not support themselves in flight. Here
then are also many recruits to people what

we should have thought a deserted habita-
tion.

168 FORMATION OF SM'ARMS.

LETTER X.

THE SAME SUBJECT CONTINUED.

To preserve greater regularity in conti-
nuing the history of swarms, I think it
proper to recapitulate, in a few words, the
principal points of the preceding letter, and
in expatiating on each, to give the result of
some new experiments, respecting which I
still have been silent.

In the first place, If^ on the return of
spring, tee examine a hive ixell peopled, .
and governed by a fe?'tile queen, we shall
see her lay a prodigious number of the.
eggs of males in the coui'se of May, and i
the workei's will choose that period for
constructing several royal cells of the kind
described by M. de Reaumur. Snch is.

FORMATION OF SWARMS.

169

the result of several long continued observa-
tions, among which there has not been the
slightest variation, and I cannot hesitate in
announcing it as a fact demonstrated. How-
ever, I should here add a necessary explana-
tion. Before a queen commences her great
laying of the eggs of males, she must be ele-
ven mouths old ; when younger she lays on-
ly those of workers. A queen hatched in
spring, perhaps will lay fifty or sixty eggs
of drones in whole, during the course of the

ensuing summer; but previous to beginning
her great laying of them, which should be
from one to two thousand, she must have
completed her eleventh month. In our series
of experiments, which more or less disturbed
the natural state of things, it often happened
^lat the queen did not attain this age until
^ ctober, and then immediately began lav-
ing male eggs. The workers, as if induced
by some emanation from them, also chose
the same time for building the royal celh
^0 swa.™ resulted thence, it is true, because
m autumn all the circumstances req^Lito
me absolute y wanting ; but it is not le!s evi .
dent, that there is a secret relation between

I

170 FORMATION OF SMARMS.

This laying commonly continues thifty
days. The bees on the twentieth or twenty-
first lay the foundation of several royal cells.
Sometimes they build sixteen or twenty;
we have had even twenty-seven. When
the cells are three or four lines high, the
queen lays those eggs in them from which
her own species will come, but not the whole
in one day. That the hive may throw seve-
ral swarms, it is essential that the young
females conducting them be not all produced
at the same time. It may be affirmed, that
the queen anticipates the fact, for she takes
care to allow at least the interval of a day
between laying every egg deposited in the
cells ; which is proved by the bees knowing
to close them at the moment when the
w'orms are ready to metamorphose to nymphs.
Now, as they close all the royal cells at dif-
ferent periods, it is evident the included
wnrms are not all of an equal age.

The queen’s belly is extremely turgid be-
fore she begins laying the eggs of drones;
but sensibly decreases as she advances, and
when the laying terminates it is very small.
Then she finds herself in a state to uuder-
take a journey which circumstances may pro-
long; therefore this is a necessary con-

rORMATION OF SWARMS.

171

dition; and as every thing is harmonious in
the laws of nature, the origin of the males
corresponds with that of the females, which
they are to fecundate.

Secondly, When the larvce hatched from
the eggs laid by the queen in the royal
cells are ready to transform to nymphs,
this queen leaves the hive, conducting a
swarm along with her; and the first
swarm that issues from the hive is uni-
formly conducted by the old queen.* I
think I can divine the reason of it.

In order that at no time there may be a
plui-ality of females in a hive, nature has in-
spired queens with an innate inveteracy
against each other; they never meet with-
out endeavouring to fight, and to accomplish
their mutual destruction. Thus, when nearly
of equal age, the chance of the contest is
equal between them, and fortune will decide
to which the empire shall pertain. Hut if
one combatant be older than the rest, she
N is stronger, and the advantage will be wfitli
I her. Slie will destroy her rivals successively
\ as produced. Thence, if the old queen did
I not leave the hive, before the young ones

! "»»

172 FORMATION OF SWARMS.

undergo their last metamorphosis, it could
produce no more swarms, and the species
would perish. Therefore to preserve their
race, it is necessary that the old queen shall
conduct the first swarm. But what is the
secret means employed by nature to induce
her departure ? I am ignorant of it.

In this country it is very rare, though
not without example, for the swarm led forth
by the old queen, in three weeks to produce
a new colony, which is also conducted by
the same female ; and that may happen thus:
Nature has not willed that she shall quit
the first hive before her production of male
eggs is finished. It is requisite for her to
be freed of them, that she may become
lighter. Besides, if her first occupation, on
entering a new dwelling, was laying more
male eggs, still she might perish either from
age or accident before depositing those of
w'orkers. The bees in that case w'ould have
no means of replacing her, and the colony
would go to ruin.

All these things have been anticipated
Avith infinite Avisdom. The first operation
of the bees of a SAvarm is constructing the '
cells of Avorkers. They labour at them Avith
great ardour; and as the ovaries of the queen I

FORMATION OF SWARMS.

173

have been ilisposecl with admirable foresight,
the first eggs she has to lay in her new abode
are those of workers. Commonly her laying
continues ten or eleven days ; and at this
time portions of comb containing large cells
are fabricated. We may affirm, that the
bees know that their queen will lay the eggs
of drones also ; she actually does begin to
deposit some, though in much smaller num-
ber than at first ; enough, however, to en-
courage the bees to construct royal cells.
Now, if in these circumstances the weather

be favourable, it is not impossible that a
second colony may be formed, and that the
queen may depart at the head of it within
thiee ueeks aftei conducting the first swarm
But I repeat it, the fact is rare in our climate.

Let me now return to the hives from
which the queen has led the first colony.

Thirdly, Jfter the old queen has con-
ducted the first sxmrmfrom the hive, the
remaining bees take particidar care of
the royal cells, and prevent the young
queens successively hatched from leaving
them, unless at an interval of several days
between the departure of each.

In the preceding letter, I have given you
the detail and proof of this fact, and I shall

174 FORMATION OF SWARMS.

here add some reflections. During the pe-
riod of swarming, the conduct or instinct of
bees seems to receive a particular modifica-
tion. At all other times, when they have
lost their queen, they appropriate workers’
worms to replace her ; they prolong and en-
large the cells of these worms; they supply
them with a greater portion of aliment, and
of a more pungent taste ; and by this alter-
ation the worms, that would have changed
to common bees, are transformed to queens.
We have seen twenty-seven royal cells of
this kind constructed at once ; but when
finished the bees no longer endeavour to
preserve the young females from the attacks
of their enemies. One perhaps may leave
her cell, and assail all the other royal cells
successively, which she will tear open to de-
stroy her rivals, without the workers taking
any part in their defence. Sliould several
queens be hatched at once, they will pursue
each other, and fight until the throne re-
main with her that is victorious. Far from
opposing such duels, the other bees rather
seem to excite the combatants.

IMattcrs are quite reversed during the pe-
riod of swarming. The royal cells then
constructed arc of a different figure from |

FORMATION OF SWARMS. 175

the former, being fashioned like stalactites,
and at first resembling the cup of an acorn.
The bees assiduously guard them when the
young queens are ready to undergo their
last meta?norphosis. At length the female
hatched from the first egg laid by the old
queen leaves her cell; the workers then treat
lier with indifference. But she, immediate-
ly yielding to the instinct which urges her
to destroy her rivals, seeks the cells where
they are enclosed ; yet no sooner does she
approach than the bees bite, pull, and ha-
rass her, so that she is forced to remove ;
but the royal cells being numerous, scarce
can she find a place of rest. Incessantly
animated with the desire of attacking the
other queens, and as incessantly repelled,
she becomes agitated, and hastily traverses
the different groups of workers, to which
she communicates her disorder. At this
moment numbers of bees rush towards the
aperture of the hive, and, accompanied by

the young queen, forsake it to seek another
habitation.

After departure of the colony, the re-
maining workers set another queen at liber-
ty, and treat her with equal indifference as
the first, They drive her from the royal

176 FORMATION OF SWARMS.

cells ; she also, perpetually harassed, be-
comes agitated, departs, and carries a new
swarm along with her. In a populous hive
this scene is repeated with the same circum-
stances three or four times during summer.
The numher of bees being then so much re-
duced that they are no longer capable of
preserving a strict watch over the royal
cells, several females are enabled to leave
their confinement at once ; they seek each
other, fight, and the queen, at last victori-
ous, reigns peaceably over the republic.

The longest intervals we have observed
between the departure of each natural swarm
have been from seven to nine days. This
is the time that usually elapses from the
period of the first colony being led out by
the old queen, until the next swarm is con-
ducted by the first young queen set at liber-
ty. The interval between the second and
third is still shorter ; and the fourth some-
times departs on the day after the third. In
hives left to themselves, fifteen or eighteen
days are usually sufficient for the throwing
of the four swarms, if the weather continues
' favourable, as I shall explain.

A swarm is never seen except in a fine
day, or, to speak more correctly, at a time of

FORMATION OF SWARMS. 177

the day when the sun shines, and the air is
calm. Sometimes we have observed all the
precursors of swarming, disorder and agita-
tion, but a cloud passed before the sun, and
tranquillity was restored ; the bees thought
no more of their departure. An hour after-
wards, the sun having again appeared, the
tumult was renewed ; it rapidly augmented;
and the hive swarmed.

Bees generally seem much alarmed at the
prospect of bad weather, ^^hile ranging
in the fields, the obscuration of the sun by
a cloud induces their precipitate return.
I am led to think that they are disquieted
by the sudden diminution of light. For if
the sky is uniformly obscured, and there is
no alteration in clearness or by the clouds
dispelling, they proceed to the fields for
their ordinary collections, nor do the first
drops of a soft rain make them return with
much precipitation.

I am persuaded that the necessity of a
fine day for swarming is one reason that has
induced nature to admit of bees protracting
the captivity of their young queens in the
royal cells. I will not deny that they some-
times appear to use this privilege in an ar-
bitrary manner. However, the confinement

178 FORMATION OF SWARMS.

of the queens is always longer when bad
weather lasts several days together. Here
the final object cannot be mistaken. Were
the young females at liberty to leave their
cradles during this intemperate season, there
would be a plurality of queens in the hive,
consequently combats ; and victims would
fall. The continuance of bad weather
might admit of all the queens undergoing
their last metamorphosis at once, or of their
having attained their liberty. She proving
victorious over the whole would enjoy the
throne, and the hive, which shoidd natural-
ly produce several swarms, could give only
one. Thus the multiplication of the species
would be left to the chance of rain, or fine
weather, instead of wdiich it is rendered in-
dependent of either, by the wise disposi-
tions of nature. By allowing only a single
female to escape at once, the formation of
swarms is ensured. This explanation ap-
pears so simple, that it is superfluous to in-
sist farther on it.

But I should mention another important
circumstance resulting from the captivity of'
queens; which is, that they are in a condi-
tion to fly when the bees have given them
liberty, and by this means are capable cf

FORMATION OF SWARMS. 179

profiting of the first moment of sunshine to
depart at the head of a colony.

You well know, Sir, that neither drones
nor workers are in a state to fly for a day
or two after leaving their cells. Then they
are of a whitish colour, weak, and their
organs infirm. At least twenty-four or
thirty hoiu's must elapse before the acqui-
sition of perfect strength, and the develope-
ment of all their faculties. It would be the
same with the females, were not their con-
finement protracted after the period of trans-
formation ; but we see them appear strong,
full grown, brown, and in a better condition
for flying than at any other period. I have
elsewhere observed, that constraint is em-
ployed to retain the queens in captivity :
the bees solder the covering to the sides
of the cell by a band of wax. As I have
also explained how they are fed, it need not
be repeated here.

It is likewise a very remarkable fact, that
queens are set at liberty earlier or later ac-
cording to their age. Immediately when
the royal cells were sealed, we marked them
all by numbers, and we chose this period
because it indicated tlie age of the queens
exactly. The oldest was first liberated,

180 FORMATION OF SWARMS.

then the one immediately younger, and so
on with the rest. None of the younger
queens obtained their freedom before the
older ones.

I have a thousand times asked myself
how the bees could so accurately distinguish
the age of their captives ; and undoubtedly
I should do better to answer the question
by a simple avowal of my ignorance. At
the same time, I must be permitted to offer
a conjecture regarding it. You will admit
that, as some authors, I have not abused the
right of giving myself up to hypothesis.
May not the piping or sound emitted by
the young queens, in their cells, be one of
the methods employed by nature to instruct
the bees in the age of their females ? It is
certain that she, whose cell is first sealed, is
also the first to emit this sound. The queen
in the next cell emits it sooner than the
rest, and so on those immediately subse-
quent. I do not deny, that as tlieir capti-
vity may continue 8 or 10 days, it is possible
that the bees in this space of time may for-
get which has emitted it first ; but it is also
possible, that the queens diversify the sounds,
augmenting the loudness as tliey become
older, and that the bees can distinguish

FORMATION OF SWARMS. 181

these variations. W e have been able even
ourselves to discern differences in the sound,
either with relation to the succession of
notes, or their intensity ; and probably there
are gradations still more imperceptible, escap-
ing’ our organs, but which may be sensible
to those of the workers.

This conjecture receives weight from the
queens reared by Schirach’s method being
perfectly mute ; neither do the workers
fonn any guard around their cells, nor
do tliey retain them in captivity a moment
beyond the period of transformation ; and,
having undergone it, they are freely allowed
to combat, until one has become victorious
over all the rest. Why ? Because the ob-
ject is only to replace the lost queen. Now,
provided that among the worms reared as
queens, a single one succeeds, the fate of the
others is uninteresting to the bees ; whereas,
during the period of swarming, it is neces-
saiy to preserve a succession of queens, for
conducting the different colonies ; and to
ensure the safety of the queens, it is essen-
tial to avert the consequences of tlie mutual
antipathy by which they are animated against
each other. Behold the evident cause of all
the precautions that bees, instructed by na-

182 FORMATION OF SWARMS.

ture, take during the 'period of swarming;
behold an explanation of the captivity of
females ; and why, that the duration of their
restraint might be ascertained by the age of
the young queens, it was requisite for them
to have some method of communicating to
the workers when they should be liberated.
This method consists in the sound emitted,
and the variation they are able to give it.

In spite of all my researches, I never have
been able to discover the situation of the
organ producing this sound. But I have in-
stituted a new course of experiments on the
subject, which are still unfinished.

Another problem still remains for solu-
tion. Why are the queens reared, accord-
ing to Schirach’s method, mute, whilst
those bred iu the time of swarming have the
faculty of emitting a certain sound ? What
is the physical cause of this difference ? At
first I thought it might be ascribed to the
period of life when the worms that are to
become queens receive the royal food. While
hives swarm, the royal worms receive the
food adapted for queens from the moment
of leaving the egg ; those, on the contrary,
destined for queens, according to Schi-
rach’s method, receive it only the second or

FORMxiTION OF SWARMS,

18,3

third day of their existence. 'It appeared to
me that this circumstance might have an
influence on the different parts of organiza-
tion, and particularly on the organ of voice.
But experiment has not confirmed my con-
jecture. I constructed glass cells in perfect
imitation of royal cells, that the metamor-
phosis of the worms into nymphs, and of the
nymphs to queens, might be visible. Our
experiments are related in a preceding letter.
We introduced into one of these artificial
cells the nymph of a worm reared according
to Schirach’s method, twenty-four hours
before it should naturally undergo its last
metamorphosis ; and we replaced the glass
cell in the hive, that the nymph might have
the necessary degree of heat. On the fol-
lowing day we had the satisfaction of seeing
it divest itself of the spoil, and assume its
ultimate figure. This queen was prevented
from escaping from her prison ; but we liad
contrived an aperture for her thrusting out
her trunk, that the bees might feed her. I
expected that she woidd have been com-

Ipletely mute; but it was otherwise; she
emitted sounds similar to those already de-
scribed, therefore my conjecture was erro-
9 neons.

184

FORMATION OF SWARMS.

I next conceived that the queen, being
restrained in her motions, was induced to
emit certain sounds in her desire for liberty.
All queens, in this new point of view, should
be equally capable of emitting the sound,
but, to induce them to it, they must be under
restraint. In the natural state, the queens
converted from workers are not a single in-
stant confined; and, if they do not emit the
sound, it is because nothing impels them to
it. On the other hand, those produced at
the time of swarming may be induced to do
so by the captivity in which they are kept.
For my own part, I give little weight to
this conjecture ; and though I state it here,
it is less with a view to claim merit than to
put others on a plan of discovering some-
thing more probable.

I do not ascribe to myself the credit of
having discovered the piping of the queen
bee. Old authors speak of it. ISl. de Reau-
mur cites a Latin -work published 1671,
3Iona?'chia Feinina, by Charles Butler.*
He gives a very brief abstract of this natu-
ralist’s observations, who, we can easily sec,
has exaggerated or rather disguised the
truth, by mixing it with the most absurd fan-

* This work was published so long ago as 163 t. It is
both written and printed in the most extravagant style. — T.

FORMATION OF SWARMS. 185

cies ; but it is not the less evident that Butler
has heard this peculiar piping of queens,
and that he did not confound it with the con-
fused humming sometimes heard in hives.

Fourthly, Young queens conducting
s'coarnis from their native hive are still in
a virgin state. They generally depart in
quest of the males the day after being settled
in their new abode, which is usually the
fifth of their existence as queens ; for two or
three are passed in captivity, one in their
native hive, and a fifth in their new dwell-
ing. Those queens that come from the worm
of a worker also pass five days in the hive
before going in quest of males. So long as
in a state of virginity, both are treated with
indifference by the bees ; but after returning
with the external marks of fecundation, they
are received by their subjects with the most
distinguished respect. However, forty-six
hours elapse after fecundation before tliey
begin to lay. The old queen, which leads
the first swarm in spring, requires no farther
commerce with the males for the continu-
ance of her fecundity. Union only once is
sufficient to impregnate all the eggs that she
will lay for at least two years. — Pregnw,
8^/i September 1791.

186 FORMATION OF SWARMS.

LETTER XL

THE SAME SUBJECT CONTINUED.

My chief observations on swarms, those
most frequently repeated, and of which the
uniformity of result leads me to apprehend
no error, are collected in the two preceding
letters. I have deduced wdiat seem the most
direct consequences ; and in all the theoreti-
cal part, I have sedulously avoided going
beyond facts. What remains to be men-
tioned is more hypothetical, but it engrosses
several curious experiments.

It has been demonstrated, that the prin-
cipal motive urging the departure of the
young females when hives swarm, is their |
insuperable antipathy to each other. I have i

FORMATION OF SWARMS. 187

repeatedly observed that they cannot grati-
fy their aversion, because the workers with
the utmost care prevent them from attack-
ing the royal cells. This perpetual opposi-
tion at length creates a visible inquietude,
and excites a degree of agitation that in-
duces them to depart. All the young queens
are successively treated alike in hives that
are to swarm. But the conduct of the bees
towards the old queen, destined to conduct
the first swarm, is very diflTereut. Always
accustomed to respect fertile queens, they do
not forget what they owe to her ; they al-
low her the most uncontrolled liberty. She
is permitted to approach the royal cells ; and
if she even attempts to destroy them, no
impediment is offered by the bees. Thus
her inclinations are not obstructed, and we
cannot ascribe her flight, 'as that of the young
queens, to the opposition she suffers. There-
fore, 1 candidly confess myself ignorant of
the motives of her departure.

I Yet, on more mature reflection, it does
not appear to me that this fact affords so
strong an objection against the general rule
as I had at first conceived. It is certain at
least, that the old queens, as well as the
c young ones, entertain the greatest aversion

188 FORMATION OF SWARMS.

towards the individuals of their own sex.
This has been proved by the numerous
royal cells destroyed. You will remember,
Sir, that in my first observations on the de-
parture of old queens, seven royal cells open-
ed at one side were destroyed by the queen.
If rain continue several days, destruction
awaits the whole ; in which case there is
no swarm, as too often happens in our
climate, where spring is generally wet.
Queens never attack cells containing an egg
or a very young worm ; but only when the
worm ivS ready for transforming to a nymph,
or when it has undergone its last metamor-
phosis.

The presence of royal cells with nymphs
or worms near their change, also inspires old
queens with the utmost horror or aversion ;
but here it would be necessary to explain
why the queen does not always destroy them
though it be in her power. On this point
I am limited to conjectures. Perhaps the
great number of royal cells in a hive at
once, and the labour of opening the whole,
creates insuperable alarm in the old queen.
She commences indeed with attacking her
rivals ; but, incapable of obtaining imme-
diate success, her disquiet during the work

FORMATION OF SWARMS. 189

is roused to a terrible agitation. If tlie wea-
ther continues favourable, while she remains
in this condition, she is naturally disposed
to depart.

It may be easily understood that the
v/orkers, accustomed to respect their queen,
whose presence is a real necessity to them,
crowd after her ; and the formation of the
first swarm creates no difficulty in this view.
But undoubtedly you will ask. Sir, What
motive can induce the workers to follow their
young queens from the hive, while they mal-
treat them so much, and even, in their most
amicable moments, testify perfect indif-
ference towards them ? Probably it is to es-
cape the heat to which the hive is then ex-
posed. The extreme agitation of the fe-
males leads them to traverse the combs in
all directions. The groups of bees through
which they pass are injured and deranged ;
a kind of delirium is imparted to them, and
this tumultuous disorder raises the temper-
ature to an insupportable degree. AVe have
frequently proved it by the thermometeiv
In a populous hive it commonly stands be-
tween 90° and 97°, in a fine day of spring-
►but during the tumult which precedes swarm’
mg, It rises above 104°: and this is heat in-

190 FORMATION OF SWARMS.

tolerable to bees. ^Vhen exposed to it,
they rush impetuously towards the outlets of
the hive, and depart. lu general they can-
not endure the sudden augmentation of
heat, and then quit their dwelling ; neither
do those returning from the fields enter when
the temperature is extraordinary.

By means of direct experiments, I have
ascertained that the impetuous courses of
the queen over the combs actually throws
the workers into a state of agitation. It
\vas particularly important to learn, whether
she would communicate her disorder inde-
pendent of the time of swarming. But I
w'ished to avoid a complication of causes.

I took two females, still virgins, of above
five days old, susceptible of fecundation, and
having put them into separate glass hives,
sufficiently populous, where the air had free
circulation, effectually precluded any of the
bees from escaping. Then I prepared to
observe the hives every moment that the
fineness of the weather might invite both
males and females to go abroad, for the
purpose of fecundation. Next morning
being gloomy, no male left the hive, and i
the bees were tranquil; but towards ele-
ven of the following day, the sun shining

FOKMATION OF SWARMS. 191

bright, both queens began to run about seek-
ing an exit from every part of their dwell-
ing ; and from their inability to find one,
traversed the combs with the most evident
symptoms of disquiet and agitation. The
bees soon participated of the same disorder;
they crowded towards the lower part of the
hive, where the openings were situated ; un-
able to escape, they ascended with equal rapid-
ity, and hurried heedlessly over the cells un-
til four in the afternoon. It is nearly about
this hour that the sun declining in the ho-
rizon recalls the males ; queens requiring fe-
cundation never remain later abroad. The
two females became calmer, and tranquillity
in a short time was restored. This was re-
peated several subsequent days with perfect
similarity ; and I am now convinced that
there is nothing singular in the agitation of
bees while swarming, but that they are al-
ways in a tumultuous state when the queen
herself is in agitation.

I have but one fact more to mention. It
has been already observed, that on losing
[ the female, bees give the larvae of simple
I workers the royal treatment, and, accordinc.
to Schirach, in five or six days they repair
8 the loss of their queen. In this case there

192 FORMATION OF SWARMS.

are no swarms. All the females leave their
cells almost at the same moment, and, after
a sanguinary combat, the throne remains
wdth the most fortunate.

I can comprehend very well that the ob-
ject of nature is to replace the lost queen ;
but as bees are at liberty to choose either
the eggs or worms of workers, during the
first three days of existence, to supply her
place, why do they give the royal treatment
to worms, all of nearly an equal age, and
which must undergo their last metamor-
phosis almost at the same period ? Since
they are enabled to retain the young females
in their cells, why do they allow all the
queens, reared according to Schirach’s me-
thod, to escape at once? By prolonging
their captivity more or less, they would ful-
fil two most important objects at the same
time, in repairing the loss of their females,
and in preserving a succession of queens to
conduct several sw'arms.

At first it was my opinion that this dif-
ference of conduct proceeded from the dif-
ference of circumstances in -which they find
themselves situated. They are induced to
make all their dispositions relative to swarm-
ing only when in great numbers, and when

rORMATION OF SWAHMS. 193

tliey have a queen occupied with her pr iiici-
pal laying of male eggs ; whereas, having
lost their female, the eggs of drones are no
longer in the combs to influence their in-
stinct. They are in a certain degree rest-
less and discouraged.

Oil this account, after removing tlie queeu
from a hive, I thought of rendering all the
other circumstances as similar as possible to
the condition of bees preparing to swarm.
By introducing a great many workers, I in-
ci eased the population to excess, and sup-
plied them with combs of male brood in
every stage. Their first occupation was con-
structing royal cells after Schirach’s method,
mid rearing common worms with royal food!
They also began some stalactite cells, as if
the presence of the male brood had inspired
them to It ; but these they discontinued, as
there was no queen to deposit her eggs. Fi-

f f ^ ^'oyal cells,

taken .nd.ffeiontly from hives preparing to

swarm. However, all my preeantions were
untlcs.s; the bees were occupied only with
veplacng their lost queen ; they nejeeted
the royal cells entrusted to their care • the
t^luded queens came out at the onlinlr;

[ ’ ’''"'g detained prisoners I

; K

194 FORMATION OF SWARMS.

moment ; they engaged in several combats,
and there were no swarms.

Recurring to subtleties, we may perhaps
suggest the cause of this apparent contra-
diction. But the more we admire the wise
dispositions of the Author of nature, in the
laws he has prescribed to the industry of
animals, the greater reserve is necessary in
admitting any theory adverse to this beauti-
ful system, and the more must we distrust
that facility of imagination from which we
think by embellishment to attain the eluci-
dation of facts.

Naturalists in general, who have made
animals the subject of long observation, and
those in particular who have chose insects
for their favourite study, have too readily
ascribed to them our sentiments and pas-
sions, and even our designs. Yielding to
admiration, and disgusted perhaps by the
contempt with which insects are treated,
they have conceived themselves obliged to
justify the consumption of time bestowed on
this pursuit, and they have depicted differ-
ent traits of the industry of these minute
animals, with the colours inspired by a warm
imagination : Nor is the celebrated Reau-

mur even to be acquitted of such a charge.

FORMATION OF SWARMS.

195

He frequently ascribes combined intentions
to bees ; love, anticipation, and other facul-
ties of too elevated a kind. I think I can
discern that, although he formed a very just
conception of their operations, he would be
well pleased that his reader should admit that
they are sensible of their own interest. He
is a painter who by a happy prejudice flatters
the original, whose features he depicts. On
the other hand, Buffon unjustly considers
bees as mere automatons. It was reserved
for you, Sir, to establish the theory of animal
industry on the most philosophical principles
and to demonstrate that those actions that
have a moral appearance depend on an as-
sociation of ideas simply sensible. It is not
my object here to penetrate such depths or
to insist on the details.

But, on the whole, the facts relative to
tlie foimatiou of swarms, perhaps present

MroSe r f" than another

pait of tlie history of bees; whence I think

't proper to state, in a few words, the sim

phcity of the methods by which the wisdom of

nature guides their instinct. It cannot -ill

hem the slightest portion of nnderstolhw

196 FORMATION OF SWARMS.

seiisorium being adapted to the different
operations with which they are charged, it
is the impidse of pleasure which leads them
on. She has therefore pre-ordained all that
is relative to the succession of their different
labours; and to each operation she has united
an agreeable sensation. Thus, when bees
construct cells, watch over their larvse, and
collect provisions, we must not ascribe it to
method, affection, or foresight. The only
inducement must be sought for in some
pleasing sensation attached to each of these
functions. I address a philosojiher ; and as
these are his own opinions applied to new
facts, I believe my language will be easily
understood. But I request my readers to
peruse and to reflect on that part of your
w'orks which treats of the industry of ani-
mals. Let me add but another sentence.
The impulse of pleasure is not the sole
agent ; there is another principle, the ex-
traordinary influence of which, at least with
regard to bees, has been hitherto unknown,
that is, the sentiment of aversion continu-
allv felt by all females against each other ;
a sentiment whose existence is so fully do- ■
monstrated by my experiments, and which
explains many important facts in the theory |
of swarms. — Pregny, IQthSeptcmhei'llQl. I

THE EGGS OF MALES.

197

LETTER XII.

additional observations on QUEENS

LAYING ONLY THE EGGS OF DRONES,
AND ON THOSE DEPRIVED OF THE AN-
TENNiE.

In relating my first observations on queens
laying male eggs alone, I have proved that
they deposit them in cells of all dimensions
indifferently, and even in royal cells. It is
also proved that the same treatment is given
to male worms hatched from eggs laid in
the royal cells, as if they were actually to be
transformed to queens : and I have added,
that in this instance the instinct of the work-
ers appeared defective. It is indeed most
singular, that bees, knowing the worms of

198

QUEENS LAYING ONLY

males so well when the eggs are laid in small
cells, and never failing to give them a con-
vex covering when about to transform to
nymphs, should no longer recognise the same
species of worms when the eggs are laid in
royal cells, and treat them exactly as if they
should change to queens. This inconsist-
ence depends on something incomprehensi-
ble by me.*

* I have ascertained, from new observations, that bees
recognise the larvae of males, as well when the eggs pro-
ducing them have been laid in royal cells by queens of re-
tarded fecundation, as when deposited in common cells.

The former, it will be recollected, resemble a pear with
the small end downwards, and are an inch and a quarter, or
little more in length. When only sketched, and very hke
the cup of an acorn, queens lay in them.

Bees, indeed, give the same shape and dimensions to
those serving as receptacles for the males, but when the
transformation of the larvae approaches, it is easy to discover
that they have not taken them for royal worms. Instead of
closing their cells in a pointed form, as they do invariably if
containing the larva? of queens, they widen the extremity, aqd
adding a cylincMcal tube, shut them with a convex covering
differing in nothing from that which they are accustomed
to put on the cells of males. But as this tube is of the same
capacity as the hexagonal cells of the smallest diameter, the
worm, which is forced by the bees to descend into it, and
which must there undergo its final metamorphosis, becomes
a drone of the smallest size. The total length of these ex-
traordinary cells is from 20 to 22 lines (I3 to liinch):
yet bees do not always add a cylinder to a pyramidal cell ;
they arc content with giving the lower part a little enlarge-

THE EGGS OF MALES.

199

In revising what is said on the subject, I
observed an interesting experiment still
wanting to complete the history of queens
laying only the eggs of drones. I had to
investigate whether they could themselves
distinguish that the eggs which they de-
posit in the royal cells would not produce
queens. I have already remarked that they
do not endeavour to destroy these cells when
close ; and I thence concluded, that in ge-
neral the presence of royal cells in their hive
does not inspire them with the same aver-
sion to females whose fecundation has been
retarded ; but to ascertain the fact more cor-
rectly, it was essential to examine how the
presence of a cell containing a royal nymph

ment: and here the growing larva may become a large-
drone.

I am ignorant what is the cause of a difference sometimes
observed in the form of these cells ; but it appears very cer-
tain to me, that the bees never are deceived in them : thus
affording a great proof of the instinct with which they arc
endowed.

Bees being intrusted by nature with bringing up their
young, and with the care of providing them witli the ali-
ment proper to tlieir age, or even to their sex, should Ije
.also endowedwith the faculty of recognising them. Tliere
is so little resemblance between the adult male and woi kers,
that some difference should subsist between their respective
larvoe. Doubtless the workers distinguish it, though esc.ajiing
our notice.

200 QUEENS LAYING ONLY^

would affect a queen that had never laid
any other than the eggs of drones.

The experiment was easy; and I put it
in practice on the fourth of September, in a
hive some time deprived of its queen, where
the bees had not failed to construct several
royal cells for replacing her. I chose this
opportunity to supply them with a female,
whose fecundation had been retarded to the
twenty-eighth day, and which laid none but
the eggs of males. At the same time, I
removed all the royal cells, except one that
had been sealed five days. A single cell
remaining was enough to show the impres-
sion it would make on the stranger queen
introduced : if she endeavoured to destroy
it, in my opinion, this would prove that she
anticipated the origin of a dangerous rival.
You must admit the use I now make of
the word anticipate ; it is to avoid circum-
locution ; I feel its impropriety. If, on the
contrary, she did not attack the cell, I would
thence infer that the postponed fecun-
dation, which deprived her of the power of
laying workers’ eggs, had also impaired her
instinct. The latter was the fact; the queen
passed several times over the royal cell, both

THE EGGS OF MALES. 201

the first and the subsequent day, without
seeming to distinguish it from the rest.
She quietly laid in the surrounding cells ;
notwithstanding the cares incessantly be-
stowed by the bees upon it, she never one
moment appeared to suspect the danger with
which the included royal nymph threatened
her. Besides, the workers treated their new
queen as well as they would have treated
any other female. They were lavish of ho-
ney and respect, and formed those regular
circles around her that seem an expression
of homage.

Thus, independent of the derangement
occasioned by retarded impregnation in the
sexual organs of queens, it certainly impairs
their instinct. Aversion or jealousy against
their own sex in the nymphine state ceases,
nor do they endeavour to destroy them in
their cradles.

My readers will be surprised that queens,
whose fecundation has been retarded, and
whose fecundity is so useless to bees, never-
theless .should be so well treated, and be-
come as dear to them as females laying both
kinds of eggs. But I remember to liave
observed a fact more astonishing still. I
have seen workers bestow every attention

202 QUEENS LAYING ONLY

on a queen though sterile ; and after her
death treat her dead body as they had
treated herself when alive, and long pre-
fer it, though inanimate, to the most fertile
queens I had offered them. This sentiment,
which assumes the appearance of lively af-
fection, is probably the effect of some agree-
able sensation communicated to bees by their
queen, independent of fertility. Those lay-
ing only the eggs of males undoubtedly ex-
cite the same sensation in the workers.

I now recollect that the celebrated Swam-
merdam somewhere observes, that w'hen a
queen is blind, sterile, or mutilated, she
ceases to lay, and the workers of her hive no
longer continue their labours or make any
collections, as if aware that it is useless to
do so. But in advancing this as a fact, he
cites no experiments that led him to the dis-
covery. Those made by myself have afforded
some very singular results.

I frequently amputated the four wings of
queens ; yet not only did they continue lay-
ing, but the same cousidcration of them was
testified by the workers as before. There-
fore, Swammerdam has no foundation for
asserting, that mutilated queens cease to
lay. Indeed, from his ignorance of fccun-

I

203

the eggs of males.

tlatioii taking place without the hive, it is
. possible that he cut the wings oli virgin
queens, and they, becoming incapable of
bight, remained barren from inability to
seek the males in the air. Thus amputa-
tion of the wings does not produce sterility.

t

I have frequently cut olF one of the an-
tennae to recognise a queen the more easily,
and it was not prejudicial to her either in
fecundity or instinct, nor did it affect the
attention paid to her by the bees. It is
true, that as another still remained, the mu-
tilation was imperfect ; and nothing was de-
cided by the experiment. But amputation
of both antennae produced most singular ef-
fects. On the fifth of September, I cut them
off a queen that laid the eggs of males only,
and put her into the hive immediately after
the operation. From this moment there
was a great alteration in her conduct. She
traversed the combs with extraordinary vi-
vacity : scarcely had the workers time to se-
parate and recede before her, and she drop-
ped her eggs, without the precaution of de-
positing them in any cell. The hive not
I being very populous, part wanted combs.

: Hither she seemed particularly earnest to

5204.

EFFECTS OF

repair, and remained long motionless. She
appeared to avoid the bees ; however, seve-
ral workers followed her into this solitude,
and treated her with the most evident re-
spect. She seldom required honey from
them, but, when that occurred, she directed
her trunk with an uncertain kind of feeling,
sometimes on the head and sometimes on
the limbs of the workers, and if it did reach
their mouths, it was by chance. At other
times she returned upon the combs, then
quitted them to traverse the glass sides of
the hive : and always dropped eggs during
her various motions. Sometimes she ap-
peared tormented with the desire of leaving
her habitation, and rushing towards the
opening, entered the glass tube adapted
there ; but the external orifice being too
small, after fruitless exertion, she returned.
Notwithstanding these symptoms of deliri-
um, the bees did not cease to render her the
same attention as they ever pay to their
queens, hut this one received it with indif-
ference. All that I describe appeared to me
the consequence of amputating the antennse.
However, her organisation having already
suffered from retarded fecundation, and as I
had observed her instinct in some degree

MUTILATING QUEENS.

205

impaired, both causes might possibly concur
in operating the same effect. To distinguish
what peculiarly belonged to privation of
the antennas, repetition of the experiment
was necessary on a queen otherwise well or-
ganised, and capable of laying both kinds
of eggs.

This was practised on the sixth of Septem-
ber. I amputated both the antennae of a fe-
male which had been several months the sub-
ject of observation, and, being of great fecun-
dity, had already laid a considerable number
of workers’ eggs, and those of males. I put
her into the same hive where the queen of
the preceding experiment still remained, and
she exhibited precisely the like symptoms
of delirium and agitation, which I think it
needless to repeat. I shall only add that,
to judge better of the effect produced by
privation of the antennae on the industry
and instinct of bees, I attentively consider-
ed the manner in which these two mutilat-
ed queens treated each other. You cannot
1 have forgot, Sir, the animosity with which
queens, possessing all their organs, combat ;
therefore, it became extremely interesting
to learn whether a similar reciprocal aversion
nould subsist after losing their antennas.

206

EFFECTS OF

We studied these females a long time ; they
met several times in their courses, and with-
out exhibiting the smallest resentment.
This last instance is, in my opinion, the
most decisive evidence of a change operated
in their instinct.

Another very remarkable circumstance,
which the same experiment gave me occa-
sion to observe, consisted in the good recep-
tion given by the bees to the stranger queen,
while they still preserved the first. Hav-
ing so often seen the symptoms of discon-
tent occasioned by a plurality of queens, af-
ter having witnessed the clusters formed
around such supernumeraries to confine
them, I could not expect that they would
pay the same homage to a second mutilated
one which they still testified towards the
first. Was it from losing the aiitennas that
these queens no longer retained any charac-
teristic distinguishing the one from the
other ?

I was the more inclined to admit this con-
jecture, from the bad reception of a third
fertile queen preserving her antenna;, intro-
duced into the same hive. The bees seized,
bit lier, and restrained her so closely that
.slie could hardly breathe or move. There-

MUTILATING QUEENS.

207

fore, if they treat two females deprived of the
antemiEC equally well in the same hive, it is
probably because they experience the same
sensation from both, and want the means of
continuing to distinguish them from each
other.

From all this, I conclude, that the an-
tenna are not a frivolous ornament to in-
sects, but, according to every appearance,
are the organs of touch or smell. Yet I
cannot affirm which of these senses reside
in them. It is not impossible that they
are organized in such a manner as to fulfil
both functions at once.

As in the course of the last experiment
the two mutilated females constantly endea-
voured to escape from the hive, I wished to
see what they would do if set at liberty,
and whether the bees would accompany
them in their flight. Therefore, leaving
the fertile mutilated one, I removed the
first and third from the hive, and then en-
larged the entrance.

That day the queen left her habitation.
At fiist she tried to fly, but, her belly being
full of eggs, she fell down, and never at-
tempted it again. No workers accompanied
lior. Wliy, after rendering the queen so

208

EFFECTS OF

rnucli attention while she lived among them,
did they abandon her now on her departure ?
You know, Sir, that queens governing a
weak swarm are sometimes discouraged, and
fly away, candying all their little colony along
with them. In like manner sterile queens,
and those whose dwelling is ravaged by
weevils, depart, and are followed by all their
bees. Why therefore in this experiment
did the workers allow their mutilated queen
to depart alone? All that 1 can hazard on
the question is a conjecture. It appears
that bees are induced to quit the hives from
the increased heat, which occasions the agi-
tation of their queen, and the tumultuous
disorder which she communicates to them.
A mutilated queen, notwithstanding her
delirium, does not agitate the workers, be-
cause she seeks the glass panes and the un-
inhabited parts of the hive: she hurries over
clusters of bees, but the shock resembles
that of any other body, and produces only
a local and instantaneous motion. The sen-
sation arising from it, is not communicated
from one place to another, like that occa-
sioned by a queen which in the natural state
wishes to abandon her hive and lead out a
swarm ; there is no augmented temperature,

MUTILATING QUEENS. 209

consequently nothing that renders the hive
insupportable to the bees.

This conjecture, which affords a tolerable
explanation why bees persist in remaining
in the hive, though the mutilated queen has
left it, does not account for the motive in-
ducing the queen herself to depart. Her
: instinct is altered ; but that is all of which
I am sensible. I can discern nothing more.
It is very fortunate for the hive that she-
quits it, for the bees incessantly attend her ;
nor do they ever think of procuring ano-
ther queen while she remains; and if she
I were long of leaving them, it would be
i impossible to replace her ; for the workers’
f worms would exceed the term at which they
I are convertible into royal worms, and the
i hive would perish. Observe, that the eggs
I dropped by the mutilated queen can never
I serve to replace her, for, not being deposit-
i ed in cells, they wither and produce nothing.

I have yet to say a few words on the fe-
ll males laying male eggs only. Schirach
i supposes that one branch of their double

ovary suffers some alteration. He seems to
U think that the eggs of males are contained
in one of the branches, while the other has
none but common eggs ; and as he ascribes

210

MUTILATED QUEENS.

the incapacity of certain queens to lay the
latter to some disease, his hypothesis cer-
tainly is very plausible. In fact, if the eggs
of males and workers are indiscriminately
mixed in both branches of the ovary, it ap-
pears, at first sight, that whatever may be •
the cause acting on this organ, it should
affect both species of eggs equally. If, on
the contrary, one branch is occupied by the
eggs of drones only, and the other contains i
none but common eggs, we may conceive i
how disease affects the one, while the other'
remains untouched. Notmthstanding the-
probability of this conjecture, it is confuted,
by observation. We lately dissected queens!
which laid none but male eggs, and found i|
both branches of the ovary equally well ex--
panded, and equally sound, if I may use the’
expression. The only difference of which
we were sensible consisted in the eggs not
lying so close together, apparently, in these
two branches, as in the ovaries of queens by
which both kinds are laid. — Peegny, 12f/i
September 1791-

ECONOMICAL CONSIDERATIONS. 211

LETTER XIII.

ECONOMICAL CONSIDERATIONS ON BEES.

In this letter I shall treat of the advantages
that may he derived from the new invented
hives, called book or leafloives, in promoting
the economical knowledge of bees.

It is needless to relate the different me-

Ithods hitherto employed in compelling these
insects to yield up a portion of their honey
and wax ; all correspond in being cruel and
ill understood.

It is evident, that when bees are culti-
vated for the purpose of sharing the produce
of their labours, we must endeavour to mul-
tiply them as much as the nature of the
country admits ; consequently we ought to
regard their lives at the time we plunder
t em. riicicforc it is an absurd proceeding

212

ECONOMICAL

to sacrifice whole hives to get at the riches
they contain. The inhabitants of this coun-
try, who follow no other method, annually
lose immense numbers of hives ; and spring
being generally unfavourable to swarms, the
loss is irreparable. I know well that at first
they will not adopt a different plan ; they
are too much attached to prejudices and old
customs. But naturalists and intelligent
cultivators of bees will be sensible of the
utility of the method I propose ; and if they
apply it to use, I hope their example will
extend and perfect the culture of bees.

It is not more difficult to lodge a natural
swarm in a leaf hive than in any other of a
different shape. But there is one precaution
essential to success, which I should not omit
adverting to. Though bees are indifferent
as to the position of their combs, and their
greater or lesser size, they are obliged to
construct them perpendicular to the hori-
zon, and parallel ,to each other. Therefore, '
if left entirely to themselves, when esta- :
blishing a colony in one of these new hives, '
they will frequently construct several small
combs, parallel indeed, but perpendicular to
the plane of the frames or leaves, or per-
haps they will build them on the place where

CONSIDERATIONS.

213

two are joined together, and by this disposi-
tion prevent the advantages which I think
to derive from the figure of my hives, since
they could not he opened without breaking
the combs.* Thus they must previously
have a guide ; the cultivator himself can lay
the foundation of their edifices, and that by
a simple method. A portion of comb must
be fixed securely in the upper part of some of
the frames composing the hive ; the bees will
extend it ; and, in prosecution of their work,
will accurately follow the plan already given
them.f Therefore, on opening the hive, no
obstacle is to be removed, nor are stings to
be dreaded, for one of the most singular and
valuable properties attending this construc-
: tiou, is its rendering the bees tractable. I
appeal to you. Sir, for the truth of what I
say. In your presence I have opened all the
divisions of the most populous hives, when
the tranquillity of the bees has given you
great surprise. I can desire no other evi-
dence of my assertion. It is in the facility

* The author’s general meaning is, the bees will build
across the hive, or parallel to the narrow sides of <hc
frames. — T.

t If the upper part of a frame form a salient angle, it
•is said that bees establish their original work with grea"ter
'regularity. — T.

214

economical

of opening these hives at pleasure that all
the advantages are concentrated, which I
expect in perfecting the economical know-
ledge of bees.

^Vhen I observe that bees may be ren-
dered tractable,! conceive it need not be add-
ed that I do not arrogate to myself the
absurd pretence of taming them, for this ex-
cites a vague idea of deception ; and I would
willingly avoid the hazard of exposing my-
self to any such reproach. I ascribe their
tranquillity, on opening the hives, to the
manner in which they are affected by the
sudden introduction of light ; then, they
appear rather to testify fear than anger.
Many retire, and entering the cells seem to
conceal themselves. My conjecture is con-
firmed by their being less tractable during
night or after sunset than through the day.
Thus, we must open the hives, while the
sun is above the horizon, cautiously, and
without any sudden shock. Separating the
divisions slowly, and taking care to wound
none of the bees. If they cluster too much
on the combs, it is necessary to brush them
off with a feather ; and breathing on them
is to be anxiously avoided. Tlic air which i
we expne seems to excite their fury ; it cer-

CONSIDERATIONS.

215

taiiily possesses some irritating quality, for
if bellows be used, they are rather inclined
to escape than to sting.

Respecting the advantages of leaf hives,
I shall remark, that they are extremely con-
venient for forming artificial swarms. In
the history of natural swarms, I have shown
how many favourable circumstances are re-
quisite for their success. From experience
I know that they very often fail in our cli-
mate ; and even when a hive is disposed to
swarm, it frequently happens that the colony
is lost, either because the moment of its de-
paiture has not been foreseen, because it
rises out of sight, or settles on inaccessible
places. Instructing the cultivators of bees
I how to make artificial swarms is a real bene-
I fit, and the form of my hives renders this an
easy process. But it demands farther illus-
ti'ation.

Since bees, according to Schirach’s disco-
1 very, can procure another queen after having
• lost their own, provided there be workers’

! brood in the combs not above three days
' old, It results that we can produce queens
at pleasure, by removing the reigning one.

' nerefore, if a hive sufficiently populous be
divided in two, one half will retain the old

216

ECONOMICAL

queen, and the other will not be long of ob-
taining a new one. But to ensure success,
we must choose a propitious moment, which
is never certain but in leaf hives. In these
we can see whether the population is suffi-
cient to admit of division, if the brood is of
the proper age, and if males exist or are
ready to be produced for impregnating the
young queens.

Supposing the concurrence of all these
conditions, the following is the method to
be pursued. The leaf hive is to be separated
through the middle without any shock, and
two empty divisions insinuated between the
halves, which, when exactly applied to each
other, are close on the outside. The queen
inust be sought in one of the halves, and
marked to avoid mistake. If she by chance i
remains in the division with most brood, she
is to be transferred to the other with less,
that the bees may have every possible op-
portunity of obtaining another female.
Next, it is necessary to connect the halves
together, by a cord tied tight around them;
and care must be taken that they are set on
the same board that the hive previously oc-
cupied. The old entrance, now become use-
less, will be shut up ; but as each half re-

COXSIDERATIOXS.

217

quires a new one, it ought to be made at the
bottom of the first and twelfth division, on
2)ui]iose that they may be as far asunder as
jmssible. Both entrances should not be
opened on the same day. The bees in the
half deprived of the queen are to be confin-
ed twenty-four hours, and no aperture made
before that time elapses except for the ad
mission of air. Without this precaution’
they would soon search after their queen,
and infallibly finding her in the other divi
Sion, they lyoiild then retire in great num-
bers from their own, until too few remained
to perform the necessary labours. But this
will not ensue if they be confined twenty-
four hours, provided that interval is suffi-
cieiit to make them forget thp m,

When all circumstances \re tourbTe
the bees, in the division wanting her, will
begin the same day to labour in prociirino-
another, and ten or fifteen days after the
operation their loss will be repaired. T o
young fem.ale they have reared soon issues
foith to seek impregnation, and in two days

minences the laying of workers’ oggs.

thing more « wanting to the bees of tht

bive, and the success of the artifipi l
i swarm is ensured. aitificial

L

ECONOMICAL

SJ18

It is to Scliiracli that we are indebted for
sucli an ingenious method of forming swarms.
He supposes that, by producing young
queens in the beginning of spring, early
swarms might be procured, which certainly
would he advantageous in most situations.
But unfortunately this is impossible. Schi-
racli believed that queens were fecundated
of themselves; consequently he thought that,
after being artificially produced, they would
lay and give birth to a numerous posterity.
Now, this is an error ; the females, to be-
come fertile, require the concourse of the
males, and if not impregnated within a few
days of their origin, their laying, as I have
observed, is completely deranged. There-
fore, if a swarm were artificially formed be-
fore the usual time of the males originating,
the bees would be discouraged by the steri-
lity of the young female : Or should they
remain faithful to her, awaiting the period
of fecundation, as she could not receive the
approaches of the male for tliree or four
Aveeks, she would lay eggs producing males
only, and the hive in that case would perish.
Thus the natural order must not be deran-
ged, but the division of hives delayed until
males arc about to originate or actually exist.

CONSIDERATIONS. 219

Besides, if Schirach did succeed in ob-
taining artificial swarms, notwithstanding
the great inconvenience of his hives, it was
owing to his singular address and unremit-
ting assiduity. He had some pupils in the
art, by whom it w'as communicated to others;
and there are people now in Saxony who
tia\ erse the country practising this operation.
Those versant in the matter can alone ven-
tuie to undertake it wfith common hives,
whereas every cultivator can do it himself
with the leaf hives.

In this construction, another very essen-
tial advantage will be found. Bees can be
torced to work in wax ; and here I am led
to what I believe is a new observation,
w bile naturalists have directed our admi-
ration to the parallel position of the combs
they have overlooked another trait in the
industry of bees, namely, the equal distance
uniformly preserved between them ()„
measuring the interval by which they are
separated, in general it will be found a third
of an nicli. Were they too distant, it is
•my evident that the bees would bo greatly
spersed, and unable to communicate t i
eat reciprocally; whence the brood could
not receive sufficient warmth. AVere tl e

2£0

ECONOMICAL

combs too close, on the contrary, the bees
could not freely traverse the intervals, and
the work of the hive would suffer. There-
fore, a certain distance always uniform is re-
quisite, which is equally adapted to the ser-
vice of the hive, and the care necessary for
the worms. Nature, which has taught bees
so much, has instructed them in the regular
preservation of this distance. On the ap-
proach of winter, they sometimes elongate
the cells which are to contain honey, and
thus contract the intervals between the
combs. But this expedient is in preparation
for a season, when it is important to have
copious magazines, and when their activity
being relaxed, it is unnecessary for their com-
munications to be so spacious and free. On
the return of spring, the bees hasten to con-
tract the elongated cells, that they may be-
come fit for receiving the eggs which the
queen will lay, and thus re-establish the
suitable distance which nature has ordained.

This being premised, bees may be forced i
to work in wax, or, which is the same thing, |
to construct new combs. To'accomplish the
object, it is only necessary to separate those
already built so far asunder that they may •
construct others in the interval. Suppose.

CONSIDERATIONS.

221

that an artificial swarm is lodged in a leaf
hive, composed of six divisions, each contain-
ing a comb, if the young queen is as fertile
as she ought to be, the bees will be very ac-
tive in their labours, and disposed to make
great collections of wax. To induce them
towards it, an empty frame or division must
be inserted between two others, each con-
taining a comb. As all the frames are of

equal dimensions, and of the necessary width
for receiving a comb, the bees having suffi-
cient space for constructing a new one in the
empty division introduced into the hive, will
not fail to build it, because they are under
the necessity of never leaving more than a
third of an inch between them. Without
any guide, this new comb will be parallel to
the old ones, to preserve that law which es-
tablishes an equal distance throughout the
whole surface.

If the hive is strong and the weather good,
three empty divisions may be left at first
between the old combs; one between the
hist and second, another between the third
and fourth and the last between the fifth and
BWh Ihe bees will fill them in seven or
eight days, and the hive will then contain

mne combs. Should the temperature of the

222

ECONOMICAL

weather continue favourable, three new leaves
or divisions may be introduced ; consequent-
ly in fifteen days, or three weeks, the bees
will have been forced to construct six new
combs. The experiment may be extended
farther in warm climates, and in those where
flowers perpetually blow. But in our coun-
try, I have reason to think that the labour
should not be forced more during the first
year. Plate I. fig. 1, 2.

From these details, you are sensible. Sir,
how preferable leaf hives are to those of any
other construction, and even to the ingenious
stages described by 31. Palteau ; for the bees
cannot be compelled by means of the latter
to labour more in wax than they would if
left to themselves; whereas they are obliged
to do it by inserting empty divisions in the
former. Neither can the combs constructed
on those stages be removed without destroy-
ing considerable portions of brood, deranging
the bees, and creating real disorder in the
hive.

IMine have also this advantage, that what
passes within may be daily observed, whence
we can judge of the most favourable mo-
ments for depriving the bees of part of their
stores. AMth all the combs before us, we

CONSIDERATIONS.

223

can distinguish those containing brood only,
and what it is proper to preserve. The
scarcity or abundance of provisions is ex-
posed, whence the portion that is suitable
may be taken away.

I should protract this letter too much
%yere I to give an account of all my observa-
tions on the time proper for inspecting hives,
on the rules to be followed in the different

seasons, and the proportion to be observed
in dividing the riches of bees with them-
selves. I he subject would require a sepa-
rate vyork ; and, perhaps, I may one day en-
gage in it ; but until that arrives, I shall al-
ways feel gratincation in communicating to
cultivators, vvishing to follow my method,
those directions of which long practice has
demonstrated the utility.

Here I shall only observe, that we hazard
absolute rum of the liives. by robbing them
ot too great a in-oportion of honey and wax
Jn my opinion, the art of cultivating those
insects consists in moderately exercising the
privilege of sharing their labours; but in
coinpcns.atiou of it, every method must be
employed which promotes the multiplica-

sire to ^ ‘f ^ de-

t liocuiea certain quantity of honov

224

ECONOMICAL

and wax annually, it will be better to seek
it in a number of hives, managed with dis-
cretion, than to plunder a few of a great pro-
portion of their treasures.

It is indubitable that the multiplication
of these industrious creatures is greatly injur-
ed by privation of several combs, in a season
unfavourable to the collection of wax, be-
cause the time occupied in replacing them
is taken from that which should be conse-
crated to the care of the eggs and worms,
and by this means the brood suffers. Be-
sides, they must always have a sufficient pro-
vision of honey left for winter ; for although
less is consumed during that season, they do
consume a certain quantity ; because they
are not torpid, as some authors have con-
ceived.* Therefore if they have not enough,

* So far from being torpid in winter, when the thermo-
meter in the open air is several degrees below freezing, it
stands at 86 and 88° in hives sufficiently populous. The
bees then cluster together, and move to preserve their heat.

Swammerdam was of the same opinion, and observes, that
the heat of a hive is such, even in the middle of winter,
that the honey never crystalizes, or assumes a granulated
consistence, unless the numbers of tlie bees be too much
reduced. Farther, that when their queens are very fertile
they feed their young with honey also in the middle of
winter, watching over and heating them, and also heating
each other. “ I know not,” he continues, » whether there
are other insects liaving similar properties in common with

CONSIDERATIONS.

22.5

they must be supplied with it, which re-
quires a very exact proportion. I admit,
that in determining to what extent hives
may be multiplied in a particular country,
it is necessary first to know how many that
country can support, which is a problem yet
unsolved. It also depends on another pro-
blem, the solution of which is as little known,
namely, the greatest distance that bees fly

bees; for hornets, wasps, and humble bees, as well as flies
and butterflies, remain torpid and motionless during tl;e
whole winter.”

In the month of January M. de Reaumur found brood of
all ages in certain hives. The same thing happened to my-
self, when the thermometer stood in my hives at about 93°.

Now that I am on the subject of thermometrical observa-
tions, I may cursorily remark, thatM. Dubost of Bourg en
Bresse, in a memoir otherwise valuable, is of opinion '’tliat
the larv® cannot be hatched below 10f°. I have repeatedly
made the experiment with the most accurate thermometers
and obtained a very different result. When the thermome.^
ter rises to lOl, the heat is so much greater than the eggs
require, that it is intolerable to the bees. M. Dubost has
been deceived, I imagine, by introducing his thermometer ••
too suddenly into a cluster of bees, and the mercury has
rose higher, from the agitation excited among them than
t should naturally do. Had he kept it there, he would
lave soon seen it fall to between 95 and 97, which is
e usual tempera^ture of hives in summer. In August

94^7did'noTrL the open air stood at

ThVbJ had it 1 ’

226

ECONOMICAL

to in collecting their provender. Different
authors maintain, that they can fly several
leagues from the hive. But according to
the few observations I have been able to
make, this distance seems greatly exag-
gerated. It appears to me that the radius of
the circle they traverse does not exceed half
a league ; and as they return to the hive
with the greatest precipitation whenever a
cloud passes before the sun, it is probable
that they do not fly far. Nature, which has
inspired them with such terror for a storm,
and even for rain, nndonhtedly restrains
them from going so far as to be too much
exposed to the injuries of the weather. I
have endeavoured to ascertain the fact more
positively, by transporting to various dis-
tances bees with the thorax painted, that
they might he again recognised. But none
ever returned that I had carried for twenty-
five or thirty minutes from their dwelling,
while those at a shorter distance liave
found their way hack. 1 do not state
this experiment as decisive. Though bees
generally do not fly above half a league, it
is very possible that they go much farther
when flowers are scarce in their own vicini-
ty. An experiment to he conclusive must

CONSIDERATIONS.

be made in vast arid or sandy plains, sepa-
rated by a known distance from a fertile re-
gion.

Thus the question yet remains undecided.
But without pronouncing' on the number
of hives that any district can maintain, I
shall remark, that certain vegetable produc-
tions are much more favourable to bees than
others. ISIore hives, for example, may be
kept in a country abounding in meadows,
and where black grain is cultivated, than in
a district of vineyards or corn.

Plere I terminate my observations on bees.
Though I have had the good fortune to
make some interesting discoveries, I am far
from considering my labour finished. Se-
veral problems concerning the history of these
insects still remain unsolved. Experiments
which I now project may perhaps throw
some light on them ; and I shall be ani-
mated with much greater hopes of success,
if you. Sir, will continue your counsels and
direction. I am, with every sentiment of
gratitude and respect, Francis PIuber.

Fregny, 1st October 1791.

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PART II.

CHAPTER L

SCHIRACH’S DISCOVERY CONFIRMED,

Perhaps it may seem extraordinary to re-
cur to facts vitli v’liicli the reader has been
already occupied, and those apparently cor-
roborated by our own experiments. But
the interests of truth call for the defence of
a faithful observer, to whom our knowledge
of bees is indebted for its greatest progress,
and whose -reputation has been so outrage-
ously assailed.

Schirach began to undermine the opinion
of Swammerdam, wherein both IMaraldi and
leaumur concurred, that workers were not
only sterile, but absolutely neuter. How-

230

schirach’s discovery

ever, by repeated experiments, lie proved
that bees at all times can procure a queen
to themselves, provided they have comb con-
taining larvae three days old in the common
cells ; whence he concludes, that workers are
originally of the female sex, and that no-
thing but certain important conditions, such
as a particular kind of food, and more spaci-
ous lodgement, are requisite for their con-
version to queens.

Views so adverse to those generally enter-
tained were received with enthusiasm, on
the one hand, and with distrust on the other.
Notwithstanding it was admitted, that bees
might obtain a queen, because Schirach had
shown this result from experiments scrupu-
lously executed, and in the presence of in-
telligent persons, it was maintained that the
eggs had been laid in common cells ; and the
conversion of a worker’s worm into a royal
larva was denied. His experiments and
his arguments were resumed in vain : The

same objection was always urged against him,
though observers, aided by the best micro-
scopes, were unable to discern any difference
among the larvEe, from which he could at
pleasure produce cither a common bee or a
queen.

CONFIRMED.

231

iM. Bonnet, whose sanction, as that of a
great naturalist, was anxiously desired by
Schirach, requested me to repeat his expe-
liments; and in doing so, I recognised the
truth of the Lusatian observer’s assertion. I
added new illustrations, along with very con-
vincing proofs of the disputed conversion;
but I felt with him, that the establishment
of such important facts rested on ascertain-
log the sex of the workers.

. I reasoned thus Iliem’s discovery of
fertile common hees, which has been con-
firmed by my own observations, leads me
to anticipate that the whole class of work-
ers belong to the female sex. Nature does

00 mg y leaps. Fertile workers lay none
hut male eggs; m this resembling queens
whose fecundation has been retarfed 'Td!
vincing anotlier step, they may remain ab-
solutely sterile while they are not the less fe-
males originally. Too many valuable pro-

1 tics, too much industry and activity be-
' "S to common bees, to permit my^eom
c uc mg them of monstrous conformation

t ct”::;^, a™ their z

stmet and their structure, to permit mvcoii.

“ t'><! outcasts of the species.

232 schirach’s discovery

or imperfect beings, when compared with
queens.

Nothing is more repugnant to reason
than an actual metamorphosis ; and the
theory of it once entertained is now ex-
plained by evolutions. If the queen has
organs corresponding to those of the com-
mon bee, they are modified in such a man-
ner as not to fulfil any of the same func-
tions ; and in saying the conversion of the
one to the other is improbable, we shall
be right. But in admitting the two in-
sects to be of only one kind originally, and
that their species is of the same individu-
ality, we are justified in believing they may
become either workers or queens. Some
may imagine that a queen in the egg has
become a worker from certain circumstances;
others may equally affirm that the worker
was originally there, from which certain
modifications have formed a queen ; for we
cannot avoid thinking that the faculties
and organs peculiar to the common bee
pre-exist their evolution. We are thence
bound to conclude, that this being, whicli
as yet is neither worker nor queen; that
the worm, before three days elapse, contains .

CONFIUMED.

233

alike the germs of the insect which shall
prove industrious, and of the insect suscep-
tible of prodigious fecundity; — the germs
of the organs of the two animals, the in-
stinct of tlie common bee, and that of the
queen not developed, but capable of being
so, according tq the mode in which they
shall be reared, may operate. In the one
case the productive faculties will be repress-
ed, or will remain without being unfolded ;
in the other the industrial properties will
continue in a similar condition.

Perhaps nature presents beings of an in-
termediate kind between these two extremes,
participating of the essence of queens, and
of the qualities of workers ; and hence the
fertile common bees, and the small queens
observed by Needham. It is easier to com-
prehend how certain faculties, and their cor-
responding organs, maybe annihilated, than
how they may be spontaneously created ; and
It is on this my theory is founded.

_ It may be opposed, indeed, by the ojDposite
instincts of the workers and the queen in
the same hive, in relation to other queens ;
tor the workers entertain a kind of love for
then- mother, and render her the most as-
I sit nous attention, whereas queens are re-

234

SCHIRACll’S DISCOVERY

ciprocally aiiiinatecl by the most implacable
animosity. But do we knoAv how far any
sensation may be evolved in insects by cir-
cumstances ? Let me cite one example.

Very remarkable facts came under our
notice in a republic of humble bees, where, as
well as among domestic bees, there are three
kinds of individuals. Several of the workers
which, until a certain epoch, had lived in
the best intelligence with the mother of the
colony, having become fertile, they exhibit-
ed symptoms of the most violent jealousy.
Some fell victims to the rage of others; and
we saw the principal female perish by the
stings of those which she had j^roduced.
Therefore, if such rivalship can arise among
workers after participating of fecundation ;
if their affection for their companions and
for their mother is convertible in an instant
to hatred, the objection deducible from the
different instinct of queens and workers, the
strongest that may be urged against their
primitive identity, is reduced to its proper
estimate. After such a trait, who will deny
that instinct is susceptible of change ?

INIy speculations regarding the sex of
workers at length received the most uueX'
pected confirmation,

CONFIRMED.

235

History of JBlcicli Sees. — In the year
1809 we remarked something particular in
the treatment of certain bees by their com-
panions at the entrance of a hive. On the
20th of June, a cluster in such irritation,
that we durst not attempt to separate the
bees, attracted our attention. Night ap-
proaching, prevented us from ascertaining
the cause of this assemblage ; but on tlie
subsequent days, we frequently observed the
bees occupied in defending the same hive
against the entrance of some individuals
whose external appearance was not absolute-
ly similar to that of ordinary workers. Seizing
several of them, we found that their differ-

ence consisted only in colour ; that they were
less downy on the thorax and abdomen,
which gave them a blacker aspect ; but as
to the limbs, antennas, body, and size, the
whole external form presented perfect re-
semblance to common bees.

Their daily presence at the entrance of
the hive plainly denoted that they were ex
relied from it by the workers ; and in the
combats between them, the latter always had
he better. The common bee killed its ad-
1-ediicing it to such a state of
akness as to be incapable of resistance,

236

schirach’s discovery

carried it off in its teeth to a great distance
from the hive.

We caught a number of the black bees,
and introducing them into a vase, they
speedily darted on each other, and were re-
ciprocally killed. Some were confined in a
glass vessel along with workers from the
same hive ; but no sooner had the latter ob-
served them, than they were attacked and
destroyed.

Every day the proscribed bees appeared
more numerous. Once driven from the
hive, they never returned, for when the
sting spared them, they perished of hunger.

This singular scene continued during the
whole remaining part of summer. Some-
times the black bees were not so cruelly
treated by the workers, and then they seem-
ed modified a little differently from the for-
mer: their animosities were abated, nor did.
they repeat their mutual encounters. But.
the rigour of the common bees was very
soon resumed against them, and they were
expelled anew.

A'Ve could not ascertain whether all the'
brood of this hive was attacked by that ma-
lady or particular state, rendering the black .j
bees odious to their companions: and ob--i

CONFIRMED.

23.7

serving their number augmenting successive-
ly during some weeks, we had reason to be-
lieve that the whole oiFspring of the queen
was affected. Black bees, however, w’ere no
longer seen in the end of September. The
hive had apparently suffered from the exile
of so many individuals : it was w'eaker than
previously ; yet we were encouraged as to
the condition of the colony, when satisfied
that the queen had not lost the faculty of
laying eggs which produced workers perfect
in every respect.

From the month of April in the follow-
ing year this hive became the subject of
observation ; not a black bee appeared. The
increase of workers was so great as to pro-
mise a swarm ; but here we were disa2ipoint-
ed: therefore, our opinion was corroborated of
the anomaly, whatever it might be, having
affected only part of the eggs.

Was the queen entirely cured of this
disposition to produce individuals of peculiar
conformation ? Was the vice hereditary,
and what effect would it have on her fe-
male j)rogeny ?

It was proved that she had not been entire-
ly cured without relajise ; for in ]811, that
IS, two years after the bees originated, they

238

schirach’s discovery

re-appeared in great numbers under the same
character and circumstances. During 1812
the hive threw a very fine swarm ; and as
the old queen always accompanies the new
colony, we were not long of remarking
the defective bees at the entrance of her ha-
bitation.

But it was more singular still to witness
the same phenomenon in both hives at once.
The old one had swarmed on the third of
June, and on the second of July we per-
ceived black bees, at the entrance, which
evidently could not belong to the brood of
the queen that had jiassed into the new
hive. Thus, the progeny inherited the
vice of the parent. We again observed the
maltreatment of black bees at the entrance
of the same two hives in 1813, but there
were few of them.

Only a sketch of this subject is given
here. Our object is to arouse the attention of
naturalists, that a combination of fiicts may
be obtained, for enlarging the history of im-
perfect bees.

I was anxious to investigate whether any
thing external or internal indicated tlic
evolution of sexual organs in the proscribed .
insects ; for I conceived that if they were

COXFIRMED.

239

real females, their presence might disquiet
the common bees in respect to their queen,
and perhaps their expulsion from the hive
was to protect her from such rivals.

Dissection alone could verify the fact, but
it lequiied to be skilfully done. young
lady. Miss Jurine, who had devoted her
time and the liberal gifts of nature to simi-
lar studies, undertook this delicate task for
me. ^ Already did she rival Lyonet and
INIerian, yet we had soon to deplore her
loss.

IMiss Jurine, with her peculiar sagacity,
proceeded to examine whether any difference
from common workers was exhibited by the
external organization of the black bees;
but their identity seemed complete, except-
ing, as had occurred to ourselves, in a smaller
quantity of down on the thorax and abdo-
men. However, when this expert naturalist
extended her researches farther, and adopted
suitable preparations, she discovered two
distinct ovaries wherein no eggs were per-
ceptible indeed, but analogous in form and
.substance to those of queens, though not
so manifest to the observer. Plate II. fia- s
At first wc concluded that a solution

ot the question had been obtained.

240

schirach’s discovery

did not anticipate a more important dis-
covery, which was to overturn our conjec-
tures as to the persecution of the black bees,
and which would unveil a mystery long ob-
scuring the history of this tribe of insects.

A great number of common bees being
taken indiscriminately from before the en-
trance of their hive, ovaries resembling
those of the black bees were found in the
body of each, after having been kept two
days in brandy to darken the transparent
membranes.

The value of this discovery must be felt
by those who have followed the progress
and the vicissitudes of the history of bees ;
and by those who have read the objections
against Schirach’s theory, which are found-
ed exclusively on the absence of ovaries in
workers. Thus, the belief of neuters among
bees entirely vanishes, at the same time
that the most interesting phenomena of
their organization is exposed.*

The like should extend to all insects,
among which neuters have been described,

* M. Cuvier thinks he has observed minute chaplets
in common bees resembling those in the oviducts of queens,
u’hich confirms our opinion of their being females, whose
organization is not developed. Logons d' anatomic com- •
■jiarce, tom. v. p. 198.

’Sx.

Ri', J

£w.^y Tf'MJ^izarj-

CONFIRMED.

241

tliat is humble bees, wasps, and ants ; for,
according to the observation of a great na-
turalist, the more important any organ is
to the animal economy, the more general
ought to be its existence.

AVe were not entitled to decide on the sex
of the whole species from the fertility of in-
dividuals, until discovering the ovaries of
working bees. But the concurrence of these
two peculiarities being established, wher-
ever one shall manifest itself under the
same circumstances, the existence of the other
may be presumed from analogy.

According to the observationsof Biem, and
in the former part of this work I think to have
proved the fact, there are sometimes fertile
workers in a hive, but never laying any except
Uie eggs of males. I have shown that the
cause to which we should ascribe their ex-
istence centers in a certain nutriment, re-
sembling the sustenance of queens, operatin<>'
^ remarkable a change in their constitution
Their external characters are the same as
those of tlic workers, but the smallness of
their number renders it almost impossible
tor us to observe the conduct of the latter
towards them. I’erhaps, by obtaining some
in the frames used by Schiracli for produ-

M

242

schirach’s discovery

cing queens, and removing the royal cells at
a suitable period, their habits, amidst a very
few workers, might he studied. Then also
would be the time to examine whether the
fecundation of workers is attended by the
same circumstances as that of queens.

Elsewhere we have explained the exist-
ence of fertile workers among humble bees,
and described the jealousy awakened by
maturity, their rivalship and resentments.^
We have not recognised any other difference
than size between the small mothers and
real females ; and having been then unable
to ascertain whether the fertile workers had
a progeny of both sexes, we lately engaged
in researches to investigate the point.

A nest of red and black humble bees, the
hcemorhoidalis of Linnasus, was established
in an ordinary box in a window. The mo-
tion and agitation of the bees in the after-
noon, their rivalship, and, finally, their lay-
ing, soon demonstrated that fertility was not
restricted to the mother of the colony alone.
But as the latter frequently laid her eggs in
the same cells as the workers, the result of
their fecundity could not be ascertained

* Transactions of the Linnsean Society, Vol. VI.

CONFIRMED.

243

i

i

I

without an absolute separation, whereby the
product of each might be known.

Therefore, a fragment, containing nothing
but brood, was detached, and left in a box
where the humble bees were accustomed to
retire into their nest, and the mother, toge-
ther with the workers remaining at home,
were carried with the other portion to a dis-
tant window. I calculated, on the return
of the absent workers now foraging in the
country, to people the portion wanting in-
habitants. In fact they did so, and lodged
themselves on the isolated fragment, whicli
had been substituted for combs and brood,
though they did not seem insensible of the
change. I was in hopes, also, that some of
these workers might prove fertile ; nor were
my prospects disappointed : for, in the after-
noon of the same day of forming this separated
or artmcial swarm, they prepared a cell for
receiving their eggs, and I saw many of them
lay. 1 he number of eggs multiplied daily •
larval were soon hatched, and they trans-
formed to nymphs, which, in a month, be-
came humble bees. Having most carefully
exammed them as they proceeded from
their coccooiis, I found that all were males

244 schirach’s discover!^

They resembled those originating from the
eggs of the female in every respect ; they
were equally large, and coloured after the
same manner.

The red and black humble bee had been
selected for this experiment, because bands
of green hair on the thorax, and a green spot
on the front, more easily distinguish the
males than any others.

I could not be deceived regarding the
fact ; and I can affirm, that neither worker
nor female was produced in the one nest,
while the other had as many females as
males. Almost the whole workers were fer-
tile in this nest except a few very small
ones; at least, I did not ascertain 'that
these were so.

Here, therefore, is seen a great analogy
between humble bees and the honey bee.

An additional example of the generality
of the law of nature against neuters was
obtained by INI. Perrot. Studying a hang-
ing wasps’ nest with that scrupulous at-
tention which denotes the genuine natu-
ralist, he observed some of the w'orkers suc-
cessively laying eggs. Although an acci-
dent prevented him from following out the

coxnr.MED.

245

complete evolution of the nymphs, careful
inspection convinced him that all were of
the male sex.

Ants likewise afford a striking analogy
to what has been described. We never have
seen the workers lay indeed, but we have
witnessed their sexual union, as can he at-
tested by several members of the Geneva
Society of Natural History. The death of
the workers always followed the approaches
of the male; therefore, their conformation
does not admit of their becoming mothers ;
but the instinct of the males proves them to
be females.

All these facts concur in demonstrating
that there are no real neuters in this class
of insects, which would have interrupted
the chain of nature, for I know not that it
exists in any other kind. Sometimes both
sexes are conjoined in one individual, but

it seems adverse to nature that there should
be neuters.

How IS It to be explained why the work-
ers among a republic of insects, when fecun-
dated can lay none but the eggs of males ?
hat reason shall we give for it ? They
ovaries similar to those of the queen,
which they have sprung, yet they pos-

have

from

246 SCHIRACH’S DISCOVERY

sess only a partial fertility. It is equally diffi-
cult to understand why those queens impreg-
nated later than three weeks after their orio:iii
produce the eggs of males alone. Doubtless
some connection subsistsbetweenthetwo facts.

In attempting to explain Schirach’s dis-
covery, M. Bonnet conjectures that the evo-
lution of germs is effected by the action of
the prolific matter as a stimulant, as a sub-
stantial nutriment suitable for that purpose :
and he supposes that a certain quality of food,
which is administered more copiously, may
unfold those organs in the larvae of bees that
never would have appeared without it.

Is it impossible that this kind of subsist-
ence, so different from what is received by
common larvae, being administered too late
or too sparingly to] the larvae originating near
< the royal cells, may have consequences simi-
lar to the retarded fecundation of queens?
Have not the fibres of the ovaries attained
too great rigidity to be affected by the pro-
lific matter on some of the eggs ? Or does
not the royal aliment, by an opposite opera-
tion, produce such an energy as to destroy
this equilibrium ?

In a work on the 3[anagement of Bees
in Favignana, by IMonticelli, a Xcapoli-

CONFIIIMED.

247

tan professor, Schirach is reproached with
claiming the formation of artificial swarms
as his invention, and with having borrowed
it from the customs of a small colony inhabit-
ing a rock in the Mediterranean, near the
coast of Sicily. Schirach, however, was far
from giving himself out as the author of a
method practised long anterior to his own
time in the country where he lived. Prac-
tice has always preceded theory; its success
leads to discovery of the truths whereon it
is founded ; and acquaintance with these
truths in its turn establishes the waverine:

o

advances of cultivators.

Schirach relates, that being obliged to
use a great quantity of smoke for driving
bees to the top of their hive, in order that
he might cut out the brood, they were so
much annoyed by it, as to depart in num-
bers unobserved, and accompanied by their
queen. He sought the neighbourhood in
vain for the fugitives ; and abandoning the
hive as lost for want of a queen, he resolved
to rear a new one, by introducing a comb
consisting of three kinds of brood, such as that
of which he had just deprived them. Next
morning, preparing to clean the hives which
he had divided on the preceding evening.

24S

SCIURACH’s DISCOVERi"

liG observed a cluster of bees the size of an
apple, on the prop of the one whose queen
had fled. Here he discovered a queen ; and
having carried her to the entrance of the
hive which had lost its own, she was imme-
diately surrounded by the bees, and treated
in such a manner as plainly announced that
she was their queen. “ What was my
astonishment,” he proceeds, “ when wishing
to introduce her among the combs, I saw
that the bees remaining had already planned
and almost finished three royal cells. Struck
with the activity and the sagacity of these
creatures to save themselves from impend-
ing destruction, I was filled with admira-
tion, and adored the infinite goodness of
God, in the care taken to perpetuate his
works ! Having carried away two of the
cells, to ascertain whether the bees would
continue their operations, I beheld next
morning, with the utmost surprise, that they
had removed all the food from around the
third worm left behind, on purpose to pre-
vent its conversion to a queen.”

Discovery of this kind of transformation
soon rendered the art of making artificial
swarms easier and less expensive. Sehi-
rach showed that its success depended on

CONFIRMED.

249

having a single cell occupied by a v\orm
three days old, instead of providing the
bees with large combs containing the three
kinds of brood, as before supposed necessary.
He proposed several improvements in procur-
ing artificial swarms, but he never thought
of claiming the invention.

The Italian author, wishing to gain cre-
dit to his country, and forgetting that the
great republic of science is engaged rather
with the utility and perfecting of discove-
ries, than involved in disputes [about their
origin, openly charges Schirach with borrow-
ing the plan of artificial swarms from Fa-
vignana. He affirms that the practice is of
such ancient date, that even the Latin names
are preserved by the inhabitants in their
procedure: but he acknowledges, at the same
time, that both the Turks and Greeks of the
islands in the Ionian Sea are acquainted
with it. T wo methods are followed ; the one
on a larger scale, more general and complete,
which was unknown to Schirach who imi-
tates the other. The hives of the Favigna-
nese are oblong wooden boxes, botli ends
of wliich are moveable. As spring is mucli
earlier in their region tlian witli us, tlicy
arc able to commence their operations in

250

schirach’s discovery

the month of March. The hive, therefore,
being transported to some distance from
the apiary, the farther end is opened, and
the bees are driven forward by means of
smoke, Avhen some of the corahs commonly
containing honey are cut out. Next they
are driven back to the farther end, and a
certain number of the combs are taken
from the anterior part, some empty, and
some full of brood of all ages. The latter
are then put into a new hive, and by means
of rods inserted through them from without,
they are established in the same order that
they held in the old hive. After this is
done, the new hive is conveyed to the sta-
tion which was occupied by the old one,
and the latter is removed fifty paces from
the apiary. The bees now returning from
the field, and finding a hive like that which
they had left, lodge themselves there, rear
the brood, and prosper.*

The success of this operation, which very
much resembles Schirach’s, amply confirms
his theory : and we must reject JMonticelli’s
prejudices, for his work is otherwise a good

* Probably tbcsc obloug boxes are laid horizontally on
their props, and the entrance is made at tlie lower part of
tlic end in front. — T.

CONFIRMED.

251

practical treatise on the management of
bees, and the formation of artificial SAvarms.

We shall here take an opportunity of
speaking of a very ingenious method, though
somewhat different, followed by M. Lombard,
a great cultivator of bees, and the author of
an excellent treatise on their practical eco-
nomy. It is the reverse of that adopted in
Favignana to form an artificial swarm ; for
he obtains, so to speak, a natural precocious
swarm. The hive destined for that purpose
is carried to a dark place, already containing
another, in which the new swarm is to be re-
ceived. Part of the bees, together with their
queen, are forced to take refuge in this new
lodging by means of smoke ; after which the
old hive is replaced in its former situation,
that it may be repeopled by all those return-
ing from the fields ; and the new one is es-
tablished at a suitable distance from it in
the apiary. This colony, possessing a queen,
can prosper without any foreign aid, and
enjoys the advantage of the flourish of the
fruit trees, which is sometimes denied to the
earliest natural swarms. The cylindrical
form of M. Lombard’s straw hives facili-
tates the process. For tlie details I shall

252 schirach’s discovery confirmed.

refer to his work itself, which is of essential
utility to every cultivator of bees.

This method being founded on the pro-
duction of a queen in a hive containing only
brood, is a farther corroboration of Schirach’s
doctrine, for its long practice has been al-
ways attended with success.

Experiment and theory, therefore, concur
in demonstrating that the larva of a bee
may become a queen or a worker, according
to circumstances. A female always appears,
whether possessing the physical qualities of
maternity as the fecundity of queens, or
the conservative properties displayed by
w'orkers in regard for the young. This par-
tition of industry and courage, on the one
hand, and of prodigious fertility, on the
other — this partition, originating from the
mysterious rearing of the larvae, is among
the finest subjects for contemplation which
natural history affords. Thus we owe one of
the most curious discoveries which has em-.
bellished science to the penetration and per-»
severance of Schirach.

SENSES OF BEES.

253

CHAPTER II.

ON THE SENSES OF BEES, AND PARTICU-
LARLY THAT OF SMELLING.

T HE infinite varieties exhibited by the dif-
ferent tribes of insects and other animals,
excite oiir belief that the same impressions
are not produced on their sensations as on
man. Their faculties and nature not ad-
mitting the light of reason, they ought to
be^ directed by other agents. Perhaps the
opinions which we form of their senses,
founded on those peculiar to ourselves, are
incorrect; for some of a more subtile kind,
or modified otherwise, may present objects
under an aspect unknown to us, and occa-
sion impressions that we are strangers to.

254>

SENSES OF BEES.

organization suitable to its tastes anti habits,
has the power of modifying its senses be-
yond any knowledge of them that w^e can
acquire ? Cannot the Author, who framed
those five grand avenues W'hereby all our
notions of the physical world are gained,
open up channels of another kind to beings
less favoured with judgment? The assist-
ance of art enables us to determine regard-
ing objects not so immediately within the
sphere of the senses, as chemistry and
physics prove in a thousand examples. Ther-
mometers, menstrua, and re-agents, by wdiose
means w^e detect the nature of things elu-
ding our senses, are so many new organs.
Thus there may be other sensations and
means of viewing objects; and it is not re-
pugnant to admit the existence of such iu
beings so differently constituted from us in
many respects.

Insects living in a republic, among which
bees certainly occupy the highest rank, often
exhibit traits inexplicable, even admitting
them to be endow ed with the same senses as
ourselves, which renders it so difficult to
penetrate the secret impulse actuating them.
Yet they possess sensations of a less subtile
description ; and as it is expedient to gain “as

SENSES OF BEES.

255

t

much knowledge of their powers as possible,
Ave should be wrong to neglect what comes
within our reach, and whereby we can judge
at least of their appetites and aversions.

Sight, feeling, smell, and taste, are the
senses commonly ascribed to bees ; for hither-
to we have no proof that they enjoy the
sense of hearing, although a custom very
prevalent in the country seems to support
a contrary opinion : I allude to the practice
of striking a sonorous body at the moment
of departure, in order to restrain a swarm
from flying aAvay. But, in compensation,
how great is the perfection of their organ of
sight ! Since from a distance the bee recog-
nizes its habitation amidst an apiary of nume-
rous others resembling it, and returns in a
straight line with great velocity, we must
suppose that it is distinguished by marks
escaping our notice. The bee departs and
flies straight to the most flowery field : hav-
ing ascertained its course, it is seen tra-
versing it as directly as the flight of a cannon
or a musquet ball. Its collection being made,
it rises aloft in the air to reconnoitre its hive,
and returns with the rapidity of lightning.*

j * I lie sense of vision among insects seems of an obscure
i'omphcatc'cl nature, and that of the tribes of niollusca and

256

SENSES OF BEES.

The sense of feeling is still more admir-
able : it is substituted for, and completely
supplies the want of sight in the interior
of the hive. The bee constructs its combs
in darkness ; it pours its honey into the
magazines, feeds its young, judges of their
age and necessities, recognizes its queen, all
by means of the antennse, which are much
less adapted for becoming acquainted with
objects than our hands. Therefore, shall
we not grant to this sense modifications
and perfections unknown to the touch of
man ?

Perhaps the sense of taste is the least
perfect of those enjoyed by bees ; for this is
one which in general seems to admit of se-
lection. But, contrary to the received opi-
nion, the bee certainly displays little choice
in the honey that it collects.. It is not
repulsed by plants, which to us are the
most hateful and odious : poisonous flowers
are not excluded from its search ; and the

vermes is still more indefinite. Naturalists generally be-
lieve that in the former it resides both in the large reticu-
lated or granulated eyes on each side of the head, and in-
the three ocelli or specks on its upper part. It is remark-
able that the eyes of bees are covered with hair. These
insects, blinded by an opaque varnish, fly at random, or risa
vertically, until disappearing in the air. — T.

SENSES OF BEES.

257

honey of some of the American provinces is
even said to be of the most deleterious na-
ture. Xeither do bees despise the secre-
tions of aphides, notwithstanding of so im-
pure an origin : nor do they testify greater
nicety in the quality of their wnter, for the
most corrupted marshes and ditches seem
prefeiTed to the most limpid streams, nay,
to dev/ itself.

Nothing, therefore, is more unequal than
the quality of honey. The produce of one
district differs from that of another: the
honey of spring is unlike that of autumn :
nor do the contents of a certain hive al-
ways resemble those of the hive adjoining.

If bees have little choice iii their nutri-
ment, and are not delicate in regard to the
quality of honey, they are far from testify-
ing the same indifference as to the quantity
contained in flowers. They constantly re-
sort where most is to be found. They quit
their hive much less in regard to the fine-
: ness or temperature of the weather, than ac-
cording to their prospects of a more plenti-
ful or scanty collection. When the lime-
tree and black grain blossom, they brave
the rain, tliey depart before sun-rise, and
I return later than ordinary. But this acti-

258

SENSES OF BEES.

vity relaxes after the flowers have faded;
and when the enamel adorning the mea-
dows has fallen under the scythe, the bees
remain in their dwelling, how^ever brilliant
the sun-shine. To what shall we ascribe
their perceptions of the abundance or defi-
ciency of honey in the flowers, which the
whole colony appears to possess without
leaving their habitation ? Does one sense,
more subtile than the others, namely smell-
ing, warn them of it ?

There are some odours, such as the smoke
of tobacco, and every kind of smoke, to w'hich
bees are averse, and some that are agree-
able to them; but of all odorous substan-
ces, honey is that which has the most power-
ful attraction. Perhaps others have a si-
milar influence, only in as far as announ-
cing the presence of what seems so valu-
able.

To ascertain whether the appearance of
the flowers or the odour of the honey ap-
prises bees of its presence, we placed honey
in a window, near a hive, "where the shut-
ters, almost close, still allowed their passage
if they chose. A\^ithin a quarter of an hour
four bees and a butterfly had insinuated
themselves, and we found them feeding on

SENSES OF BEES.

259

it. Although this was sufficiently con-
clusive of my opinion, I wished it better
confirmed ; therefore I had four boxes,
different in size, shape, and colour, made,
(Witli small card valves, corresponding to
apertures in the covering. Honey being
put into them, they w^ere placed at the
distance of two hundred paces from my api-
ary. In half an hour bees were seen arriv-
ing ; they carefully traversed the boxes, and
soon discovering the openings whereby they
might introduce themselves, they pressed
against the valves and reached the honey.

We may thence judge of the extreme
delicacy of smelling in these insects. Not
only was the honey quite concealed from ,
[ view, but its emanations, from it being co-
i vered and disguised in the experiment, could
I not be much diffused.

Flowers frequently exhibit an organization
I resembling that of the valves we allude to.

The nectarium of several classes is situated at
1 the bottom of a tube, inclosed or concealed
among the petals ; nevertheless the bee finds it
out. Still its instinct, less refined than that
of the humble bee, affords fewer resources.
The latter, when unable to penetrate the
flowers by their natural cavity, knows to

260

SENSES OF EEES.

make an aperture at the base of the corolla,
or even of the calyx, and to insinuate its
proboscis to the place where nature has si-
tuated the reservoir of honey. By means
of this stratagem, and favoured by the length
of its organ, the humble bee can obtain
honey where the common bee would reach
it with great difficulty. From the differ-
ence of the honey produced by these two
insects, we may conjecture that their collec-
tions are not made from the same flowers.
Common bees, however, are as much attract-
ed by the honey of the humble bee as by
their own. We have seen them pillage one
of their nests in time of scarcity, which had
been placed in an open box near an api-
ary, and carry the contents almost totally
away. Some of the humble bees remaining
after this disaster, still repaired to the fields,
and returned with the surplus of their pro-
vender to their ancient asylum. But the
plunderers, posting in pursuit, accompanied
them home, and never quitted them until
having obtained the fruits of their labour.
They licked the others, offered their trunks,
surrounded them, and departed only after
they had rendered up the saccharine fluid of ’
which they were the depositories. Ivo in-

SENSES OF BEES.

261

jury was done to the insect which thus af-
forded the bees their repast : the sting
never was unsheathed, and the humble bee
itself, accustomed to such exactions, yielded
up its honey and resumed its flight. This
remarkable kind of intercourse lasted above
three weeks. Wasps, invited by the same
attraction, did not become so familiar with
the original proprietors of the nest. At
night it was occupied by the humble bees
alone. At length they disappeared, nor did
the parasite insects return.

We have been assured that the same scene
may be witnessed between predatory bees
and those of weak hives, which is less sur-
prising.

Not only have these insects a very acute
sense of smelling, but to this property is
added the recollection of sensations. Honey
I had been placed in a window in autumn,

I where the bees resorted to it in multitudes!

It was removed, and the shutters closed diir-
I ing \\inter; but when opened again, on the
I return of spring, the bees came back, though
uo honey was there. Undoubtedly they
remembered it; therefore an interval of
I several weeks did not obliterate the impres-
I sion they had received.

262

SENSES OF BEES.

Let US now inqirire into the site or the
organ of this sense, whose existence has been
so well established.

Nostrils have not been yet recognised in
insects, nor do we know in what part of the
body they, or any organs corresponding to
them, are placed. Probably odours reach the
sensorium through the medium of a me-
chanism similar to our own : that is, the air
is introduced into some opening at the ter-
mination of the olfactory nerves : whence
we should examine if the stigmata do not
perform this function; or whether the or-
gan we are in quest of be not situated in the
head, or in some other part of the body.*
With the view of elucidating the matter,,
we made the following experiments : —

1. A pencil dipped in oil of turpentine,,
one of the substances most formidable to in--
sects,'was presented successively to all parts-
of the body of a bee, which did not appear;
in the least affected, whether on approach--
ing the thorax, abdomen, or stigmata of the
thorax. 2. We then took a very fine pencil,
that it might reach every point of the head,

* Certain apertures, generally called stigmata, appear
on each side of the body of insects, •which naturalists be- ■
lieve to be approjiriated exclusively for respiration. — 7’.

SENSES OF BEES.

263

and brought it near the antennae, the eyes,
and protruded trunk of a bee, in the act of
feeding, but in vain. It was otherwise on
carrying it near the cavity of the mouth,
above the insertion of the proboscis. At that
instant the bee receded, left the honey, and
beating its wings, while moving about in
much agitation, it would have taken flight
had not the pencil been withdrawn. Having
renewed its repast, we resumed the applica-
tion, always carrying the impregnated por-
tion near the mouth. The bee now quitting
the honey, fixed on the table, and fanned it-
self during some minutes. The organ of
smelling, therefore, seems to reside in the
mouth itself, or in the parts depending on it.

Bees not occupied in feeding appeared
more sensible of the odour of the turpentine.
They were affected by it at a greater distance,

S and speedily took flight, whereas, when so
; engaged with the trunk immersed in honey,
i several parts of the body might be touched
by the pencil without their withdrawing.

cie they absorbed by their greediness of
the honey, and distracted by its smell, or
were their organs less exposed ? This could
I be ascertained in two ways ; cither by covct-
I mg all parts of the body with a varnish,

264

SENSES OF EEES,

and leaving the sensible organ free, or leav-
ing the whole parts untouched excepting
that wherein the sense was supposed to re^
side.

The latter method appearing the more
practicable and decisive, we seized several
bees and compelling them to unfold the
trunk, filled the mouth with flour paste.
A\’'hen this obstruction was dry enough to
prevent them from divesting themselves of
it, they were released; none seemed to suffer
any inconvenience. They breathed and
moved with the same facility as their com-
panions. Honey did not attract them ;
they neither approached it, nor were they
affected, apparently, by odours otherwise
offensive to them. Pencils were dipped in
the oil of turpentine and cloves, in ether,
in fixed and volatile alkalis, and their points
insinuated very near the mouth. But the
odour of these fluids, which would have-
occasioned a sudden shock to bees in their
natural state, had no sensible effect on them.
On the contrary, several mounted on the
impregnated pencils, and traversed them as>
if unhurt ; therefore, they had at once losttlj
the sense of smelling, the site of which itJr
appeared to us resided in the mouth. ' i

SENSES OF BEES.

265

111 the next place, we wished to investi-
gate how bees were affected by different
-odours.

JMmeral acids and volatile alkali present-
ed on a pencil to the opening of the mouth,
made the same impression as the spirit of
turpentine, but acted with greater energy.
The effect of some other substances was not
so decided. On our presenting musk to bees
feeding before the entrance of their hive
they ceased, and dispersed a little, but with-
out precipitation or beating their wings.
'^^"e threw it pulverized on a drop of honey ;
into which the bees thrust their trunks,
but as if by stealth, and keeping at as great
a distance as possible. Had the drop not
been covered with musk, it would have disap-
peared in a few minutes ; now it had suffer-
ed no sensible diminution in a quarter of an
hour, although the bees had often plunged
their trunks amongst it.

Experiments were made with substances
which do not undergo any perceptible alter-
ation ; F or certain odours vitiate the air,
without directly affecting the nerves.

S. 1 ounded assa‘-fetida, whose odour is in-
supportable to us, being mixed with honey,
and put at the entrance of a hive, did not

N

266

SENSES OF BEES.

seem to annoy the bees. They greedily
absorbed all the honey in contact with the
extraneous particles ; they neither attempt-
ed to withdraw, nor vibrated their wings ;
and of the mixture only the molecules of
assa-fetida at last remained.

4. I remarked that bees going out to
the fields, and coming home, turned aside
in the air to avoid passing immediately
over a piece of camphor laid before the en-
trance of their hive.. Some being attracted
by honey on a card, I brought the camphor
tow'ards the mouth, while their trunks were
immersed among it. All took flight, but
after flying about some time in my cabi-
net, they settled beside the honey. While
absorbing it, I threw fragments of camphor
on the surface : they receded a little, still
keeping the end of the trunk inserted;
and w'e observed that at first they took
only that which was not covered with the
camphor. One vibrated its wings as -it
fed : Some agitated them seldom, and

others not at all. I covered the honey en-
tirely with camphor, and at that instant all
flew aw'ay. This card w'as carried to my
hives, and pure honey also put within reach i
of the bees on another. The latter w'as soon i

SENSES OF BEES.

267

discovered, and the honey consumed in a
few minutes : but an hour elapsed, before a
single worker approached the camphorated
card, when two resting on it, thrust their
trunks into the edge of the drop of honey.
Their number gradually augmented, and
two hours after it was covered by them, the
whole carried off, and only the camphor re-
mained.

These experiments prove, that if camphor
be disagreeable to bees, their repugnance
is surmounted by the attraction of honey ;
and that there are odours which repel them
to a certain degree without vitiating the air.

It is shown by many experiments also,
that the influence of odours on the nervous
system of bees is incomparably more active
in a close vessel than in the open air.

5. A bee having been satiated with honey
was put under a receiver containing a small
glass of spirit of wine, allowed to evaporate.
Immediately endeavouring to escape, it in-
cessantly vibrated its wings and sought an
exit for an hour. A constant tremor of the
limbs, wings, and proboscis, then became
perceptible: the insect, unable to stand, lay
I supine, and in this position advanced in a
j strange manner, using the four wings like

268 SENSES OF BEES,

oars or feet ; and all the honev swallowed
before exposure to the effluvia of the spirit
of wine was disgorged. Immersion twice
in cold water was beneficial to the bee, and
vinegar seemed to revive it, but it soon pe-
rished.

^Vindow flies and wood lice were destroy-
ed by the same effluvia ; however, a large
spider did not appear to be affected.

C, It was curious to ascertain the impres-
sion that the penetrating odour of the poi-
son of bees would have On the animals
themselves : therefore, the sting of one, with
its numerous appendages, being extracted
with pincers, was presented to workers in
a tranquil state, before the entrance of their
dwelling. Emotion instantly spread a-
inong them : none fled away, but two or
three darted against the sting, and one
wrathfully attacked ourselves. It was not
the threatening apparatus of the experi^
inent, however, which had irritated them;
for when the poison had coagulated on the
point of the sting, and on its appendages,
they seemed insensible of its presence, and
it might be offered with impunity. It is |
the odour of the poison alone that enrages !
them. Having put some bees into a glass i

SENSES UF BEES.

269

tube, close at only one end, we rendered
them half torpid, in order to prevent their
escape by the other. They were gradually
revived by the heat of the sun, and then
irritated with the beard of an ear of wheat.
All jn-otruded the sting, on the extremity
of which appeared drops of poison.

Their first symptoms of life, therefore,
w'ere attended by demonstrations of anger ;
and I doubt not, had they been at liberty,
they would have pierced each other or flown
on the observer. But they were retained in
the tube.

1 took them out one by one with pincers,
and confined them in a receiver, that my ex-
periment might not be disturbed. ^The
poison darted against the inside of the tube,
had left an unpleasant smell. When the
bees became sensible of this, from my pre-
senting the open extremity to groups before
their hive, they were agitated ; but their
agitation was not the result of fear, for they

testified anger in the same way as on the
former trial.

Ihus, certain odours not only act physi-
cally, but have an influence on the inclina-
tions of these insects ,■ ami licrc there doubt-
1®S commences a particular class of sciisa-

270

SENSES OF BEES.

tions eluding our researches, and of which
we can form but a confused conception.
Wliat a variety of impressions are produced
by smell on dogs of the chase !

In accounting for the conduct of insects
under special circumstances, the influence of
different sensations, modifying their habits
at the moment, must be appreciated.

Bees are impelled to flight by too high
a temperature, or repulsive odours. But if
invited by some other cause, as the attrac-
tion of honey, to remain, they know to pre-
serve their present enjoyments, and shel-
ter themselves from disagreeable sensations,
by agitating the air. -Hetained in their
hive by all the attractions which natiwe has
there combined for them, and unable to with-
draw from the surrounding mephitic gaz,
without abandoning their young together
with their stores, they have recourse to
ventilation for renewal of the air. To
w’hat can we ascribe the general tranquil-
lity, when a few individuals are occupied in
obtaining a salubrious atmosphere? We \
cannot suppose only a portion of them i
affected by a cause which has no operation i
on the greater number, ’\^"hen the air was H
not renewed in the manner desirable, how- I-

SENSES OF BEES.

271

ever, we have seen all vibrating their wings
at once, though this never occurs in the na-
tural state, when the vibrations of a few are
sufficient for ventilation.

Some insects of -the same species are more
immediately affected than others by the
same cause; which may ensue from circum-
stances or their occupations. Therefore,
after the air has been rendered of sufficient
salubrity by a certain number of ventilators,
the rest, not experiencing the sensation of its
impurity to an equal extent, abstain from
vibrating their wings, and yield to more
urgent pursuits. Should the number of
ventilating bees be suddenly reduced, the
workers first experiencing the alteration of
the air, will begin to vibrate, and their num-
ber will augment, until their united exer-
tions restore to this element the degree of
purity essential to respiration.

Such we conceive is the mode of establish-
ing the perpetual series of ventilating bees, for
no communication is observed betw’’een them.

I T. his hypothesis presumes an organization
i sufficiently delicate to detect the smallest al-
I teiation of the respired fluid, which may lose
I many degiees of its purity before we arc a-
I ware of it, though proving very noxious to us.

272

USE OF THE ANTENNA

CHAPTER III.

•v^ % -X ^-v%- % -v-v

RESEARCHES ON THE USE OF THE AN-
TENNA IN SOME COMPLICATED OPE-
RATIONS OF BEES.

E have examined the general relation of
the senses of bees to objects of immediate
utility. But it is extremely probable that
the sphere of their activity is not restricted
to distinguishing odours and the substances
these insects have to collect. The art of pro-
curing their materials and putting them in
operation, is only one branch of the history
of bees. Their conduct as a great society,
whose prosperity depends on variable ele-
ments, should offer civil connections, so to
speak, between all its members.

IN OPERATIONS OF BEES. 273

As their senses undoubtedly must have a
large share in the operations resulting from
this state of things, it is important to in-
quire what influence should be ascribed^ to
them when instinct seems adapted to the
most complicated circumstances.

One of the facts which appears the most
deserving of meditation and research, is the
formation of a queen in a hive that has lost
its own. Whether the worker he confident
or not of attaining its purpose, by changing
the food and form of the cell destined for a
royal worm, its conduct certainly displays a
refinement of instinct, of which we could
not believe an insect capable.

In given circumstances, though very rare-
ly, the colony runs the hazard of destruc-
I tion by losing its queen. The precautions
I of the bees 'avert this disaster. But how
1 are they invited to it ? How can the absence
• of the queen guide tiiem to so complex
1 and remarkable a proceeding, as choosing
I larvs of that age, which will admit of what
is to be accomplished ?

^ Hid the absence of the queen alone pro-
duce these effects, we should observe new
! cells constructed immediately after her dis-
i appeal ancc. But the bees do not seem to

S74 USE OF THE ANTENNiE

miss her ; the works of every kind advance ;
order and tranquillity are uninterrupted ;
nor is it until an hour from her departure
that disquiet testifies itself among the work-
ers, when they hurriedly traverse the hive,
and are no longer engrossed by care of
the young. Yet the first symptoms of
agitation are not felt every where at once.
They originate on a single portion of the
combs ; the disturbed bees soon quit their
little circle, and meeting their companions,
the antennae are reciprocally crossed, and
they slightly strike them. Those receiving
the blow of the antennae become agitated in
their turn, and carry trouble and confu-
sion to other places ; the disorder augment-
ing in rapid progression, gains the opposite
side of the comb, and is at length dissemi-
nated throughout the colony. Then are
the workers seen running over the combs,
rushing against each other, and impetuous-
ly hurrying towards the outlets of the hive.
From thence they disperse themselves
around. They return and depart repeated-
ly ; the buzzing is very great within, and it
increases with the disorder of the bees.
This continues, however, only two or three *1
hours, seldom four or five, but never longer. .|

IN OPERATIONS OF BEES. 275

What is the source of sucli an efferves-
cence ? How do tlie bees gradually reco-
ver their natural state, and resume an in-
terest in all that seemed to have become in-
different to them ? How are they recalled
spontaneously to their young, which for
some hours they had abandoned ? And
whence is their inducement afterwards to
visit the larvag of different ages, to select
from among them some to he reared as
queens ?

In twenty-four hours subsequent to the
departure of the common mother, we see
that the bees have been labouring to repair
her loss. The cells occupied by the favour-
ed larvse are always of the smallest diame-
ter; their form has not yet been altered, but
they are already distinguished by the quan-
I tity of paste in them, which infinitely ex-
I ceeds the portion devoted to the larva? of
woikers. It follows that, by this profusion,
instead of being lodged at the bottom of the
cells, they are now brought very near the ori-
I fice. Probably it is to give these larva? an
elevated position that the bees accumulate
the paste behind them ; and it is proved not
to be for food, by our finding it entire when
the worm has descended into the pyramidal

276 USE OF THE ANTENNA

prolongation by whieli the workers terminate
their abode.

W e, therefore, know what larvas are des-
tined for queens by the aspect of the cells
occupied by them, even previous to their en-
largement and alteration of shape ; and by
attending to this, we discover easily, at the
end of twenty-four hours, whether the bees
have resolved to repair the loss of their
queen.

In whatever manner they ascertain her
removal, it is evidently known to all within
an hour ; they feel themselves in a distress-
ing condition, and seek for the object of
their solicitude. It may be asked, IVhether
it is by means of smell, touch, or any un-
known sense; or whether from the bees
communicating w'ith each other by repre-
sentative signs? Experiment might dis-
close the fact.

I divided a hive into two portions by
means of a grating, executing the operation
with such expedition and delicacy, that the
smallest agitation was imperceptible, nor
Avas a single bee injured. The bars of the ;
grating admitted the free circulation of air, jj
but were too close for the reciprocal passage ij
of the bees. I did not know which half

IN OPERATIONS OF BEES. 277

' contained the queen, but the tumult and
buzzing in No. 1 soon assured me that she
was in the other, No. 2, where quietness
and tranquillity prevailed. Still preserving
the circulation of the air, I closed the en-
trance of both, that the bees, seeking for
their queen, should not find her. In two
hours they calmed, and the wonted order
was restored in all respects.

On the fourteenth (day of the month
that this experiment was made) we saw
the commencement of three royal cells in
No. 1. The entrance of both halves was
opened on the fifteenth ; the bees went out
to forage, and on their return we observed
that they kept to their respective hives with-
out intermixing. Two queens were ob-
served dead at the entrance of No. 1, on
the twenty-fourth; and in examining the
combs, we found her that had killed them.
She left the hive on the thirtieth, was fe-
cundated, and from thenceforward the pros-
perity of the colony was ensured.

The apertures in the division between the
halves allowed the communication of tlie
bees of No. 1, with this queen, by means of
smelling, hearing, or any unknown sense.
They were separated by an interval not

278 USE OF THE ANTENNAE

exceeding a third or a fourth part of an
inch, which they could not pass: yet the
same bees became agitated ; they construct-
ed royal cells, and reared young queens:
therefore they conducted themselves just as
if their queen had been truly carried away,
and lost to them for ever.

This observation proves that it was nei-
ther from sight, hearing, nor smell, that the
bees were sensible of the presence of their
queen, and that the aid of another sense
was interposed. The division inserted be-
tween the halves of the hive having deprived
them of nothing but contact with her, w’as
it not very probable that her presence had
to be learned by touching her with the an-
tennse? It is by means of these organs that
bees gain the knowledge of their combs,
their young, their companions, and also of
their queen, all communicated by the sense
of feeling.

To be satisfied on the point, a frame w'as
removed from one of my glass hives, and re-
placed by a box of the same dimensions,
covered within by a grating, which allowed
transmission of the antenna;, but too small
for the heads of the bees. A moveable glass
frame closed the other side.

IN OPERATIONS OF BEES.

279

Waiting until the queen placed herself
on the front of a comb in view, we opened
the frame, and took her from amidst her
companions, without alarming them. She
was immediately confined in the glass box
destined to receive her ; but that she might
not suffer too much from a situation so dif-
ferent from what she had been accustomed
to, some bees of the same hive w^ere intro-
duced along with her, from which she expe-
rienced the wonted attentions.

We remarked from the first, that the dis-
tress commonly following the departure of a
queen was not manifested on this occasion.
Every thing remained in order ; the bees
did not forsake their brood during a single
moment ; their labours were uninterrupted ;
nor did w^e see the rudiments of any royal
I cells forty-eight hours afterwards on open-
I ing the hive; no arrangements had been
made for obtaining another queen. Thus all
the bees knew that they had no need to re-
place her, that she was not lost ; neither did
they treat her as a stranger on being re-
stored ; they seemed to recognise her imme-
diately. She soon began to lay amidst the
I surrounding circle of workers.

280

USE OF THE ANTENNA

'The means of communicating with this
queen, which the bees adopted during her se-
clusion, were very singular. An infinite num-
ber of antennae thrust through the grating,
and turning in all directions, plainly indicated
that they were occupied with her. She ac-
knowledged the interest which -was taken
in the most decided manner, by always re-
maining fixed on the grating, and crossing
her antennae with those so evidently em-
ployed in ascertaining her presence. The bees
attempted to pull her through to themselves,
for her legs were seized and firmly held by
theirs passing to the other side. Their
trunks, likewise, were observed to be intro-
duced to the queen’s division; and while a
captive, she was fed by her subjects from
within the hive.

How can we doubt, after this experiment,
that the workers and the queen preserved a
communication by the mutual touch of the
autennaE! ; but knowing she was so near,
the latter felt that there was no necessity
for providing themselves with another.

To prove that it is not by smell that bees
are sensible of her presence, I put the queen
of one of my leaf hives into a box with a

IN OPEIlx\.TIONS OF BEES.

281

double grating, too distant for the operation
of the antennae. We had foreseen the re-
sult. After remaining calm an hour, the
bees became agitated ; they abandoned their
labours, and forsook their young ; left the
hive, and returned : but in two or three
hours tranquillity was restored. Next morn-
ing we recognised the rudiments of eight or
ten royal cells, which had been commenced
since the preceding evening. This proved
to demonstration, that the bees had believed
their queen lost, though she resided among
them. Her emanations, therefore, were not
sufficient to undeceive them, and they requir-
ed actual contact with her to be assured of
her presence. As every bee cannot be in
all parts of the hive, we must admit likewise
that they mutually communicate their dis-
i quiet, and that they labour in common to
1 repair their loss.

Perhaps the experiments made on the
! antennae of the queen will be recollected :
t That the amputation of one did not affect
: her instinct ; but if both were cut off near
: the root, that these beings so privileged, these
mothers so much the object of considera-
tion, lost all their influence ; even the in-
stinct of maternity disappeared : their eggs

282

USE OF THE AXTENNvE

were not deposited in cells ; their mutual
animosities also were forgot ; they passed
close together without recognising each
other ; and the workers themselves seemed
to participate in their indifference, as if no-
thing except the agitation of the queen ren-
dered them aware of danger to their colony.

We amputated the antennse of two hun-
dred males and three hundred workers. The
latter being released, speedily revisited the
hive : but we observed that they did not as-
cend the combs as usual, that they no lon-
ger shared in the common operations, and
pertinaciously remained below where some
light was admitted from the entrance. This
proved their sole attraction. They soon
departed to return no more.

The same effects were produced on the
males. They likewise retreated to their
habitations ; but now they could not find
the internal passages, and they rushed to-
wards the side where an open shutter ad-
mitted the light, there finding an exit.
Some were seen requiring honey from the
workers, but in vain : they could no longer
guide the proboscis ; it was directed blind-
ly towards the head or the thorax, there-
fore, they obtained no relief. On excluding

IN OPERATIONS OF BEES. 283

the light, they hurried out of the hive though
it was six o’clock, when no males left the
others. Thus their departure must be
ascribed to the loss of that sense which is
employed to guide them in the dark.

As the amputation of only one of the an-
tennae had no perceptible effect on the in-
stinct of queens, neither had a similar muti-
lation any influence on that of males or
workers. Privation of a small portion of
these organs did not impair their faculty
of recognising objects, for they remained in
the hive, and pursued their wonted labours.
Therefore the conduct of bees deprived of
the antennae cannot be ascribed to pain ;
it ensues from their incapacity to guide
themselves and communicate with other
members of the colony.

]\Iy conjectures are supported by the use
to which bees, particularly in the night,
apply the antennae. Let us follow their
operations by moon-shine, when they keep
watch at tlie opening of the hive, to pre-
vent the intrusion of moths then on the wins',
and we shall be convinced of the fact. It
I is curious to observe how artfully the moth
knows to profit to the disadvantage of the
bees, which require much light for seeing

284 USE OF THE ANTENNAE

objects, and the precautions taken by the
latter in reconnoitering and expelling so
dangerous an enemy. Like vigilant cen-
tinels, they patrole around their habitation,
with their antennae stretched out straight
before them, or turning to right and left :
wo to the moth if it cannot escape their
contact. It tries to glide along between the
guards, carefully avoiding their flexible or-
gans, as if aware that its safety depended
on its caution.

We have not affirmed that bees possess
the sense of hearing ; yet we shall avow that
we have been tempted to believe it so. Fre-
quently we hear an abrupt agitated sound
among those watching the entrance of the
hive during the night; but if a stranger in-
sect, or any enemy happens to touch their
antenujc, the guard is roused, the sound as-
sumes a different character from that pro-
duced by the bees while buzzing or in flight,
and the enemy is assailed by several work-
ers from the interior.

Should we strike the board of a hive, the
bees soon put their wings in motion ; but
' on breathing through a cleft of their habi-
tation, we hear some of them, by means of
their wings, produce sharp and interrupted

IK OPEIlAflONS OF BEES. 285

sounds ; and then other workers are ob-
served in agitation, and directing themselves
f]'om the side where the air has entered.

These facts seem to correspond with the
sound emitted by queens, in leading us to
admit a sense in bees analagous to hearing.
Yet it must be remarked that they receive
no impression from sounds which are not
relative to their instinct. They do not seem
to he affected by thunder, or the report of
fire arms. Thus if the sense of hearing does
subsist among them, it is differently modi-
fied from the same sense among beings of a
higher order.

We shall restrict ourselves to observing,
that certain sounds produced by bees ap-
parently serve as a signal to their com-
panions, and are followed by regular conse-
quences. They may be additional means of
communication to those afforded by the an-
tenna?.

There is nothing repugnant to our con-
ceiving the subsistence of a language among
beings whose instinct is so enlarged as that
of bees ; whose conduct is compounded of a
thousand proceedings, and which, combined
in great numbers, cannot share their respec-
tive labours, or aid each other suitably,

286

USE OF THE ANTENNA.

without the means of mutual communica-
tion. The like observation applies to all in-
sects dwelling in society, as well as those
large animals regulated by the same con-
ditions.

RESPIRATION OF BEES.

287

CHAPTER IV,

ON THE RESPIRATION OF BEES.

Ihe resiiiration of insects accumulated
i together in a confined space, where the air
can be renewed with difficulty, offers a new
problem to the naturalist. Such is the case
with regard to bees. Their hive, whose di-
mensions do not exceed one or two cubic feet,
contains a multitude of individuals, all ani-*
mated, active, and laborious. Its entrance,
which is constantly very restricted, and often
obstructed, by crowds of bees departing and
.. arriving during the heats of summer, is the
I' only opening admitting the air ; yet it suffi-
^ ces for their exigencies. The hive, besides
■■ being internally plastered over with wax
; and propolis by its inhabitants themselves.

S88

RESPIRATION OF BEES.

and closed up with lime from without by the
cultivator, has none of the conditions neces-
sary for preserving a current of natural air.

If a lighted taper be placed in a vessel
of equal capacity as a hive with an aperture
larger in proportion than its entrance, the
flame grows pale in a few minutes, then
burns bluish, and is extinguished. Ani-
mals in the same situation would soon ex-
pire, yet how do the bees survive in their
dwelling ?

Article I. Proofs of Respiration. —
Experiment 1. Bees in the receiver of an
air pump w'ere not affected by the first strokes
of the piston ; but when the mercury sunk
to a quarter of an inch above the level of ‘
the cistern, they fell down motionless. Ex-
posure to the open air revived them.

2. I took three empty flagons, each ca-
pable of holding 16 ounces of water, and in-
troduced 2,50 workers into the first, the
same number into the second, and 150 males
into the third. The first and last were |
shut close, the second only restrained the ;l
escape of the bees, that they might serve for •'
comparison. In a quarter of an hour the
workers in the close vessel began to testify
signs of uneasiness ; their rings contract-

RESPIUATION OF BEES.

289

eel and dilated with greater rapidity; they
jierspired copiously, and seemed strongly
affected, because they licked the humid sides
of their vessel. In another quarter of an
hour, a cluster, which had formed around a
bit of straw, suddenly separated, and each of
the bees fell to the bottom of the vessel, in-
capable of rising. All became asphyxiated
in three quarters of an hour nevertheless,
when removed and exposed to the air, they
recovered. The males were affected more
fatally, for none survived, but the bees in the
vessel admitting the air did not suffer. We
found the air of the others greatly altered ;
the oxygen gaz was almost totally consumed,
und bees now introduced into it perished.

3, 4. Experiment demonstrated, that a
new supply of oxygen gaz restored the
asphyxiated bees. When this gaz was pure,
some of them put among it lived eight times
as long as in common air ; but all were suffo-
cated by its conversion into carbonic acid gaz.

! ^ 6, 7. Bees became insensible instantly

I in carbonic acid gaz obtained from chalk,

I though they quickly revived in the open

Asphixy is that state of suffocation rendering animated
beings motionless and insensible, and which would tenni,
nate in death, if protracted. — T.

0

i

290 respiratiox of bees.

air ; but they perished instantly and ir-
recoverably in that obtained from a mix-
ture of sulphur and iron filings, and the same
succeeded in hydrogen gaz obtained from zinc.

8, 9. Bees had expired in an hour in a
mixture of three parts of hydrogen gaz and
one of vital air ; the whole volume of both
equalling that of six ounces of water. They
perished as instantaneously in an atmos-
phere composed of three parts of azotic gaz
from sulphur and iron filings moistened, and
one of air.

10. To ascertain the effect of vitiated air
' on these insects in a torpid state, we sur-
rounded a receiver containing them with
pounded ice. The thermometer fell from
65°, the temperature of the atmosphere, to
45°, when their torpidity commenced. Be-
ing transferred into tubes full of the gaz,
which had been so fatal to those of the-
preceding experiment, and kept among it
three hours, they revived afterwards by the
heat of the hand. It is not contact of the
mephitic gaz, therefore, but its introduction
into the respiratory organs, that is fatal.

11, 12,13. The result of experiments on
the eggs, larv83, and nymphs of bees was si-
milar, proving the consumption of oxygen

RESPIRATION OF BEES.

291

and the fonnation of carbonic acid gaz. The
larvffi consumed more than the eggs, and
less than the nymphs, but the last alone
perished. Larvfe resisted the pernicious in-
fluence of carbonic acid gaz better, during^
some seconds, than adult bees would have
done. Nymphs died almost instantly.

14. Eggs put in air vitiated by the respi-
lation of bees, lost the faculty of evolution ;
but larvas and nymphs rendered torpid en-
duied exposure to deleterious gazes several
hours without inconvenience,

T. he lespiration of bees at an early age
was thus proved to be regulated by the same
laws as that of adults, as was to be ex-
pected from Swammerdam having observed
three pair of stigmata on the thorax and
seven on the abdomen of nymphs. But-1
considered it of some importance to ascer-
tain whether these organs are preserved in
the adult insect. The wonted metliod of
immersion was employed ; and to avoid any
complication which might ensue from tor-
pidity, the water was slightly heated.

1.5, 16, 17. Mnicn only tlic head of a
bee was plunged m mercury, or in water,

1 did not seem to suffer: but if the head
alone remained out of the fluid, the insect

292 EESPIRATIOX OF BEES*

displayed its trunk, and soon became asphyxia
ated ; if the head and thorax were immersed,
leaving the abdomen free, the bee struggled
a few seconds, and quickly died.

18. The orifices of the respiratory organs
being apparently in the thorax, it was left
free, while the head and abdomen were im-
mersed. A bee supported tli,e experiment
very patiently, and took flight when re-
leased.

19, 20. The action of the stigmata can
be best observed by the suffocation of bees
in water. Four air bubbles then become
conspicuous : two between the origin of the
neck and the root of the wings, a third on
the neck at the origin of the proboscis, and
the fourth on the opposite extremity of the
thorax, close to the pedicle uniting it with
the abdomen. The bee seems to have some
power in their retention, as they do not rise
to the surfiice of the water, until acquiring
sufficient size to overcome the resistance of
inspiration, or adherence to the sides of the

' cavities. By the last two bubbles, the ex-
istence of stigmata, which Swammerdam
overlooked, is indicated. As other experi-
ments showed that one orifice remaining
li-ce is sufficient for carrying on respiration,

RESPIRATION OF BEES. 293

some internal communication must subsist
between the stigmata.

21. We ascertained from the precipita-
tion of lime from lime water by air bubbles
escaping from the body of bees, that the
formation of the carbonic acid gaz is chiefly
owing to their resjiiration.

Article II. On the air of hives.—
Experiment 1. It could not be supposed
that the atmospliere surrounding 25,000 or
30,000 bees, or sometimes more, preserved a
sufficient degree of purity for their respira-
tion. Yet, to prove it, we prepared a large
tubular receiver for a hive ; and after lodg-
ing a swarm there, allowed time for con-
structing some combs, that the condition of
the bees might be assimilated to their or-
dinary state. In one experiment^ the air
of the receiver, taken at different times of
the day,^ proved nearly as pure as atmos-
pheric air, and equally so in another.

found, besides, that wax and pollen rather
vitiated its purity.

2. If the bees had any sources of vital
air in their hive, capable of supporting then*
necessities, it would be indifferent whether
the entrance were close or open. Therefore
while great activity prevailed, and their

29-i RESPIRATION OF BEES.

buzzing in the receiver was audible at the
distance of ten paces, I shut up the entrance
at three o’clock on a rainy day, when the
whole bees were collected within. In a
quarter of an hour they began to testify some
uneasiness; for, until that time, they seemed
unconscious of their imprisonment. But
their labours were now suspended, and the
hive assumed quite a different aspect; all
the bees, those covering the surface of the
combs, as well as those clustering together,
quitted their stations, and vibrated their
wings in great agitation. This ferment con-
tinued about ten minutes, when the motion
of the wings gradually relaxed, and became
less incessant. At 37 minutes past three,
the workers had lost their strength : they
could not cling by the legs, and speedily
fell down. The number of falling bees al-
ways augmented, until thousands strewed
the board of the hive. Not one remained
in the combs, and three minutes later the
whole colony was suffocated. The hive
cooled suddenly, the thermometer in it sink-
ing from 95° to the temperature of the at-
mosphere. In hopes that the admission of
purer air would restore heat and animation,
we opened a stopcock adapted to the hive,

RESPIRATION OP^ BEES.

295

and also its entrance. The effects of the
current introduced were unequivocal. In a
few minutes the bees were in a condition to
respire : the rings of the abdomen began to
plav: the vibration of their wiup-s com-
menced simultaneously ; a very remarkable
fact, which, we had previously remarked, at
the moment privation of the external air
had been felt. The bees in a short time
re-ascended their combs; the temperature
rose to the degree which these insects know
to preserve habitually, and in a few hours
order was established in their dwelling.

o

Undoubted evidence was thus obtained
that bees have no substitute in their hive for
external air.

Article III. Researches on the mode of
renewing the air in hives. — While investi-
gating all the faculties of bees themselves,
i which might effect renewal of the internal
, air, ue nere struck with the vibration oftlie
' wings. W e suspected that,'having sufficient
1 action to produce the continual buzzin.o-
. heard within, it might be for the purpose of
i displacing the air vitiated by respiration.

During fine weather, a certain number of
bees always a]ipear before the entrance of
the hive occupied in this manner, but still

296

RESPIRATION OF BEES.

more are found to be engaged in ventilating
the interior. The ordinary place of ventila-
tion is on the board ; those outside of the
entrance have their heads turned in towards
it ; those within have them turned in the
opposite direction.

We may affirm, that they arrange them-
selves regularly to ventilate more at ease.
Thus they form files, terminating at the en-
trance of the hive, and sometimes disposed
like so many diverging rays. But this or-
der is not uniform. Probably it is owing to
the necessity for the ventilating bees giving
way to those going and coming, whose rapid
course compels them to range themselves in
a file, to avoid being hurt or thrown over
every instant.

Sometimes above twenty bees ventilate at
the, bottom of a hive; at other times their
number is more circumscribed, and their em-
ployment of various duration. We have
seen them engaged in it during twenty-five
minutes, only taking breath, as it were, by
the shortest interruption of the vibration.
On ceasing, they are succeeded by others ;
so that there never is any intermission of
the buzzing in a populous hive.

TlESPirtATION OF BEES.

297

If under the necessity of ventilating dur-
ing winter, near the centre of the mass then
united towards the top of their dwelling,
doubtless the bees perform this important
function among the vacuities of the irregu-
lar combs, where there is room for their wings
to expand, as at least half an inch is requi-
site for them to play freely.

Is ventilation as- essential to bees in their
natural state, when inhabiting hollow trees,
or the more spacious cavities of rocks, as in
their domesticated condition ? In imitation
ol such situations, we lodged a swarm in a
glass hive, five feet high, which on all sides
exposed their pyramidal cluster, suspended
from the combs above. A common entrance
was made at the bottom. AVe observed that
very few were occupied here in fanning
themselves : it was always on the side of the
same quarter where the greatest numbers of
the swarm had accumulated : they kept at a
little distance from each other, and in the
way of those returning from the fields.

riie ventilation of the bees, or the buz-
zing which denotes it, seems to me more ac-
tive during winter than at any other time.
It was easy to prove that this operation es-
tablished a current of air; for anemometers

298 ' EESPIRATION OF BEES.

of light paper or cotton, hung by a thread,
were impelled towards the entrance, and re-
pelled from it with equal rapidity. The ac-
tion on them never was entirely interrupted,
and its force appeared proportional to the
number of bees fanning themselves.

If some cultivators shut up the entrance
of their hives in winter without prejudice to
the bees, it must be considered that the air
will penetrate, nevertheless, through the
straw composing them. I confided an ex-
periment on this head to Burnens, then at a
distance from me, who ‘communicated the
result in a letter, dated Oldens, Sd Feh,
1797. Having closed down a very populous
straw hive fast on its board, he found that
a piece of the finest paper, suspended by a
hair before the entrance, oscillated above an
inch off the perpendicular line. He pour-
ed liquid honey through an opening in the
top, when a buzzing soon began, and the tu-
mult increasing within, several bees depart-
ed. The oscillations now became stronger
and more frequent. His experiments were
made at three o’clock, the sun shining bright,
and the thermometer in the shade standing
at 44".

RESPIRATION OF BEES.

299

Article IV. Proofs from an artificial
ventilator. — Although the purpose of ven-
tilation is evident from the preceding experi-
ments, ]\I. Senebier advised me to obtain a
similar effect from an artificial ventilator.
INIr. Schwepp, who is both'an expert mecha-
nic, and an ingenious naturalist, aided me
in constructing a small wind-mill, with
eighteen tin vanes, put into a box, on the
top of which a large cylindrical vessel was
adapted.

Experiment 1, 2. An aperture, repre-
senting the entrance of a hive was left free,
but a taper in the vessel was extinguished
in eight minutes, though its capacity was
equivalent to 3228 inches. A single aper-
ture, therefore, does not promote the renewal
of air, unless it be agitated from some ex-
ternal cause.

3. Fresh air was restored, and a taper in-
troduced, when the ventilator being put in
action, several anemometers, suspended be-
fore the entrance, proved the establishment
of two currents. During the whole course
of the experiment, which may be prolong-
ed indefinitely, the brillianey of the light
remained undiminished. A thermometer,
at the bottom of the apparatus, stood at

300 RESPIRATION OF BEES.

122°: the temperature in the receiver above
evidently was higher.

4. 'Wishing to try whether my ventila-
tor would correct the vitiation of the air
by two tapers, I found that they burnt
fifteen minutes, and that both went out at
once. In another experiment, when the
ventilator was still, they were extinguished
in three minutes.

5. Contrary to our expectations, a more
powerful current of air was not obtained
by multiplying the apertures.

These experiments, showing that/the air
can be renewed in a vessel with apertures
on only one side, when it is displaced by a
mechanical cause, seem to confirm our con-
jectures regarding the ventilation which
bees can produce in their hive.

Article V. Immediate souree of
ventilation. — In striking the air with their
wings, bees know little of the real object
attained. Perhaps some inclination or some
very simple necessity induces them to put
organs in motion, apparently bestowed on
them only for flight. Doubtless it is to re-
lieve an immediate sensation, for w^e cannot
grant them the same knowledge that would
induce us to act after a relative mannen

RESPIRATION OF BEES. 301

Nevertheless, it is curious to learn those
propensities, however gross, by means of
which nature enables them to accomplish
an object.

The bees fanning themselves for coolness,
may be one immediate cause of ventilation.
A^^e remarked that the rays of the sun, hav-
ing darted on combs covered with bees, from
opening the shutter of a glass hive, those
in the shade remained in tranquillity, while
those sensible of the heat began buzzing.
Likewise, the bees of clusters formed be-
fore the hive, in summer, fan themselves
vigorously if incommoded by the heat of
the sun ; but the ventilation ceases on shad-
ing part of the clusters, though it continues
in the rest.

The same may be observed in humble
bees, an analogous genus. Those kept with
their nest in a window, peaceable in gene-
ral, began to be very noisy as the rays of
the sun fell on the box containing them.
All vibrated their wings and buzzed loud-
ly. Sometimes, also, we hear this sound in
I the nests of wasps and hornets. Thus
i it appears certain that it is heat which
i induces bees and other insects to fan thcm-
: selves.

302

HESPIEATION OF BEES.

But it is singular that they ventilate,
likewise, during the cold of winter, when
heat may be only a secondary cause.

W e found that the influence of obnoxious
odours led them to fan themselves. Having
separated some bees from their hive by the
attraction of honey, we brought cotton, dip-
ped in spirit of wine, near the head. Its
effects could not be mistaken: they dispersed,
agitating their wings, and then drew toge-
ther again to resume their repast. When
completely engaged, the experiment was re-
peated, and they dispersed anew, hut with-
out retracting the proboscis entirely: they
were satisfied with vibrating the wings, and
continued feeding.

When too much affected by disagi-eeable
sensations, they precipitately withdrew, and
took flight. Frequently a bee turned away
from the honey, and began to fan itself, un-
til the sensation or its cause were by this
means abated, and then returned to feed.

Such experiments never are more success-
ful than at the entrance of the hive ; be-
cause, the bees retained by the united at-
traction of honey and their home, are less
disposed to retreat from external imi>rcs-
sions. Humble bees adopt the same me-

RESPIRATION OF BEES. ~ 303

tliocl of dispelling pernicious odours. But
what is very remarkable, and may partly
show the importance of vibrating the wings,
neither their males nor those of domestic
bees, though very sensible of similar emana-
tions, know to protect themselves, like the
workers, against them.

Ventilation, therefore, is one of the in-
dustrial operations peculiar to the workers.

The Author of Nature, in assigning a
dwelling to these insects where the air can
hardly penetrate, bestows the means of avert-
ing the fatal effects which might result from
the vitiation of their atmosphere. Perhaps
the bee is the only creature entrusted with
so important a function, and which indi-
cates, thus to speak, such delicacy in its
organization.

An indirect consequence of ventilation is
the elevated temperature of the hive, pre-
served by the bees without any exertion,
and resulting, like the heat of all animals,
\ from their respiration. This, which one
i author gratuitously ascribes to the fermen-
il tation of the honey, certainly is derived
I from the accumulation of a great many bees
i in the same place. It is so essential to tlicm
and their young, that it ought to be in-

304

RESPIRATION OF BEES.

dependent of the temperature of the atmos-
phere. The existence of bees thus depends
in more than one point on the constancy of
ventilation. Yet while called on to execute
so many diflPereut labours, each member of
the colony cannot be occupied, on its own
account, with the care of preserving the air
at the necessary degree of purity. This
function being exercised by a few indivi-
duals alternately, does not abridge the other
branches of industry indispensible.

Thus the condition of bees admitting of
their fulfilling by turns the different func-'
tions imposed on the whole society, corres-
ponds to the beneficent views of the Crea-
tor, and supplies the place of the arrange-
ments which we ourselves would make for
our own advantage.

A NEW ENEMY OF BEES.

805

CHAPTER V.

ON A NEM' ENEMY OF BEES.

Among the labours of insects, those which
concern the defence of their habitations are
not the least deserving the attention of man,
who is so often called on to fortify himself
against the enterprises of his enemies. It
is here that nature unfolds the most unex-
pected resources, where she admits of the
greatest latitude ; for the chances of war are
the object of one of those general laws concur-
ring to the preservation of universal order.
Without the alternatives of success and re-
verse, how could an equilibrium be main-
tained ? One species endowed with superior
strength would annihilate another. Never-
theless the most timid have subsisted since

306 A NEW ENEMY OF BEES.

the origin of things : their tactics, their in-
dustry, their fecundity, or other circum-
stances peculiar to their kind, have enabled
them to escape that extinction which seems
to menace them.

ximong bees, as with the greater part of
their class, the ordinary means of resistance
are those poisoned weapons with which they
wound their enemies. The fate of w^ar
would be more in their hivour, were not se-
veral of their antagonists armed still better
than themselves ; if others had not the art
of shunning their vigilance by sheltering
themselves under a covering ; and were there
not also some which can profit, by the weak-
ness of an ill-peopled hive, to gain a surrep-
titious introduction into it.

AVasps, hornets, the larvae of moths, and
mice, have been known from all antiquity
by tlieir ravages among hives ; and having
nothing to add to what is generally said re-
specting them, I shall confine myself to
pointing out a new enemy of bees.

Towards the close of summer, after hav-
ing stored up part of their collections, we
sometimes hear a surprising noise in their
habitations. A multitude of workers de-
part through the night, and lose themselves

A NEW ENEMY OF BEES. 307

in the air : the tumult frequently continues
during several hours, and on examining into
the consequences of so great an agitation in
the morning, we find numbers dead before
the hive. For the most part, the honey is
observed to be exhausted, and occasionally
the hive is quite deserted.

In the year 1804, many of my neighbour-
ing cultivators came to consult me on an
occurrence of this nature. But I could
give them no explanations ; notwithstand-
ing my long practice in studying bees, I
never had seen any thing similar.

On visiting the scene, I found the phe-
nomenon still subsisting, and that it had
been very accurately described ; but it was
assigned to the introduction of bats into the
hives, which I had difficulty in crediting.
These creatures are satisfied with seizins'
nocturnal insects in their flight, which are
never wanting in summer. They do not feed
on honey ; why, therefore, should they at-
tack bees in their habitations, for the pur-
pose of pillaging their stores ?

However, it might be some other animal.

I Therefore, having put my people iu ambush,
I they soon brought me intelligence, not of
|bats indeed, but of the sphinx atropos, or

308

A NEW ENEMY OF BEES.

death’s head, a large moth flying in num-
bers about the hives ; and one was caught
at the moment of attempting to penetrate
a hive among the least populous, where it
evidently designed living at the expense of
the bees.

From all quarters I had information of
similar ravages committed by bats, as sup-
posed : Cultivators, who expected a plentiful
harvest, found their hives as light as in the
first days of spring: thongh recently well
provisioned, they were now reduced to the
weight of wax.

At length, the gigantic moth which had
occasioned the desertion of the bees, was
surprised in several hives.

Such reiterated proofs were requisite to
persuade me that a butterfly, an insect want-
ing a sting, unprovided with any shield or
other means of defence, could contend vic-
toriously against thousands of bees ; but this
year they were so common, that it was easy
to be convinced of the fact.

As the enterprises of the sphinx con-
stantly became more fatal to the bees, we
resolved to prevent its access, by contracting
the entrance of the hive by a kind of grat-
ing, large enough to admit none but its pro-

A NEAV ENEMY OF BEES.

809

Jier tenants. This plan had complete suc-
cess, quiet was restored, and the devasta-'
tion ceased.

The same precautions had not been uni-
versally taken ; but we observed that the
bees, left to themselves, had provided for
their own security. Without any foreign
aid, they had barricaded , themselves, by
means of a thick wall of propolis and wax,
rising behind the entrance of the hive,
sometimes in the entrance itself, and com-
pletely obstructing it, but penetrated by
passages for one or two workers at a time.

The operations of man and of the insect
had completely coincided.

The works which the bees had established
were of very various formation. Here was
a single wall, whose opening arcades were
disposed in its higher part : there several
bulwarks behind each other recalled the bas-
tions of our citadels. Gateways, masked by
walls in front, opened on the face of those of
the second row, Avhile they did not corres-
pond with the apertures of the first. Some-
times a scries of intersecting arcades permit-
ted free egress to the bees, but refused admit-
tance to their enemies. These fortifications
weremassy; their substance firm and compact.

310 A NEW ENEMY OF BEES.

As such casemated gates are not con-
structed by bees without urgent necessity,
we cannot ascribe their proceedings to any
of those demonstrations of prudence prepared
to obviate incouveniencies, which the insect
neither can know nor anticipate. It is only
when danger is present, when it is pressing
and immediate, that, compelled to seek pro-
tection, it employs this last resource. Thus
it is curious to observe an insect, so well
armed and supported by the advantage of
numbers, securing itself by an admirable
combination against the inefficiency of its
weapons and its courage. The art of war-
fare among bees, therefore, is not restricted
to attacking their enemies : they know also
to construct ramparts, as shelter from their
enterprises : from the part of simple soldiers,
they pass to that of engineers.

But it is not against the sphinx alone that
they must be guarded. Weak hives are
sometimes attacked by stranger bees, at-
tracted by the odour of the honey, and the
hopes of easy pillage. Those beset being
unable to defend themselves from this inva-
sion, are known to have recourse to a mea-
sure resembling that employed against the
hostile moth. Then, also, they raise walls,

A XEAV EXEMY OF BEES. 311

leaving only narrow oiienings for the pas-
sage of a single bee at a time, and which
therefore can be easily protected.

But a period arrives when these galleries
are no longer suitable to the bees. At the
time that their harvest is abundant, their
hive excessively populous, and the formation
of new colonies approaches, they demolish the
gateways which had been erected in the hour
of danger, and which now restrain their im-
petuosity. Such safeguards have become
inconvenient, and they are removed, until
new alarms demand their re-construction.

The entrances formed in 1804 were de-
stroyed in spring 1805. The sphinx did
not appear that year, nor was it seen in the
year following; but it returned in great
numbers in autumn 1807. By speedily
barricading themselves, the bees prevented
^tlieii thieatened ravages ; but, before the
departure of swarms in I\Iay 1808, they de-
molished the fortifications, whose narrow pas-
sages prohibited free egress to the multi-
tude.

AVhen the entrance to the hive is itself re-
stricted, or care being taken to contract it soon
enough to prevent the devastation of their
efiemics, bees dispense with walling them-

I

312

A NEW ENEMY OF BEES.

selves in. We cannot explain tlieir conduct
otherwise, than by admitting the evolution
of their instinct according as excited by cir-.
cumstances.

But how can the sphinx alarm colonies so
warlike? How can a moth, the dread of
superstitious people, also exercise a secret
influence over insects, and have the faculty
of paralyzing their courage ? Does it emit
any emanation pernicious to bees ?

Other species of sphinxes subsist on the
nectar of flowers alone : they have a long,
slender, flexible, spiral trunk, and seek
their food at sun-set. But the atxopos is
later on the wing ; nor does it hover about
the hives until night is farther advanced.
It is provided with a thick, short proboscis ;
is endowed with great strength ; and when
seized, some unknown organ emits an acute
stridulous sound. May not this, which in-
spires the vulgar with sinister ideas, be also
the dread of bees? ]\Iay not its resem-
blance to that emitted by the queen in
her captivity, which has the faculty of sus-
pending the vigilance of the workers, ex-
plain the disorder observed in their hive on
the approach of the sphinx ? But this is
only a conjecture, founded on the analogy

A NEW ENEMY OF BEES.

S13

of soimcls, to which I cattach no importance.
Meantime, were any piercing notes ob-
served to proceed from the sphinx during
its assaults, and that the bees then yielded
without resistance, my conjecture would ac-
Tjuire some weight.*

The introduction of a butterfly so large
and recognisable as the sphinx atropos in-
to a well peopled hive, and the extraordi-
nary consequences thence resulting, are phe-
nomena of the more difficult explanation,
from nothing in the organization of the in-
sect, indicating that it is screened from the
sting of bees.

^ We have been anxious to witness this
singular contest in glass hives, but no op-
portunity has hitherto offered. However,
to solve some of my doubts, I have made a
few experiments on the mode in which the
sphinx is received by humble bees.

Having procured some of the largest size
I introduced them at night-fall into a glass

Reaumur ascribes the sound to tlie friction of the
timnk agmust its sheaths, hut we have ascertained that this
organ has no share m it. Though many naturidists have
mvestigated its source, notliing satisfactory is known on tlie
ubjcct. I. „ undoubted .hot the sphinx ctnits , Z

P

I

SI 4 A NEW ENEMY OF BEES.

box, where a colony of small brov/n humble
bees (muscoriim) had been established.
The first carried thither did not appear to
be aflPected by the smell of the honey: it re-
mained quiet in a corner. Beginning to ap-
proach the nest and its inhabitants, it soon
became the object, not of the dread, but of
the wrath of the workers. Repeatedly as-
sailing it with fury, they gave it frequent
stings: it sought to escape: it run quickly;
and at last raising the glass cover of the
apparatus by a violent exertion, succeeded.
It seemed to suffer very little from its
wounds : remained tranquil all the night ;
and several days after was wonderfully well.

i\nother sphinx, very vigorous and lively,
which often emitted the sound peculiar
to its species, was confined along with the*
same colony. Its activity only served tO'
render it sooner the victim of their rage.
Immediately on approaching the nest, which
nevertheless it seemed to have no desire to
enter, all the workers darted on it at once,
with their stings, and harassed the crea-
ture so incessantly as to compel it to retreat. '
The moth defended itself only by the vio-
lent agitation of its wings, hut it could not
evade the attacks of the humble bees under

A NEW ENEMY OF BEES. 315

the belly, where it seemed most sensible of
their weapons. At length, after an hour of
suffering, it perished under many wounds.

I was unwilling to carry this cruel expe-
riment farther. Captivity, or some other
circumstance, evidently reduced the insect to
too great an inferiority to the humble bees.
Yet, after the experiment, it became still
more difficidt for me to understand how it
could introduce itself into the hives of com-
mon bees, whose stings are so much more dan-
gerous, as well as their numbers incomparably
greater. Had the light of a torch been an
obstacle to the sphinx exercising its means of
attack? Possibly the success of its enter-
prises on hives results from the fiiculty of
seeing during the night, like other moths of
the same genus.

Offering honey to these insects was equal-
ly fruitless, fhey remained during a week
beside a comb without touching it : we un-
folded the proboscis, and dipped it among
honey, in vain. This experiment, though
succeeding perfectly with day butterflies,
proved abortive with the sphinx atronos.

Had I not obtained proofs of its avi-
’ dity while in the natural state, T miglit
ihave entertained doubts of the predilection

316 A NEW ENEMY OF BEES.

for honey. Besides, the facts above related
are supported by my having recently dissect-
ed a large sphinx taken in the open air, and
found the abdomen quite full of pure honey,
of the same taste and consistence as that of
bees. The quantity would have filled a table
spoon ; and what appeared very singular, was
its not being contained in any particular in-
testine, but occupying the cavity usually re^
served for air in the body of these insects.
All the vertical delicate membranes di-
viding the abdomen into so many compart-
ments had disappeared. I cannot affirm po-
sitively whether they had been ruptured by
the quantity of honey gorging the sphinx,
or by ourselves ; but one thing is undoubted,
that in others of the sphinx atropos, opened
by us, we have always seen the receptacles
entire though empty.

From facts illustrating the history of the
sphinx, let us return to the means of preserv-
ing bees from so dangerous an enemy.

This may be accomplished by the aid of
a small slider at the entrance of the hive,
penetrated by three kinds of apertures, pro--
portioned to the necessities of the bees, and i
adapted to use in a manner corresponding:
with their own plans of security.

A NEW ENEMY OF BEES.

317

As they destroy their fortifications in
spring, previous to the flight of the swarms,
then we ought to leave the entrance free.
They have few enemies to dread, and their
community well peopled can defend itself.
After departure of the swarms the entrance
is to be contracted, because the hive being
weakened, stranger bees or moths may in-
troduce themselves ; and our proceedings
are pointed out by the works of the bees
when menaced with pillage.* In the middle
of July these entrances are so much enlarged
by them, that two or three of their number
can pass at once, and they allow the free
egress of the males, which are of larger size
than the workers, At this epoch, therefore.

* The author elsewhere observes, that on tlie 9th of
July 1804 some dead and wounded bees were found on the

board of the hive, which were recognised to have come for
the purpose of pillage. On the 11th, however, the en-
trance was built up by the bees with wax, two openings
being left at the part farthest above the board, which wouid
allow only a single bee to pass at a time. Thus, two of
their own number always were enough to guard it. These
apertures were enlarged agaipst the 22d, so as to admit the
passage of two or three bees at once ; and another opcniim
was made in the course of September. The author advises
mlu,ut„r, 10 visii ilioir M,,., frciucMly ; a.ul on fin, lino
them pillage,!, to feed the beea with honey or syr,,,,, withoiU
nch he never was able to preserve them. Daily supplies
melted sltould be introduced in putsheUs from belund.-r.

318

A NEW ENEMY OF BEES.

it is necessary to advance the slider so far as
to bring the apertures, whose convexity
should be downwards, opposite to the en-
trance of the hive. Finally, the collection
being in its full force in the months of Au-
gust and September, it is essential that the
bees ought not to be too much confined.
Those whose example we follow opened a
third passage in the waxen wall, resembling
a very low vault, which w'e should imitate
in the third row of apertures. By this
means the access of the sphinx is precluded,
Avhile the bees can find an easy exit. If the
slider be made of tin instead of wood, it will
exclude mice, another most dangerous enemy
to bees.^

In subduing animals, man in some mea-
sure impairs the equilibrium established by
nature among rival species, and more or less
diminishes their energy and vigilance. The
features of their instinct can be then dis-
covered only by judging of their peculiarities;
and, in his turn, he should partly compensate
the advantages of which they have been de-
prived. He must do more, if wishing to aug-

, * In September 1802, M. Lombard found the entrance
of all his hives aln^ost totally closed up by the bees ; but
at that time he did not know to what he should ascribe it.
3'fmnicl dcs rropritiuiri’s d AhcUles, p. 32, T-

A NEVv" ENEMY OF EEES. 319

mciit tlieii; products, for he has to contend
with nature, wdiich assigns limits to the
multiplication of individuals. But this de-
mands a very profound knowledge of the
wants of the creatures subjected to his do-
minion, and of the resources which Provi-
dence has put within their reach ; for it is
from themselves that the art of managing
them must be learned.

I

320

NEW OPINIONS ON WAX.

CHAPTER VI.

NEW OPINIONS ON WAX.

Since the time of Reaumur aucl tie CTeer,
whose works have inspired a general taste
for entomology, great advances have been
made in this science ; all its branches are
extended, and the history of bees, in parti-
cular, has been enriched.

Schirach and Riem have opened a new
path ; and, perhaps, we have ourselves con-
tributed to clear it of the prejudices which
clogged its progress, by establishing the
facts announced in a more rigorous manner.

Some observations, also, have been pub-
lished in other countries, hut so inaccurate-
ly, that they would sink into oblivion did I
we not endeavour to support tliem by facts.

XE^V OPINIONS ON M"AX. 321

Naturalists have principally directed their
attention to wax, and chemists likewise
have attempted an analysis of it : But the
result of their labours presents so little co-
incidence, as to prove the insufficient dis-
cussion of the subject, and that it requires
new examination.

AVhen JNI. Bonnet wrote, it was the gene-
ral opinion that the pollen, farina or dust of
thestaminaof flowers wascouverted into wax;
and it is interesting to peruse his details of
its collection, the manner in which the bees
load themselves, and how- they store it up
for preservation. All these facts, and the
utility of the farina, had been scrupulously
observed by Reaumur, INIaraldi, and other
learned men ; but is this substance truly
the elementary principle of wax ?

Reaumur entertained some hesitation re-
garding it, from the great difference between
the pollen and the wax ; however, he in-
clined to believe that the former, by receiv-
ing some peculiar elaboration from the bees,
was converted into real wax in the sto-
mach, and disgorged under the appearance
of a kind of paste.

A Lusatian cultivator, whose name has
not reached us, observes, that althougli it

322 NEW OPINIONS ON WAX.

had been previously supposed that wax is
discharged from the mouth, it actually comes
from the rings of the abdomen ; and that
this is evident by withdrawing a bee from
the cell where it works in wax, when the
wax in the form of scales appears by the
extension of the body.*

In the year 1793, we were greatly asto-
nished at finding scales of a substance ana-
logous to it under the rings. They exhibited
its real characteristics on being applied to
the flame of a taper, and we showed them
to some of our friends.

John Hunter, an Englishman of high re-
putation, engaged in observations on bees at
the same period, discovered the actual recep-
tacles of wax under the belly of these insects.
He found a flexible matter in them, which he
recognised as such; he ascertained the differ-
ence between the farina or pollen of flowers
and the composition of the combs, and as-
signed a new property to the pellets curried
home on the legs of bees : But he could
only offer conjectures on the use of the pol-
len ; nor had he witnessed the application
of the scales which, according to his suppo-

* Scliirncli’s History of the Queen Bee.

NEW OPINIONS ON WAX.

323

sition, transuded from the body. Our in-
quiries were carried farther.

Workers alone have the property of se-
creting wax : scales of it, ranged in pairs,
are contained in minute receptacles under
the lower segments of the abdomen, and si-
tuated to right and left of the angular pro-
jection. The conformation of the same part
of queens and males is very different, below
the rings of which no scales subsist.

Nothing but what is common to the ab-
domen of wasps. and many other hymenop-
tera appears externally in that of the bee,
being half segments partially covering each
other. But they are not flat below as in
most analogous insects, for the abdomen cf
the bee is traversed by an angular promi-
nence. Plate II. fig. 10, a, b. By gently
drawing out the abdomen, the concealed
parts are disclosed, fig. 9, 10, 11, 12.

What should be considered the base of
each ring, because it adheres to the body of
the insect, occupies at least two thirds of
I the segment, and is of a yellowish wliitc,
j soft, transparent, membranaceous substance.

I Plate III. fig. 13, c, d, e, g. It is divided
I in two by a small horny prominence, a, b.

324) NEW OPINIONS ON WAX.

corresponding exactly with the horny pro-
jection of the abdomen, and forms two areas
bounded by a solid edge on the surface of
the membrane, n, c, b, a, d, — m, g, a, b, e.
The scales of wax are deposited in these two
areas, and assume the same conformation :
being of an irregular pentagonal figure.
Only eight scales belong to each individual
bee, for the first and last ring, constituted dif-
ferently from the others, afford none. Their
size decreases with the diameter of the rings
whereon they are moulded : the largest are
under the third, and the smallest under the
fifth. All are not alike in every bee, for a
difference is perceptible in consistence,
sliape, and thickness ; some are so thin and
transparent as to require a magnifier to be
recognised ; or we have been able to discover
nothing but spiculte similar to those of wa-
ter freezing.

Neither the spiculse nor scales rest im-
mediately on the membrane ; a slight li-
quid medium is interposed, serving to lu-
bricate the junctures of the rings, or to
render the extraction of the scales easier, as
otherwise they might adhere too firmly to
the sides of the receptacles.

XEW OPINIONS ON WAX.

325

Finally, we have seen the scales so large
as to project beyond the rings, being visible
without stretching the segments, and of a !

whitish yellow from greater thickness les- ;

selling their transparency. ;

These shades of difference in the scales of
various bees, their enlarged dimensions, the
fluid interposed beneath them, the corres-
pondence between the scale and the size
and form of the receptacles, seem to infer
the transudation of its substance through
the membranes whereon it is moulded.

We 'w'ere confirmed in this opinion by j

the escape of a transparent fluid, on piercing
the membrane, whose internal surface seemed i

to be applied to the soft parts of the belly. J

It coagulated in cooling, when it resembled
wax, and again liquified on exposure to heat.

The scales themselves also melted and
coagulated like wax. ■

In prosecuting our experiments farther
on the analogy of the tw^o substances, we
found, 1. That scales, thrown into spirit of i

turpentine, dissolved and disappeared before S

reaching the bottom of the vessel, without i-

rendering the fluid turbid. But an equal I

quantity of the spirit could neither dissolve !i

I

326

NEW OPINIONS ON WAX.

some of tlie whitest fragments of worked
wax taken from new combs so quickly nor
so entirely. Many particles remained sus-
pended among it.

2. Taking two vessels of sulphuric ether,
we appropriated one for scales from the rings
of bees, the other for wax from their combs,
equivalent in weight to the scales. Scarcely
had fragments of the wax touched the ether,
when they divided, and were precipitated in
powder to the bottom of the vessel : hut the
scales were preserved entire, and only lost
their transparency, becoming of a dull white.
No change ensued in either vessel during
several days. Oh evaporating the ether
from each, a thin stratum of wax was found
on the glass. Frequent repetition of this
experiment presented the same result : frag-
ments of the combs always were reduced to
powder: the scales, on the contrary, were
not broke down : and after the lapse of se-
veral months, only a very small portion of
them had been dissolved by the ether.

We thence concluded that the wax of
the rings was less compound than that made
into cells, since the latter dissolved in ether,
while the former remained entire, and as i
the one dissolved hut partially in the spirit

NEW OPINIONS ON WAX. 327

of turpentine, whereas the other was held in
complete solution.

If this substance laying under the rings
be truly the elements of wax, it undergoes
some preparation in leaving its receptacles,
and the bees are capable of impregnating it
with matter, imparting the whiteness and
ductility of real wax. Hitherto we are ac-
quainted only with its fusibility, but such
being the chief property of the wax of the
combs, we cannot doubt that the scales en-
ter their composition.

We have been disappointed of reaching
the sources of the waxy matter by dissec-
tions, though executed by the skilful hand of
JMiss Jurine. No direct communication be-
tween the receptacles and the abdomen could
be discovered ; nor were any vessels found,
except some tracheae, certainly for the intro-
duction of air. But the membrane of the
receptacles is covered with a reticulation of
hexagonal meshes, to which we should ascribe
a certain function connected with the secre-
tion of wax. Plate III. fig. 15, 16*. This
is wanting in males, but it exists in queens,
under a modified texture, and occupying
two thirds of each segment.

328 NEW OPINIONS ON WAX.

A reticulation quite similar to that of the
working bee, and occupying the whole an-
terior part of the segments, is found in hum-
ble bees producing wax: but the receptacles
are imperceptible : their abdomen being
formed like that of the hymenoptera of the
same genus.

The stomach and other internal parts are
separated from the reticulation by a greyish
membrane entirely covering the abdominal
cavity of the common bees. When the sto-
mach is full of the juices elaborated by it,
they may transude its very thin integuments,
and traversing the greyish membrane, which
is not thick, they may come in contact with
the reticulation. It is not impossible that
the reticulation operating a re-absorptiou
and a kind of digestion of these juices, the
secretion of wax follows.

Though direct evidence be yet wanting,
we may admit that this substance is pro-
duced by a particular organ, after the man-
ner of other secretions.

ORIGIN OF WAX,'

329

CHAPTER VII,

ON THE ORIGIN OF WAX.

The existence of the organs above de-
scribed, and the scales seen under different
gradations, induce us to believe them ap-
propriated for the secretion of wax. But,
ill common with other animal and vegeta-
ble secretions, the means by which this is ac-
complished appears to be carefully veiled by
nature.

Oiu* researches by simple observation
thus being obstructed, we felt it essential to
adopt other methods for ascertaining whe-
1 ther wax actually is a secretion, or the coB
-i lection of a particular substance.

S30

ORIGIN OF WAX.

Providing it were the former, we had first
to verify the opinion of IXeaumur, who con-
jectured that it came from an elaboration
of pollen in the stomach, though we did not
coincide with him, that bees then disgorged
it by the mouth. Neither were we disposed
to adopt his sentiments regarding its origin;
for, like Hunter, it had struck us that swarms
newly settled in empty hives do not bring
home pollen, notwithstanding they construct
combs, while the bees of old hives, having
no cells to build, gather it abundantly.

W e had, therefore, to learn whether bees,
deprived of pollen for a series of time, would
make wax, and all that this required was con-
finement.

On the 24th of INiay we lodged a swarm
which had just left the parent stock in a
straw hive, with as much honey and water
as necessary for the consumption of the
bees, and closed the entrances so as to pre-
vent all possibility of escape, leaving ac-
cess for renewal of the air.

At first the bees were greatly agitated ;
but we succeeded in calming them by carrying
the hive to a cool dark jdace, where their cap-
tivity lasted five days. They were then allow-
ed to take flight in an apartment, the windows

ORIGIN OF AVAX.

331

of which were carefully shut, and where the
hive could be examined conveniently. The
bees had consumed their whole provision of
honey; hut their dwelling, which did not
contain an atom of w'ax when we established
them in it, had now acquired five combs of
the most beautiful wax suspended from its
arch, of a pure white, and very brittle.

We did not expect so speedy a solution
of the problem ; but before concluding that
the bees had derived the faculty of produ-
cing wax from the honey on which they
fed, a second experiment, susceptible of no
other explanation, was necessary.

The workers, though in captivity, had
been able to collect farina ; while they
were at liberty they might have obtained
provisions on the eve or on the day itself of
their imprisonment, and enough might have
been in the stomach or on the limbs to en-
able them to extract the wax from it that we
had/ound in the hive. But if it actually came
from the farina previously collected, this
source was not inexhaustible ; and the bees
being unable to obtain more, would cease to
construct combs, and would fall into abso-
lute inaction,

332

OmGIN OF WAX.

Before proceeding to the second experi-
ment, which was to consist in prolonging
their captivity, we took care to remove all
the combs they had formed in that preced-
ing. Burnens made them return to the
hive, and confined them again with a new
portion of honey. The experiment was not
tedious. From the evening of the subse-
quent day we observed them w^orking in
wax anew ; and on examining the hive on
the third day, we actually found five combs
as regular as those they had made during
their first imprisonment.

The combs were removed five times succes-
sively, but always under precaution of the
escape of the bees from the apartment being
prevented ; and during this long interval,
the same insects were preserved and fed with
honey exclusively. Undoubtedly, the ex-
periment, had we deemed it necessary, might
have been prolonged with equal success.
On each occasion that we supplietl them
with honey they produced new combs, which
puts it beyond dispute that this substanco
effected the secretion of wax in their bodies,
without the aid of pollen. As the reverse of
the preceding experiment would prove whe-i
thcr the pollen itself had the same property,

ORIGIN OF WAX.

333

instead of suppl}dng our bees with honey,
we fed them on nothing except fruit and
farina. Tliey w^re kept eight days in cap-
tivity, under a glass bell with a comb, hav-
ing only farina in the cells : Yet they nei-
ther made wax, nor were scales seen under
the rings. Could any doubt exist as to
the real origin of wax ?— We entertained
none.

It was not improbable that wax might be
contained in honey, and reserved for use by
the bees as requiring it, for some particles
always rise to the surface of honey diluted
with water. But the microscope indicated
them to be fragments of cells previously
made.

However, to obviate any objection, and to
ascertain whether the saccharine principle
was the real source of wax, we supplied a
swarm confined in a glass hive with a pound
of refined sugar reduced to syrup. In order
to render the experiment more instructive, by
comparison, two swarms were introduced
into other two hives, one of which we fed

with very dark brown sugar, and the other
witli honey.

1 he result proved as satisfactory as could
be expected : wax was obtained in all the

334

ORIGIN OF WAX.

three. Those bees that had been fed with
the different kinds of sugar produced it
sooner, and in greater abundance, than those
that had subsisted on honey.

A pound of refined sugar, reduced to
syrup, and clarified with eggs, produced 10
drams 52 grains of wax, darker than that
extracted by the bees from honey. An
equal weight of dark brown sugar produced
22 drams of verv white wax; the like came
from sugar of the maple.*

We repeated this experiment seven
times successively with the same bees: wax
was always obtained, and nearly in the same
proportions as above. Thus it is demon-
strated that sugar, and the saccharine part
of honey, enable bees to produce wax, a
property denied to farina.

Although there was no uncertainty regard-
ing these facts, w'hich soon received a more
favourable confirmation, it was essential to
learn whether bees, in the natural state, pur-
sued the same course as tliose held captive.

A long series of observations, of w'hicii
only a sketch shall be given, have established,

* That is, two ounces and tliree quarters was the
greatest quantity of wax obtained from a pound of sugar,
or nearly one-sixth of the weight. — T.

OKIGIN OF WAX.

335

tliat when the state of the country affords
a copious collection of honey, it is anxiously
stored up by the workers of old hives, while
new swarms convert it into wax.

At a time when I had not a great num-
ber of hives myself, those of the neighbour-
ing peasantry served for comparison, though
made of straw, and wanting the facilities of
my own. Certain remarks, which we had
made on the combs and on the bees them-
selves when working in wax, enabled us to
avail ourselves of them.

Wax is originally white : but the cells
soon become yellow: they grow brown in
time ; and the combs of very old hives have
a blackish hue. Thence we can immediately
discover whether bees are working in wax,
or whether their labours are suspended,
merely by raising the hives to see the lower
edge of the combs. '

The following observations, founded on
the fact of there being two kinds of workers
in a hive, which was unknown to my pre-
cuisois, also may afford some indications of
the piesence of honey in the flowers. One
of these is, in general, destined for the
elaboration of wax, and its size is consider-
ably enlarged when full of honey: The

S36

ORIGIN OF WAX.

other immediately imparts what it has col-
lected to its companions : its abdomen under-
goes no sensible change ; or it retains only
the honey necessary for its own subsistence.
The particular function of the bees of this
kind is to take care of the young, for they
are not charged with provisioning the hive.
In opposition to the wax workers, we shall
call them 'small bees or nurses.

Although the external difference be in-
considerable, this is not an imaginary dis-
tinction. Anatomical observations prove
that the capacity of the stomach is not the
same : Experiments have ascertained that
one of the species cannot fulfil all the func-
tions shared among the workers of a hive.
^^Ve painted those of each class with differ-
ent colours, in order to study their proceed-
ings ; and these were not interchanged. In
another experiment, after supplying a hive
deprived of a queen with brood and pollen,
we saw the small bees quickly occupied in
nutrition of the larvre, while those of the
wax working class neglected them.

When hives are full of combs, the wax
workers disgorge their honey into the ordi-
nary magazines, making no wax : but if
they want a reservoir for its reception, and

ORIGIN OF WAX.

S37

if their qiieeu does not find cells ready made,
wherein to lay her eggs, they retain the
honey in the stomach ; and in twenty-four
hours wax exudes through the rings. Then
the labour of constructing combs begins.

Perhaps it will be supposed that, when
the country does not afford honey, the wax
workers may consume the provision stored
up in the hive. But they are not permitted i

to touch it. A portion of honey is care-
fully preserved; and the cells containing it
are protected by a waxen covering, which
never is removed except in case of extreme
necessity, and when honey is not to be I

otherwise procured. The cells are at no |

time opened during summer; other reser- (

voirs always exposed contribute to the daily I

use of the community; each bee, however, I

supplying itself from them with nothing but
what is required by present exigence. j

AVax workers appear with large bellies at '■

the entiance of their hiv'e only when the ''

country affords a copious collection of honey. i

^ T rom what we have said, it may be con- i

I eluded that production of the waxy matter I

depends on a concurrence of circumstances i

( not invariably subsisting. (

I

338

ORIGIN OF -VVAN;.

Small bees also produce wax, but in a
my inferior quantity to what is elaborated
by the real wax workers.

Another characteristic, whereby an atten-
tive observer can determine the moment of
bees collecting sufficient honey to produce
wax, is the strong odour of both these sub-
stances from the hive, which is not equally
intense at any other time.

From such data, it was easy for us to dis-
cover whether the bees worked in wax in
our own hives, and in those of the other
cultivators of the district.

The inclemency of the season in 1793
had retarded the departure of swarms until
after the 24th of May. INIost of the hives
swarmed in the middle of June, the country
being covered with flowers. INIuch honey
was collected by the bees, and the new
swarms worked actively in wax.

Burnens inspected sixty-five hives on the
18th, and saw wax workers before the en-
trance of all. Those returning to the old
hives, speedily stored up their collections,
without coustructing combs; but those of
the swarms converted their honey to wax,
and hastened to prepare receptacles for the
eggs of their queen.

ORIGIN OF WAX.

339

The 19tli was showery; and although the
bees went out, they brought home nothing
but pollen. The weather continued chill and
rainy until the 27th, when we became de-
sirous of learning what had resulted from
it. Eurnens, raising all the hives on the
28th, found that the labour of the bees had
been interrupted ; that the combs measured
on the 9th had received no accession ; they
were of citron yellow, nor were white cells
in any of the hives.

Wax workers re-appeared on the first of
July, the weather being milder, and the
chesnut and elm having come in flower
They collected much honey; the swarms
enlarged their combs; great activity pre-
vailed ; and these labours continued until
the middle of the month. But on the 16th,
the thermometer rising above 77 drought
commenced: the flowers of the meadows,
together with those of the chesnut and elni,
had faded entirely. No longer containing
honey, their pollen alone attracted the bees,
and was copiously collected ; but they pro-
I duced no wax. The combs were not en-

I larged : those of the swarms remained sta-
: tionary.

1

340

ORIGIN OF wan.

Previous to the 10th of August it had
not rained for six weeks, neither did noc-
turnal dews temper the extraordinary heati
The bees found pollen alone, but no honey,
in the flowers of black grain, which now had
been spread during several days. After some
hours rain on the 10th, they exhaled the
odour of honey, and we actually saw it glit-
teriog on the faded flourish. The bees found
enough for their subsistence, but too little
to induce them to work in wax.

The drought returning on the 1 4th, con-
tinued until the end of the month ; when
we examined the sixty-five hives for the last
time. It appeared that the bees had prepared
no wax from the middle of July ; that they
had stored up much pollen ; that the pro-
vision of honey had diminished considerably
in the old hives, and that there was scarcely
any in the new.

The year, therefore, was very unfavour-
able to the labour of bees ; which I ascribe
partly to the atmosphere not being charged
with electricity, which has great influence
on the secretion of honey in the nectarium
of flowers. I have remarked, that the col-
lection by these creatures is never more abun-
dant, nor their operations in wax more active,

OEIGIN OF "WAX.

341

than when the wind is from the south, the air
moist and warm, and a storm approaching.
Heat too long protracted, however, and its
concomitant drought, chill rains, and a north
wind, entirely suspend the elaboration of
honey in vegetables, and consequently the
work of bees.

When bees were confined, for the purpose
of discovering whether honey was sufficient
for the production of wax, they supported
their captivity patiently, and showed un-
common perseverance in rebuilding their
combs according as we removed them. Had
part of the, combs been left, the queen would
have laid in the cells : we would have seen in
what manner the workers conducted them-
selves towards the young, and what the lat-
ter suffered from total privation of the fa-
rina. But occupied at the time solelv re-
garding the origin of wax, we preferred a
separate inquiry into the subject of rearing
their young.

Our experiments here required the pro-
sence of larvae ; honey and water had to be
provided : the bees were to be supplied with
combs containing brood, and at the same

342

ORIGIN OF WAX.

time it was necessary to confine them, that
they might not seek pollen abroad. Hav-
ing a swarm by chance, which had become
useless from sterility of the queen, we
devoted it for our investigation in one of
my leaf hives, which was glazed on both
sides.

We removed the queen, and substituted
combs full of brood for those of the first and
last division, that is, containing eggs and
young larvse, but no cell with farina ; the
smallest particles of tlie substance, which
John Hunter conjectured to he the basis of
the nutriment of the young, were even taken
away.

Nothing remarkable occurred during the
first and second day : the bees brooded over
the young, and seemed to take an interest
in them ; but at sunset, on the third aloud
noise was heard in the hive. Impatient to
discover the reason, we opened a shutter,

. and saw all in confusion: the brood 'was
abandoned : the workers ran in disorder
over the combs : thousands rushed towards
the lower part of the hive ; and those about
the entrance gnawed at its grating. Their
design was not equivocal: they wished to

ORIGIN OF WAX.

343

quit their prison. Some imperious neces-
sity evidently obliged them to seek else-
where what they could not find in the hive;
and apprehensive that they might perish if
I restrained them longer from yielding to
their instinct, I set them at liberty.

The wdiole swarm escaped ; hut the hour
being unfavourable for their collections, they
flew around the hive, and did not depart far
from it. Increasing darkness and the cool-
ness of the air compelled them very soon
to return. Probably these circumstances
calmed their agitation, for we observed them
peaceably remounting their combs ; order
seemed re-established, and we profited of
this moment to close the hive.

Next day, the 19th of July, we saw the
rudiments of two royal cells, which the bees
had formed on one of the brood combs.

This evening, at the same hour as on the
preceding, we again heard a loud buzzing in
the close hive; agitation and disorder rose
to the highest degree, and we were again ob-
liged to let the swarm escape. The bees
t|i did not remain long absent from their habi-
^ tation; they quieted and returned as before.

* AN e remarked on the 20th, that the royal
cells had not been continued, as would have

344

ORIGIN OF WAX.

been the case in the ordinary state of things.
A great tumult took place in the evening,
the bees appeared to be in a delirium ; we
set them at liberty, and order was restored
on their return.

Their captivity having endured five days,
we thought it needless to protract it farther ;
besides, we were desirous of kno’fv’ing whe-
ther the brood was in a suitable condition,
if it had made the usual progress ; and we
wished also to try to discover what might be
the cause of the periodical agitation of the
bees.

Burnens, therefore, having exposed the
two brood combs, the royal cells were im-
mediately recognised; but it was obvious
that they had not been enlarged. Why
should they ? Neither eggs, worms, nor that
kind of paste peculiar to the individuals of
their species were there ! The other cells
were vacant likewise, no brood, not an atom
of paste was in them. Thus the worms had
died of hunger.

Had we precluded the bees from all means
of sustenance, by removing the farina?

To decide this point, it was necessary to
confide other brood to the care of the same
insects, now giving them abundance of pollen.

OIIIGIX OF WAX.

345

They had not been enabled to mate any
collections while we examined their combs.
On this occasion they escaped in an apart-
ment where the windows w'ere shut ; and
after substituting young worms for those
they had allowed to perish, we returned
them to their prison.

Next day wt remarked that they had re-
sumed courage : they had consolidated the
combs, and remained on the brood. They
were then provided with fragments of combs,
where other workers had stored up farina ;
and to be able to observe w'hat they did with
it, w'e took this substance from some of their
cells, and spread it on the board of the hive.

The bees soon discovered both the farina
in the combs and what w^e had exposed.
They crowded to the cells, and also descend-
ing to the bottom of the hives, took the
pollen grain by grain in their teeth, and con-
veyed it to their mouths. Those that had
ate it most greedily mounted the combs
before the rest, and stopping on the cells of
the young worms, inserted the head, and re-
mained tliere for a certain time, llurnens
opened one of the divisions of the hive
gently, and powdered the workers, for the
purpose of recognising them when they

346

ORIGIN OF WAX.

should ascend the combs. He observed them
during several hours, and by this means as-
certained that they took so great a quantity
of pollen only to impart it to their young.

Royal cells were sketched on the 23d.
Next day, removing the bees which concealed
the brood, we found that all the young
worms had jelly as in the ordinary hives ;
that they had grown and had advanced in
their cells, and that others had been lately
closed up, probably from the period of their
metamorphosis approaching. Finally, from
observing the royal cells prolonged, we could
not doubt the restoration of order.

Then withdrawing the portions of comb
which had been placed by us on the board
of the hive, we saw that the pollen had been
sensibly diminished in quantity. They were
returned to the bees, to augment their pro-
vision still farther, for the purpose of
extending the experiment. The royal, as
well as several common cells •were soon
closed ; and on opening the hive, all the
worms were found to have prospered; Some
still had their food before them ; the cells
of others that had spun were shut with a
waxen covering.

Or.IGIX OF WAX-

34^7

This result was already very striking;
but it particularly excited our astonish-
ment, that notwithstanding such long pro-
tracted captivity, the bees did not now
seem to have any desire to go out. That
agitation, that increasing and periodical
disorder, and that common impatience mani-
fested in the first part of the experiment, had
ceased. Several bees, indeed, attempted
to escape in the course of the day, but find-
ing it impossible, they returned peaceably
to their young.

We witnessed these facts repeatedly, and
always with equal interest- They so deci-
sively prove the regard of the bees towards
the larvae which they are entrusted with
rearing, that we shall not seek for any other
explanation of their conduct.

Another fact, no less extraordinary, and
much more difficult to be accounted for, was
exhibited by bees constrained to work in
wax, several times successively, from the
syrup of sugar. Towards the close of the
experiment they ceased to feed the young,
though in the beginning these had received
the usual attention. 1 hey even frequent-
ly diagged them from their cells, and carried
them out of the hive..

348

ORIGIN OF WAX.

Ignorant what this disposition should be
ascribed to, I endeavoured to revive their
instinct, by supplying other brood for their
care : my attempts were abortive ; for not-
withstanding their stores of farina, they did
not feed the young larvae. We offered them
honey, as a more congenial means of aliment-
ing the brood, but also in vain; for the whole
perished. Perhaps the bees were incapable
of longer producing that paste which is
the subsistence of larvae. Except in this
particular, they seemed to have lost none of
their faculties ; they were alike active and
laborious. At last, influenced by motives
unknown to us, one day all deserted their
hive together, and did not return.

AVhatever is the cause affecting the in-
stinct of bees fed too long on honey, perhaps
we should consider, with admiration, that
this substance undergoes such a modification
in flowers, that bees can consume it without
inconvenience. But every thing in nature
is adapted for long continued use ; and ali-
mentary substances are combined with so
much foresight, that their action never pro-
ceeds from isolated and unabated energies.

architecture of bees. 349

CHAPTER VIII.

ON THE ARCHITECTURE OF BEES.

But it is time to examine liow bees con-
vert the substance exuding from their rings
to use, and to investigate the treatment
changing it to real wax : for it does not
leave the receptacles wherein it is moulded
in a perfect state. Likewise, it differs there
in several respects from its condition after
having been wrouglit. Fusibility alone indi-
cates it to be wax : it is short and friable,
wanting that ductility subsequently ac-
quired; and at first it is transparent as
plates of talc, whereas it appears opaque
and of a whitish yellow in the cells.

350 AUCHITECTURE OF BEES.

Perhaps it may be supposed that bees are
provided with instruments corresponding to
the angles of their cells. These can be only
their teeth or mandibles, their feet, andanten-
nse : yet there is no more resemblance between
the form of the teeth and the angles of- the
cell, than between the chisel of the sculp-
tor, and the work quitting his hands. Plate
III. fig. 17, 18, 19. Nor will the figure of
the head afford a better explanation. The
limbs are fashioned like those of most in-
sects, consisting of the haunch, fig. 20, a; of
the thigh, b ; the leg, c ; and the foot or tar-
sus, d, e. There is a hollow part in the leg
of the third pair, called the basket by
Reaumur, wherein bees deposit the farina ;
fig. 20, 21, c; triangular, smooth, with
a row of hairs on the outside, resembling a
kind of basket by their inclination. The
first articulation of the tarsus is much larger
than the rest, and in all the three pairs is of
a very different form from those of other in-
sects of the same genus ; fig. 20, 21, d.
Being employed in collecting the globules of
farina scattered over the body of the bee,
this articulation is called the brush. That
of the third pair of limbs offers some re-
markable peculiarities in the prolongation

architecture of bees. Sol

of the lower part, and in the mode of its
junction with the leg or pallet, by the figure
of which, and its own incurvature, a pair of
real pincers is formed. A row of shelly
teeth, like those of a comb, proceed from
the lower edge of the pallet, fig. 22, a, cor-
responding to bundles of very strong hairs,
with which the neighboiniug portion of the
brush is provided. When the two sides of
the pincers meet, that is, the under edge of
the pallet, and the upper edge of the brush,^
the hairs of each are incorporated together.

This organization is too obvious, not to
have a particular end. Kothiug resembling
it appears on the limbs of males or queens;
but it is seen in humble bees, a race very
near to the honey bee, and whose habits bear
some analogy.

Farther, the tarsus is composed of three
small conical articulations, together ^ith a
fourth, which is greatly elongated, and ter-
minates in two pair of claws. Perhaps Reau-
mur is right in considering this last as be-
ing truly two articulations.

The antennas of bees consist of twelve arti-
culations; the first two forming a peculiar sec-
tion, move in every direction on the base,
and serve to support the next division, com-

352 ARCHITECTUEE OF BEES.

posed of the other ten. The first articulation
of the whole is globular; the second cylindri-
cal, and much elongated: the third, which
commences the second division, is very short
and conical ; the second very long and coni-
cal ; the remainder cylindrical ; and the last
terminated by a soft point. Thus the flexi-
bility of the antennas enables them to follow
the outline of every object.

But the formation of the teeth, limbs, and
antennae cannot, in any respect, explain the
structure of the cells, though they can be
employed in it; and their effect depends en-
tirely on the object wliich the insect pro-
poses. Neither does the figure of the waxy
scales correspond with the form of the dif-
ferent parts of the cells they are appropriat-
ed to build. Hunter inferred that the bot-
tom was nearly equal in thickness to one of
them, and that they were accumulated in
fashioning the sides of the cells.

Some persons may imagine, perliaps, tliat
glass hives of four sides are sufficiently
adapted for exposing the construction of
combs : but their architecture is always con-
cealed from our view by clusters of bees,
amidst which, and in darkness, the work
goes on.

t

A li

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M

u

"4

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w- •i ■ ' ’ ^

* :

' |»^*

t

ARCHITECTURE OF BEES. 353

Having taken a large bell- shaped glass
receiver, I glued thin wooden slips to the
arch at certain intervals, because the glass
itself was too smooth to admit of the bees
supporting themselves on it. A swarm, con-
sisting of some thousand workers, several
hundred males, and a fertile queen, was in-
troduced, and they soon ascended to the
top. Those first gaining the slips fixed
themselves there by the fore feet : others
scrambling up the sides, joined them, by
holding their legs with their own, and they
thus formed a kind of chain, fastened by the
two ends to the upper parts of the receiver,
and served as ladders or a bridge to the
workers enlarging their number. The lat-
ter were united in a cluster, hanging like an
; inverted p^namid from the top to the bot-
tom of the hive.

The country then affording little honey,
I we provided the bees with syrup of sugar,
P in order to hasten their labour. They
I crowded to the edge of a manger contain-
I ing it, and having satisfied themselves, re-
I turned to the group. We were now struck
I with the absolute repose of this hive, con-
P trasted with the usual agitation of bees. All
the external stratum of the cluster consti-*

354 ARCHlTECTUllE OF BEES.

tuted a kind of curtain formed exclusively
of wax workers united together, and arran-
ged so as to represent a series of festoons in-
tersecting each other in every direction.
The fluctuation of the interior alone com-
municated motion to this cluster, where the
back of the bees generally was opposed to
the observer.

Meanwhile, the small bees alone went to
forage in the country ; they returned with
farina, kept guard at the entrance of the
hive, cleansed it, and stopped up its edges
with the odoriferous resin propolis. The
\vax workers remained motionless above fif-
teen hours; the curtain consisting always of
the same individuals, assured us that none
replaced them, Some hours later we re-
marked that almost all these individuals had
scales under the rings : and next day this
phenomenon was still more general. The
bees forming the external stratum of the
cluster, now having somewhat altered their
position, enabled us to see tbe under part
of the abdomen distinctly. By the pro-
jection of the scales, the rings seemed
edged with white : the curtain was rent in
several places : less tranquillity reigned in
liive,

architecture of bees. 355

Convinced that the combs would origi-
nate in the centre of the swarm, our whole
attention was then directed towards the
vault of the receiver. A. worker at this
time detached itself from one of the central
festoons of the cluster, separated from the
crowd, and with its head drove away the
bees at the beginning of the row in the
middle of the arch. Here turning round, it
formed a space for its free motion, an inch
or more in diameter, and fixed, itself in the
centre.^

The worker now employing the pincers
at the joint of one of the third, pair of limbs,
seized a scale projecting from a ring, and
brought it forward to the mouth with the
claws of the fore legs, where it appeared in
a vertical position, Plate IV. fig. 23, 24.

We remarked that the claws turned it
in every necessary direction ; that the edge
of the scale was immediately broke down,
and the fragments having been accumulat-
ed in the hollow of the teeth, issued forth
like a very narrow ribbon, impregnated
with a frothy liquid by the tongue. The
tongue itself assumed the most varied shape.

* This description is not quite intelligible. — T,

356

ARCHITECTURE OF BEES.

and executed the most complicated opera-
tions : and after imbuing the whole sub-
stance of the ribbon, pushed it forward, w'hen
it was dra^vn out a second time, hut in an
opposite direction.

At length, the bee applied these particles
of wax to the vault of the hive where the
gluten impregnating them promoted their
adhesion : and also communicated a white-
ness and opacity, which were wanting v/hen
the scales left the rings. Doubtless, this
process was to give the wax that ductility
and tenacity belonging to its perfect state.
The bee then separated those portions, not
yet applied to use, with its teeth, and with
the same organs afterwards arranged them
at pleasure.

The founder, a name appropriate to this
insect, continued the like manoeuvre until
all the fragments, worked up and impreg-
nated with the fluid, were attached to the
vault ; when it repeated the preceding oper-
ations on the part of the scale yet kept
apart, and united what was obtained from it
anew to the rest.

A second and third scale w^ere treated
thus by the same bee : yet the work was
only sketched ; for the worker did nothing

/

ARCHITECTURE OF BEES. 357

but accumulate the particles of wax to-
gether.

INIeanwhile, the founder, quitting its po-
sition, disappeared amidst its companionsj
Another, with wax under the rings, suc-
ceeded it ; which, suspending itself to the
same spotj withdrew a scale by the pincers
of the hind legs, and, passing it through
the teeth, prosecuted the work. Observing
to make its deposit in a line with the
former, it united their extremities. A third
worker, detaching itself from the interior of
the cluster, now came and reduced some of
the scales to paste, and put them near the
materials accumulated by its companions, but
not in a straight line. One, apparently sen-
sible of the defect, removed the misplaced
wax before our eyes, and carrying it to the
former heap, deposited it there exactly
in the order and direction pointed out.
From all these operations was produced a
block of rugged surface, depending from the
arch, without any perceptible angle, or any
traces of cells. It was a simple partition,
running in a straight line, and without the
least inflection, two-thirds of an inch in
1 length, about two-thirds of a cell high, and

358 ARCHITECTURE OF BEES.

declining towards the extremities. We
have seen other blocks from an inch to an
inch and a half long ; the form always the
same, but none ever of greater height.

The vacuity in the centre of the cluster had
permitted us to discover the first manoeuvres
of the bees, and the art with which they
laid the foundations of their edifice. How- ■
ever, it was filled up too soon for our sa-
tisfaction ; for workers collecting on both
faces of the block, obstructed our view of .’
their farther operations.

But although unable to behold all that,
we might desire, we concluded ourselves very
fortunate in having an opportunity of doing'
justice to Reaumur, who thought he observ-
ed the wax discharged from the mouth of'
bees as a paste. Unquestionably he sup-
posed the whitish frothy matter, v;ith which
the waxy substance is moistened, to be wax.

Construction of cells. — The cells of bees .
consist of two parts, a prismatic hexagonal
tube, and a pyramidal bottom. The latter,
which must be considered the most dc--
licate and essential part of the work, is.
composed of three equal lozenges, simi-

ARCHITECTURE OF BEES. 359

lar, uniting in a common centre, and form-
ing a slight cavity by their reciprocal incli-
nation.

Their depression into one face of the comb
makes a projection on the other, there cor-
responding to three cells partially common
to the whole.

It is evident that the hexagonal figure of
cells admits of their application by only one
angle to the surface of the roof, where
many are ranged laterally, and that there
must he large vacuities between the angles.
But a more solid fixture becomes the
marked solicitude of nature, at two epochs ;
1. On the formation of combs ; 2. When
these have become too heavy to be trusted
to so fragile a support.

The first row of cells, that by which the
whole comb is attached to the roof of the
hive, differs from all the rest. Instead of a
hexagon, the orifice is an irregular pentagon.
Plate IV. fig. 25, 26, 27. The cell con-
sists of four sides, with the roof of the hive
;in place of a fifth. The bottom also is dif-
ferent from that of the common cells, con-
sisting of three pieces on the face of the
comb, and on the other side of two. Only
one of these pieces is a lozenge, fig. 27,

360

ARCHITECTURE OF BEES.

28, c; the other two are of an irregular
quadrilateral figure, a, b. On the opposite
side of the comb, the bottom of the cells of
the first row is composed only of two quadri-
lateral pieces ; fig, 26, b, 29, a, b. Those of
faces are seen in front and reverse ; fig.
30, 31 *

By the simple dispositions preserved here,
the stability of the comb is completely in-
sured; for it touches the interior surface of'
support in the hive, in the greatest possible
number of points.

The innumerable crowd accumulated where
the labours of bees are conducted obstructs
the view of the observer : whence it appeared
to me, that the only method of isolating
the architects would be inducing them to
change the direction of their operations, and
work upwards.

I had a box made twelve inches square
and nine deep, with a moveable glass lid.
Plate I. fig. 5, a, b. Combs full of brood
honey and farina were next selected from one
of my leaf hives, as containing what might
interest the bees, and being cut into pieces I

* It will be observed, on referring to the engravings, i
that some of the figures are magnified in explaining the con-
struction of the cells. — T.

ARCHITECTURE OF BEES. 361

a foot long, and four inches deep, they ^ve^e
arranged vertically at the bottom of the box,
at the same intervals as the insects them-
selves usually leave between them. A small
slip of wooden lath covered the upper edge of
each. It was not probable that the bees would
attempt to found new combs on the glass
roof of the box, because its smoothness pre-
cluded the swarm from adhering to it ; there-
fore, if disposed to build, they could do so
over the slips, resting on the combs, which
left a vacuity five inches high above them.

But had it not been for the assistance of
Burnens, to wdiose skill, courage, and assi-
duity 1 have been so much indebted in
these difficult observations, my invention
would have proved of little avail. The ap-
pearance of the objects was affected by the
glass interposed, and, spite of my remon-
strances and the imminent danger, lie re-
solved to remove it. The gentleness of his
motions, and the habit of repressing his
respiration, could alone preserve him from
the wrath of such formidable insects.

As we had foreseen, the swarm with
which this box was peopled established it-
self among the combs below. ^Ye then ob-
served the small-bellied bees displaying their

R

262 ARCHITECTURE OF EEES.

natural activity. They dispersed themselves
throughout the hive, to feed the young lar-
vae, to clear out tlieir lodgment, and adapt
it for their convenience. Certainly the combs,
which were roughly cut to fit the bottom of
the box, and in some parts damaged, appeared
to them shapeless and misplaced ; for they
speedily commenced their reparation. They
beat down the old wax, kneaded it between
their teeth, and formed binding to consolidate
them. We were astonished beyond expres-
sion by such a multitude of workers employed
at once in labours to v/hich it did not appear
they should have been called, at their coin-
cidence, their zeal, and prudence.

But it was still more wonderful, that
about half the numerous population took no
part in the proceedings, remaining motion-
less, while the others fulfilled the func-
tions required. The wax w’orkers, in a state
of absolute repose, recalled our former ob-
servations. Gorged with the honey w'e had
put within their reach, and continuing in
this condition during twenty-four hours,
they had secreted that substance so long be-
lieved to be collected from the anthera; of
flowers. Th.e wax, formed under their rings,
was now ready to be put in operation ; and, !

AECHITECTURE OF BEES. 363

to our great satisfaction, we saw a little block
rising on one of the slips that w'e had pre-
pared to receive the superstructure.

Ts^o obstacle was offered to the progress
of our observations; and, for the second time,
we beheld both the undertaking of the
founder, and the successive labours of seve-
ral wax workers, in forming the block.
"Would that my readers could share the in-
terest which the view of these architects in-
spired !

The block, originally very small, was en-
larged as the work required ; and here they
excavated a hollow on one side, of about the
width of a common cell ; and on the opposite
surface two others somewhat more elongated.
The middle of the single cell corresponded
exactly to the partition separating the lat-
ter ; fig. 32, 33. The arches of these ex-
cavations, projecting by the accumulation of
wax, were converted into rectilinear promi-
nences; whence the cells of the first row
were pentagonal, considering the slip as one
side; and those of the second row hexago-
nal, fig. 34, 35.

The interior conformation of the cavities
apparently was derived from the position of
their respective outlines. It seemed that

364 ARCHITECTURE OF BEES-

the bees, endowed with an admirable deli-
cacy of feeling, directed their teeth princi-
pally to the place where the w'ax was thick-
est ; that is, the parts where other workers
on the opposite side had accumulated it:
and this explains why the bottom of the
cell is excavated in an angular direction be-
hind the projection on the sides of which
the sides of the corresponding cells are to rise.

The largest of the excavations, which was
opposite to three others, was divided into
three parts, while the excavations of the first
row on the other face applied against this one
were composed of only two.

In consequence of the manner in which
the excavations were opposed to each other,
those of the second row, and all subsequent,
partially applied to three cavities, were com-
posed of three equal lozenges, as the figures
explain. I may here remark, that each
part of the labour of bees appears the na-
tural result of wdiat has preceded it ; there-
fore chance has no share in these admir-
able combinations. But let us enter into
farther detail.

First roxo of cells. — A block rose above i
the slip like a minute vertical partition, five or
six lines long, two lines high, but only half a

AKCHITECTURE OF BEES. 86o

line in thickness ; * the edge circular, and
the surface rough. Plate IV. fig. 36, 37.
Quitting the cluster among the combs, a
small bee mounted the slip, turned around
the block, and visiting both sides, began
to work actively in the middle. It remov-
ed as much wax with its teeth as might
equal the diameter of a common cell ; and
after kneading and moistening the particles,
deposited them on the edge of the excava-
tion. This insect, having laboured some se-
conds, retired, and was soon replaced by an-
other : a third continued the w’ork, raising
the margin of the edges, now projecting from
the cavity, and with assistance of its teetli
and feet fixing the particles, so as to give
these edges a straighter form. More than
twenty bees successively participated in the
same work ; and when the cavity was little
above a line and a half in height, though
equalling a cell in width, a bee left the
swarm, and after encircling the block, com-
menced its operations on the opposite face,
where yet untouched. But its teeth acting
only on one-half of this side, the hollow

The reader is reminded that all the measurements for
this work are originally stated in lines, of which there are .
twelve in an inch. — T,

366 ARCHITECTURE OF BEES.

which it formed was opposite to only one
of the slight prominences bordering the
first cavity. Nearly at the same time an-
other worker began on the right of the face
that had been untouched, wherein both
were occupied in forming cavities, which may
be designed the second and third; and they
also w’ere replaced by substitutes. These
two latter cavities were separated only by
the common margin, framed of particles of
wax withdrawn from them ; which margin
corre.sponded with the centre of the cavity
on the opposite surface. The block itself
was still of insuflftcient dimensions to admit
the full diameter of a cell ; but while the
excavations were deepened, wax workers, ex-
tracting their scales, applied them in en-
larging its circumference : so that it rose
nearly two lines farther around the circular
arch. The small bees, which appeared more
especially charged with sculpturing tlie cells, ’
then being enabled to continue their out-
lines, prolonged the cavities, and heightened
their margins on the new addition of wax.
Plate V. fig. 38, 39.

Next, the arch, formed by the edge of
each of these cavities, was divided as by
two equal chords, in the line of which the '

3 € ^ •

Ar.ClIlTECTUIlE OF BEES.

367

bees formed stays or projecting borders or
margins meeting at an obtuse angle ; fig. -fO,
f e b : 41, i b c — c f 1. The cavities now had
four margins, two lateral and perpendicular
to the supporting slip, and two oblique, which
were shorter.

Meantime it became more difficult to
follow the operations of the bees, from their
frequently inteiqmsing the head between the
eye of the observer and the bottom of the
cell ; but the partition, whereon their teeth
laboured, had become so transparent, as to
expose what passed on the other side.

The cavities of which we speak form-
ed the bottom of the first three cells ; and
while the bees engaged w^ere advancing
them to perfection, other workers commenced
sketching a second row of cells above the
first, and partly behind those in front ; for
in general their labour proceeds by combina-
tion. We cannot say, “ when bees have
finished this cell, they will begin new ones;”
but, “ while particular workers advance a
certain portion, others carry on the adja-
cent cells.” Farther, the work begun on
one face of the comb is already the com-
mencement of that which is to follow on the
reverse. All this depends on a reciprocal

368 ARCHITECTURE OF BEES.

relation, or a mutual connection of the parts,
rendering the whole subservient to each
other. It is undoubted, therefore, that slight
irregularities on the front will affect the
form of the cells on the back of the comb.

Second row of cells. — The bottoms of
the first row of cells, composed of two tra-
peziums and a rhomb, we observed to be
larger than those of the cells opposed to
them, as these consisted of two trapeziums
only.* The space between the edge of these
back cells and the edge of the block, ad-
mitted the bottom of a common cell, fig.
42; but there was not room for a complete
bottom above those in front, as the block
rose no higher than r, fig. 43. Several bees
employed themselves in sketching the bottom
of a new cell, in the unwrought portion of
the reverse, fig. 42, proceeding in the follow-
ing manner.

They first excavated a vertical fluting,
f m b p, in the space comprised between
the oblique margins f c, b c, of two neigh-
bouring cells, fig. 44, and produced mar-
gins, by accumulating to right and left

* The unlearned reader may be apprized tiiat trapezium
here merely signifies a four-sided figure, and rhomb, a
lozenge. — T,

ARCHITECTUEE OF BEES. 569

the wax extracted from the block. The
perpendicular margins, f m, b p, formed
by the workers, rose exactly above the an-
gles f, b, of the two lower cells, a g, fig.
44. At this time the height of the block
was insufficient for the whole bottom of
a cell, and the fluting as yet was termi-
nated by a curved outline, p, r, m, fig. 44.
But other bees established two rectilinear
margins on the curvature, which, imiting as
two chords in the centre of the arch, formed
the obtuse angle m r p, fig. 42. The cavity
was then bordered by six margins of equal
length, forming the outline of a cell, but parts
of them somewhat more elevated than others.

The bees now occupied themselves in flat-
tening the back of the rhomboidal portion,
f c b e, fig. 42, confining it by the traces
f e, b e, wliich they had excavated behind
the corresponding margins on the opposite
j face. Thus this portion was the first and
I upper part of a pyramidal bottom, and it
occupied a third of the surface of the cavity,

|| there being space enough hollowed out with-
in the hexagon for two similar lozenges,
i This space, as yet only sketclied, remain-
i ed in that state until the operations on the

370 ARCHITECTURE OF BEES,

opposite face of the comb allowed the bees
to construct a vertical margin behind the
same cell, which could result only from com-
mencing a new cavity on each side of that
margin ; but it being established, a bee be-
gan to excavate the bottom of the space re-
ferred to, still rough, and formed a furrow
-also vertical in the middle, running from
the upper angle of the lozenge to the upper
angle of the hexagon. The two pieces, re-
sulting from the division, being smoothed,
we observed that they constituted two
new lozenges, f e r m, and e r b p, equal to
the preceding lozenge f c b e. Thus the
six margins of the hexagonal outline sur-
rounded three equal sized lozenges, that is,
a complete pyramidal bottom.

The first bottom of this kind was con-
structed on the posterior face of the block ;
and, during all these operations, cells were
sketched in the same manner to right and
left. Meantime the wax workers were en-
larging the block, which the punctuated line,
in fig. 43, shows to have been still inade-
quate during the construction of the poste-
rior cell of the second row, but it was large
enough for new cells of this row, when they
commenced the anterior hexagonal cell.

371

architecture of bees.

From what has been said, it is easy to
conceive that the bottom of the subsequent
cells is always commenced between the ob-
lique upper margins of two adjoining cells.
Vertical margins are formed above the angle
of each, bordering the new cavity to right
and left, and the total circumference is com-
pleted by other two oblique margins on the
upper curvature of the hollow, whereoy a
hexagon is produced.

The lower portion of these excavations
always corresponds with the intermediate
margins of the cells on the opposite side of
the comb, whereby all on the front, so to
design it, have two lozenges below and one
above, while the.cells of the reverse have one
below and two above.

Each of the six margins, environing tire
pvTamidal bottom of a cell, is destined for
one of the six sides forming the tubular part
of it. At first sight, nothing appears more
simple than adding wax to the margins;
but from the inequalities, occasioned by the
shape of the bottom, the bees must accumu-
late wax on the depressions to bring them
to a level. All the edges of the cells tlicn
offer a unifonn surface from tire commence-
ment, and before the tubes have acquired

372 ARCHITECTURE OF BEES.

their proper dimensions. But the surface
of a new comb is not quite flat, for there is a
progressive degradation in the work of bees ;
the sides of the cells are prolonged in an or-
der corresponding to the completion of the
bottom to which the tubes belong; and the
length of these tubes is so well regulated,
that there is no interruption or conspicuous
irregularity among them. Thence the sur-
face of a new comb is lenticular, its thick-
ness always decreasing to the edges, because
the latest cells are shorter than the older
ones. While the circumference of the
comb extends, this figure is preserved ; but
when the bees have no more room for its
enlargement, they make all the cells equal,
by prolonging the newer to the dimensions
of the older. It gains two flat surfaces,
which it will constantly retain.

How can we explain the combination reign-
ing throughout the work of bees? How
does instinct lead them to give a difierent
form and different dimensions to the bottom
of the first row of cells on either face ? Or
how can those posted on one side of the
block determine the space to be excavated
for the mutual relation of their bases ? They
do not visit the two surfaces alternately to

ARCHITECTURE OF BEES.

373

compare the respective position of the eavi-
ties they are to sketch ; hut feeling the part
about to be hollowed out wdth the antennae,
they seem by this means sufficiently apprised
of the mode of executing a very complicated i

work, wherein every thing is exactly ar- j

ranged. • i

Nothing is excavated without previously j

applying the antennae to the part to be
sculptured : bees do not trust to their eyes i

for any of their operations ; but amidst dark-
ness, by the aid of their antennae, they can [

fashion even those combs wffiich are de- '

servedly regarded as the most admirable -
work of insects.

Bees, therefore, seem to be regulated in |
their work by some local circumstance. We !
J have observed that, while sketching the bot-
j tom of a cell before there was any upright I
margin on the reverse, a projection arose by ‘
their pressure on the wax still soft and
„ flexible. Sometimes it occasioned a breach
1 of the partition, which was soon repaired : '

But a slight prominence always remained on ^
the opposite surface, to the right and left of i
which they placed themselves to begin a
new excavation ; and they heaped up part of I
the materials between the two flutiims form-

O

1

S74

ARCHITECTURE OF BEES.

eel by their labour. The prominence, being
converted into a real rectilinear margin, be-
comes a guide for the direction to be fol-
lowed by the bees in making the vertical
furrow of the front cell.

We have often conceived, from the bees
observing the direction of the perpendicular
margin so accurately, as to work the corres-
ponding hollow, that they were ' aware of
the thickness of the partition, from its flexi-
bility, elasticity, or some other property of
the wax. Whatever may be the case, it is
certain that, without any mechanical means
of measuring it, they give the bottom of the
cells a uniform thickness.

Perhaps the proceedings of bees may be
explained without resorting to extraordi-
nary causes. The length of the cavities,
their respective position, and the thickness
of the block, once determined, the inclination
of the oblique sides of the trapeziums of the
first row, to which that of the lozenges of the
second is subordinate, is established of itself.

Probably the mode of enlarging the block
chiefly contributes to establish the relation
between the unequal cells of the first row.
Its original height determines nearly the
vertical diameter of the posterior cells.

li

architecture of bees. 375

whicli is 0qiial to two thirds of a common
cell. But the bottom of the front cell
cannot be completed without augmenting
the block, which, on that account, is extend-
ed more than necessary for finishing the
front cell, but allowing just sufficient space
for the whole bottom of a posterior cell of
the second row: for the lozenge making part
of it, is already comprised in the interval be-
t^Yeen the cells formed of trapeziums. Still
advancing the block two thirds of the dia-
meter of a cell, the bees are enabled to con-
struct the bottom of cells of the second row on
the/ro7z/, part being previously intercepted
between the upper edges of the first cells :
but as yet there will not be room for building
the thii-d row, nor until the block shall be
enlarged anew.

The bees cannot deviate from the rule
prescribed, unless particular circumstances
alter the bases of their work : for the block
is augmented only by a uniform quantity:
and what is admirable, this is done by the
wax workers, the depositories of its first
elements ; but which are not endowed with
the faculty of sculpturing the cells.

Thus, in sharing the functions of the
hive between the wax workers and the

376 ARCHITECTURE OF BEES.

other bees, the Author of nature seems to
have distrusted the exclusive operation of
instinct.

May not we deduce from the preceding
facts, that the geometry, which apparently
embellishes the productions of these insects,
is rather the necessary result than the prin-
ciple of their proceedings ?

MODIFICATIONS OF ARCHITECTURE. 377

CHAPTER IX.

MODIFICATIONS OF THE ARCHITECTURE
OF BEES.

The vulgar commonly believe that sensa-
tions and physical necessity exercise an ab-
solute empire over animals. Unquestion-
ably they have a powerful influence; but
although it has been maintained, that sel-
fishness is the sole motive of actions, it would
be as difficult to explain the'conduct of crea-
tures subjected to instinct, by the attrac-
tions of pleasure and the dread of suffering
exclusively, as it would be unjust to ascribe
the virtues of reflecting and reasoning beings
i to interested views alone. Xeither can any
[i theory explain the apparent exceptions from

378 MODIFICATIONS OF THE

general rules in the great code of nature, or
how animals of restricted faculties can act
in some circumstances, as if they could in-
terpret the design of the Legislator,

Every thing connected with the fabrica-
tion and use of combs has been skilfidly
combined. Cells turned downwards, like
those- of wasps, would not have suited bees,
for they have to store up a fluid. Perhaps
the figure of these innumerable minute
honey-pots, covering both surfaces of the
comb, together with the affinity between their
contents and wax, retains the liquid from
escaping. The combs hang parallel to each
other, and are separated by passages but a
few lines wide. This position is not one of
the least difficult subjects of explanation;
nor would it be practicable, did we conceive
that their foundation was laid simultaneously
by a number of workers. Blocks of wax are
not reared here and there at the same time.
A single worker deposits the materials in
what seems the suitable direction : it de-
parts : another replaces it : the block rises ;
and the bees sculpture its opposite faces al-
ternately. But scarcely are some rows of
cells constructed, when other two blocks si-
milar to the first arc established, one ou

ARCHITECTURE OF BEES. 379

each side, equally removed from it, and
parallel. These very soon become small
combs, for the bees work with astonishing
rapidity ; and in a short time we discover
other two, constructed in a position parallel
to the preceding. All are extended in pro-
portion to their earlier origin: that in the
middle, being the farthest advanced, exceeds
those parallel to its two faces by some rows
of cells ; and the latter exceed those follow-
ing them by a similar quantity. Thus, the
greater part of both surfaces of the comb
are always concealed by those next to them.

I shall not attempt to explain how the
bees take such accurate measurements, and
know the direction parallel to the original
comb. But were they pennitted to rear
ditferent blocks in the roof of their hive at
the same time, it is obvious that the combs
neither would be parallel, nor at suitable
distances.

It is invariably a single bee that selects
and determines the site of the first cavity,
which being established, serves to direct all
the ulterior labours. Did each of several
workers sketch a cell at once, the symmetry
to result from their operations would be en-
dangered ; for these insects, strangers to

380

MODIFICATIONS OF THE

subordination, are subject to no discipline.
A great many bees work on the same comb
indeed, but they are not guided by a simul-
taneous impulse, as might be conceived,
without studying their proceedings from
the outset. The impulse is successive. A
single bee begins each partial operation, and
several others substitute their efforts, tend-
ing to the same end. Each apparently acts
individually, either as directed by the bees
preceding it, or by the state in which it
finds the work which it is to continue. If
any thing can presume consent almost una-
nimous, it is the inaction of the rest of the
colony while a single worker proceeds to
determine the position of the comb. After
others assist it in extending the block,
they cease ; but a single individual of a dif-
ferent profession, if the word may be allow-
ed, comes to sketch the bottom of a cell, the
preparation for another kind of work. It
is a base or foundation for establishing the
whole edifice. A delicate sense of feeling
obtains through the partition tlie situa-
tion of the margin of the opposite cavity,
which aids the worker in the proper divi-
sion of the bottom of the new cells. But
it is not by means of the margin only that

ARCHITECTUEE OF BEES. 381

they discover the direction to be followed.
IV e are convinced that they profit by va-
rious circumstances to guide themselves in
their excavations.

The bee, however, which forms the first
cell, is a remarkable exception ; it works in
a rough mass, and thus has nothing to point
out the wav. Instinct is its sole conductor.
The workers sketching the cavities of the
second row, on the contrary, can profit by
the previous margins and angles of the same
surface, as a ground or point of departure
for future operations.

Their work is executed with much less
rapidity'in ascending, than when prosecuted
in the opposite direction. This was very
favourable to our observations, otherwise it
would have been impossible to follow the
detail of their proceedings. Yet the tardi-
ness of the bees here had its inconveni-
encies : their work was sometimes interrupt-
ed during hours : the materials were want-
ing, or they were not sculptured soon after
being deposited ; or several blocks arose on
the same slip. The operations evidently
were relaxed and impeded; and we were
enabled to form an acciurate opinion of their

S82 Modifications of the

architecture only from the different small
combs established.

To ascertain whether the above described
facts succeeded under ordinary circumstances,
I constructed a new hive, the top of which
consisted of glass and wooden slips or spars
alternately. By means of a screw at each end
of the slips, any one could he raised so much
as to admit of convenient observation, and it
might be replaced without deranging the
bees. Plate I. fig. 5, b.

Having lodged a swarm here, we allowed
sufficient time for commencement of the
first cells, when turning the screws to bring
up the work, we beheld cavities similar to
those formed by the operations of bees in as-
cending. Then lowering the slip, they con-
tinued, and our observations being renewed
in some minutes, we found the work farther
advanced. The cells of the two faces were un-
like each other: the bottom had vertical trape-
ziums, but only those in front had a lozenge
in the under part. Cells of the second row
were then begun, and every thing proceeded
on the same principles, and by the same gra-
dations as when the comb was constructed
upwards.

architecture of bees. S83

It is clear, in my opinion, therefore, that
the particular configuration of the first cells
of both faces determines invariably the figure
of the pyramidal bottom of all the subse-
quent cells.

IVhen I compelled the bees to work up-
wards, they usually commenced their combs
on the flat part of the slips. But they were
not always so docile, and repeatedly extend-
ed old combs in the place where I wished
them to build new ones.

If a comb be put under a spar, and the bees
work upwards, they begin by lengthening
the cells, so as to project a little beyond the
spar, and then go on with others on the ver-
tical surface or side of the spar ; but the
bottoms are flat, being the spar itself. On
gaining the upper edge, they extend the
extremities of the last constructed cells to
the middle of the horizontal surface of the
spar, where they raise a block, and sculpture
it after the prolongation of these cells. They
now give the shape peculiar to the bottom of
the foundation row to that of the first which
they establish here, and three lozenges to
the bottom of the subsequent cells ; fig. 45 *

The author s description is exceedingly obscure. T.

384.

MODIFICATIONS OF THE

Bees can thus form cells on wood of an
hexagonal figure, without having a pyra-
midal bottom, and perpendicular margins ~
opposite to guide them* But although these
flat bottomed cells are less regular than the
common cells, and some want angular mar-
gin Sj an hexagonal character is always dis-
cernible.

Having seen bees work both up and
down, it was natural to investigate whether
we could compel them to construct their
combs in any other direction. We tried
to confound them with a hive glazed above
and below, so that they had no placQ of sup-
port but the upright sides of their dwelling,
Ijodging themselves in the upper angle,
they built their combs perpendicular to one
of these sides, and as regularly as those
which tliey usually build under a horizontal
surface. The foundations were laid on a
place which does not serve naturally for the
base. Yet, except in the difference of di-
rection, the first row of cells resembled those
in ordinary hives. The others were no less
fit for use, distributed on both faces, and the
bottoms alternately corresponded with the ^
same symmetry.

AIICHITECTURE OP BEES. 385

I put the bees still to a greater trial. As
they now testified their inclination to carry
their combs, in the shortest way, to the op-
posite side of the hive, for they prefer unit-
ing them to wood, or a surface rougher than
glass, 1 covered it with a pane. Whenever
this smooth and slippery substance was inter-
posed between them and the wood, they de-
parted from the straight line hitherto follow-
ed, and bent the structure of their comb at
a right angle, to what was already made, so
that the prolongation of the extremity might
reach another side of the hive, which had
been left free.

^"arying this experiment after several fa-
shions, I saw the bees constantly change
the direction of their combs, when I ap-
proximated a surface too smooth to admit of
their clustering on it. They always sought
the wooden sides. I thus compelled tirem
to curve the combs in the strangest shapes

by placing a pane at a certain distance from
their edges.

1 hese results indicate a degree ofiiistinct
truly wonderful. They denote even more
■than instinct : for glass is not a substance
. against wliicli bees can be warned by nature
till trees, tlieir natural abode, tliere is nothin.^

386

MODIFICATIONS OF THE

that resembles it, or with the same polish.
The most singular part of their proceeding
is changing the direction’ of the work be-
fore arriving at the surface of the glass, and
while yet at a distance suitable for doing so.
Do they anticipate the inconvenience which
would attend any other mode of building ?

No less curious is the plan adopted by
the bees for producing an angle in the
combs ; the wonted fashion of their work,
and the dimensions of the cells, must be al-
tered. Therefore, the cells on the upper
or convex side of the comb are enlarged :
they are constructed of three or four times
the width of those on the opposite surface.
How can so many insects, occupied at once
on the edges of the combs, concur in giving
them a common curvature from one extrem-
ity to the other ? How do they resolve on
establishing cells so small on one side,
while dimensions vso enlarged are bestowed
on those of the other ? And is it not still
more singular, that tliey have the art of
making a correspondence between cells of
such reciprocal discrepance ? The bottom
being common to both, the tubes alone as-
sume a taper form. Perhaps no other in-
sect has afforded a more decisive proof of

A^lCHITECTUilE OF BEES. 387

tlie resources of instinct, when compelled to
deviate from the ordinary course.

But let us study tliem in their natural
state ; and there we shall find that the dia-
meter of their cells must be adapted to the
individuals which shall be bred in them.
The cells of males have the same figure, the
same number of lozenges and sides as those
of workers, and angles of the same size.
Their diameter is 31 lines, while those of
workers are only 21.

It is rarely that the cells of males occupy
the higher part of the combs. They are
generally in the middle or on the skies, where
they are not isolated. The manner in which
they are surrounded by other cells, alone can
explain how the transition in size is effected.
AVhen the cells of males are to be fabricat-
ed under those of workers, the bees make
several rows r)f intennediate cells, whose dia-
meter augments progressively, until gaining
that proportion pro})er to the cells required ;
and in returning to those of workers, a degra-
dation is observed in a manner correspondino*.

U^ually there arc three or four rows of
intermediate cells. The first of those for
males still participate in the irregularity of
the adjrjning margins; the bottoms there

388

MODIiFICATIONS OF THE

corresponding to four cells, instead of three.
Their furrows are always in the direction of
the margins; hut on one face the side of the
cell, instead of being immediately opposite
to the centre of the cell on the reverse, di-
vides it unequally, which alters the shape of
the bottom, so that it no longer contains
three uniform lozenges, but consists of pieces
more or less irregular.'*^

The farther removed, from the transition
cells, those of the males become more regular.
Several rows are free of any defect. But
the irregularity is resumed on their opposite
confines, nor does it disappear until several
rows of workers’ cells, of an extraordinary
fashion, are interposed.

Bees, in preparing the cells of males, pre-
viously establish a block or lump of wax on
the edge of their comb, thicker than is usual-
ly employed for those of workers. It is also
made higher, otherwise the same order and
symmetry could not be preserved on a larger
scale.

Several naturalists notice the irregulari-
ties in the cells of bees as so many defects.

* This part of the subject cannot be easily understood,
■without actually inspecting combs and attending to the
subsequent explanations. — T.

architectcr-e of bees. 389

"What would have been their astonishment
had they observed that part of them are the
result of calculation. Had they followed
the imperfection of their organs, some other
means of compensating them would have been
granted to the insects. It is much more
surprising that they know to quit the ordi-
nary route, when circumstances demand the
construction of enlarged cells; and after build-
ing thirty or forty rows of them, to return to
the proper proportions from which they have
departed, by successive reductions.

It appears that the particular species of
cells to be constructed by bees is determin-
ed by the laying of the queen. They never
build the cells of males so long as she pro-
duces the eggs of workers. But when she
wants a place for depositation of the for-
mer, they very soon become sensible of it ;
they are seen forming their cells irregularly,
gradually giving them greater diameter,
and finally preparing a cradle for reception
of the whole masculine race.

There is another circumstance under
which bees augment the dimensions of their
cells, namely, when there is au opportunity
for a great collection of honey. Xot onlv
arc they then constructed of a diameter

390 MODIFICATIONS OF THE

much exceeding that of the common cells^
hut they are elongated throughout the whole-
space admitting it. A great portion of ir-
regular comb contains cells an inch, or even
an inch and a half in depth.

Bees, on the contrary, sometimes are in-
duced to shorten their cells. When wdshr
ing to prolong an old comb, whose cells have
received their full dimensions, they gradu-
ally reduce the thickness of its edges, by
gnawing down the sides of the cells, until
restoring it to its original lenticular form.*
They add a waxen block ai'ound the whole
circumference, and on the edge of the comb
construct pyramidal bottoms, such as those
fabricated on ordinary occasions. It is a
certain fact, that a comb never is extended
in any direction unless the bees have thinned
tlie edges, which are diminished throughout
a sufficient space to remove any angular pro-
jection.

The law which obliges these insects partly
to demolish the cells on the edges of the
comb before enlarging it, unquestionably
demands more profound investigation. How.

* By lenticular, the shape of a magnifier is to be uiulcr-
stoocl, or a body thicker in, the inuUUe than at the circum--
ferencc. — T,

ARCHITECTURE OF REES. 391

eau we account for instinct leading them to
undo what they have executed Avith the ut-
most care ? The wonted regular gradation,
which mav be necessary for new cells, sub-
sists among those adjoining the edges of a
comb recently constructed. But afterwards,
when those on the edge are deepened like
the cells of the rest of the siu'face, the bees
no longer preserve the decreasing gradation
which is seen in the new combs. Thus it
is evidently for the purpose of restoring the
comb to its primitive form, which prepares
the circumference for enlargement, that they
reduce the depth of the cells proportionally
to their distance from the edge.

All the anomalies exhibited in the la-
bours of bees are so well appropriated to the
object proposed, that they seem to consti-
tute part of the plan under which the crea-
tures act, and which conciu' for the ereneral
order.

Appexdix hj P. Huber to the observa-
tions on the architecture of bees.

The original foundation of a comb com-
prehends tbree or four cells, and sometimes

V

392 MODIFICATIONS OF THE

more: and after being continued of that^
breadth for two or three inches, it begins
at about three-fourths of this length to be
enlarged.

W ere tbe work prosecuted only in descend-
ing, it would form a narrow stripe, and ad-
vance slowly. But it is necessary that it
should proceed rapidly, and that numbers of
bees should be able to operate in all direc-
tions. The more the comb is enlarged be-
low, the more essential that it should rise
immediately to reach the vault of tbe hive.

It thence results, that all the cells of the
first row are not constructed at once through-
out the hive. Those alone can be consi-
dered primitive which are built before the
broadening of the comb. Nevertheless, the
cells of that row formed either by ascend-
ing or obliquely have nearly the same shape
as the primitive cells; likewise, though there
be somewhat greater confusion and irregu-
larity, the general solidity suffers nothing.

Bees work in all directions : in every case
their proceedings are uniform. JMeanwhile
we should be unable to recognise the little
original waxen block, without adverting, that
it appears like a ribband two or three lines
broad, running round the edge of the combs,
and seemingly more compact than the rest.

ARCHI TECTUEE OF BEES. 393

In this edge the bees sculpture their new
cells ; and here they deposit their scales of
wax, labouring over the whole circumfer-
ence at the same time, when they have
abundance of that substance.

Xotwithstanding the work goes on in all
points, it does not advance in the same pro-
gression. Bees build quicker downwards
than horizontally, and slowest in ascending.
Thence ensues the kind of lens or elliptic
shape of the comb at the period of its en-
largement. Thence also the greater length
than breadth ; its being more pointed at
the lower extremity, and narrower towards
the top than in the middle. The shape of
the combs, therefore, is veiy’ regular ; their
outline in general is void of any asperity :
besides, a singular harmony subsists among
all the cells in their elongation. We have
previously stated, that this is in proportion
to their priority ; but investigating the fact
with greater attention, we have obser\'ed
that in a new comb their length is propor-
tioned to their distance from the edge. Thus
the first row of cells is not the deepest ; the
cells are there shallower than those in the
middle of the comb : but when the comb
acquires a certain weight, the bees hasten

394) MODIFICATIONS OF THE

to prolong those so essential to the solidity
of the whole; sometimes making them deeper
than what follow.

The cells are not perfectly horizontal, the
orifice being almost always a little higher
than the bottom ; which enables us to as-
certain the original position of a comb,,
though detached. lienee the axis of a cell
is not perpendicular to the partition separa-
ting the two faces of the comb. This fact,
which has been hitherto overlooked, is an
insuperable bar to geometriccd calculations
regarding the figure of cells, because they
have a certain inclination from the base.
Sometimes they, deviate from the level of
the horizon above 20°, and commonly 4 or 5.

Yet, whatever be their irregularities here,
they are less conspicuous than those of the
bottom; and frequently where these last
are irregular, the tubes preserve a hexagonal
shape.

Bees ill general observe a tendency to
symmetry, not so much perhaps in small
details, as in the totality of their operations.
Sometimes, indeed, the combs exhibit a sin-
gular formation ; but on following all the
rainutijE of labour, we can, for the most
part, assign reasons for the apparent ano-

395

ARCHlTliCTUllE OF EEES»

malies. These insects are obliged to adapt
themselves to localities : one irregularity
produces another, and it usually originates
in the arrangements which we make them
adopt. The inconstancy of the temirerature
of the atmosphere affects the symmetry of
the combs, from frequently interrupting the
operations of those bees entrusted with theii
construction. AVe have remarked, that less
perfection is offered by a work resumed^ than
one of uninterrupted labour.

On allowing too little interval betwreen
the spars for receiving the foundation ot
the combs, tbe work has assumed a particu-*
lar direction. At first the bees did not seem
sensible of the defect ; but very soon they
appeared to suspect their error, and gra-
dually changing the line of the w'ork,
they gained the customary distances. This
having given the comb a recurvature, new
ones commenced opposite to its middle ne-
cessarily had tlie like deformity, which was
imparted to all the others successively.
INIeantime, however, tlie bees eiulcavoured
as' much as possible to bring them back to
the regular form. Often a comb is convex
above ; farther down the defect is rectified.

396 MODIFICATIONS OF THE

and the surfaces of the lower part are brought
to the proper figure.

We have seen their regard for symmetry
exhibited in a manner still more decided.
From a series of preceding irregularities,
having been induced to plant two blocks
instead of one on a spar, but not in the same
line, they could neither be enlarged without
interrupting each other, nor could their
edges unite, from their respective position.
However, the bees adopted a very effectual
plan : they curved the edges of the two
combs, and brought them to meet so per-
fectly, that they could continue them toge-
ther. The part above this junction diverged
greatly from a straight line ; but in pro-
portion as the united combs were prolonged,
their surface became more and more level,
and at length perfectly uniform.

comb never commences Avith the cells
of males, the first rows being formed of very
regular small cells ; But the reciprocal cor-
respondence among them soon ceases to be
so exact, and tlie bottoms are less symme-
trical. It would be impossible for the bees
to produce an absolute correspondence be-
tween unequal cells and those perfectly re-
gular ; we frequently observe little masses

ARCHITECTURE OF BEES. 397

of wax occupying the intervals between them.
By thickening the sides and rendering the
circumference more circular, they sometimes
succeed in uniting cells of different diame-
ter, for they have more than one fashion of
compensating irregularities.

The modifications of the bottom of cells
are so much more decided and constant, as to
announce a determinate plan, and also ex-
plains their progressive enlargement. Ex-
amining a comb by a vertical line from the
beginning, and passing through the middle,
we observe that the cells next to this line
enlarge without much alteration of shape ;
but the bottom of those adjacent no longer
consists of three equal lozenges. Each, in-
stead of corresponding with three others,
corresponds with four cells on the opposite
face, while their orifices are not the less hex-
agonal. The bottom is composed of four
pieces, two of which are hexagonal, and two
rhomboidal or four sided, whose size and
figure vary. These cells, which are some-
what larger than the third part of three op-
posite cells, comprehend a portion of the bot-
tom of a fourth cell in their circumference.
Under the last regular^ pyramidal bottoms
are found tliose having a bottom consisting

398

MODIFICATION'S OF THD

of four parts, three very large and one a
rhomb, very small. A great interval se-
parates the two rhombs of the cells of tran-
sition, while the two hexagonal portions ad-
join and are perfectly similar ; Plate V. fig..
40, 47. There is less inequality between
the two rhombs of the bottom, a cell farther
down : its circumference embraces more of
the fourth cell on the opposite surface. Fi-
nally, we find a great number of cells whose
bottom consists of four pieces perfectly regu-
lar; namely, two elongated hexagons, and
two lozenges equal, but smaller than those of
the pyramidal bottom ; fig. 48. Proportion-
ally as we recede from cells whose bottom
contains regular four sided portions, whe-
ther by descending or proceeding from right
to left, we observe the cells recovering their
' ordinary form ; that is, one of the lozenges
contracts and at length disappears entirely ;
fig. 49, 50. The pyramidal sliape of the
bottom is resumed, but of larger dimensions
than in the cells of the higher part of the
comb. It is preserved with the utmost re-
gularity through many rows: afterwards the
cells arc altered, and we remark the pre-
sence of four pieces in the bottom, until the
common cells arc restored.-

auchitectoie of bees. 39^)

Thus it is by encroaching a little on the
limits of those of the opposite face, that
bees come at last to give their cells the
largest dimensions. The gradation of the
transition cells being reciprocal on both
faces of the comb, it follows that the hexa-
sonal circumference on each embraces four
cells.

Having reached any degree whatever of
this progression, the bees can stop there,
and preserve it in several consecutive rows.
They seem to continue longest at the me-
dium, where we find a great many cells with
the bottom quite regularly constructed of
four pieces. Therefore, they could build the
whole comb on this plan, were not their
purpose to resume the pyramidal form tiiey
have left. In diminishing the diameter of
their cells, bees return by similar gradations,
in an inverted order.

It would be very difficult to measure the
inclination of the four-pieced bottoms ; but
they seem a little shallower than those which
are pyramidal ; and it should be so, for the
two lozenges being smaller, the line con-
necting their extremities is less depresserl.

In general, it appears that the form of
the tube of the cell is more essential than

400 MODIFICATIONS OF ARCHITECTURE.

the shape of the bottom ; as we have seen
bottoms of four irregular pieces with hex-
agonal tubes, and cells also of six sides con-
structed on glass or wood which served for
the bottom instead of wax. The figure of
the bottom depends on the intersection of
the outline of the cells on both faces, in the
direction of the margins.

The shape of the sides of the four-pieced
bottoms differs according to the facettes from
Avhich they rise : those corresponding to one
side of a lozenge and part of a hexagonal
facette,are bevelled, that they may be adapted
to each other ; while the two sides correspon-
ding to the long side of the hexagon are
rectangular parallelograms.

COMPLETION OF THE CELLS. 401

CHAPTER X.

■ XX XX XX XX XX

COMPLETION OF THE CELLS.

The substance 'whereof the cells are formed,
is in its origin of a dull white colour, semi-
transparent, soft, and uniform, without being
smooth. But in a few days it loses most
of these qualities, or rather acquires new
ones. A yellowish tint pervades the in-
terior surface of the cells : their edges, now
much thicker than at first, are far less re-
gular; and, from being incapable of resisting
the slightest pressure, they have attained a
consistence of which they did not seem sus-
ceptible.

IVe had remarked the greater propor-
tional weight of finished combs than of those

402 COMPLETION OF THE CELLS.

in a progressive state ; tliat the latter break
^ on the slightest touch, whereas the others
rather yield than fracture. Their orifices
have something glutinous at this stage ; and
white cells melt at a much lower tempera-
ture of water than what liquifies the co-
loured cells. Old combs, therefore, contain
a substance foreign to wax.

In examining the orifices of the yellow
cells, we perceived that their circumference
was surrounded by a reddish, unctuous, odo-
riferous varnish, indicating, as we thought,
the resin propolis. Afterwards it appear-
ed not to be restricted to the orifices, but
that reddish threads of it were applied
around all the interior of the sides, like a
kind of soldering, at the points of contact of
the different pieces, to strengthen them.

Bees are obliged to interrupt their labour
when they can procure no more wax, during
which time, jirobably, they varnish the edges
of their cells. On resuming their work, the
cells arc deepened, and traces of the propolis
remain pointing out where it has been ap-
plied.

In so far as we were aware, these peculia-
rities had not struck any of the authors on
bees : therefore, it was important to ascer-

403

CaMPLETIOK OF THE CELLS.

tain that propolis enters the composition of
cells.

The substance taken from the sides of a
hive, and the fragments of cells bordered
with it, imparted a golden yellow colour to
ether, spirit of ^'dne, and spirit of turpen-
tine. The brown matter ot the c-ells was
dissolved there, even when they were cold.
In the two latter fluids, the orifices still re-
tained the cellular form, and their yellow
tinge after losing the varnish. This was
also lost in ether ; and the cells quickly
whitening, disappeared with solution of the
wax.

The propohs of the orifices of cells and of
the sides of a hive, being softened by gentle
heat, could be drawn out in a thread. Ni-
trous acid, at a low heat, poured on both,
whitened the yellow w^ax in a few minutes ;
but the varnish of the orifices and masses of
propolis underwent no alteration.

AVhen some orifices were put into boiling
water, and the wax melting, fonned a cake,
the varnish remained entire above it, the
hexagonal figure of the cells being preserv'-
ed, while their diameter seemed a little en-
larged.

404 COMPLETION OF THE CELLS.

Fixed caustic alkali, which forms a kind
of soap with wax, had no effect on the var-
nish, though cells were immersed in it se-
veral months.

The substance colouring the edges of
cells and the lines on their sides, therefore,
have the greatest analogy to propolis ; and
the yellow of the cells has no relation to
the varnish at the places where the sides
unite.

Notwithstanding my confidence in such
conclusions, I felt that they w'ould become
indisputable, only by surprising the bees in
collecting propolis. For many years I had
fruitlessly endeavoured to find them on trees
producing an analogous substance, though
having seen multitudes returning laden
with it.

In July, some branches of the wild pop-
lar, which had been cut since spring, with
very large buds full of a reddish, viscous,
odoriferous matter, were brought to me, and I
planted them in vessels before hives in the way
of the bees going out to forage, so that they
could not be insensible of their presence.
Within a quarter of a hour they were visit-
ed by a bee, w'hich separating the involucra

COMPLETION OF THE CELLS. 405

of a bud with its teeth, drew out threads of
the viscous substance, and lodged a pellet
of it in one of the baskets of its limbs. From
another bud it collected another pellet lor
the opposite limb, and departed to the hive.
A second bee replaced it in a few minutes,
following the same procedure.

Young shoots of poplar, recently cut,
did not seem to attract these insects ; but
their viscous matter had less consistence
than the fomier.

Different experiments proved the iden-
tity of this substance with propolis ; and now
having only to discover how the bees applied
it to use, we peopled a hive so prepared as
to fulfil our views.

The bees, building upwards, soon reached
the glass above ; but, unable to quit their
habitation on account of supervening rains,
they were three weeks without bringing
home propolis. Their combs remained per-
fectly white until the beginning of July,
when the state of the atmosphere became
more favourable for our observations. Se-
rene warm weather engaged them to forage ;
and they returned from the fields, laden with
a resinous gum, resembling a transparent
jelly, and having the colour and lustre of

406 COMPLETION OF THE CELLS.

the gamet. It was easily distinguished
fe'om the farinaceous pellets then collected
by other bees. The workers hearing the
propolis ran over the clusters suspended from
the roof of the hive, and rested on the rods
supporting the combs, or sometimes stopped
on the sides of their dwelling, in expectation
of their companions coming to disencumber
them of their burden. We actually saw two
or three arrive, and carry the propolis from
olf the limbs of each with their teeth. The
upper part of the hive exhibited the most
animated spectacle; thither a multitude of
bees resorted from all quarters, to engage in
the predominant occupation of the collec-
tion, distribution, and application of the
propolis. Some conveyed that of which they
had unloaded the purveyors in tlreir teeth
and deposited it in heaps ; others hastened,
before hardening, to spread it out like a var-
nish, or formed it into strings, proportioned
to the interstices of the sides of the hive to
be filled up. Nothing could be more diver-
sified than the operations carried on.

The bees, apparently charged with apply-
ing the propolis within the cells, were easily
distinguished from the multitude of work-
ers, by the direction of tlicir heads towards

COMPLETION OF THE CELLS, 407

-the horizontal pane forming the roof of the
liive, and, on reaching it, they deposited
their burden nearly in the middle of inter-
vals separating the combs. Then they con-
veyed the propolis to the real place of its
destination. They suspended themselves
by the claws of the hind legs to points of
support, afforded by the viscosity of the pro-
polis on the glass ; and as if swinging them-
selves backwards and forwards, brought the
heap of this substance nearer to the cells at
each impulse. Here the bees employed
their fore feet, remaining free, to sweep up
what the teeth had detached, and to unite
the fragments scattered over the glass,
w-hich recovered all its transparency, when
the whole propolis was brought to the vi-
cinity of the cells.

After some of the bees had smoothed
down and cleaned out the glazed cells, feel-
ing the way with their antennae, one desist-
ed, and having approached a heap of pro-
l>olis, drew out a thread with its teeth.
Tlris being broke off, it was taken in the
claws of the fore feet, and the bee re-enter-
ing the cell, immediately placed it in the
angle of two ])ortions that had been smooth-
ed, in which operation the fore-feet and

408 COMPLETION OF THE CELLS.

teeth were used alternately. But probably
proving too clumsy, the thread was reduced
and polished, and we admired the accuracy
with which it was adjusted when the work
was completed. The insect did not stop
here. Returning to the cell, it prepared
other parts of it to receive a second thread :
for which we did not doubt that the heap
would be resorted to. Contrary to our expec-
tation, however, it availed itself of the por-
tion of the thread cut off on the former oc-
casion, arranged it in the appointed place,
and gave it all the solidity and finish of
which it was susceptible. Other bees con-
cluded the work which the first had begun,
and the sides of the cells were speedily se-
cured with threads of propolis, while some
were also put on the orifices. But we could
not seize the moment when they were var-
nished, though it may be easily conceived
how it is done.

Experiments demonstrate that the mat-
ter imparting the yellow colouring to wax
has no analogy with propolis. This is not
the natural hue of wax, for new cells are
white. But the whiteness gives place to a
yellow tinge, which afterwards deepens.
Sometimes the new combs yellow in two or

COMPLETION OF THE CELLS. 409

tlirG6 davs. As otlicr naturalists, I was in-
clined to ascribe the alteration to the heat
of the hives ; to the vapours disseminated in
them ; to the emanations of the honey ; or
of it and the pollen remaining in the cells:
Yet these opinions could not support rigor-
ous examination. T*^ew combs have been
observed to undergo no alteration in several
months. Sometimes one face has been of
a jonquil colour, and the other white : and
on the same surface we have observed a por-
tion consisting of cells of a very lively yel-
low, while those adjacent had lost none of
their whiteness. Likewise a single cell
has shown several of the sides yellow, others
white on one side, or both white and yellow.
Honey and pollen would have tinged all
the sides uniformly, as far as they reached ;
and the vapours disseminated throughout the
hive would have had a general influence.

In order to ascertain the effect of preserv-
ing the combs from the contact of bees, I
took a hive divided in the middle, and kept
a piece of perfectly white comb in it during
a month, liable to heat, moisture, and all at-
mospheric vapours, without the colour being
affected. Meanwhile, the combs exposed
to the contact of the bees yellowed more

T

410 COMPLETION OF THE CELLS.

and more, but this colouring was partial
and distributed irregularly, as if in stripes.
Every thing argued that it resulted from
some direct action on the part of the bees,
not from exposure in the interior of liives.

Without actual discovery of the fact, we
have been induced to ascribe it to two dif-
ferent sources ; first, to the bees rubbing
different parts of the body, their teeth or feet
on the surfaces where they seem to rest :
secondly, to the trunk sweeping from right
to left, like a fine, pliant pencil, when it
appears to leave some sprinkling of a trans-
parent liquid. We shall not affirm which
of these operations is the source of the
yellowness of wax ; but we incline to refer
it to the first, because after the bees had
rubbed certain cells with their teeth and
fore legs, we sometimes thought the colour
altered.

These insects do not restrict themselves
to painting and varnishing their cells : they
are also occupied in giving greater solidity
to the edifice itself, by means of a mortar
which they know to compose for that pur-
pose.

The ancients were acquainted with some
of the properties of propolis ; and aware that

COMPLETION OF THE CELLS. 411

the bees, under certain circumstances, mixed
it with wax, they gave the name pisso-
ceros to the compound ; and how well they
had studied the subject was proved by one
of my experiments.

I immersed some fragments of the com-
pound taken from the sides of an old hive
in ether, having already found that this fluid
held a very small portion of wax in solution.
Decanting it several times, I concluded that
the whole propolis was dissolved, when the
fluid ceased to colour. Accordingly, the
residue was found to consist of a little white
wax which the bees had mixed with it.

Pliny believed that these insects used a
mixture of wax and propolis in constructing
the fixture or basis of the comb : Reaumur,
on the contrary, thought it only pure
wax. Perhaps the opinion of such emi-
nent naturalists may be reconciled by the
following facts.

Soon after some new combs had been
finished in a hive, manifest disorder and agi-
tation prevailed among the bees. They seem-
ed to attack their own works. The primi-
tive cells, whose structure we had admired,
scarcely were recognizable. Thick and raassv
walls, heavy shapeless pillars were substi-

412 COMPLETION OF THE CELLS.

tuted for the slight partitions, previously
built with such regularity. The substance
had changed along with the form, being
composed apparently of wax and propolis.
From the perseverance of the workers in
their devastations, we suspected that they
proposed some useful alteration of their edi-
fices ; and our attention was directed to the
cells least injured. Several were yet im-
touched ; but the bees soon rushed precipi-
tately on them, destroyed the tubes, broke
down the wax, and threw off the fragments.
But we remarked, that the bottom of the
cells of the first row was spared; neither
were the corresponding parts on both faces
of the comb demolished at the same time.
The bees laboured at them alternately,
leaving some of the natm-al supports, other-
wise the comb would have fallen do^\^l, which
was not their object : they wished, on the
contrary, to provide it a more solid base,
and to secure its union to the vault of the
hive, with a substance whose adhesive pro-
perties infinitely surpassed those of ■wax.

The propolis employed on this occasion had
been deposited in a mass over a cleft of the
hive, and had hardened in drying, which pro-
bably rendered it more suitable for the pur-

COMPLETION OF THE CELT.S. 413

pose. But the bees experienced some diffi-
culty in making any impression on it ; and
we thought, as also had appeared to M. de
Reaumur, that they softened it with the same
frothy matter from the tongue which they
use to render w^ax more ductile. .

We very distinctly observed the bees mix-
ing fragments of old wax with the propolis,
kneading the two substances together to in-
corporate them; and the compound was em-
ployed in rebuilding the cells that had been
destroyed. But they did not now follow
their ordinary rules of architecture, for they
were occupied by the solidity of their edi-
fices alone. Xight intervening, suspended
our observations, but next morning confirm-
ed what we had seen.

"We find, therefore, that there is an epoch
in the labour of bees, when the upper foun-
dation of their combs is constructed simply
of wax, as Reaumur credited, and that after
all the requisite conditions have been attain-
ed, it is converted to a mixture of wax and
propolis, as remarked by Pliny so many ages
before us. Thus is the apparent contradic-
tion between these two great natm-alists ex-
plained. The first row of cells had been
temporarily established as a base, and to servo

414 COMPLETION OF THE CELLS.

as a direction for the pyramidal bottoms of
those to be subsequently constructed. So
long as the magazines were not quite full,
this was sufficient to support the edifice ; but
plates of wax so thin probably would have
been inadequate to sustain the weight of se-
veral pounds. The bees seemed to have an-
ticipated the eventual inconvenience : they
destroyed the frail sides of the first row of
cells, to form strong pillars and waxen walls
of a viscous and compact substance. Per-
haps the alteration depends on the concur-
rence of special circumstances, nor does it en-
sue at a marked and regular period. Some-
times the bees only border the sides of the
upper cells with propolis, without changing
their shape, or strengthening them.

But this is not the utmost extent of the-
foresight of these insects. When they have
plenty of wax, they make their combs the
full breadth of the hive, and solder them
to the glass or wooden sides, by structures,
more or less approaching the form of cells,
as circumstances admit. But should the
supply of wax fail before they have been
able to give sufficient diameter to the combs,
whose edges are rounded, large intervals re-
main between them and the upright sides

COMPLETION OF THE CELLS. 415

of the hive, and they are fixed only at the
top. Therefore, did not the bees provide
against it, by constructing great pieces of
^vax, mixed with propolis, in the inter-
vals, they might be borne down by the
weight of the honey. These pieces are of
irregular shape, strangely hollowed out, and
their cavities void of symmetry.

During winter, a comb in one of my bell
glass hives, having been originally insecure,
fell down, but preserving its position pa-
rallel to the rest. The bees were unable to
fill up the vacuity left above it, because
they do not build combs of old wax, and
none new could be then obtained. At a
more favourable season they would have en-
grafted a new comb on the old one; but
now their provision of honey could not be
spared for the elaboration of this substance,
which induced them to ensure the stability
of the comb by another process.

Crowds of bees taking w'ax from the lower
part of other combs, and even gnawing it from
the surface of the orifices of the deepest cells,
they constructed so many irregular pillars,
joists, or buttresses, between the sides of the
fallen comb, and the others on the glass of
the hive. All these were artificially adapted

416 COMPLETION OF THE CELLS.

to localities. Neither did they confine them-
selves to repairing the accidents which their
works had sustained. They seemed to pro-
fit by the warning, to guard against a simi-
lar casualty.

The remaining combs were not displaced ;
therefore, while solidly adhering by the
base, we were greatly surprised to see the
bees strengthen their principal fixtures mth
old wax. They rendered them much thick-
er than before, and fabricated a number of
new connections, to unite them more firmly
to each other, and to the sides of their
dwelling. All this passed in the middle of
January, a time that these insects common-
ly keep in the upper part of their hive, and •
when work is no longer seasonable.

I may restrain myself from reflections
and commentaries : but I acknowledge that
at such measures I cannot suppress, senti-
ments of admiration.

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APPENDIX

I, Generation of Bees. — The author has entered on
prolix anatomical details, in explanation of the theory
of the generation of bees, which many will think un-
interesting, and an interruption of the narrative. There-
fore, the subject, and several relative experiments, have
been abbreviated by the translator.

Swammerdam was unable to discover the external
termination of the oviduct of the queen, which the
author has ascertained to be in the last ring of the
abdomen ; also that the eggs are di-opped by it into a
kind of cavity, fi-om which they are quickly discharged.

He has discovered certain parts among the sexual
organs of the male, that are lost in commerce with
the queens; particularly fig. 7, Plate II. which he con-
siders as corresponding to the principal organ of other
animals.

^ He obtained a number of females after Schirach's
j method ; and, on the tenth of July, successively re-

leased three, which were four or five days old, and still
’ in a virgin state. 1 wo departed several times ; their
absence was short and fruitless : the third profited bet-
ter by her freedom. The first and second time lier
absence was short ; but the last continued thirty-five

418

APPENDIX.

minutes, when she returned in a very different condi-
tion, and such as admitted no doubt of what had hap-
pened, for she exhibited the organs of the male that
had rendered her a mother. They wei'e disengaged by
herself : she was allowed to enter her hive, and its en-
trance adapted so that she could not leave it unnoticed.
No eggs were found in it on the seventeenth ; the
queen was as slender as on the first day ; therefore she
had not been fructified. Being again set at hberty, she
departed twice, and returned with evidence of a second
union. The eggs which she laid after this event showed
that it had been more successful; but a repetition' is
rarely requisite. It was remarked on only two occa-
sions.

On the eighteenth, a virgin queen twenty-seven days
old was released ; she departed twice. Her second ab-
sence lasted twenty-eight minutes, when she returned
with the proofs of her union. On the igth another
queen, four days old, was set at liberty ; she also de-
parted twice. Her first absence was short : the second
continued thirty minutes, when she returned with the
evidence of fecundation.

In a subsequent experiment, two virgin queens were
liberated on the twentieth. The first had been abroad
on the preceding days, but the scarcity of males had
prevented her impregnation. She now returned with
the wonted organs, of which she divested herself with
her feet. The second queen departed twice. Her first
absence was short as usual ; the next lasted half an
hour, when she returned impregnated.

It appeared to the author that additional evidence of
the preceding facts might be obtained, b}" seizing the
male on its return which had rendered a queen a mother;
and this could be expected only if sudden death did not
follow. His assistant believing that he could easily

appendix.

419

distinguish those dying immutilated by their com-
merce, engaged in examining the whole found dead at
the hives during the season of swarming. After a
painful research, some were actually discovered dead,
and clearly in a mutilated state. None of their usual
organs remained; or only some decayed portions of
them. It was thence established that the males perish,
but that mutilation is not followed by immediate deatli,
as they have time to return to the site of the hive.

II. Secretion of JFax. — The author had committed
the dissection of bees to Miss Jurine, a young lady dis-
tinguished by her talents, and her taste for natural
history, for the purpose of investigating the organs
destined for secretion of wax.

On carefully removing the four wax segments of a
worker, this skilful anatomist found an adipose mem-
brane, interspersed with tracheae, and perfectly re-
sembhng that which had been recognised by Swam-
merdam, below the upper segments of the abdomen.
As it exists beneath all the segments, while wax ap-
pears only in the lower ones, it is not to be presumed
the secretory organ; and from examining the abdo-
men of the violet bee Qsylocopa violacea,] and of two
species of wasps, she was satisfied of the fact.

A whitish membrane, covering the internal surfiice
of that portion of the wax segments provided -with
the receptacles, was afterwai’ds discovered. It could
be easily removed by maceration ; and, presented to
the microscope, appeared to consist of a beautiful net,
of very small hexagonal meshes, full of a liquid as
thick as syrup. It was also found under the same
segments of the abdomen of humble bees, but occu-

420

APPENDIX.

pying all their inferior half. This membrane is most
perceptible when bees are building their combs : at
other times it is indistinct, but then it is so full of the
whitish matter, that it might be taken for scales of
wax.

Detaching the reticulation from the ring, to ascer-
tain whether it contained real wax, or only prepara-
tions for it, and taking the scales of wax themselves,
M. Jurine put each into a separate vessel, and poured
boiling water over them. The latter melted, but no
waxy particle escaped from the former.

She likewise endeavoured to discover the vessels
communicating with the reticulation, and bow the
wax transuded from the abdomen. However, on dis-
secting a number of bees, nothing except minute
tracheae, immediately communicating with the reticu-
lation, were discernible. Following another plan, she
fed bees for some days on honey, coloured with lac ;
yet this substance did not reach farther than the di-
gestive organs: and injections of mercury were at-
tempted as unsuccessfully.

M. Jurine conjectured that wax might be formed by
a transudation through the stomach, which is very
much gorged while bees work in that substance. She
found, by gently pressing the stomach of several wax
workers, half the liquid it contained escaped into the
abdominal cavity. Its taste was saccharine; and
the same bees having been afterwards exposed to mo-
derate heat, it assumed the consistence of dried syrup.
Fees having several methods of obtaining such a pres-
sure, she asks, may it not be attended with similar ef-
fects : and may not the liquid transuding the stomach,
reach the hexagonal reticulation, where it receives a
preparation suitable for its conversion to wax }

APPENDIX.

421

Unable to discern any channels of communication,
M. Jiirine stretched out the abdomen of several work-
ers having wax scales, to remove them with more fa-
cility. Then pressing the wax segments with the
head of a needle, she saw their receptacles insensibly
moistened with a liquid, of the consistence of syrup,
observed nowhere besides. It acquired still greater
consistence, but no waxy appearance, when the bees
in this condition were exposed to heat.

I'he same experiment having been repeated on bees
that had been dead several days, and were somewhat
dried, the wax scales broke into fragments on attempts
to remove them. By the simple expedient of pressing the
wax segments several times, the scales could be obtained
entire : nor was this to. be ascribed to any thing but
the oozing of the matter like syrup into their recepta-
cles, as had been remarked in the preceding experi-
ment.

The reticulated membrane, which occupies none ex-
cepting the wax receptacles in workers, is replaced in
queens by one extending over two-thirds of the ante-
rior part of each segment. Its fine and delicate tex-
ture is ascertained by means of the microscope. Re-
moving this membrane, M. Jurine remarked, that the
organization of the scale exhibited a liexagonal tissue,
much more decided in the half of the segment corres-
ponding to the wax receptacles of the worker, than to
tlie posterior half. Being thence led to narrower in-
spection of the shell ofj the wax segment of workers,
slie found it perfectly smooth in the part where the
receptacles are situated, and resembling that of the
queen throughout the rest of the segment.

Males are altogether destitute of the adipose mem-
brane and the hexagonal reticulation ; and muscular
fibres with tracheae, disposed as in the worker appears

422

APPENDIX.

in their place. The organization of the shell of the
rings displays the same hexagonal tissue as that of the
queens.

III. Origin of wax, — The author gives an extract
from a Memoir by the late Dr. John Hunter, wherein
preceding opinions on the formation of wax are ques«
tioned.

He observes, that the materials forming the comb are
in a different state as a composition, from what they
appear in any vegetable where they are found. The
pellets brought home on the legs of bees, which had
been previously supposed the source of wax, are only
the farina of flowers. Endeavouring to ascertain
whether they contained a quantity of oil sufficient to
account for the quantity of wax formed from them, and
to learn whether they contained oil, he held one near a
candle : it burnt, but without the smell of wax, and
emitted the same smell as farina did i!i burning.

Dr. Hunter remarked that the pellets are of different
colours on different bees, but ahvays of the same colour
on both legs of the same bee ; whereas the shade of
new made comb is uniform. The farina is collected
with greater avidity for old hives, where the comb is
complete, than for those where it is only begun, which
would hardly be the case were it the elements of wa.x.
Two days usually elapse before the bees of a new co-
lony bring home pellets on their legs, and then the
number of gatherers increase, because some cells arc
now formed to receive it, and some eggs are laid
whose young will soon require it for food.

When the weather was so cold or so wet in June as
to prevent a young swarm from going abroad, as much

APPENDIX,

423

comb was constructed, nevertheless, as had been made
in an equal time when the weather was favourable.

Wax, in Dr. Hunter’s opinion, is formed by the bees
themselves : it may be called an external secretion of
oil, and originates between each plate of the under side
of the belly. On first observing it, in examining a
worker, he was embarrassed to explain its appearance,
and doubted whether new plates were forming, or
wliether bees cast the old ones like the shell of lobsters
and other animals. He found it nowhere but between
the rings below. Most of the bees climbing the panes
of a glass hive exhibited this substance ; and it gave
the lower or posterior edge of the segment the resem-
blance of being double, or there seemed to be double
scales ; but it was loose, and did not adhere.

Having proved that the pellets of the limbs were
fai'ina, apparently destined for the larvae and not for
wax, and having seen nothing else which could be
I supposed the source of it, he conceived that the scales
I might be wax itself. Several taken on the point of a
I needle, and held to a candle, melted into a globule,
which confirmed his opinion of the fact ; and he was
tlie more satisfied of the truth, fi’om not finding the
scales, unless during the season that bees construct
their combs. However, he was unable to seize the
moment of the bees detaching the scales from them-
selves, though he conjectured they might be taken up
by others, if shaken out from between the rings. It is
with these scales that the bees construct their comb,
but perhaps when the secretion is not in abundance,
they mix a little farina with it.

Dr. Hunter, besides, makes several interesting obser-
vations on the structure of combs. Fhi/osophical
Transactions, for 1792, p. 14'3.

424

APPENDIX.

IV. Explanation of the Plates. — Plate I. fig. 1. The
leaf or book hive, consisting of twelve vertical frames,
such as fig. 3, applied parallel to each other, and con-
nected together. The sides f g,f g, fig. 3, are twelve
inches long: the cross spars g g,ff, nine or ten; the
thickness of both the sides and the spars an inch, and the
breadth of the sides of the frame an inch and a quarter,
which last dimensions should be exactly observed. A
piece of comb, a a, is introduced to guide the bees in
their work, supported on a moveable slider c, which may
be elevated or depressed by the pegs e e. The comb is
secured by the pegs b b ; and in fig. 4, the position of
those corresponding on the opposite sides, as well as
the whole apparatus, is illustrated.

The twelve frames, all numbered fig. 1, are covered
by two shutters, b b, at the ends. Between 6 and
7, two frames, a a, each Avith a shutter, are inserted to
separate the bees in forming an artificial SAvarra. The
entrance at the bottom of each frame may be opened at
pleasure; but the whole, except the first and last, should
be kept close. It is explained by fig 2, showing the
hive partly open, how the component frames may be
connected by means of hinges, and opened like the
leaves of a book.

Fig. 5, a glass box, a, for exposing the work of bees
while enlarging their combs upwards. Fig. 5, b, a
box with moveable Avooden spars, Avhich can be raised
and depressed by a screAv at each end, s s, for expos-
ing the progress of the cells.

Plate If. All the figures are magnified. Fig. 6. the
sexual organs of the drone, as particularly described
by Swammerdam and Reaumur. Last segment of the
abdomen a; a membranaceous sac m; a lens-formed
portion i /, communicating by a duct e r, with the tAVO

appendix.

425

vesiculae seminales s s, which are connected with the
vasa deferentia d d, and the latter with the testes 1 1.
A pallet shaped substance : two appendages c c. Fig.
7, a separate view of the portion resembling a lens i I,
after rupture of the duct. It is composed of two shelly
plates, i e, i e, and two cartilaginous n n, together
w ith what is supposed the principal organ r.

Fig. 8. Unexpanded'sexual organs of the workers:
ovaries a a; portions of the spinal maiTow h b; the
sting, with its appendages c, and poison bag d.

Fig. 9. A worker magnified, showing the position ot
the scales of wax under the segments of the abdomen.

Fig. 10, 11, 12. Comparative configuration of the
abdomen of workers, males, and queens.

Plate III. All the figures are magnified. Fig. 13,

: Segment of the abdomen of a worker, showing the re-
! ceptacles wherein the scales of wax are produced.

Fig. 14. Scales extracted from tlie receptacles.

Fig. 15. A reticulation lying under the scales in
the segment, fig, l6.

Fig. 17, 18, 19. Views of the teeth or mandibles
of bees.

Fig. 20, 21, 22. A limb of the third pair, and its
different parts. The leg c, where the pollen is col-
lected in a corbiculus or basket. The shelly teeth,
a, fig. 22, at the junction of the leg with the foot, em.?

I ployed as pincers by bees to extract the scales from
' under their rings.

Plate IV. fig. 23, 24. A bee extracting a scale of
I wax, and preparing it for work.

Fig. 25. A cell of the first or foundation row, having
a pentagonal orifice.

Fig. 26. Front view of the first and second row
t of cells, a. Reverse of the same portion,

426

APPENDIX.

Fig. 27. Foundation and second row of cells mag-
nified to illustrate the diffei’ent structure of the bottom.

Fig. 28. The bottom of a front cell of the first row,
consisting of two trapeziums, a b, and a lozenge c :
also shown in fig. 27, a b c.

Fig. 29. The bottom of a reverse cell of the first
I’ow, consisting of two trapeziums, a b.

Fig. 30, 31. Front and reverse of the bottom of the
first and second row of cells. Fig. 30, fa, bg, the al-
ternate quadrilateral pieces of the former ; gb, a f,
those of the latter : c d e, the lozenges forming the bot-
tom of the second row.

Fig. 32. Commencement of the first cell.

Fig. 33. Commencement of the first cell on the op-
posite side of the comb.

Fig. 34. Foundation cell on the front.

Fig. 35. Foundation cells on the reverse, and one
of the second row, showing that the former are penta-
gonal, and the latter hexagonal.

Fig. 36, 37. Blocks of wax in which the cells are
excavated.

Plate V. All the figures excepting 45 are magnified.

Fig. 38, 39, 40. Progress of a foundation cell of the
first row. The real position of these figures and of some
of the others is inverted, though represented upright
in the engraving, as bees work downwards.

Fig. 40, g d a. The base of the cell or roof of the
hive.

Fig. 41. Two foundation cells behind the former,
also attached to h d fc, the roof.

Fig. 42, 43. Front and reverse of the first and
second row of cells in progress ; according as additions
are made to the block of wax.

iVrPENDIX.

427

Fiff. 44. Formation of a cell of the second row,
showing the curvature of the margin p r m, to be con-
verted to the rectilinear margins p r m of fig. 42.

Fig. 45. Comb begun under a spar, and continued
upwards above it.

Fig. 46, 47, 48, 49, 50. The bottom of transition
cells, consisting of four pieces, and their return to three
lozenges.

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ANALYTICAL INDEX.

The form of Reaumur’s hive unfavourable
Hive invented by the author,

Very thin and flat hives indispensible, -
Swammerdam on the propagation of bees.
Sentiments of M. de Reaumur,

Mr. Debraw’s opinion, - - -

Refuted by the author, - - . .

Hattorf’s opinion, . . _ i

Refuted, - . «

Difficulty of the subject, -

Experiments to discover the fact.

Propagation is attended with mutilation of the
drones, - - * _

Disproportionate number of males.

Suggestions by M. Bonnet, - , .

Commerce of bees takes place in the air.
Probably the males perish.

Page.

2

3

5

6

7

8

10

14

16

17

18

21

24

26

33

35

Retarded impregnation affects the ovaries of the
queen, - * , - 36

She then lays only eggs producing males, sg

Queens leave the hive only once for propagation^

Experiments retarding fecundation, i 40

f The first eggs in the natural state are those of
workers, . . _ _

430

INDEX.

Page.

ii

44

47

51

52

56

57

5S

62

Next are those of males.

Eggs to be laid for at least two years are fertilized
at once, - . _ .

Retarded impregnation of other animals occasions
sterility, - - - -

Piefrigeration of the atmosphere suspends laying,

Great fecundity of a queen, - - -

Bees do not transport the eggs to different cells.

They sometimes eat them.

The instinct of queens affected by retarded fecun-
dation, - _ - .

Eggs producing males sometimes laid in i*oyal cells, 59
\\ hole bi'ood consisting of drones, - - 60

Drones are spared longer in hives where the queen
lays only eggs of males, - -

The queen being lost, a common worm may be
converted to a new one.

Operations of the bees to accomplish it,

A worm 48 hours old is susceptible of the change.

The bees in a few hours endeavour to replace the
lost queen, -

Reaumur believes that queens are produced natur-
ally, . - - -

Schirach thinks that there are no eggs to produce
them naturally, _ _ _

Queens lay eggs in royal cells which will become
queens, -

Fertile workers sometimes exist.

Experiments to discover them.

They have ovaries, - -

But lay only the eggs of males.

The royal cells are destroyed where workers lay
male eggs, - - -

All common bees are originally females.

64

65
67

69

70

71

74

76

78

SO

SI

82

J

INDEX.

431

Page,

Experiments to prove the source of evolution of
their ovaries, - - - 8-t

It is from receiving the royal food while larva?, 87
Queens have an instinctive jealousy of them still
in the cell> - - - S9

Combats of queens, - - . 90

Almost all M. de Reaumur’s conjectures are right, —
Obervations by Reaumur, Schirach, and Riem, 9I
Enmity of queens against others while nymphs, Q2
Mutual enmity of queens, - - 94

There ought to be only one in a hive, - —

All might perish if their destruction were commit-
ted to workers, - _ . 95

The common bees seem to promote the combat^ 96

Queens tear open other royal cells, - 97

A stranger and a native queen confined by the
common bees, unless when inclined to fight, 99

Combat between a fertile and a virgin queen, 100

Of several queens, one alone survives, » 101

i Bees keep constant guard at the entrance of the

hive, - - i. JQ2

i A stranger queen entering is instantly in restraint,

' She is scarcely ever stung, - . 203

' She perishes from hunger or suffocation, -

f Riem believed the workers killed supernumerary
I queens, - - . 204

1 Disorderof bees when missing their queen,

It does not abate on substitution of a stran^^er
queen, - - . - 105

|i If twenty-four hours elapse the stranger is well

received, - . _ 2 of)

1 The bees supply her with honey, - 107

I They desist from constructing cells to replace tlie
I lost queen, - . , 2 08

INDEX.

432

<

Page.

Their former queen is forgot in twenty-four or
thirty hours, - . . 109

Massacre of the drones, - - 1 1 0

Those of six hives destroyed at the same hour, 1 1 1

They are stung to death, - - 112

Males are never destroyed in hives wanting
queens, - - - 113

Reception of a stranger queen, - 114

Different results from those of Reaumur, ^ 115

A stranger is ill received if substituted early, —

Reaumur’s observations, - * 1 1 6

A plurality of queens never is tolerated in a hive, 1 1 8
Experiments to ascertain whether the queen be
oviparous) >■ - * 122

The workers seem occasionally to repose, - 124

Eggs require no care, > - - 125

Drones do not enter the cells to repose, - 126

Interval between production of the egg and per-

fect state, - - - —

Larvas move in the cellS) - - 127

Mode of spinning the coccoon, - - 12S

That of the queen is left open, - - 129

It is owing to her cell, - - 132

Experiments to ascertain whether the size of bees
is affected by the size of their cells, - 133

The queen selects cells suitable for her eggs, 134

Eggs of different kinds are not indiscrimmately
mixed in her ovaries, - - 136

None of the different bees attain larger size from
greater capacity of the cells, - 140

Their growth may be cramped by small cells, —
Formation of swarms, - - - 143

Reaumur supposed that the first swarm was head-
ed by a young queen, - - •“

INDEX.

433

Page.

Experiments proving that it is always the old queen, 145
The old queen conducts the swanu of a subsequent

year, - - - -

She never quits the hive before laying in the royal
cells, ...

Weak hives do not swarm.

History of royal cells left by the old queen.
Agitation of the queen befoi’e swarming is com-
municated to the bees.

Bees reduce the wax of the royal cells.

They retain the queens there for some time.
Singular effect of a sound emitted by perfect queens.
Bees feed the young queens retained in the cells.
Conduct of queens successively hatched.

The bees guard the royal cells from their attacks.
Combats between young queens.

The bees testify indifl'erence to a virgin queen.
How a hive can throw several successive swarms.
Royal cells built when the queen lays the eggs of
males, - - - _

Closed when the larvae are about to transform to
nymphs, - - -

The queen should lay before swarming to lighten
her for flight, - - - .

She quits the hive when the royal larvae are ready
to transform to nymphs.

Her first eggs in her new hive are those of

148

149

150

152

155

156

157

158

159
161

163

164

165

169

170

171

workers, - - - ,

The young queens in the old hive are prevented
from all leaving tlieir cells at once.

Conduct of the first queen quitting her cell.

Bees are alarmed at the prospect of bad weather,
and swarm only in a fine day.

Young queens are liberated from their cells ac-
cording to their age,

U

173

175

176
179

434

INDEX.

Page.

Probably the bees detennine this from the sound
emitted, - - - - 180

Queens reared from the larvae of workers are
mute, - - - - 181

Tliose bred during the time of swarming are
otherwise, - - 183

The young queen conducting a swarm is in a
virgin state, - - - 185

The conduct of bees to the old queen is peculiar, 187
Royal cells with eggs or very young larv$ never
are attacked by the queen, - - 188

Conjectures on the cause of swarming, - iqo
The formation of swarms is attended with singular
facts, - - - - 195

Bees are aware of the species of eggs deposited in
the different cells, - - - 19S

The instinct of queens is affected by retarded im-
pregnation, - - - 201

Workers regard a dead queen, - - 202

Singular effects of amputating the queen’s an-
tennae, . _ - « 203

It obliterates their mutual enmity, - - 206

Use of the antennae, - 207

Mutilated females leave the hive alone, - —

It is absurd to destroy hives to reap their pro-
ducts, - - - - 211

Advantages of the book or leaf hive, - - 212

It renders the bees tractable, - - - 213

4'hey are affected by the sudden introduction of
light,

Process of forming artificial swarms.

It is the invention of Schirach,

Bees can be forced to work in wax.

Uniform distance between the combs.

Bees always fill up intervals, <

21 1
215
2IS

219

220
221

INDEX.

435

Page.

Only a moderate portion of their stores is to be
removed, - - - ■

Natural heat of bees, . - -

l^istance of their excursions.

Opposite views of Schirach’s discovery.

The autlior’s hypothesis, - •

Metamorphosis is repugnant to reason,

I'he faculties and organs of the common bee may
be unfolded by peculiar treatment.

Mutual enmity of female humble bees awakened
by fecundation, - - -

History of black bees in hives,

I’heir enmities, - - -

They are seen in the same colony in successive

223

224

225

230

231

232

233

234
2.35

years, - - - - 237

All common bees are females, - - 240

Fertile workers lay only the eggs of males, 241

The w'orkers among humble bees and w'asps are
also restricted to them, - - 243

The workers of ants perish after union, - 245

Schirach unjustly accused of plagiary, - 24()

Formation of artificial swarms, - - 247

I^ombard’s method, - - - 251

Senses of bees, - - - - 253

Their sight is very acute, - - 255

Their sense of taste the least pei-fect, - 257

1 The odour of honey their most powerful attraction, 258
I The instinct of the humble bee is more refined, 259
I Singular intercourse between humble and domestic

bees, - 260

I Sense of smelling resides in or near the mouth, 26.3
I Influence of different odours on bees, - 26.5

I The odour of their own poison excites their anger, 268
‘ How they repel disagreeable sensations, - 270

J I'se of the antennae, ... o'-o

436

INDEX.

T ■ . , . Page.

Instinct in replacing a lost queen, - - 273

She is not missed immediately, - - 274

In twenty-four hours bees have prepared to re-
place her, - - - - 275

Experiments to discover how her presence is as-
certained, - - _ oyy

It is by the antennae exclusively, - - 280

When these cannot touch her, bees proceed as if
she were lost, - - - 281

They guide bees in all their operations, - 283

Their use during the night to exclude strange in-
sects, - - - -

Probably bees have hearing, - - 284

They are not affected by sounds bearing no rela-
tion to their instincts, - - - 285

Experiments on bees in rarefied and vitiated air, 288
Males suffer most, - - - 289

All perish from inhaling carbonic acid gaz, 291

The stigmata carry on respiration, - 292

Two new ones discovered, - - —

Bees preserve the air of their hives pure, - 293

They are suffocated by shutting up the entrance, 294
They have no substitute for external air, - 293

Bees ventilate their hive, - - —

Ventilation establishes a current of air, - 297

Experiments by Burnens, - - 298

An artificial ventilator preserves a light burning, 299
Bees ventilate or fan themselves to obtain coolness, 301
Heat and noxious odours induce them to do so, 302
It is not practised by the males of domestic or

humble bees, ~ - - 303

Bees in succession are occupied in ventilating, 304
A new enemy of bees, . - . 305

The purpose of warfare is to preserve an equili-
brium, - - 305

INDEX.

437

Page.

Devastation among hives.

The sphinx atropos destructive to bees,

Hives artificially protected by a grating.

Bees themselves barricade their entrance with wax.
It is only when danger is imminent.

Weak hives construct a defence against invaders.
The defences are removed before swarming, ^ -
No precautions taken, if the entrance be artificially
restricted, -

Peculiar sound emitted by the sphinx.

It may affect the bees, , . -

The sphinx introduced among humble bees is de-
stroyed, -

It is not observed to consume honey.

The abdomen found full of it.

Bees destroy their own fortifications in spring.

307

308

309

310

311

312

31.3

315

316

317

Predatory bees, - -

The study of animal instinct is essential, - 318

Late advances in entomology, - - 320

Reaumur’s opinion on wax, - - 321

Hunter discovered the receptacles of wax, - 322

It is produced by workers only, - - 32.3

The receptacles described, - - 32i

Eight waxen scales are produced by each bee, - 32-1*

These ascertained to be wax, - - 325

Some difference subsists between them and the
wax of the combs, - - - 326

They undergo a preparation, - - 327

Bees producing them are peculiarly organised, 328
Wax probably transudes from the stomach to the
rings, - - . .

It is produced from honey, - - 331

Combs removed, five hives were replaced by the

same bees, - - - 332

Wax is obtained from sugar, - - . 333

438

INDEX.

Page.

The coarsest sugar produces the finest wax, - 334

Proportional quantity obtained, - -

VV'ax is originally white, - - . 33.^

Two kinds of workers are in a liive, - - —

Their anatomical structure is different, - 336

Wax secreted in twenty-four hours after bees have
fed, - - 337

The honey of hives is never consumed but in ne-
cessity, - - - - —

Combs are constructed only when honey is abun-
dant in the flowers, . _ _ 333

Circumstances under which bees are most active, 340
Experiments proving that pollen does not enter
wax, - - - . - 341

Bees supply their young with it, - - 345

Anomalies when they were forced to ■work in
wax from sugar, - - - 347

Every thing in nature is adapted for permanent use, 348
Architecture of bees, - . - 349

Their organization, . - - 35Q

Jt has no reference to the cells, - - — ■

Teeth or mandibles, - —

Peculiarity in the third pair of the limbs of workers, 35 1
Antennae, - - - - —

Secretion of wax in bees, - - 353

While going on in one class of bees, another ad-
vances the works of the hive, - - 354

Origin of the combs, , , - 355

Workers exti’act their own scales of wax, - — >

These are accumulated for use in a block, 357

Construction of cells, ... 358

I’eculiar figure of those of the first row, - 359

Bees avoid very smooth surfaces, - - 3(il

Secretion of wax, _ . - 362

Size of the waxen block, . , - 3(i4

INDEX.

439

Excavation of tlie first cell.

Pagk.

.

365

Enlargement of the block.

-

- 366

Formation of the second row

of cells.

* 367

The surface of a new comb

is elevated in the

middle.

-

- 372

Bees are guided by the antennae in their operations.
The wax workers enlarge the block, but do not
sculpture the cells, - . .

The ordinary conformation of combs is alone
adapted to the nature of bees.

All originate from tlie work of a single bee,

A different kind of bee selects the site of the first
cell, - . - _ .

The figure of the bottom of the first row of cells
determines that of all the rest.

Instinct of bees, ...

Formation of an angle in the combs.

Diameter of the different cells,

Apparent irregularities the result of design.

Cells are enlarged when honey is very abundant.
They are sometimes shortened.

Original foundation of combs.

The work advances slowest in ascending.

Cells are not horizontal, - . . '

I. ess symmetry in the detail than in the totality of
the operations of bees, - _ .

Symmetry is affected by interruptions of tlie work.
Accidental imperfections are corrected.

Combs never are commenced with the cells of
males, - - _

Modifications of the bottom of cells.

Departure from three lozenges and return.

The original qualities of wax alter, i

Finished combs are heavier in proportion than
when in progress.

0*^0
O iO

375

378

379

380

383

384

386

387
383

389

390

39 1
393
S94

395

396

397
399

401

440

INDEX.

Page.

Old combs contain propolis, - - 402

It is obtained from plants, « . - 404

Operations of bees with it, - - 405

Those laden are disburdened by their companions, 406
Instinct in resorting to fragments, - - 408

New cells are white, - - —

Cause of their subsequent yellowmess, - 410

Bees compound wax and propolis, - - —

Opinions of Pliny and P.eaumur, - - 411

Bees destroy the foundations of their combs to
renew and strengthen them, - - —

Proj)olis probably is softened with a fluid, 413

Combs are strengthened wdien wax abounds, 414
Means whereby bees secured a fallen comb, 415
Sometimes the combs are strengthened in winter, 41 6
Appendix, - - - - 417

Propagation of bees, - - - —

The males are mutilated, - - 418

Secretion of wax, - - - - 41 9

Dissections by Miss Jurine, - - —

Wax supposed to transude from the stomach, 420
Organization of the receptacles of wax, - 421

Mules have nothing similar, - - —

Origin of wax, . _ - - 422

The pellets on the limbs of bees are only farina, —
M’ax is secreted by bees, - - - 423

Scales of it are found only during the construction
of combs, - - - —

THE EXn.

EmN’iiuiioii:

Printed by J, C. Miiirhcad,

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