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+The Project Gutenberg EBook of New observations on the natural history of
+bees, by Francis Huber
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: New observations on the natural history of bees
+
+Author: Francis Huber
+
+Translator: Anonymous
+
+Release Date: August 28, 2008 [EBook #26457]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW OBSERVATIONS ON BEES ***
+
+
+
+
+Produced by Louise Pryor, Steven Giacomelli and the Online
+Distributed Proofreading Team at https://www.pgdp.net (This
+file was produced from images produced by Core Historical
+Literature in Agriculture (CHLA), Cornell University)
+
+
+
+
+
+{Transcriber's note
+
+The spelling in the original is sometimes idiosyncratic. It has not
+been changed, but a few obvious errors have been corrected. The
+corrections are listed at the end of this etext.}
+
+
+{Illustration: The figures that are referred to in the text}
+
+
+
+
+  NEW OBSERVATIONS
+  ON THE
+  NATURAL HISTORY
+  OF
+  BEES,
+
+  BY
+  FRANCIS HUBER.
+
+  TRANSLATED FROM THE ORIGINAL.
+
+       *       *       *       *       *
+
+  EDINBURGH:
+
+  PRINTED FOR JOHN ANDERSON,
+  AND SOLD BY
+  LONGMAN, HURST, REES, AND ORME,
+  LONDON.
+
+  ALEX SMELLIE, Printer.
+
+  1806.
+
+
+
+
+  _To
+  SIR JOSEPH BANKS, BART._
+
+  _KNIGHT OF THE MOST HONOURABLE ORDER
+  OF THE BATH, A PRIVY COUNCILLOR,
+  PRESIDENT OF THE ROYAL
+  SOCIETY OF LONDON,
+  &c. &c._
+
+  _THIS TRANSLATION
+  IS INSCRIBED._
+
+
+
+
+CONTENTS.
+
+
+  LETTER 1.--On the impregnation of the queen bee         page 1
+
+  LETTER 2.--Sequel of observations on the impregnation
+             of the queen bee                                 41
+
+  LETTER 3.--The same subject continued; observations
+             on retarding the fecundation of queens           44
+
+  LETTER 4.--On M. Schirach's discovery                       76
+
+  LETTER 5.--Experiments proving that there are
+             sometimes common bees which lay fertile eggs     89
+
+  LETTER 6.--On the combats of queens; the massacre
+             of the males; and what succeeds in a hive
+             where a stranger queen is substituted for
+             the natural one                                 108
+
+  LETTER 7.--Sequel of observations on the reception
+             of a stranger queen; M. de Reaumur's
+             observations on the subject                     137
+
+  LETTER 8.--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 coccoons 145
+
+  LETTER 9.--On the formation of swarms                      171
+
+  LETTER 10.--The same subject continued                     201
+
+  LETTER 11.--The same subject continued                     223
+
+  LETTER 12.--Additional observations on queens
+             that lay only the eggs of drones, and on
+             those deprived of the antennæ                   237
+
+  LETTER 13.--Economical considerations on bees              253
+
+  APPENDIX                                                   275
+
+
+
+
+TRANSLATOR'S PREFACE
+
+
+The facts contained in this volume are deeply interesting to the
+Naturalist. They not only elucidate the history of those industrious
+animals, whose nature is the peculiar subject of investigation, but they
+present some singular features in physiology which have hitherto been
+unknown.
+
+The industry of bees has proved a fertile source of admiration in all
+countries and in every age; and mankind have endeavoured to render it
+subservient to their gratifications or emolument. Hence innumerable
+theories, experiments, and observations have ensued, and uncommon
+patience has been displayed in prosecuting the enquiry. But although
+many interesting peculiarities have been discovered, they are so much
+interwoven with errors, that no subject has given birth to more
+absurdities than investigations into the history of bees: and
+unfortunately those treatises which are most easily attained, and the
+most popular, only serve to give such absurdities a wider range, and
+render it infinitely more difficult to eradicate them. A considerable
+portion of the following work is devoted to this purpose. The reader
+will judge of the success which results from the experiments that have
+been employed.
+
+Perhaps this is not the proper place to bestow an encomium on a treatise
+from which so much entertainment and instruction will be derived.
+However, to testify the estimation in which it is held in other nations,
+the remarks upon it by the French philosopher Sue, may be quoted, 'The
+observations are so consistent, and the consequences seem so just, that
+while perusing this work, it appears as if we had assisted the author
+in each experiment, and pursued it with equal zeal and interest. Let us
+invite the admirers of nature to read these observations; few are equal
+to them in excellence, or so faithfully describe the nature, the habits,
+and inclinations of the insects of which they treat.'
+
+It is a remarkable circumstance that the author laboured under a defect
+in the organs of vision, which obliged him to employ an assistant in his
+experiments. Thus these discoveries may be said to acquire double
+authority. But independent of this the experiments are so judiciously
+adapted to the purposes in view, and the conclusions so strictly
+logical, that there is evidently very little room for error. The talents
+of _Francis Burnens_, this philosophic assistant, had long been devoted
+to the service of the author, who, after being many successive years in
+this manner aided in his researches, was at last deprived of him by some
+unfortunate accident.
+
+Whether the author has prosecuted his investigation farther does not
+appear, as no other production of his pen is known in this island.
+
+It is vain to attempt a translation of any work without being to a
+certain degree skilled in the subject of which it treats. Some parts of
+the original of the following treatise, it must be acknowledged, are so
+confused, and some so minute, that it is extremely difficult to give an
+exact interpretation. But the general tenor, though not elegant, is
+plain and perspicuous; and such has it been here retained.
+
+
+
+
+LETTER I.
+
+_ON THE IMPREGNATION OF THE QUEEN BEE._
+
+
+  SIR,
+
+When I had the honour at Genthod of giving you an account of my
+principal experiments on bees, you desired me to transmit a written
+detail, that you might consider them with greater attention. I hasten,
+therefore, to extract the following observations from my journal.--As
+nothing can be more flattering to me than the interest you take in my
+researches, permit me to remind you of your promise to suggest new
+experiments{A}.
+
+After having long studied bees in glass hives constructed on M. de
+Reaumur's principle, you have found the form unfavourable to an
+observer. The hives being too wide, two parallel combs were made by the
+bees, consequently whatever passed between them escaped observation.
+From this inconvenience, which I have experienced, you recommended much
+thinner hives to naturalists, where the panes should be so near each
+other, that only a single row of combs could be erected between them. I
+have followed your admonitions, Sir, and provided hives only eighteen
+lines in width, in which I have found no difficulty to establish swarms.
+However, bees must not be entrusted with the charge of constructing a
+single comb: Nature has taught them to make parallel ones, which is a
+law they never derogate from, unless when constrained by some particular
+arrangement. Therefore, if left to themselves in these thin hives, as
+they cannot form two combs parallel to the plane of the hive, they will
+form several small ones perpendicular to it, and, in that case, all is
+equally lost to the observer. Thus it became essential previously to
+arrange the position of the combs. I forced the bees to build them
+perpendicular to the horizon, and so that the lateral surfaces were
+three or four lines from the panes of the hive. This distance allows the
+bees sufficient liberty, but prevents them from collecting in too large
+clusters on the surface of the comb. By such precautions, bees are
+easily established in very thin hives. There they pursue their labours
+with the same assiduity and regularity; and, every cell being exposed,
+none of their motions can be concealed.
+
+It is true, that by compelling these insects to a habitation where they
+could construct only a single row of combs, I had, in a certain
+measure, changed their natural situation, and this circumstance might
+possibly have affected their instinct. Therefore, to obviate every
+objection, I invented a kind of hives, which, without losing the
+advantages of those very thin, at the same time approached the figure of
+common hives where bees form several rows of combs.
+
+I took several small fir boxes, a foot square and fifteen lines wide,
+and joined them together by hinges, so that they could be opened and
+shut like the leaves of a book{B}. When using a hive of this
+description, we took care to fix a comb in each frame, and then
+introduced all the bees necessary for each particular experiment. By
+opening the different divisions successively, we daily inspected both
+surfaces of every comb. There was not a single cell where we could not
+distinctly see what passed at all times, nor a single bee, I may almost
+say, with which we were not particularly acquainted. Indeed, this
+construction is nothing more than the union of several very flat hives
+which may be separated. Bees, in such habitations, must not be visited
+before their combs are securely fixed in the frames, otherwise, by
+falling out, they may kill or hurt them, as also irritate them to that
+degree that the observer cannot escape stinging, which is always
+painful, and sometimes dangerous: but they soon become accustomed to
+their situation, and in some measure tamed by it; and, 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 exhibiting too formidable
+symptoms of displeasure. You will remember, Sir, that on visiting my
+retreat, I shewed you a hive of this kind that had been a long time in
+experiment, and how much you were surprised that the bees so quietly
+allowed us to open it.
+
+In these hives, I have repeated all my observations, 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
+inconvenience 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 error; and it also appears, that some
+advantages are found in these which may be called _Book_ or
+_Leaf-hives_, as they prove extremely useful in the economical treatment
+of bees, which shall afterwards be detailed.
+
+I now come to the particular object of this letter, the fecundation of
+the queen bee; and I shall, in a few words, examine the different
+opinions of naturalists on this singular problem. Next I shall state the
+most remarkable observations which their conjectures have induced me to
+make, and then describe the new experiments by which I think I have
+solved the problem{C}.
+
+Swammerdam, who studied bees with unremitting attention, and who never
+could see a real copulation between a drone and a queen, was satisfied
+that copulation was unnecessary for fecundation of the eggs: but having
+remarked that, at certain times, the drones exhaled a very strong odour,
+he thought this odour was an emanation of the _aura seminalis_, or the
+_aura seminalis_ itself, which operated fecundation by penetrating the
+body of the female. His conjecture was confirmed on dissecting the male
+organs of generation; for he was so much struck with the disproportion
+between them and those of the female, that he did not believe copulation
+possible. His opinion, concerning the influence of the odour, had this
+farther advantage, that it afforded a good reason for the prodigious
+number of the males. There are frequently fifteen hundred or two
+thousand in a hive; and, according to Swammerdam, it is necessary they
+should be numerous, that the emanation proceeding from them may have an
+intensity or energy sufficient to effect impregnation.
+
+Though M. de Reaumur has refuted this hypothesis by just and conclusive
+reasoning, he has failed to make the sole experiment that could support
+or overturn it. This was to confine all the drones of a hive in a tin
+case, perforated with minute holes, which might allow the emanation of
+the odour to escape, but prevent the organs of generation from passing
+through. Then, this case should have been placed in a hive well
+inhabited, but completely deprived of males, both of large and small
+size, and the consequence attended to. It is evident, 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
+confuted had she produced no eggs, or only sterile ones. However the
+experiment has been made by us, and the queen remained barren;
+therefore, it is undoubted, that the emanation of the odour of the males
+does not impregnate bees.
+
+M. de Reaumur was of a different opinion. He thought that the queen's
+fecundation followed actual copulation. He confined several drones in a
+glass vessel along with a virgin queen: he saw the female make many
+advances to the males; but, unable to observe any union so intimate
+that it could be denominated copulation, he leaves the question
+undecided. We have repeated this experiment: we have frequently confined
+virgin queens with drones of all ages: we have done so at every season,
+and witnessed all their advances and solicitations to the males: we have
+even believed we saw a kind of union between them, but so short and
+imperfect that it was unlikely to effect impregnation. Yet, to neglect
+nothing, we confined the virgin queen, that had suffered the approaches
+of the male, to her hive. During a month that her imprisonment
+continued, she did not lay a single egg; therefore, these momentary
+junctions do not accomplish fecundation.
+
+In the _Contemplation de la Nature_, you have cited the observations of
+the English naturalist Mr Debraw. They appear correct, and at last to
+elucidate the mystery. Favoured by chance, the observer one day
+perceived at the bottom of cells containing eggs, a whitish fluid,
+apparently spermatic, at least, very different from the substance or
+jelly which bees commonly collect around their new hatched worms.
+Solicitous to learn its origin, and conjecturing that it might be the
+male prolific fluid, he began to watch the motions of every drone in the
+hive, on purpose to seize the moment when they would bedew the eggs. He
+assures us, that he saw several insinuate the posterior part of the body
+into the cells, and there deposit the fluid. After frequent repetition
+of the first, he entered on a long series of 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 no brood.
+He saw the queen lay eggs, which were bedewed by the males, and from
+which larvæ were hatched, consequently, he could not hesitate advancing
+as a fact demonstrated, that male bees fecundate the queen's eggs in
+the manner of frogs and fishes, 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 number of males in a hive. At the same time,
+the author had neglected to answer one strong objection. Larvæ appear
+when there are no drones. From the month of September until April, hives
+are generally destitute of males, yet, notwithstanding their absence,
+the queen then lays fertile eggs. Thus, the prolific fluid cannot be
+required to impregnate them, unless we can suppose that it is necessary
+at a certain time of the year, while at every other season it is
+useless.
+
+To discover the truth amidst these facts apparently so contradictory, I
+wished to repeat Mr Debraw's experiments, and to observe more precaution
+than he himself had done. First, I sought for the fluid, which he
+supposes the seminal, in cells containing eggs. Several were actually
+found with that appearance; and, during the first days of observation,
+neither my assistant nor myself doubted the reality of the discovery.
+But we afterwards found it an illusion arising from the reflection of
+the light, for nothing like a fluid was visible, except when the solar
+rays reached the bottom of the cells. Fragments of the coccoons of
+worms, successively hatched, commonly cover the bottom; and, as they are
+shining, it may easily be conceived that, when much illuminated, an
+illusory effect results from the light. We proved it by the strictest
+examination, for no vestiges of a fluid were perceptible when the cells
+were detached and cut asunder.
+
+Though the first observation inspired us with some distrust of Mr
+Debraw's discovery, we repeated his other experiments with the utmost
+care. On the 6. of August 1787, we immersed a hive, and, with scrupulous
+attention, examined the whole bees while in the bath. We ascertained
+that there was no male, either large or small; and having examined all
+the combs, we found neither male nymph, nor worm. When the bees were
+dry, we replaced them all, 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 necessary that no male should enter the hive during the
+experiment, a glass tube was adapted 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 removed any male that appeared,
+that the result of the experiment might be undisturbed, and I can
+positively affirm that not one was seen. However, from the first day,
+which was the sixth 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 where it was impossible one could be
+introduced, since these eggs, I say, were fertile, it becomes
+indubitable that the fluid of the males is not required for their
+exclusion.
+
+Though it did not appear that any reasonable objection could be started
+against this conclusion, yet, as I had been accustomed in all my
+experiments to seek for the most trifling difficulties that could arise,
+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 fluid to their own habitations for depositing it
+on the eggs.
+
+It was easy to appreciate the force of this objection, for all that was
+necessary was a repetition of the former experiments, and to confine the
+bees so closely to their hives that none could possibly escape. You
+very well know, Sir, that these animals can live three or four months
+confined in a hive well stored with honey and wax, and if apertures are
+left for circulation of the air. This experiment 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 larvæ.
+
+I extended the precautions so far as to immerse this hive a second time,
+to assure myself that no male had escaped my researches. 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; and that was, his using queens whose
+history he was unacquainted with from their origin. When he observed
+the eggs produced by a queen, confined along with males, were fertile,
+he thence concluded that they had been bedewed by the prolific fluid in
+the cells: but to render his conclusion just, he should first have
+ascertained that the female never had copulated, and this he neglected.
+The truth is, that, without knowing it, he had used, in his experiments,
+a queen after she had commerce with the male. Had he taken a virgin
+queen the moment she came from the royal cell, and confined her along
+with drones in his vessels, the result would have been opposite; for,
+even amidst a seraglio of males, this young queen would never have laid,
+as I shall afterwards prove.
+
+The Lusatian observers, and M. Hattorf in particular, thought the queen
+was fecundated by herself, without concourse with the males. I shall
+here give an abstract of the experiment on which that opinion is
+founded.{D}
+
+M. Hattorf took a queen whose virginity he could not doubt. He excluded
+all the males both of the large and small species, and, in several days,
+he 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
+absent, the queen laid eggs, from which came worms: whence he considers
+she is impregnated by herself.
+
+Reflecting on this experiment, I do not find it sufficiently accurate.
+Males pass with great facility from hive to hive; and M. Hattorf took no
+precaution that none was introduced into his. He says, indeed, there was
+no male, but is silent respecting the means he adopted to prove the
+fact. Though he might be satisfied of no large drone being there, 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, and without greater 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. For several days I
+visited the hive, and found new hatched worms in it. Here then is the
+same result as M. Hattorf obtained? But before deducing the same
+consequence from it, we had to ascertain beyond dispute that no male had
+entered the hive. Thus, it was necessary to immerse the bees, and
+examine each separately. By this operation, we actually found four small
+males. Therefore, to render the experiment decisive, not only was it
+requisite to remove all the drones, but also, by some infallible method,
+to prevent any from being introduced, which the German naturalist had
+neglected.
+
+I prepared to repair this omission, by putting a virgin queen into a
+hive, from which the whole males were carefully removed; and to be
+physically certain that none should enter, a glass tube was adapted at
+the entrance of such dimensions that the working bees could freely pass
+and repass, but too narrow for the smallest male. Matters continued thus
+for thirty days, the workers departing and returning performed their
+usual labours: but the queen remained sterile. At the expiration of this
+time, her belly was equally slender as at the moment of her origin. I
+repeated the experiment several times, and always with the same
+consequence.
+
+Therefore, as a queen, rigorously separated from all commerce with the
+male, remains sterile, it is evident she cannot impregnate herself, and
+M. 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 acquired the
+knowledge of new facts, but these were apparently so contradictory as to
+render the solution of the problem still more difficult. While
+examining Mr. Debraw's hypothesis, I confined a queen in a hive, from
+which all the drones were removed; the queen nevertheless was fertile.
+When considering the opinion of M. Hattorf on the contrary, I put a
+queen, of whose virginity I was perfectly satisfied, in the same
+situation, she remained sterile.
+
+Embarrassed by so many difficulties, I was on the point of abandoning
+the subject of my researches, when at length by more attentive
+reflection, I thought these contradictions might arise from experiments
+made indifferently on virgin queens, and on those with whose history I
+was not acquainted from the origin, and which had perhaps been
+impregnated unknown to me. Impressed with this idea, I undertook a new
+method of observation not on queens fortuitously taken from the hive,
+but on females decidedly in a virgin state, and whose history I knew
+from the instant they left the cell.
+
+From a very great number of hives, I removed all the virgin females,
+and substituted for each a queen taken at the moment of her birth. The
+hives were then divided into two classes. From the first, I took the
+whole males both large and small, and adapted a glass tube at the
+entrance, so narrow, 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 all the drones belonging to them, and even introduced more; and to
+prevent them from escaping, a glass tube, also too narrow for the males,
+was adapted to the entrance of these hives.
+
+For more than a month, I carefully watched this experiment, made on a
+large scale; but much to my surprise, all the queens remained 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 necessary 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 29. June 1788.
+
+Aware, that in summer the males usually leave the hive at the warmest
+time of the day, it was natural for me to conclude that if the queens
+were also obliged to go out for impregnation, instinct would induce them
+to do so at the same time as the males.
+
+At eleven in the forenoon, we placed ourselves opposite a hive
+containing an unimpregnated queen five days old. The sun had shone from
+his rising; the air was very warm; and the males began to leave the
+hives. We then enlarged the entrance of that which we wished to observe,
+and paid great attention to the bees that entered and departed. The
+males appeared, and immediately took flight. Soon afterwards, the young
+queen appeared at the entrance; at first she did not fly, but brushed
+her belly with her hind legs, and traversed the board a little; neither
+workers nor males paid any attention to 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 departure, perhaps judging this precaution
+necessary to recognize it; she then flew away, describing horizontal
+circles twelve or fifteen feet above the earth. We contracted the
+entrance 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 observe all her motions. But she did not remain
+long in a situation favourable for us, and rapidly rose out of sight. We
+resumed our place before the hive; and in seven minutes, the young queen
+returned to the entrance of a habitation which she had left for the
+first time. Having found no external appearance of fecundation, we
+allowed her to enter. 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. Her second
+absence was much longer than the first; twenty-seven minutes elapsed
+before she came back. We then found her in a state very different from
+that in which she was after her first excursion. The sexual organs were
+distended by a white substance, thick and hard, very much resembling the
+fluid in the vessels of the male, completely similar to it indeed in
+colour and consistence{E}.
+
+But more evidence than mere resemblance was requisite to establish that
+the female had returned with the prolific fluid of the males. We allowed
+this queen to enter the hive, and confined her there. In two days, we
+found her belly swoln; and she had already laid near an hundred eggs in
+the worker's cells.
+
+To confirm our discovery, we made several other experiments, and with
+the same success. I shall continue to transcribe my journal.
+
+On the second of July, the weather being very fine, numbers of males
+left the hives. We set at liberty an unimpregnated young queen, eleven
+days old, whose hive had always been deprived of males. Having quickly
+left the hive, she returned to examine it, and then rose out of sight.
+In a few minutes, she returned without any external marks of
+impregnation. In a quarter of an hour, she departed again, but her
+flight was so rapid that we could scarcely follow her a moment. This
+absence continued thirty minutes. On returning, the last ring of the
+body was open, and the sexual organs full of the whitish substance
+already mentioned. She was then replaced in the hive from which all the
+males were excluded. In two days, we found her impregnated.
+
+These observations at length demonstrate why M. Hattorf obtained results
+so different from ours. His queens, though in hives deprived of males,
+had been fecundated, and he thence concludes that sexual intercourse is
+not requisite for their impregnation. But he did not confine the queens
+to their hives, and they had profited by their liberty to unite with the
+males. We, on the contrary, have surrounded our queens with a number of
+males; but they continued sterile; because the precaution of confining
+the males to their hives had also prevented the queens from departing to
+seek that fecundation without, which they could not obtain within.
+
+These experiments were repeated on queens, twenty, twenty-five, and
+thirty days old. All became fertile after a single impregnation;
+however, we have remarked some essential peculiarities in the fecundity
+of those unimpregnated until the twentieth day of their existence; but
+we shall defer speaking of the fact until we can present naturalists
+with observations sufficiently secure and numerous to merit their
+attention: Yet let me add a few words more. Though neither my assistant
+nor myself have witnessed the copulation of a queen and a drone, we
+think that, after the detail which has just been commenced, no doubt of
+it can remain, or of the necessity of copulation to effect impregnation.
+The sequel of experiments, made with every possible precaution, appears
+demonstrative. The uniform sterility of queens in hives wanting males,
+and in those where they were confined along with them; the departure of
+these queens from the hives; and the very conspicuous evidence of
+impregnation with which they return, 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 copulation.
+
+Naturalists have always been very much embarrassed to account for the
+number of males found 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 that extent. As fecundation
+cannot be accomplished within, and as the queen is obliged to traverse
+the expanse of the atmosphere, 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 drones in each hive, there would be little probability
+of their departure at the same instant with the queen, or that they
+would meet in their excursions; and most of the females would thus
+remain sterile.
+
+But why has nature prohibited copulation within the hives? This is a
+secret still unknown to us. It is possible, however, that some
+favourable circumstance may enable us to penetrate it in the course of
+our observations. Various conjectures may be formed; but at this day we
+require facts, and reject gratuitous suppositions. It should be
+remembered, that bees do not form the sole republic among insects
+presenting a similar phenomenon; female ants are also obliged to leave
+the ant-hills previous to fecundation.
+
+I cannot request, Sir, that you will communicate the reflections which
+your genius will excite concerning the facts I have related. This is a
+favour to which I am not yet entitled. But as new experiments will
+unquestionably occur to you, whether on the impregnation 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 encouragement that the
+continuance of my labours can receive.
+
+  _Pregny, 13th August 1789._
+
+       *       *       *       *       *
+
+_Letter from M. Bonnet to M. Huber._
+
+You have most agreeably surprised me, Sir, with your interesting
+discovery of the impregnation of the queen bee. It was a fortunate idea,
+that she left the hive to be fecundated, and your method of ascertaining
+the fact was extremely judicious and well adapted to the object in view.
+
+Let me remind you, that male and female ants copulate in the air; and
+that after impregnation the females return to the ant hills to deposit
+their eggs. _Contemplation de la Nature, Part 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 a copulation in the air. If you have satisfactory evidence
+that the fluid bedewing the last rings of the female is the same with
+that of the male, it is more than mere presumption in favour of
+copulation. Perhaps it may be necessary that the male should seize the
+female under the belly, which cannot easily be done but in the air. The
+large opening at the extremity of the queen, which you have observed in
+so particular a condition, seems to correspond to the singular size of
+the sexual parts of the male.
+
+You wish, my dear Sir, that I should suggest some new experiments on
+these industrious republicans. In doing so, I shall take the greater
+pleasure and interest, as I know to what extent you possess the valuable
+art of combining ideas, and of deducing from this combination 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
+queen, by introducing a little of the male's prolific fluid with a
+pencil, and at the same time observing every precaution to avoid error.
+Artificial fecundation, you are aware, has already succeeded in 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 paint the thorax with some varnish that resists humidity. It will
+also be right to paint the thorax of a considerable number of workers in
+order to discover the duration of their life. This is a more secure
+method than slight mutilations.
+
+For hatching the worm, the egg must be fixed almost vertically by one
+end near the bottom of the cell. Is it true, that it is unproductive
+unless fixed in this manner? I cannot determine the fact; and therefore
+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 queens in the cells, and my
+inclination to suppose them minute worms not yet begun to expand. Their
+elongated figure seems to favour my suspicions. It would therefore be
+proper to watch them with the utmost assiduity, from the instant of
+production until the period of exclusion. If the integument bursts,
+there can be no doubt that these minute substances are real eggs.
+
+I return to the mode of operating copulation. The height that the queen
+and the males rise to in the air prevent us from seeing what passes
+between them. On that account, the hive should be put into an apartment
+with a very lofty ceiling. M. de Reaumur's experiment of confining a
+queen with several males in a glass vessel, merits repetition; and if,
+instead of a vessel, a glass tube, some inches in diameter and several
+feet long, were used, perhaps something satisfactory might be
+discovered.
+
+You have had the fortune to observe the small queens mentioned by the
+Abbe Needham, but which he never saw. It will be of great importance to
+dissect them for the purpose of finding their ovaries. When M. Reims
+informed me that he had confined three hundred workers, along with a
+comb containing no eggs, and afterwards found hundreds in it, I strongly
+recommended that he should dissect the workers. He did so; and informed
+me that eggs were found in three. Probably without being aware of it, he
+has dissected small queens. As small drones exist, it is not surprising
+if small queens are produced also, and undoubtedly by the same external
+causes.
+
+It is of much consequence to be intimately acquainted with this species
+of queens, for they may have great influence on different experiments
+and embarrass the observer: we should ascertain whether they inhabit
+pyramidal cells smaller than the common, or hexagonal ones.
+
+M. Schirach's famous experiment on the supposed conversion of a common
+worm into a royal one, cannot be too often repeated, though the Lusatian
+observers have already done it frequently. I could wish to learn
+whether, as the discoverer maintains, the experiment will succeed only
+with worms, three or four days old, and never with simple eggs.
+
+The Lusatian observers, and those of the Palatinate, affirm, that when
+common bees are confined with combs absolutely void of eggs, they then
+lay none but the eggs of drones. Thus, there must 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 ovaries contain male eggs alone?
+
+According to M. de Reaumur, the life of chrysalids may be prolonged by
+keeping them in a cold situation, such as an ice-house. The same
+experiment should be made on the eggs of a queen; on the nymphs of
+drones and workers.
+
+Another interesting experiment would be to take away all the combs
+composing the common cells, and leave none but those destined for the
+larvæ 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.
+Nevertheless, it is likely, by taking away only part of the common
+cells, the workers may be forced to lay common eggs in the cells of
+drones.
+
+I should also wish to have the young larvæ 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 influence 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
+adopted to instruct us how bees can regulate their labours, and apply
+them to existing circumstances. This may enable us to discover
+particular facts which we cannot foresee.
+
+The royal eggs and those producing drones, have not yet been carefully
+compared with the eggs from which workers come. But they ought to be so,
+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 endeavour, with the point of
+a pencil, to remove 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 this experiment.
+
+Various facts, which require corroboration, were collected in my Memoirs
+on Bees; of this number are my own observations. You can select what is
+proper, my dear Sir. 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. _Genthod, 18. August 1789._
+
+
+FOOTNOTES:
+
+{A} All these letters are addressed to the celebrated naturalist M.
+Bonnet.--_T._
+
+{B} The leaf or book hive consists of twelve vertical frames or boxes,
+parallel to each other, and joined together. Fig. 1. the sides, f f. f
+g. should be twelve inches long, and the cross spars, f f. g g. nine or
+ten; the thickness of these spars an inch, and their breadth fifteen
+lines. It is necessary that this last measure should be accurate; a a. a
+piece of comb which guides the bees in their work; d. a moveable slider
+supporting the lower part; b b. pegs to keep the comb properly in the
+frame or box; four are in the opposite side; e e. pegs in the sides
+under the moveable slider to support it.
+
+A book hive, consisting of twelve frames, all numbered, is represented
+fig. 2. Between 6 and 7 are two cases with lids, that divide the hive
+into two equal parts, and should only be used to separate the bees for
+forming an artificial swarm; a a. two frames which shut up the two sides
+of the hive, have sliders, b. b.
+
+The entrance appears at the bottom of each frame. All should be close
+but 1 and 12. However it is necessary that they should open at pleasure.
+
+The hive is partly open, fig. 3. and shews how the component parts may
+be united by hinges, and open as the leaves of a book. The two covers
+closing up the sides, a. a.
+
+Fig. 4. is another view of fig. 1. a a. a piece of comb to guide the
+bees; b b. pegs disposed so as to retain the comb properly in the frame;
+c c. parts of two shelves; the one above is fixed, and keeps the comb in
+a vertical position; the under one, which is moveable, supports it
+below.
+
+{C} I cannot insist that my readers, the better to comprehend what is
+here said, shall peruse the Memoirs of M. de Reaumur on Bees, and those
+of the Lusace Society; but I must request them to examine the extracts
+in M. Bonnet's works, tom. 5. 4to edit. and tom. 10. 8vo, where they
+will find a short and distinct abstract of all that naturalists have
+hitherto discovered on the subject.
+
+{D} Vide M. Schirach's History of Bees, in a memoir by M. Hattorf,
+entitled, _Physical Researches whether the Queen Bee requires
+fecundation by Drones?_
+
+{E} It will afterwards appear that what we took for the generative
+fluid, was the male organs of generation, left by copulation in the body
+of the female. This discovery we owe to a circumstance that shall
+immediately be related. Perhaps I should avoid prolixity, by suppressing
+all my first observations on the impregnation of the queen, and by
+passing directly to the experiments 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 think service is
+done to the reader by a candid avowal of my errors. This is an
+additional proof to so many others, of the absolute necessity that an
+observer should repeat all his experiments a thousand times, to obtain
+the certainty of seeing facts as they really exist.
+
+
+
+
+LETTER II.
+
+_SEQUEL OF OBSERVATIONS ON THE IMPREGNATION OF THE QUEEN BEE._
+
+
+  SIR,
+
+All the experiments, related in my preceding letter, were made in 1787
+and 1788. They seem to establish two facts, which had previously been
+the subject of vague conjecture: 1. The queen bee is not impregnated of
+herself, but is fecundated by copulation with the male. 2. Copulation is
+accomplished without the hive, and in the air.
+
+The latter appeared so extraordinary, that notwithstanding all the
+evidence obtained of it, we eagerly desired to take the queen in the
+fact; but, as she always rises to a great height, we never could see
+what passed. On that account you advised us to cut part off the wings of
+virgin queens. We endeavoured to benefit by your advice, in every
+possible manner; but to our great regret, when the wings lost much, the
+bees could no longer fly; and, by cutting off only an inconsiderable
+portion, we did not diminish the rapidity of their flight. Probably
+there is a medium, but we were unable to attain it. On your suggestion,
+we tried to render their vision less acute, by covering the eyes with an
+opaque varnish, which was an experiment equally fruitless.
+
+We likewise attempted artificial fecundation, and took every possible
+precaution to insure success. Yet the result was always unsatisfactory.
+Several queens were the victims of our curiosity; and those surviving
+remained sterile. Though these different experiments were unsuccessful,
+it was proved that queens leave their hives to seek the males, and that
+they return with undoubted evidence of fecundation. Satisfied with this,
+we could only trust to time or accident for decisive proof of an actual
+copulation. We were far from suspecting a most singular discovery, which
+we made in July this year, and which affords complete demonstration of
+the supposed event, namely, that the sexual organs of the male remain
+with the female.{F}
+
+
+FOOTNOTES:
+
+{F} The remainder of this Chapter chiefly consists of anatomical
+details. These may rather be considered an interruption of the
+narrative; and the Translator has judged it expedient to transfer them
+to an Appendix.
+
+
+
+
+LETTER III.
+
+_THE SAME SUBJECT CONTINUED.--OBSERVATIONS ON RETARDING THE FECUNDATION
+OF QUEENS._
+
+
+In my first letter, I remarked, that when queens were prevented from
+receiving the approaches of the male until the twenty-fifth or thirtieth
+day of their existence, the result presented very interesting
+peculiarities. My experiments at that time were not sufficiently
+numerous; but they have since been so often repeated, and the result so
+uniform, that I no longer hesitate to announce, as a certain discovery,
+the singularities which retarded fecundation, produces on the ovaries of
+the queen. If she receives the male during the first fifteen days of her
+life, she remains capable of laying both the eggs of workers and of
+drones; but should fecundation be retarded until the twenty-second day,
+her ovaries are vitiated in such a manner that she becomes unfit for
+laying the eggs of workers, and will produce only those of drones.
+
+In June 1787, being occupied in researches 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. When a hive is ready to swarm, I had before
+observed, that the moment of swarming is always preceded by a very
+lively agitation, which first affects the queen, is then communicated to
+the workers, and excites such a tumult among them, that they abandon
+their labours, and rush in disorder to the outlets of the hive. I then
+knew very well the cause of the queen's agitation, and it is described
+in the history of swarms, but I was ignorant how the delirium
+communicated to the workers; and this difficulty interrupted my
+researches. I therefore thought of investigating, by direct experiments,
+whether at all times, when the queen was greatly agitated, even not in
+the time of the hive swarming, her agitation would in like manner be
+communicated to the workers. The moment a queen 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 it
+would produce a kind of delirium. I had the patience to observe this
+queen thirty-four days. Every morning about eleven o'clock, when the
+weather was fine and the sunshine invited the males to leave their
+hives, I saw her impetuously traverse every corner of her habitation,
+seeking to escape. Her fruitless efforts threw her into an uncommon
+agitation, the symptoms of which I shall elsewhere describe, and all the
+common bees were affected by it. As she never was out all this time, she
+could not be impregnated. At length, on the thirty-sixth day, I 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 this experiment, I was far from
+any hopes that it would lead to the knowledge of another very remarkable
+fact; how great was my astonishment, therefore, on finding that this
+female, which, as usual, began to lay forty-six hours after copulation,
+laid the eggs of drones, but none of workers, and that she continued
+ever afterwards to lay those of drones only.
+
+At first, I exhausted myself with conjectures on this singular fact; the
+more I reflected on it, the more did it seem inexplicable. At length, by
+attentively meditating on the circumstances of the experiment it
+appeared there were two principles, the influence of which I should
+first of all endeavour to appreciate separately. On the one hand, this
+queen had suffered long confinement; on the other, her fecundation had
+been extremely retarded. You know, Sir, that queens generally receive
+the males about the fifth or sixth day, and this queen had not copulated
+until the thirty-sixth. Little weight could be given to the supposition,
+that the peculiarity could be occasioned by confinement. Queens, in the
+natural state, leave their hives only once to seek the males. All the
+rest of their life they remain voluntary prisoners. Thus, it was
+improbable that captivity could produce the effect I wished to explain.
+At the same time, as it was essential to neglect nothing in a subject so
+new, I wished to ascertain whether it was owing to the length of
+confinement, or to retarded fecundation.
+
+Investigating this was no easy matter. To discover whether captivity,
+and not retarded fecundation, vitiated the ovaries, it was necessary to
+allow a female to receive the approaches of a male, and also to keep her
+imprisoned. Now this could not be, for bees never copulate in hives. On
+the same account, it was impossible to retard the copulation of a queen
+without keeping her in confinement. I was long embarrassed by the
+difficulty. At length, I contrived an apparatus, which, though
+imperfect, nearly fulfilled my purpose.
+
+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 allowed free passage 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 the queen could at all times come and fly about, enjoying a
+purer air than was to be 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 between them. By the
+experiments related in my first letter, copulation takes place high in
+the air only: therefore, in this apparatus, I found the advantage of
+retarding fecundation, while the liberty the queen now had, did not
+render her situation too remote from the natural state. I attended to
+the experiment fifteen days. Every fine morning, the young captive left
+her hive; 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 aloft in the air, and soon returned with full evidence of
+impregnation. In two days, she laid, first the eggs of workers, and
+afterwards as many as the most fertile queens.
+
+It thence followed, 1. That captivity did not alter the organs of
+queens. 2. When fecundation took place within the first sixteen days,
+she produced both species of eggs.
+
+This was an important experiment. It rendered my labours much more
+simple, by clearly pointing out the method to be pursued: it absolutely
+precluded the supposed influence of captivity; and left nothing for
+investigation but the consequences of retarded fecundation.
+
+With this view, I repeated the experiment; but, instead of giving the
+virgin queen liberty on the sixteenth day, I retained her until the
+twenty-first. She departed, rose high in the air, was fecundated, and
+returned. Thirty-six hours afterwards, she began to lay: but it was the
+eggs of males only, and, although very fruitful afterwards, she laid no
+other kind.
+
+I occupied myself the remainder of 1787, and the two subsequent years,
+with experiments on retarded fecundation, and had constantly the same
+results. It is undoubted, therefore, that when the copulation of queens
+is retarded beyond the twentieth day, only an imperfect impregnation is
+operated: instead of laying the eggs of workers and males equally, they
+will lay none but those of males.
+
+I do not aspire to the honour of explaining this singular fact. When the
+course of my experiments led me to observe 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
+fecundation. My evidence is demonstrative, for I can always prevent
+queens from laying the eggs of workers, by retarding their fecundation
+until the twenty-second or twenty-third day. But, what is the remote
+cause of this peculiarity; or, in other words, why does the delay of
+impregnation render queens incapable of laying 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 delayed. The
+queen then lays the eggs of workers forty-six hours after copulation,
+and continues for the subsequent eleven months to lay these alone: and
+it is only after this period that a considerable and uninterrupted
+laying of the eggs of drones commences. When, on the contrary,
+impregnation is retarded after the twentieth day, the queen begins, from
+the forty-sixth hour, to lay the eggs of males, and no other kind during
+her whole life. As, in the natural state, she lays the eggs of workers
+only, during the first eleven months, it is clear that these, and the
+male eggs, are not indiscriminately mixed in the oviducts. Undoubtedly
+they occupy a situation corresponding to the principles that regulate
+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 inverted by retarded copulation? How does it happen
+that all the workers eggs which the queen ought to lay, if fecundation
+was in due time, now wither and disappear, yet do not, impede the
+passage of the eggs of drones, which occupy only the second place in the
+ovaries. Nor is this all. I have satisfied myself that a single
+copulation 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 a single copulation will impregnate all the eggs that she will lay
+during her whole life: but I want absolute proof for more than two
+years. This, which is truly a very singular fact in itself, renders the
+influence of retarded fecundation still more difficult to be accounted
+for. Since a single copulation suffices, it is clear that the male fluid
+acts from the first moment on all the eggs that the queen will lay in
+two years. It gives them, according to your principles, that degree of
+_animation_ that afterwards effects their successive expansion. Having
+received the first impressions of life, they grow, they mature, so to
+speak, until the day they are laid: and as the laws of laying are
+constant, because the eggs of the first eleven months are always those
+of workers, it is evident that those which appear first are also the
+eggs that come soonest to maturity. Thus, in the natural state, the
+space of eleven months is necessary for the male eggs to acquire that
+degree of increment they must have attained when laid. This consequence,
+which to me seems immediate, renders the problem insoluble. How can the
+eggs, which should grow slowly for eleven months, suddenly acquire their
+full expansion in forty-eight hours, when fecundation has been retarded
+twenty-one days, and by the effect of this retardation alone? Observe, I
+beseech you, that the hypothesis of successive expansion is not
+gratuitous; it rests on the principles of sound philosophy. Besides, for
+conviction that it is well founded, we have only to look at the figures
+given by Swammerdam of the ovaries of the queen bee. There we see eggs
+in that part of the oviducts contiguous to the vulva, much farther
+advanced, and larger than those contained in the opposite part.
+Therefore the difficulty remains in full force: it is an abyss where I
+am lost.
+
+The only known fact bearing any relation to that now described, is the
+state of certain vegetable seeds, which, although extremely well
+preserved, lose the faculty of germination from age. The eggs of workers
+may also preserve, only for a very short time, the property of being
+fecundated by the seminal fluid; and, after this period, which is about
+fifteen or eighteen days, become disorganised to that degree, that they
+can no longer be animated by it. I am sensible that the comparison is
+very imperfect; besides, it explains nothing, nor does it even put us
+on the way of making any new experiments. I shall add but one reflection
+more.
+
+Hitherto no other effect has been observed from the retarded
+impregnation of animals, but that of rendering them absolutely sterile.
+The first instance of a female still preserving the faculty of
+engendering males, is presented by the queen bee. But as no fact in
+nature is unique, it is most probable that the same peculiarity will
+also be found in other animals. An extremely curious object of research
+would be to consider insects in this new point of view, I say _insects_,
+for I do not conceive that any thing analogous will be found in other
+species of animals. The experiments now suggested would necessarily
+begin with insects the most analogous to bees; as wasps, humble bees,
+mason bees, all species of flies, and the like. Some experiments might
+also be made on butterflies; and, perhaps, an animal might be found
+whose retarded fecundation would be attended with the same effects as
+that of queen bees. Should the animal be larger, dissection will be more
+easily accomplished; and we may discover what happens to the eggs when
+retarded fecundation prevents their expansion. At least, we might hope
+that some fortunate circumstance would lead to solution of the
+problem{G}.
+
+Let us now return to my experiments. In May 1789, I took two queens just
+when they had undergone the last metamorphosis: one was put in a _leaf
+hive_, well provided with honey and wax, and sufficiently inhabited by
+workers and males. The other was put 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, but the common
+bees enjoyed perfect liberty. The queens were imprisoned thirty days;
+and being then set at liberty, they departed, and returned impregnated.
+Visiting the hives in the beginning of July, I found much brood, but
+wholly consisting of the worms and nymphs of males. There actually was
+not a single worker's worm or nymph. Both queens laid uninterruptedly
+until autumn, and constantly the eggs of drones. Their laying ended in
+the first week of November, as that of my other queens.
+
+I was very earnest to learn what would become of them in the subsequent
+spring, whether they would resume laying, or if new fecundation would be
+necessary; and if they did lay, of what species the eggs would be.
+However, the hives being very weak, I dreaded they might perish during
+winter. Fortunately, we were able to preserve them; and from April 1790,
+they recommenced laying. The precautions we had taken prevented them
+from receiving any new approaches of the male. Their eggs were still
+those of males.
+
+It would have been extremely interesting to have followed the history of
+these two females still farther, but, to my great regret, the workers
+abandoned their hives on the fourth of May, and that same day I found
+both queens dead. No weevils were in the hive, which could disturb the
+bees; and the honey was still very plentiful: but as no workers had been
+been produced in the course of the preceding year, and winter had
+destroyed many, they were too few in spring to engage in their wonted
+labours, and, from discouragement, deserted their habitation to occupy
+the neighbouring hives.
+
+In my Journal, I find a detail of many experiments on the retarded
+impregnation of queen bees, so many, that transcribing the whole would
+be tedious. I may repeat, however, that there was not the least
+variation in the principle, and that whenever the copulation of queens
+was postponed beyond the twenty-first day, the eggs of males only were
+produced. Therefore, I shall limit my narrative to those experiments
+that have taught me some remarkable facts.
+
+A queen being hatched on the fourth of October 1789, we put her into a
+leaf-hive. Though the season was well advanced, a considerable number of
+males was still in the hive; and it here became important to learn,
+whether, at this period of the year, they could equally effect
+fecundation; also, in case it succeeded, whether a laying, begun in the
+middle of autumn, would be interrupted or continued during winter. Thus,
+we allowed the queen to leave the hive. She departed, indeed, but made
+four and twenty fruitless attempts before returning with the evidence of
+fecundation. Finally, on the thirty-first of October, she was more
+fortunate: She departed, and returned with the most undoubted proof of
+the success of her amours: She was now twenty-seven days old,
+consequently fecundation had been retarded. She ought to have begun
+laying within forty-six hours, but the weather was cold, and she did not
+lay; which proves, as we may cursorily remark, that refrigeration of the
+atmosphere is the principal agent that suspends the laying of queens
+during winter. I was excessively impatient to learn whether, on the
+return of spring, she would prove fertile, without a new copulation. The
+means of ascertaining the fact was easy; for the entrances of the hives
+only required contraction, so as to prevent her from escaping. She was
+confined from the end of October until May. In the middle of March, we
+visited the combs, and found a considerable number of eggs, but, none
+being yet hatched, we could not know whether they would produce workers
+or males. On the fourth of April, having again examined the state of the
+hive, we found a prodigious quantity of nymphs and worms, all of
+drones; nor had this queen laid a single worker's egg.
+
+Here, as well as in the preceding experiment, 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
+copulation 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
+begin to lay in spring without requiring new copulation.
+
+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, already flies, two thousand four hundred and
+thirty-eight cells, containing either eggs or nymphs of drones. Thus,
+she had laid more than three thousand male eggs during March and April,
+which is above fifty each day. Her death soon afterwards unfortunately
+interrupted my observation, I intended to calculate the total number of
+male eggs that she should lay throughout the year, and compare it with
+those 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 considerable, commences in August, also in the
+interval, that they produce the eggs of workers almost solely. But it is
+otherwise with the females whose copulation has been retarded: they
+produce no workers' eggs. For four or five months following, they lay
+the eggs of males without interruption, and in such numbers, that, in
+this short time, I suppose one queen gives birth to more drones than a
+female, whose fecundation has not been retarded, produces in the course
+of two years. It gives me much regret, that I have not been able to
+verify this conjecture.
+
+I should also describe the very remarkable manner in which queens, that
+lay only the eggs of drones, sometimes deposit them in the cells.
+Instead of being placed in the lozenges forming the bottom, they are
+frequently deposited on the lower side of the cells, two lines from the
+mouth. This arises from the body of such queens being shorter than that
+of those whose fecundation has not been retarded. The extremity remains
+slender, while the first two rings next the thorax are uncommonly swoln.
+Thus, in disposing themselves for laying, the extremity cannot reach the
+bottom of the cells on account of the swoln rings; consequently the eggs
+must remain attached to the part that the extremity reaches. The worms
+proceeding from them pass their vermicular state in the same place where
+the eggs were deposited, which proves that bees are not charged with the
+care of transporting the eggs as has been supposed. But here they
+follow another plan. They extend beyond the surface of the comb those
+cells where they observe the eggs deposited, two lines from the mouth.
+
+Permit me, Sir, to digress a moment from the subject, to give the result
+of an experiment which seems interesting. Bees, I say, are not charged
+with the care of transporting into cells, the eggs misplaced by the
+queen: and, judging by the single instance I have related, you will
+think me well entitled to deny this feature of their industry. However,
+as several authors have maintained the reverse, and even demanded our
+admiration of them in conveying the eggs, I should explain clearly that
+they are deceived.
+
+I had a glass hive constructed of two stages; the higher was filled with
+combs of large cells, and the lower with those of common ones. A kind of
+division, or diaphraghm, separated these two stages from
+each other, having at each side an opening 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 had only those of workers to lay, and
+she was obliged to deposit them in the surrounding large cells from the
+want of others. My object in this arrangement will already be
+anticipated. My reasoning was simple. If the queen laid workers' eggs in
+the large cells, and the bees were charged with transporting them if
+misplaced, they would infallibly take advantage of the liberty allowed
+to pass from either stage: they would seek the eggs deposited in the
+large cells, and carry them down to the lower stage containing the cells
+adapted for that species. If, on the contrary, they left the common eggs
+in the large cells, I should obtain certain proof that they had not the
+charge of transporting them.
+
+The result of this experiment excited my curiosity extremely. We
+observed the queen several days without intermission. During the first
+twenty-four hours, she persisted in not laying a single egg in the
+surrounding cells; she examined them one after another, but passed on
+without insinuating her belly into one. She was restless, and traversed
+the combs in all directions: her eggs appeared an oppressive burden, but
+she persisted 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 mother. I was amused to observe, when she
+approached the edges of the division separating the two stages, that she
+gnawed at them to enlarge the passage: the workers approached her, and
+also laboured with their teeth, and made every exertion to enlarge the
+entrance 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. Then we conceived that the bees would convey them into the
+small cells of the lower stage, and we sought them there with the utmost
+assiduity; but I can safely affirm there was not one. The eggs that the
+queen still laid the third day disappeared as the first. We again sought
+them in the small cells, but none were there. The fact is, they are ate
+by the workers; and this is what has deceived the naturalists, who
+supposed them carried away. They have observed the misplaced eggs
+disappear, and, without farther investigation, have asserted that the
+bees convey them elsewhere: they take them, indeed, not to convey them
+any where, but to devour them. Thus nature has not charged bees with the
+care of placing the eggs in the cells appropriated for them, but she has
+inspired females themselves with sufficient instinct to know the species
+of eggs they are about to lay, and to deposit them in suitable cells.
+This has already been observed by M. de Reaumur, and here my
+observations correspond with his. Thus it is certain that in the natural
+state, when fecundation takes place at the proper time, and the queen
+has suffered from nothing, she is never deceived in the choice of the
+cells where her eggs are to be deposited; she never fails to lay those
+of workers in small cells, and those of males in large ones. The
+distinction is important, for the same certainty of instinct is no
+longer conspicuous in the conduct of those females whose impregnation
+has been deferred. I was oftener than once deceived respecting the eggs
+that such queens laid, for they were deposited indiscriminately in small
+cells and those of drones; and not aware of their instinct having
+suffered, I conceived that the eggs in small cells would produce
+workers; therefore I was very much surprised, when, at the moment they
+should have been hatched, the bees closed up the cells, and
+demonstrated, by anticipation, 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 experiments on queens that lay only the eggs of males, not to
+be deceived by these circumstances, and expect that eggs of males will
+be deposited in the workers cells.
+
+It is a singular fact, that the females, whose fecundation has been
+retarded, sometimes lay the eggs of males in royal cells. I shall prove,
+in the history of swarms, that immediately when queens, in the natural
+state, begin their great laying of male eggs, the workers construct
+numerous royal cells. Undoubtedly, there is some secret relation between
+the appearance of male eggs and the construction of these cells; for it
+is a law of nature from which bees never derogate. It is not surprising,
+therefore, that such cells are constructed in hives governed by queens
+laying the eggs 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 take exactly the same care of the male eggs
+deposited in royal cells, as of those that should become queens. They
+provide them more plentifully with food, they build up the cells as if
+containing a royal worm; in a word, they labour with such regularity
+that we have frequently been deceived. More than once, in the firm
+persuasion of finding royal nymphs, we have opened the cells after they
+were sealed, yet the nymph of a drone always appeared. Here the instinct
+of the workers seemed defective. In the natural state, they can
+accurately distinguish the male worms from those of common bees, as they
+never fail giving 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 to investigate the instinct 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 de la Reine des Abeilles_, translated from the
+German by _Blassiere_, there is printed a letter from M. Schirach to
+you, dated 15 April 1771, where he speaks of some hives, in which the
+whole brood changed into drones. You will remember that he ascribes this
+circumstance to some unknown vice in the ovaries of the queen; but he
+was far from suspecting that retarded fecundation had been the cause of
+vitiation. He justly felicitated himself on discovering a method to
+prevent the destruction of hives in this situation, which was simple,
+for it consisted in removing the queen that laid the eggs of males only,
+and substituting one for her whose ovaries were not impaired. But to
+make the substitution effectual, it was necessary to procure queens at
+pleasure; a secret reserved for M. Schirach, and of which I shall speak
+in the following letter. You observe that the whole experiments of the
+German naturalist tended to the preservation of the hives whose queens
+laid none except male eggs; and that he did not attempt to discover the
+cause of the vice evident in their ovaries.
+
+M. de Reaumur also says a few words, somewhere, of a hive containing
+many more drones than workers, but advances no conjectures on the cause.
+However, he adds, as a remarkable circumstance, that the males were
+tolerated in this hive until the subsequent spring. It is true that bees
+governed 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. I can ascribe no reason for it, but it is a fact I have
+several times witnessed during my long course of observations on
+retarded impregnation. In general it has appeared that while the queen
+lays male eggs, bees do not massacre the males already perfect in the
+hive. PREGNY, _21. August 1791_.
+
+
+FOOTNOTES:
+
+{G} The experiments suggested in this paragraph, recall a singular
+reflection of M. de Reaumur. Where treating of oviparous flies, he says,
+it would not be impossible for a hen to produce a living chicken, if,
+after fecundation, the eggs she should first lay could by any means be
+retained twenty-one days in the oviducts. _Mem. sur. les Insect. tom. 4.
+mem. 10._
+
+
+
+
+LETTER IV.
+
+_ON M. SCHIRACH'S DISCOVERY._
+
+
+When you found it necessary, Sir, in the new edition of your works, to
+give an account of M. Schirach'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
+testimonies. For this reason, I hasten to inform you that all my
+researches establish the reality of the discovery. During ten years
+that I have studied bees, I have repeated M. Schirach's experiment so
+often, and with such uniform success, that I can no longer have the
+least doubt on the subject. Therefore, I consider it an established
+fact, when bees lose their queen, and 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, but a greater quantity of it,
+and 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 a fact, and the philosophical consequences
+you have deduced from it. _Contemplation de la Nature, part. II, chap.
+27._
+
+In this letter I shall content myself with some account of the figure of
+the royal cells constructed by bees around those worms that are destined
+for the royal state, and terminate with discussing some points wherein
+my observations differ from those of M. Schirach.
+
+Bees soon become sensible of having lost their queen, and in a few hours
+commence the labour necessary to repair their loss. First, they select
+the young common worms, which the requisite treatment is to convert into
+queens, and immediately begin with enlarging the cells where they are
+deposited. Their mode of proceeding is curious; and the 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.
+Having chosen a worm, they sacrifice three of the contiguous cells:
+next, they supply it with food, and raise a cylindrical inclosure
+around, by which the cell becomes a perfect tube, with a rhomboidal
+bottom; for the parts forming the bottom are left untouched. If the bees
+damaged it, they would lay open three corresponding cells on the
+opposite surface of the comb, and, consequently, destroy their worms,
+which would be an unnecessary sacrifice, and Nature has opposed it.
+Therefore, leaving the bottom rhomboidal, they are satisfied with
+raising a cylindrical tube around the worm, which, like the other cells
+in the comb, is horizontal. But this habitation remains suitable to the
+worm called to the royal state only during the first three days of its
+existence: another situation is requisite for the other two days it is a
+worm. Then, which is so small a portion of its life, it must inhabit a
+cell nearly of a pyramidal figure, and hanging perpendicularly; we may
+say the workers know it; for, after the worm has completed the third
+day, they prepare the place to be occupied by its new lodging. They gnaw
+away the cells surrounding the cylindrical tube, mercilessly sacrifice
+their worms, and use the wax in constructing a new pyramidal tube, which
+they solder at right angles to the first, and work it downwards. The
+diameter of this pyramid decreases insensibly from the base, which is
+very wide, to the point. During the two days that it is inhabited by
+the worm, a bee constantly keeps its head more or less inserted into the
+cell, and, when this worker quits it, another comes to occupy its place.
+In proportion as the worm grows, the bees labour in extending the cell,
+and bring food, which they place before its mouth, and around its body,
+forming a kind of cord around it. The worm, which can move only in a
+spiral direction, turns incessantly to take the food before its head: it
+insensibly descends, and at length arrives at the orifice of the cell.
+Now is the time of transformation to a nymph. As any farther care is
+unnecessary, the bees close the cell with a peculiar substance
+appropriated for it, and there the worm undergoes both its
+metamorphoses.
+
+Though M. Schirach supposes that none but worms three days old are
+selected for the royal treatment, I am certain of the contrary; and that
+the operation succeeds equally well on those of two days only. I must
+be permitted to relate at length the evidence I have of the fact, which
+will both demonstrate the reality of common worms being converted into
+queens, and the little influence which their age has on the effect of
+the operation.
+
+I put some pieces of comb, with some workers eggs, in the cells, and of
+the same kind as those already hatched, into a hive deprived of the
+queen. The same day several cells were enlarged by the bees, and
+converted into royal cells, and the worms supplied with a thick bed of
+jelly. Five were then removed from those cells, and five common worms,
+which, forty-eight hours before we had seen come from the egg
+substituted for them. The bees did not seem aware of the change; they
+watched over the new worms the same as over those chosen by themselves;
+they continued enlarging the cells, and closed them at the usual time.
+When they had hatched on them seven days{H}, we removed the cells to
+see the queens that were to 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 having elapsed, and no queen appearing, we
+opened them. In one, was a dead queen, but still a nymph; the other two
+were empty. The worms had spun their silk coccoons, but died before
+passing into their nymphine state, and presented only a dry skin. I can
+conceive nothing more conclusive than this experiment. It demonstrates
+that bees have the power of converting the worms of workers into queens;
+since they succeeded in procuring queens, by operating on the worms
+which we ourselves had selected. It is equally demonstrated, that the
+success of the operation does not depend on the worms being three days
+old, as those entrusted to the bees were only two. Nor is this all; bees
+can convert worms still younger into queens. The following experiment
+showed, that when the queen is lost, they destine worms only a few
+hours old to replace her.
+
+I was in possession of a hive, which being long deprived of the female,
+had neither egg nor worm. I provided a queen of the greatest fertility;
+and she immediately began laying in the cells of workers. I removed this
+female before being quite three days in the hive, and before any of her
+eggs were hatched. The following morning, that is, 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 depositing around them a much more abundant provision
+of food than is supplied to common worms. Next day, the worms were near
+forty hours old: the bees had enlarged and converted their hexagonal
+cells into cylindrical ones of the greatest capacity. During the
+subsequent days, they still laboured at them, and closed them on the
+fifth from the origin of the worms. Seven days after sealing of the
+first of these royal cells, a queen of the largest size proceeded from
+it. She immediately rushed towards the other royal cells, and
+endeavoured to destroy their nymphs and worms. In another letter, I
+shall recount the effects of her fury.
+
+From these details, you will observe, Sir, that M. Schirach's
+experiments had not been sufficiently diversified when he affirmed that
+it was essential for the conversion of common 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 existence.
+
+After my researches to corroborate M. Schirach's discovery, I was
+desirous of learning whether, as this observer conceives, the only means
+which the bees have of procuring a queen, is giving the common worms a
+certain kind of aliment, and rearing them in the largest cells. You
+will remember, that M. de Reaumur's sentiments are very different: "The
+mother should lay, and she does lay, eggs from which flies fit for being
+mothers must in their turn proceed. She does 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 peculiar interest in the eggs that one is to
+proceed from, and to consider them of the greatest value. They construct
+particular cells where they are to be deposited.--The figure of a royal
+cell only begun, very much resembles a cup, or, more correctly speaking,
+the cup that has lost its acorn."
+
+M. de Reaumur, though he did not suspect 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 M. Schirach, on the other hand,
+bees always having the power of procuring a queen by bringing up worms
+three days old in a particular manner, it would be needless for nature
+to grant females the faculty of laying royal eggs. Such prodigality is,
+in his eyes, inconsistent with the ordinary laws of nature. Therefore he
+maintains, in direct terms, that she 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 would always be
+too long for the belly of the mother to reach the bottom.
+
+I admit that M. de Reaumur no where says he has seen the queen lay in
+the royal cell. However he did not doubt the fact; and, after all my
+observations, I must esteem his opinion just. It is quite certain that,
+at particular periods of the year, the bees prepare royal cells; that
+the females deposit their eggs in them; and that worms, which shall
+became queens, proceed from these eggs.
+
+M. Schirach's objection, concerning the length of the cells, proves
+nothing; for the queen does not delay depositing her egg till they are
+finished. While only sketched and shaped like the cup of an acorn, she
+lays it. This naturalist, dazzled by the brilliancy of his discovery,
+saw only part of the truth. He was the first to find out the resource
+granted to bees by nature, for repairing the loss of their queen; and
+too soon persuaded himself that she had provided no other resource for
+the production of females. This error arose from not observing bees in
+very flat hives: had he used such as mine, he would have found, on
+opening them in spring, a confirmation of M. de Reaumur's opinion. Then,
+which is the season of swarming, hives in good condition are governed by
+a very fruitful queen: there are royal cells of a figure widely
+different from those constructed around the worms destined by the bees
+for queens. They are large, attached to the comb by a stalk, and
+hanging vertically like stalactites, such, in short, as M. de Reaumur
+has described them. The females lay in them before completion. 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 the egg has been
+laid. In proportion as the worm grows, they are enlarged, and closed by
+the bees when the first transformation approaches. Thus it is true,
+that, in spring, the queen deposits in royal cells, previously prepared,
+eggs from which flies of her own species are to come. Nature has,
+therefore, provided a double means for the multiplication and
+conservation of their race.
+
+  _PREGNY, 24. August 1791._
+
+
+FOOTNOTES:
+
+{H} The author's meaning here is obscure.--T.
+
+
+
+
+LETTER V.
+
+_EXPERIMENTS PROVING THAT THERE ARE SOMETIMES COMMON BEES WHICH LAY
+FERTILE EGGS._
+
+
+The singular discovery of M. Riems, concerning the existence of fertile
+workers, has appeared very doubtful to you, Sir. You have suspected that
+the eggs ascribed to workers by this naturalist had actually been
+produced by small queens, which, on account of their size, were
+confounded with common bees. But you do not positively insist that M.
+Riems is deceived; and, in the letter which you did me the honour to
+address to me, you requested me to investigate, by new experiments,
+whether there are actually working bees capable of laying fertile eggs.
+I have made these experiments with great care: and it is for you to
+judge of the confidence they merit.
+
+On the fifth of August 1788, we found the eggs and worms of large drones
+in two hives, which had both been some time deprived of queens. We also
+observed the rudiments of some royal cells appended like stalactites to
+the edges of the combs. The eggs of males were in them. Being perfectly
+secure that there was no queen of large size among the bees of these two
+hives, the eggs, which daily became more numerous, were evidently laid
+either by queens of small size or by fertile workers. I had reason to
+believe it was actually by common bees, for we had frequently observed
+them inserting the posterior part into the cells; and assuming the same
+attitude as the queen when laying. But, not withstanding every exertion,
+we had never been able to seize one in this situation, 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 observations were continued with equal assiduity, in hopes that, by
+some fortunate chance, or in a moment of address, we could secure one of
+them. More than a month all our endeavours were abortive.
+
+My assistant then offered to perform an operation that required both
+courage and patience, and which I could not resolve to suggest, though
+the same expedient had occurred to myself. He proposed to examine each
+bee in the hive separately, to discover whether some small queen had not
+insinuated herself among them, and escaped our first researches. This
+was an important experiment; for, should no small queen be found, it
+would be demonstrative evidence that the eggs had been laid by simple
+workers.
+
+To perform this operation with all possible exactness, immersing the
+bees was not enough. You know, Sir, that the contact of water stiffens
+their organs, that it produces a certain alteration of their external
+figure: and, from the resemblance of small queens to workers, the
+slightest alteration of shape would prevent us from distinguishing with
+sufficient accuracy to what species those immersed might belong.
+Therefore it was necessary to seize the whole bees of both hives,
+notwithstanding their irritation, and examine their specific character
+with the utmost 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, but what the relief of
+his eyes required. He took every bee in his hand; he attentively
+examined the trunk, the hind limbs, and the sting: there was not one
+without the characteristics of the common bee, that is, the little
+basket on the hind legs, the long trunk, and the straight sting. He had
+previously prepared glass cases containing combs. Into these, he put
+each bee after examination. It is superfluous to observe they were
+confined, which was a precaution indispensible until termination of the
+experiment. Neither was it enough to establish that the whole were
+workers; we had also to continue the experiment, and observe whether any
+would produce eggs. Thus we examined the cells for several days, and
+soon observed new laid eggs, from which the worms of drones came at the
+proper time. My assistant held in his hands the bees that produced them;
+and as he was perfectly certain they were common ones, it is proved that
+there are sometimes fertile workers in hives.
+
+Having ascertained M. Schirach's discovery, by so decisive an
+experiment, we replaced all the bees examined, in very thin glass hives,
+being only eighteen lines thick, and capable of containing but a single
+row of combs, and thus were extremely favourable to the observer. We
+thought, by strictly persisting to watch the bees, we might surprise a
+fertile one in the act of laying, seize and dissect her. This we were
+desirous of doing, for the purpose of comparing her ovaries with those
+of queens, and to ascertain the difference. At length, on the eighth of
+September, we had the good fortune to succeed.
+
+A bee appeared in the position of a female laying. Before she had time
+to leave the cell, we suddenly opened the hive and seized her. She
+presented all the external characteristics of common bees; the only
+difference we could recognise, and that was a very slight one, consisted
+in the belly seeming less and more slender than that of workers. On
+dissection, her ovaries were found more fragile, smaller and composed of
+fewer oviducts than the ovaries of queens. The filaments containing the
+eggs were extremely fine, and exhibited swellings at equal distances. We
+counted eleven eggs of sensible size, some of which appeared ripe for
+laying. This ovary was double like that of queens.
+
+On the ninth of September, we seized another fertile worker the instant
+she laid, and dissected her. The ovary was still less expanded than that
+of the preceding bee, and only four eggs had attained maturity. My
+assistant extracted one from the oviducts, and succeeded in fixing it by
+an end on a glass slider. We may take this opportunity of remarking,
+that it is in the oviducts themselves the eggs are imbued with the
+viscous liquid, with which they are produced, and not in passing through
+the spherical sac as Swammerdam believed. During the remainder of this
+month, we found ten fertile workers in the same hives, and dissected
+them all. In most, the ovaries were easily distinguished, but in some we
+could not discern the faintest traces of them. In these last, the
+oviducts to all appearance were but imperfectly developed, and more
+address than we had acquired in dissection was necessary to distinguish
+them.
+
+Fertile workers never lay the eggs of common bees; they produce none but
+those of males. M. Riems had already observed this singular fact; and
+here all my observations correspond with his. I shall only add to what
+he says, that fertile workers are not absolutely indifferent in the
+choice of cells for depositing their eggs. They always prefer large
+ones; and only use small cells when unable to find those of larger
+diameter. But they so far correspond with queens whose impregnation has
+been retarded, that they sometimes lay in royal cells.
+
+Speaking of females laying male eggs alone, I have already expressed my
+surprise that bees bestow, on those deposited in royal cells, such care
+and attention as to feed the worms proceeding from them, and, at the
+period of transformation, to close them up. But I know not, Sir, why I
+omitted to observe 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 fertile workers
+lay the eggs of drones is very different. They begin indeed with
+bestowing every care on their eggs and worms; they close the cells at a
+suitable time, but never fail to destroy them three days afterwards.
+
+Having finished these first experiments with success, I had still to
+discover the cause of the expansion of the sexual organs of fertile
+workers. M. Riems had not engaged in this interesting problem; and at
+first I dreaded that I should have no other guide towards its solution
+than conjecture. Yet from serious reflection, it appeared, that, by
+connecting the facts contained in this letter, there was some light that
+might elucidate my procedure in this new research.
+
+From M. Schirach's elegant discoveries, it is beyond all doubt that
+common bees are originally of the female sex. They have received from
+nature the germs of an ovary, but she has allowed 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 get 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 larvæ destined to replace her.
+Therefore, if fertile workers are produced in this situation alone, it
+is evident their origin is only in those hives where bees prepare the
+royal jelly. Towards this circumstance, I bent all my attention. It
+induced me to suspect that when bees give the _royal treatment_ to
+certain worms, they either by accident or a particular instinct, the
+principle of which is unknown to me, drop some particles of royal jelly
+into cells contiguous to those containing the worms destined for queens.
+The larvæ of workers that have accidentally received portions of so
+active an aliment, must be more or less affected by it; and their
+ovaries should acquire a degree of expansion. But this expansion will be
+imperfect; why? because the royal food has been administered only in
+small portions, and, besides, the larvæ having lived in cells of the
+smallest dimensions, their parts cannot extend beyond the ordinary
+proportions. Thus, the bees produced by them will resemble common
+workers in size and all the external characteristics. Added to that,
+they will have the faculty of laying some eggs, solely from the effect
+of the trifling portion of royal jelly mixed with their aliment.
+
+That we may judge of the justness of this explanation, it is necessary
+to consider fertile workers from their origin; to investigate whether
+the cells, where they are brought up, are constantly in the vicinity of
+the royal cells, and if their food is mixed with particles of the royal
+jelly. Unfortunately, the execution of these experiments is very
+difficult. When pure, the royal jelly is recognised by its sharp and
+pungent taste; but, when mixed with other substances, the peculiar
+savour is very 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{J}.
+
+In June 1790, I observed that one of my thinnest hives had wanted the
+queen several days, and that the bees had no mean 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 worms destined for
+queens, in 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
+destined for replacing the queens, it was very interesting to follow
+their history, and necessary to watch the moment of their last
+transformation. I removed the nineteen cells into a grated box, which
+was introduced 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 experiment. But here another
+precaution was also requisite. It was to be feared, that the bees being
+deprived of the produce of their labour, and the object of their hope
+might be totally discouraged; therefore, I supplied them with another
+piece of comb, containing the brood of workers, reserving power to
+destroy the young brood when necessary. This plan succeeded admirably.
+The bees, in bestowing all their attention on these last worms, forgot
+those that had been removed.
+
+When the moment of transformation of the nymphs in the nineteen cells
+arrived, I examined the grated box frequently every day, and at length
+found six bees exactly similar to _common bees_. The worms of the
+remaining thirteen had perished without changing.
+
+The portion of brood comb that had been put into the hive to prevent the
+discouragement of the bees was then removed. I put 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 six
+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 produced by some of the six bees?
+But, to attain absolute certainty, it was necessary to take them in the
+act of laying. Some ineffaceable mark was also required for
+distinguishing them in particular.
+
+This proceeding was attended with the most ample success. We 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
+perceived one introducing the posterior part into 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 privation of the right antenna, instantly
+recognised that it was one of the six that had passed to the vermicular
+state in the vicinity of the royal cells.
+
+I could no longer doubt the truth of my conjecture; at the same time, I
+know not whether the truth will appear as rigorous to you, Sir, as it
+does to myself. But I reason in the following manner: If it is certain
+that fertile workers are always produced in the vicinity of royal cells,
+it is no less true, that in itself, the vicinity is indifferent; for the
+size and figure of these cells can produce no effect on the worms in
+those surrounding them; there must be something more; we 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 the worms in the
+adjacent cells have had a portion of the same food. This is what they
+gain, therefore, by vicinity to the royal cells. The bees, in their
+course thither, will pass in numbers over them, stop and drop some
+portion of the jelly destined for the royal larvæ. This reasoning, I
+presume, is consistent with the principles of sound logic.
+
+I have repeated the experiment now described so often, and weighed all
+the concomitant circumstances with so much care, that whenever I please,
+I can produce fertile workers in my hives. The method is simple. 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
+supplying the included worms with the royal jelly. Portions of this
+aliment also fall on the young larvæ deposited in the adjacent cells,
+and it unfolds the ovaries to a certain degree. 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. Therefore, to
+save them, all their enemies must be removed, and the larvæ of the royal
+cells taken away before undergoing their last metamorphoses. Then the
+fertile workers, being without rivals at the time of their origin, will
+be well received, and, by taking the precaution to mark them, it will be
+seen, in a few days, that they produce the eggs of males. Thus, the
+whole secret of this proceeding consists in removing the royal cells at
+the proper time; that is, after being sealed, and previous to the young
+queens leaving them{K}.
+
+I shall add but a few words to this long letter. There is nothing so
+very surprising in the production of fertile workers, when we consider
+the consequences of M. Shirach's beautiful discovery. But why do they
+lay male eggs only? I can conceive, indeed, that the reason of their
+laying few is from their ovaries being but imperfectly expanded, but I
+can form no idea why all the eggs should be those of males, neither can
+I any better account for their use in hives; and hitherto, I have made
+no experiments on their mode of fecundation.
+
+  _PREGNY, 25. August 1791._
+
+
+FOOTNOTES:
+
+{I} It is difficult to discover whether the author thinks, as some
+naturalists, that bees are instrumental in hatching the eggs.--T.
+
+{J} The original is extremely confused in the preceding passages.--T.
+
+{K} I have frequently seen queens, at the moment of production, begin
+first by attacking the royal cells and then the common ones beside them.
+As I had not seen fertile workers when I first observed this fact, I
+could not conceive from what motive the fury of the queen was thus
+directed towards the common cells. But now I know they can distinguish
+the species included, and have the same instinctive jealousy or aversion
+towards them as against the nymphs of queens properly so denominated.
+
+
+
+
+LETTER VI.
+
+_ON THE COMBATS OF QUEENS: THE MASSACRE OF THE MALES: AND WHAT SUCCEEDS
+IN A HIVE WHERE A STRANGER QUEEN IS SUBSTITUTED FOR THE NATURAL ONE._
+
+
+M. de Reaumur had not witnessed every thing relative to bees when he
+composed his history of these industrious animals. Several observers,
+and those of Lusaçe in particular, have discovered many important facts
+that escaped him; and I, in my turn, have made various observations of
+which he had no suspicion: at the same time, and this is a very
+remarkable circumstance, not only has all that he expressly declares he
+saw been verified by succeeding naturalists, but all his conjectures are
+found just. The German naturalists, Schirach, Hattorf, and Riems
+sometimes contradict him, indeed, in their memoirs; but I can maintain
+that, while combating the opinion of M. de Reaumur, it is they who are
+almost always wrong; of which several instances might be adduced.
+
+What I shall now proceed to say will give me an opportunity of detailing
+some interesting facts.
+
+It was observed by M. de Reaumur, that when any supernumerary queen is
+either produced in a hive, or comes into it, one of the two soon
+perishes. He has not actually witnessed 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. M. Schirach, on the other
+hand, and, after him, M. Riems, thinks that the working bees assail the
+stranger, and sting her to death. I cannot comprehend by what means they
+have been able to make this observation: as they used very thick hives
+only, with several rows of combs, they could at most but observe the
+commencement of hostilities. While the combat lasts, the bees move with
+great rapidity; they fly on all sides; and, gliding between 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 queens and
+workers, but I have very often beheld one between the queens themselves.
+
+In one of my hives in particular, there were five or six royal cells,
+each including a nymph. The eldest first underwent its transformation.
+Scarcely did ten minutes elapse from the time of this young queen
+leaving her cradle, when she visited the other royal cells still close.
+She furiously attacked the nearest; and, by dint of labour, succeeded
+in opening the top: we saw her tearing the silk of the coccoon with her
+teeth; but her efforts were probably inadequate to the object, for she
+abandoned this end of the cell, and began at the other, where she
+accomplished a larger aperture. When it was sufficiently enlarged, she
+endeavoured to introduce her belly, and made many exertions until she
+succeeded in giving her rival a deadly wound with her sting. Then having
+left the cell, all the bees that had hitherto been spectators of her
+labour, began to increase the opening, and drew out the dead body of a
+queen scarcely come from its envelope of a nymph.
+
+Meanwhile, the victorious young queen attacked another royal cell, but
+did not endeavour to introduce her extremity into it. There was only a
+royal nymph, and no queen, come to maturity, as in the first cell. In
+all probability, nymphs of queens inspire their rivals with less
+animosity; still they do not escape destruction: because, whenever a
+royal cell has been opened before the proper time, the bees extract the
+contents in whatever form they may be, whether worm, nymph, or queen.
+Lastly, the young queen attacked the third cell, but could not succeed
+in penetrating it. She laboured languidly, and appeared as if exhausted
+by her first exertions. As we now required queens for some particular
+experiments, we resolved to remove the other royal cells, yet in safety,
+to secure them from her fury.
+
+After this observation, we wished to see what ensued on two queens
+leaving their cells at the same time, and in what manner one perished. I
+find an observation on this head in my Journal, 15. May 1790.
+
+In one of our thinnest hives, two queens left their cells almost at the
+same moment. Whenever they observed each other, they rushed together,
+apparently with great fury, and were in such a position that the antennæ
+of each was seized by the teeth of the other: the head, breast, and
+belly of the one were exposed to the head, breast, and belly of the
+other: the extremity of their bodies were curved; they were reciprocally
+pierced with the stings; and both fell dead at the same instant. But it
+seems as if nature has not ordained that both combatants should perish
+in the duel; but rather that, when finding themselves in the situation
+described, namely, opposite, and belly to belly, they fly at that moment
+with the utmost precipitation. Thus, when these two rivals felt the
+extremities about to meet, they disengaged 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 think that we may here
+penetrate the intention of nature.
+
+There ought to be none but one queen in a hive: therefore it is
+necessary, if by chance a second is either produced or comes into the
+hive, that one of the two must be destroyed. This cannot be committed to
+the working bees, because, in a republic composed of so many
+individuals, an equal consent cannot be supposed always to exist; it
+might frequently happen that one group of bees destroyed one of the
+queens, while a second would massacre the other; and the hive thus be
+deprived of queens. Therefore it was necessary that the queens
+themselves should be entrusted with the destruction of their rivals: but
+as, in these combats, nature demands but 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 trifling facts. But here the object and the means
+seem so plain, that I have ventured to advance my conjectures. You will
+judge better than I can, whether they are well founded.--Let me now
+return from this digression.
+
+A few minutes after the two queens separated, their terror ceased, and
+they again began to seek each other. Immediately on coming in sight,
+they rushed together, seized one another, and resumed exactly their
+former position. The result of this encounter was the same. When their
+bellies approached, they hastily disengaged themselves, and fled with
+precipitation. During all this time, the workers seemed in great
+agitation; and the tumult appeared to increase when the adversaries
+separated. Two different times, we observed them stop the flight of the
+queens, seize their limbs, and retain them prisoners above a minute. At
+last, the queen, which was either the strongest 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, brought the
+extremity of her own body under the belly of the other; and, by this
+means, easily pierced her with the sting. Then she withdrew her sting
+after losing hold of the wing. The vanquished 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 fecundated, and mothers, had the
+same animosity.
+
+On the 22. of July, we selected a flat hive, containing a very fertile
+queen: and being curious to learn whether, as virgin queens, she would
+destroy the royal cells, three were introduced into the middle of the
+comb. Whenever she observed this, _she_ sprung forward on the whole, and
+pierced them towards the bottom; nor did she desist until the included
+nymphs were exposed. The workers which had hitherto been spectators of
+this destruction, now came to carry the nymphs away. They greedily
+devoured the food remaining at the bottom of the cells, and also sucked
+the fluid from the abdomen of the nymphs: and then terminated with
+destroying the cells from which they had been drawn.
+
+In the next place, we introduced a very fertile queen into this hive;
+after painting the thorax to distinguish her from the reigning queen. A
+circle of bees quickly formed around the stranger, but their intention
+was not to caress and receive her well; for they insensibly accumulated
+so much, and surrounded her so closely, that in scarcely a minute she
+lost her liberty and became a prisoner. It is a remarkable circumstance,
+that other workers at the same time collected round the reigning queen
+and restrained all her motions; we instantly saw her confined like the
+stranger. Perhaps it may be said, 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 way to allow her full liberty for the attack; then if the queens
+testified a disposition to fly, they returned to enclose them.
+
+We have repeatedly witnessed this fact, but it presents so new and
+singular a characteristic in the policy of bees, that it must be seen
+again a thousand times before any positive assertion can be made on the
+subject. I would therefore recommend that naturalists should attentively
+examine the combat of queens, and particularly ascertain what part is
+taken by the workers. Is their object to accelerate the combat? Do they
+by any secret means excite the fury of the combatants? Whence does it
+happen that accustomed to bestow every care on their queen, in certain
+circumstances, they oppose her preparations to avoid impending danger?
+
+A long series of observations are necessary to solve these problems. It
+is an immense field for experiment, which will afford infinitely curious
+results. I intreat you to pardon my frequent digressions. The subject is
+deeply philosophical, genius such as your's is required to treat it
+properly; and I shall now be satisfied with proceeding in the
+description of the combat.
+
+The cluster of bees that surrounded the reigning queen having allowed
+her some freedom, she seemed to advance towards that part of the comb
+where her rival stood; then, all the bees receded before her, the
+multitude of workers, separating the two adversaries, gradually
+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 combination, we had still to examine
+whether a combat would ensue between two queens, one impregnated, and
+the other a virgin; and what circumstances attended it.
+
+On the 18. of September, we introduced a very fruitful queen into a
+glass hive, already containing a virgin queen, and put her on the
+opposite side of the comb, that we might have time to see how the
+workers would receive her. She was immediately surrounded, but they
+confined her only a moment. Being oppressed with the necessity of
+laying, she dropped some eggs; however, 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 advanced towards the edge of the
+comb, and soon approached very near the virgin queen. When in sight,
+they rushed together; the virgin queen got on the back of the other, and
+gave her several stings in the belly, but, having aimed at the scaly
+part, they did not injure her, and the combatants separated. In a few
+minutes, they returned to the charge; but this time the impregnated
+queen mounted on her rival; however, she sought in vain to pierce her,
+for the sting did not enter; the virgin queen then disengaged herself
+and fled; she also succeeded in escaping another attack, where her
+adversary 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 successful exertion, the virgin queen
+mortally wounded the stranger, and she expired in a moment. The sting
+had penetrated so far that the victorious queen was unable to extract
+it, and she was overthrown by the fall of her enemy. She made great
+exertions to disengage the sting: but could succeed by no other means
+than turning on the extremity of the belly, as on a pivot. Probably the
+barbs of the sting fell by this motion, and, closing like a spiral
+around the stem, came more easily from the wound.
+
+These observations, Sir, I think will satisfy you, respecting the
+conjecture of our celebrated 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 will
+at no time attempt to employ their stings against the stranger queen. I
+can conceive what has misled M. Riems and Schirach; but it is necessary
+for 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 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 supernumerary queens; and if an observer wishes 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 a natural state,
+for the following 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
+antennæ every individual endeavouring to penetrate the hive, and also
+the various substances put within their reach; which affords us an
+opportunity of observing that the antennæ are certainly the organs of
+feeling. If a stranger queen appears, she is instantly seized by the
+bees on guard, which prevent her entry by laying hold of her legs or
+wings with their teeth, and crowd so closely around her, that she cannot
+move. Other bees, from the interior of the hive, gradually come to their
+assistance, and confine her still more narrowly, all having their heads
+towards the centre where the queen is inclosed; and they remain with
+such evident anxiety, eagerness, and attention, that the cluster they
+form may be carried about for some time, without their being sensible
+of it. A stranger queen, so closely confined and hemmed in, cannot
+possibly penetrate the hive. If the bees retain her too long imprisoned,
+she perishes. Her death probably ensues from hunger, or the privation of
+air; it is undoubted, at least, that she is never stung. We never saw
+the bees direct their stings against her, except a single time, and then
+it was owing to ourselves. We endeavoured, from compassion for a queen's
+situation, to remove her from the center 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 several of the workers were themselves killed; and
+surely they could not intend destroying one another. Had we not
+interfered, they would have been content with confining the queen, and
+would not have massacred her.
+
+It was in similar circumstances that M. Riems saw the workers anxiously
+pursue a queen. He thought they designed to sting her, and thence
+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 II, chap. 27, note 7_. But you are sensible, Sir, from
+these details, that he has been mistaken. He did not know the attention
+that bees bestow on what passes at the entrance of their hive, and he
+was entirely ignorant of the means they take to prevent supernumerary
+queens from penetrating it.
+
+       *       *       *       *       *
+
+After ascertaining that the workers in no situation sting the
+supernumerary queens, we were curious to learn how a stranger queen
+would be received in a hive wanting a 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.
+
+Bees do not immediately observe the removal of their queen; their
+labours are uninterrupted; they watch over the young, and perform all
+their ordinary occupations. 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 a
+delirious impetuosity. Then they discover their queen is no longer among
+them. But how do they become sensible of it? How do the bees on the
+surface of the comb discover that the queen is not on the next comb? In
+treating of another characteristic of these animals, you have yourself,
+Sir, proposed the same question; I am incapable of answering it indeed,
+but I have collected some facts, that may perhaps facilitate the
+elucidation of this 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
+interpret this expression strictly. Substitution of another queen is not
+attended with the same effect, if she is introduced into the hive within
+the first twelve hours after removal of the reigning one. Here the
+agitation continues; and the bees treat the stranger the same as when
+the presence of their own leaves them nothing to desire. They surround,
+seize, and keep her captive, a very long time, in an impenetrable
+cluster; and she commonly dies either from hunger or privation of air.
+
+If eighteen hours elapse before substitution of a stranger queen for the
+native one removed, she is at first treated 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 moves
+languidly; and sometimes 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 where they had originally been ill
+received.
+
+If, before substituting the stranger queen, twenty-four hours elapse,
+she 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 word reception is very
+indefinite, it is proper to enter into some detail for explaining the
+exact sense in which I use it. On the 15. of August, I introduced a
+fertile queen, eleven months old, into a glass hive. The bees were
+twenty-four hours deprived of their queen, and had already begun the
+construction of twelve royal-cells, such as described in the preceding
+chapter. Immediately on placing this female stranger on the comb, the
+workers near her touched her with their antennæ, and, passing their
+trunks over every part of her body, they gave her honey. Then these gave
+place to others that treated her exactly in the same manner. All
+vibrated their wings at once, and ranged themselves in a circle around
+their sovereign. Hence resulted a kind of agitation which gradually
+communicated to the workers situated on the same surface of the comb,
+and induced them to come and reconnoitre, in their turn, what was going
+on. They soon arrived; and, having broke through the circle formed by
+the first, approached the queen, touched her with the antennæ, and gave
+her honey. After this little ceremony they retired; and, placing
+themselves behind the others, enlarged the circle. There they vibrated
+their wings, and buzzed without tumult or disorder, and as if
+experiencing some very agreeable sensation. The queen had not yet moved
+from the place where I had put her, but in a quarter of an hour she
+began to move. The bees, far from opposing her, opened the circle at
+that part to which she turned, followed her, and formed a guard around.
+She was oppressed with the necessity of laying, and dropped eggs.
+Finally, after four hours abode, she began to deposit male eggs in the
+cells she met.
+
+While these events passed on the surface of the comb where the queen
+stood, all was quiet on the other side. Here the workers were apparently
+ignorant of a queen's arrival in the hive. They laboured with great
+activity at the royal cells, as if ignorant that they no longer stood in
+need of them: they watched over the royal worms, supplied them with
+jelly and the like. But the queen having at length come to this side,
+she was received with the same respect that she had experienced from
+their companions on the other side of the comb. They encompassed her;
+gave her honey; and touched her with their antennæ: and what proved
+better that they treated her as a mother, was their immediately
+desisting from 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 that bees receive
+a stranger queen; when they have time to forget their own, she is
+treated exactly as if she was their natural one, except that there is
+perhaps at first greater interest testified in her, or more conspicuous
+demonstrations of it. I am sensible of the impropriety of these
+expressions, but M. de Reaumur in some respect authorises them. He does
+not scruple to say, that bees pay _attention_, _homage_, and _respect_,
+to their queen, and from his example the like expressions have escaped
+most authors that 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 full of combats and disastrous
+scenes, I should, perhaps, give you an account of some more pleasing
+and interesting facts relative to their industry. However, to avoid
+returning to duels and massacres, I shall here subjoin my observations
+on the massacre of the males.
+
+You will remember, Sir, it is agreed by all observers, that at a certain
+period of the year, the workers kill and expel the drones. M. de Reaumur
+speaks of these executions as a horrible massacre. He does not expressly
+affirm, indeed, that he has himself witnessed it, but what we have seen
+corresponds so well with his account, that there can be no doubt he has
+beheld the peculiarities of the massacre.
+
+It is usually in the months of July and August, that the bees free
+themselves of the males. Then they are drove away and pursued to the
+inmost parts of the hive, where they collect in numbers; and as at the
+same time we find many dead drones on the ground before the hives, it is
+indubitable that after being expelled, the bees sting them to death.
+Yet on the surface of the comb, we do not see the sting used against
+them; there the bees are content to pursue and drive them away. You
+observe this, Sir, yourself, in the new notes added to _la Contemplation
+de la Nature_; and you seem disposed to think, that the drones forced to
+retire to the extremity of the hive, perish from hunger. Your conjecture
+was extremely probable. Still it was possible the carnage might take
+place in the bottom of the hive, and had been unobserved, because that
+part is dark, and escapes the observer's eye.
+
+To appreciate the justice of this suspicion, we thought of making the
+support of the hive of glass, and of placing ourselves below to see what
+passed in the scene of action. Therefore, a glass table was constructed,
+on which were put six hives with swarms of the same year; and, lying
+under it, we endeavoured to discover how the drones were destroyed. The
+invention succeeded to admiration. On the 4 of July, we saw the workers
+actually massacre the males, in the whole six swarms, at the same hour,
+and with the same peculiarities.
+
+The glass table was covered with bees full of animation, which flew upon
+the drones, as they came from the bottom of the hive; seized them by the
+antennæ, the limbs, and the wings, and after having dragged them about,
+or, so to speak, after quartering them, they killed them by repeated
+stings directed between the rings of the belly. The moment that this
+formidable weapon reached them, was the last of their existence; they
+stretched 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 stuck
+the sting so deep, that it could hardly be withdrawn, and these bees
+were obliged to turn upon themselves before the stings could be
+disengaged.
+
+Next day, having resumed our former position, we witnessed new scenes of
+carnage. During three hours, the bees furiously destroyed the males.
+They had massacred all their own on the preceding evening, but now
+attacked those which, driven from the neighbouring hives, had taken
+refuge amongst them. We saw them also tear some remaining nymphs from
+the combs; they greedily sucked all the fluid from the abdomen, and then
+carried them away. The following days no drones remained in the hives.
+
+These two observations seem to me decisive. It is incontestible that
+nature has charged the workers with the destruction of the males at
+certain seasons of the year. But what means does she use to excite their
+fury against them? This is a question that I cannot pretend to answer.
+However, an observation I have made may one day lead to solution of the
+problem. The males are never destroyed in hives deprived of queens, on
+the contrary, while a savage massacre prevails in other places, they
+there find an asylum. They are tolerated and fed, and many are seen even
+in the middle of January. They are also preserved in 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 but in hives where the queens are completely
+fertile, and it never begins until the season of swarming is past.
+
+  _PREGNY, 28 August 1791._
+
+
+
+
+LETTER VII.
+
+_SEQUEL OF EXPERIMENTS ON THE RECEPTION 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 a sensible pleasure in acknowledging that
+if I have made any progress in the art of observation, I am indebted for
+it to profound study of the works of this naturalist. In general his
+authority has such weight, that I can scarcely trust my own experiments
+when the results are different from his. Likewise, on finding myself in
+opposition to the _historian of bees_, I repeat my experiments. I vary
+the mode of conducting them; I examine with the utmost caution all the
+circumstances that might mislead me, and never are my labours
+interrupted before acquiring the moral certainty of avoiding error. With
+the aid of these precautions, I have discovered the justice of M. de
+Reaumur's suggestions, and I have a thousand times seen, if certain
+experiments seemed to combat them, it was from incorrectness of
+execution. Yet I must except some facts where my results have constantly
+been different from his. Those respecting the reception of a stranger
+queen substituted for the natural one, are of the number.
+
+If, after removing the natural queen, a stranger is immediately
+substituted, the usurper is ill received. I never could succeed in
+making them adopt her, but by allowing an interval of twenty or
+twenty-four hours to elapse. Then they seemed to have forgot their own
+queen; and respectfully received any female put in her place. M. de
+Reaumur, on the contrary, asserts, that should the original queen be
+removed, and another presented, this new one will be perfectly well
+received from the beginning. As evidence of this assertion, he gives the
+detail of an experiment which must be read in his work, for I shall here
+give only an extract of it{L}. 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 this moment, the
+tumult ceased, and the stranger queen was received with all respect.
+
+I do not dispute the truth of this experiment; but, in my opinion, it
+does not warrant the conclusion that M. de Reaumur deduces from it. His
+apparatus removed the bees too much from their natural condition, to
+allow him to judge of their instinct and dispositions. In other
+situations, he has himself observed, that these animals, reduced to
+small numbers, lost their industry and activity, and feebly continued
+their ordinary labours. Thus their instinct is affected by every
+operation that too much diminishes their number. To render such an
+experiment truly conclusive, it must be made in a populous hive; and on
+removing the native queen, a stranger must immediately be substituted in
+her place. Had this been done, I am fully persuaded, that M. de Reaumur
+would have seen the bees imprison the usurper, confine her at least
+twelve or fifteen hours among them, and frequently suffocate her: nor
+would he have witnessed any favourable reception before an interval of
+twenty-four hours after removal of the original queen. No variation has
+occurred in my experiments regarding this fact. Their number, and the
+attention bestowed on them, make me presume they merit your confidence.
+
+M. de Reaumur, in another passage of the same Memoir, affirms, that
+_bees, which have a queen they are satisfied with, are nevertheless
+disposed to give the best possible reception to any female that seeks
+refuge among them_. In the preceding letter, I have related my
+experiments on this head: their success has been very different from
+that of M. de Reaumur's. I have proved that the workers never employ
+their stings against the queen; but this cannot be called the welcome
+reception of a stranger. They retain her within their ranks, and seem to
+allow her liberty only when she prepares to combat the reigning queen.
+This observation cannot be made except in the thinnest hives. Those used
+by M. de Reaumur had always two parallel combs at least, which must
+have prevented him from observing some very important circumstances that
+influence the conduct of workers when supplied with several females. The
+first circles formed around a stranger queen he has taken for caresses;
+and, from the little that this queen could advance between the combs, it
+must have been impossible for him to observe that the circles, which
+always continued contracting, ended in restraint of the females there
+inclosed. Had he used thinner hives, he would have discovered that what
+he supposed indication of a favourable reception was the prelude of
+actual imprisonment.
+
+I feel reluctant to assert that M. de Reaumur was deceived. Yet I cannot
+admit that, on certain occasions, bees tolerate a plurality of females
+in their hives. The experiment on which this affirmation rests will not
+be considered decisive. In the month of December, he introduced a
+stranger queen into a glass hive, in his cabinet, and confined her
+there. The bees had no opportunity of going out. This stranger was well
+received; her presence awakened the workers from their lethargic state,
+into which they did not relapse; she excited no carnage; 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 queen was living when the
+new one was introduced into the hive: however the author neglected this;
+and it is very probable 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 observations coincide with his, and still
+more, when my experiments justify his conjectures. But I think it
+proper to point out those cases where the imperfections of his hives
+have led him into error, and to explain from what causes I have not seen
+certain facts in the same manner he did. I feel particular anxiety to
+merit your confidence, and I am aware that the greatest exertions are
+necessary, when I have to combat the historian of bees. I confide in
+your judgment; and pray you to be assured of my respect.
+
+  _PREGNY, 30. August 1791._
+
+
+FOOTNOTES:
+
+{L} Edit. 4to, Tom. V. p. 258.
+
+
+
+
+LETTER 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 COCCOONS._
+
+
+In this letter I shall collect some isolated observations relative to
+various points in the history of bees, concerning which you wished me to
+engage.
+
+You desired me to investigate whether the queen is really _oviparous_.
+M. de Reaumur leaves this question undecided. He observes, that he has
+never seen the worm hatched; and he only asserts that worms are found in
+those cells where eggs have been deposited three days preceding. If we
+attempt to catch the moment when the worm leaves the egg, we must extend
+our observations beyond the interior of the hive; for there the
+continual motion of the bees obscures what passes at the bottom of
+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 to hatch the worms, should the eggs
+be too soon deprived of it they wither and perish. The sole method of
+succeeding in seeing the worm come out, consists in watching the queen
+while she lays, in marking the egg so as to be recognised, and removing
+it from the hive to the microscope only an hour or two before the three
+days elapse. The worm will certainly be hatched, provided it has been
+exposed as long as possible to the full degree of heat. Such is the
+course I have pursued; and the following are 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 the cell, and put
+the pyramidal bottom, where the egg was fixed, on a glass slider. Slight
+motions were soon perceptible in the eggs. At first, we could observe no
+external organization: the worm was entirely concealed from us by its
+pellicle. We then prepared to examine the egg with a powerful magnifier;
+however, during the interval, the worm burst its surrounding membrane,
+and cast off part of the envelope, which was torn and ragged on
+different parts of the body, and more evidently so towards the last
+rings. The worm alternately curved and stretched itself, with very
+lively action. Twenty minutes were occupied in casting off the spoil;
+when this exertion ceased: the worm lay down, curved, and seemed to take
+that rest which it required. An egg laid in a worker's cell produced
+this animal, which would have become a worker itself.
+
+We next directed our attention to the moment when a male worm would be
+hatched. An egg was exposed to the sun on a glass slider; and, with a
+good magnifier, nine rings of the worm were perceptible within the
+transparent pellicle. This membrane was still entire, and the worm
+perfectly motionless. The two longitudinal lines of tracheæ were visible
+on the surface, and many ramifications. We never lost sight of the egg a
+single instant, and now succeeded in observing the first motions of the
+worm. The thick end alternately straightened and curved, and almost
+reached the part where the sharp extremity was fixed. These exertions
+burst the membrane, first on the upper part, towards the head, then on
+the back, and afterwards on all the rest successively. The ragged
+pellicle remained in folds on different parts of the body, and then fell
+off. Thus it is beyond dispute, that the queen is oviparous.
+
+Some observers affirm, that the workers attend to the eggs before the
+worms are hatched; and it is certain that, at whatever time a hive is
+examined, we always see some workers with the head and thorax inserted
+into cells containing eggs, and remaining motionless several minutes in
+this position. It is impossible to discover what they do, for the
+interior of the cell is concealed by their 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 in a strange hive, where there is the necessary degree of
+heat, the worms come out at the usual time, just as if they had been
+left in the cells. Thus no extraordinary aid or attention is required
+for their exclusion.
+
+When the workers penetrate the cells, and remain fifteen or twenty
+minutes motionless, I have reason to believe, it is only to repose from
+their labours. My observations on the subject seem correct. You know,
+Sir, that a kind of irregular shaped cells, are frequently constructed
+on the panes of the hive. These, being glass on one side, are
+exceedingly convenient to the observer, since all that passes within is
+exposed. I have often seen bees enter these cells when nothing could
+attract them. The cells contained neither eggs nor honey, nor did they
+need further completion. Therefore the workers repaired thither only to
+enjoy some moments of repose. Indeed, they were fifteen or twenty
+minutes so perfectly motionless, that had not the dilatation of the
+rings shewed their respiration, we might have concluded them dead. The
+queen also sometimes penetrates the large cells of the males, and
+continues very long motionless in them. Her position prevents the bees
+from paying their full homage to her, yet even then the workers do not
+fail to form a circle around her, and brush the part of her belly that
+remains exposed.
+
+The drones do not enter the cells while reposing, but cluster together
+on the combs; and sometimes retain this position eighteen or twenty
+hours without the slightest motion.
+
+As it is important, in many experiments, to know the exact time that the
+three species of bees exist before assuming their ultimate form, I shall
+here subjoin my own observations 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. The worm
+now begins spinning its coccoon, in which operation thirty-six hours are
+consumed. In three days, it changes to a nymph, and passes six days in
+this form. It is only on the twentieth day of its existence, counting
+from the moment the egg is laid, that it attains the fly state.
+
+The royal worm also passes three days in the egg, and is five a worm;
+the bees then close its cell; and it immediately begins spinning the
+coccoon, which occupies twenty-four hours. The tenth and eleventh day it
+remains in complete repose, and even sixteen hours of the twelfth. Then
+the transformation to a nymph takes place, in which state four days and
+a third are passed. Thus it is not before the sixteenth day that the
+perfect state of queen is attained.
+
+The male worm passes three days in the egg, six and a half as a worm,
+and metamorphoses into a fly on the twenty-fourth day after the egg is
+laid.
+
+Though the larvæ of bees are apodal, they are not condemned to absolute
+immobility in their cells; for they can move by a spiral motion. During
+the first three days, this motion is so slow as scarcely to be
+perceptible, but it afterwards becomes more evident. I have then
+observed them perform two complete revolutions in an hour and three
+quarters. When the period of transformation arrives, they are only two
+lines from the orifice of the cells. As their position is constantly the
+same, bent in an arc, those in the workers' and drones' cells are
+perpendicular to the horizon, while those in the royal cells lie
+horizontally. It might be thought, that the difference of position has
+much influence on the increment of the different larvæ; yet it has none.
+By reversing combs containing common cells full of brood, I have put the
+worms in a horizontal position; but they were not injured. I have also
+turned the royal cells, so that the worms came into a horizontal
+direction; however their increment was neither slower nor less perfect.
+
+       *       *       *       *       *
+
+I have been much surprised at the mode of bees spinning their coccoons,
+and 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. The royal larvæ,
+on the other hand, spin imperfect coccoons, open behind, and enveloping
+only the head, thorax, and first ring of the abdomen. The discovery of
+this difference, which at first may seem trifling, has given me extreme
+pleasure, for it evidently demonstrates the admirable art with which
+nature connects the various characteristics in the industry of bees.
+
+You will remember, Sir, the evidence I gave you of the mutual aversion
+of queens, of the combats in which they engage, 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 a complete coccoon, she 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 retract it, would
+become the victim of her own fury. Thus, that the queen might destroy
+her rivals, it was necessary the last rings of the body should remain
+uncovered, therefore the royal nymphs must only form imperfect 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 in her care
+of preserving and multiplying the species. But from the facts I relate,
+we must admire her precautions in exposing certain individuals to a
+mortal danger.
+
+The detail on which I have just entered clearly indicates the final
+cause of the opening left by the royal worms in their coccoons; but it
+does not shew whether it is in consequence of a particular instinct that
+they leave this opening, or whether the wideness of their cells prevents
+them from stretching the thread up to the top. This question interested
+me very much; the only method of deciding it was to observe the worms
+while spinning, which cannot be done in their opaque cells. 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 operation
+consisted in extracting worms and introducing them here; but my
+assistant accomplished it with much address. He opened several sealed
+royal cells, where we knew the larvæ were about to begin their coccoons,
+and, taking them gently out, introduced one into each of my glass cells
+without the smallest injury.
+
+They soon prepared to work; and commenced by stretching the anterior
+part of the body in a straight line, while the other was bent in a
+curve. This formed a curve of which the longitudinal sides of the cells
+were tangents, and afforded two points of support. The head was next
+conducted to the different parts of the cell which it could reach, and
+it carpeted the surface with a thick bed of silk. We remarked that the
+threads were not carried from one side to another, and that this would
+have been impracticable, for the worms being obliged to support
+themselves, and to keep the posterior rings curved, the free and
+moveable part of the body was not long enough for the mouth to reach the
+sides diametrically opposite, and fix the threads to them. You will
+remember, Sir, that the royal cells are of a pyramidal form, with a wide
+base, and a long contracted top. These cells hang perpendicularly in the
+hive, the point downwards, from which position the royal worm can be
+supported in the cell, only when the curvature of the posterior part
+forms two points of support; and that it cannot obtain this support
+without resting on the lower part, or towards the extremity. Therefore
+if it attempted to stretch out and spin towards the wide end of the
+cell, it could not reach both sides from being too distant. One part
+would be touched by its extremity, the other by its back, and it would
+consequently 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
+support themselves.
+
+These first experiments obviated the suspicion of any particular
+instinct in the royal worms. They proved, if the worms spun incomplete
+coccoons, it was because they were forced to do so by the figure of
+their cells. However, I wished to have evidence still more direct. I put
+them into cylindrical glass cells, or portions of glass tubes resembling
+common cells, and I had the satisfaction of seeing them spin complete
+coccoons, as the worms of workers do. Lastly, I put common worms in very
+wide cells, and they left the coccoon open. Thus it is demonstrated,
+that the royal worms, and those of workers, have the same instinct and
+the same industry, or in other words, when situated in the same
+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 coccoons, undergo all their metamorphoses equally well. Thus
+the necessity imposed on them by nature, of having the coccoons open, is
+not necessary for their increment; nor has it any other object than that
+of exposing them to the certainty of perishing by the wounds of their
+natural enemy; an observation new and truly singular.
+
+       *       *       *       *       *
+
+I ought to 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 find 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, influence their size. With this view, you have
+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 large cells produced workers of
+larger size, we were bound to conclude that the size of the cells had a
+sensible influence on the size of the bees. The first time I made this
+experiment, it did not succeed, because weevils lodged in the hive
+discouraged the bees. But I repeated it afterwards, and the result was
+very remarkable.
+
+I removed the whole comb, consisting of common cells, from one of my
+best glass hives, and left that composed of males' cells alone: and to
+avoid vacuities, I supplied others of the same kind. This was in June,
+the season most favourable to bees. I expected that the bees would
+quickly have repaired the ravages produced by this operation 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 they would resume their activity, I continued
+observing them several days; however, my hopes were disappointed. Their
+homage to the queen was not interrupted indeed; but except in this,
+their conduct to the queen was quite different from what it usually is;
+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 eggs, hesitated long before depositing them in the
+large cells; she chose rather to drop them at random than lay in cells
+unsuitable. However, on the second day, we found six that had been
+deposited there with all regularity. The worms were hatched three days
+afterwards, and then we began to study their history. Though the bees
+provided them with food, they did not carefully attend to it; yet I was
+in hopes they might be reared. I was again disappointed; for next
+morning all the worms had disappeared, and their cells were left empty.
+Profound silence reigned in the hive; few bees left it, and these
+returned without pellets of wax on the limbs; all was cold and
+inanimate. To promote a little motion, I thought of supplying the hive
+with a comb, composed of large cells, full of male brood of all ages.
+The bees, which had twelve days obstinately refused working in wax, did
+not unite this comb to their own. However, their industry was awakened
+in a way that I had not anticipated. They removed all the brood from
+this comb, cleaned out the whole cells, and prepared them for receiving
+new eggs. I cannot determine whether they expected the queen 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 removed 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, and
+this order cannot be inverted. We have seen that another fact led me to
+the same consequence; and as that was extremely important, I am
+delighted to have it confirmed by a new observation. 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 certain 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 large cells; for, by the
+experiment just described, 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 has instructed her that their size is
+neither proportioned to the size nor necessities of common worms, would
+not she also have been instructed not to lay 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 bees is not very conspicuous. Here the most prominent
+feature of industry appears in the common bees. When I supplied them
+with a comb of small cells, full of male brood, their activity was
+awakened; but instead of bestowing the necessary care on this brood, as
+they would have done in every other situation, they destroyed the whole
+nymphs and larvæ, and cleaned out their cells, that the queen, now
+oppressed with the necessity of laying, might suffer no delay in
+depositing her eggs. Could we allow them either reason or reflection,
+this would be an interesting proof of their affection for her.
+
+The experiment, now detailed at length, not having fulfilled my object
+in determining the influence of the size of the cells on that of the
+worms, I invented another which proved more successful.
+
+Having selected a comb of large cells, containing the eggs and worms of
+males, I removed all the worms from their farina, and my assistant
+substituted those of workers a day old in their place. Then he
+introduced this comb into a hive that had the queen. The bees did not
+abandon these substituted worms; they covered their cells with a top
+almost flat, a kind quite different from what is put on the cells of
+males; which proves, that they were well aware that these, though
+inhabiting large cells, were not males. This comb remained eight days
+in the hive, counting from the time the cells were sealed. I then
+removed it to examine the included nymphs, which proved those of workers
+in different stages of advancement; but, as to size and figure, they
+perfectly resembled what had grown in the smallest cells. I thence
+concluded, that the larvæ of workers do not acquire greater size in
+large than in small cells. Although this 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; undoubtedly
+she has ordained that their organs should be fully expanded, and there
+is sufficient space for that purpose; therefore more would be useless.
+Their expansion ought to be no greater in the most spacious cells than
+in those appropriated 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 will probably be less than that of common workers,
+because they have been cramped in the cells; but it does not thence
+ensue, that a larger cell will admit of them growing 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 larvæ
+of workers in the same circumstances. The large cells of males are
+sufficiently capacious for the perfect expansion of their organs. Thus,
+although reared in cells of still greater capacity, they will grow no
+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 them, and reared 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
+will be checked, they will not attain the usual size, of which there is
+proof in the following experiment. I had a comb consisting of the cell
+of large drones, and one with those of workers, which also served for
+the male worms. Of these, my assistant took a certain number from the
+smallest cells, and deposited them on a quantity of food purposely
+prepared in the large ones; and in return he introduced into the small
+cells the worms that had been hatched in the other, and then committed
+both to the care of the workers 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 metamorphosis arrived,
+gave both kinds that convex covering usually put on those of the 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.
+
+You suggested another experiment which I carefully made, but it met with
+an unforeseen obstacle. To appreciate the influence of the royal food on
+the expansion 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
+without success. Nor do I think it can ever succeed. 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 deprived of a queen,
+they change the cells containing worms into cells of the largest kind.
+Then the worms will infallibly be converted to queens.
+
+But there is another situation where we can judge of the influence of
+the royal food administered to worms in common cells. I have spoken at
+great length in my letter on the existence of fertile workers. You
+cannot forget, Sir, that the expansion of their sexual organs is owing
+to the reception of some particles of royal jelly, while in the
+vermicular form. For want of new observations, I must refer you to what
+is previously said on the subject.
+
+  _PREGNY, 4 September 1791._
+
+
+
+
+LETTER IX.
+
+_ON THE FORMATION OF SWARMS._
+
+
+I can add but a few facts to the information M. de Reaumur has
+communicated relative to swarms.
+
+A young queen, according to this celebrated naturalist, is always or
+almost always at the head of a swarm; but he does not assert the fact
+positively, and had some doubts on the subject. "Is it certain," says
+he "as we have hitherto supposed, in coincidence with all who have
+treated of bees, that the new colony is always conducted by a young
+mother? May not the old mother be disgusted with her habitation? or may
+she not be influenced by some particular 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 conducted the young swarms. By the following
+details, you will observe, that his suspicions 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 related.
+
+But previous to entering on this subject, I should repeat what has
+already been frequently observed, that the _leaf_ or flat hives are
+indispensible in studying the industry and instinct of bees. When they
+are left at liberty to conduct several rows of parallel combs, we can no
+longer observe what is continually passing between them, or they must be
+dislodged by water or smoke, for examining what has been constructed; a
+violent proceeding, which has a material influence on their instinct,
+and consequently exposes an observer to the risk of supposing simple
+accidents permanent laws.
+
+I now proceed to experiments proving that an old queen always conducts
+the first swarm.
+
+One of my glass hives consisted of three parallel combs, placed in
+squares that opened like the leaves of a book. It was well peopled and
+abundantly provided with honey, wax, and brood, of every age. On the
+fifth of May 1788, I removed the queen, and on the sixth, transferred
+all the bees into another hive, with a fertile queen at least a year
+old. They entered easily and without fighting, and were in general 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 formed regular
+circles around her. However, there was a little agitation in the
+evening, but confined to the surface of the comb where we had put the
+queen, and which she had not quitted. All was perfectly quiet on the
+other side of this comb.
+
+In the morning of the seventh, the bees had destroyed the twelve royal
+cells, but, independent of that, order continued prevalent in the hive;
+the queen laid the eggs of males in the large cells, and those of
+workers 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
+perpendicularly 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, and
+the longest were only about two lines and a half in depth from the
+bottom to the orifice.
+
+On the thirteenth, the queen seemed already more slender than when
+introduced into the hive; however she still laid some eggs, both in
+common cells and those of males. We also surprised her this day laying
+in a royal cell: she first dislodged the worker there employed, by
+pushing it away with her head, and then supported herself by the
+adjoining cells while depositing the egg.
+
+On the fifteenth, the queen was still more slender: the bees continued
+their attention to the royal cells, which were all unequally advanced;
+some to three or four lines in height, while others were already an inch
+long; which proved 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 hastened to collect
+and put the bees into a hive purposely prepared. Though we had
+overlooked the facts attending the departure of the swarm, the object of
+this experiment was fulfilled; for, on examination of all the bees, we
+were convinced they had been conducted by the old queen; by that we
+introduced on the sixth of the month, and which had been deprived of one
+of the antennæ. Observe, there was no other queen in this colony. In the
+hive she had left, we found seven 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.
+
+The new swarm next became the object of our attention: we observed it
+during the rest of the year, during winter and the subsequent spring;
+and, in April, we had the satisfaction of seeing a new swarm depart with
+the same queen at its head that had conducted the former swarm in May
+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 royal cells; she laid in
+one of them before us; and in the last place led forth the swarm.
+
+We have several times repeated the same experiment with equal success.
+Thus it appears incontestible, that the old queen always 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.
+The bees prepare these cells only while the queen lays male eggs; and a
+remarkable fact attends it, that after this laying terminates, her belly
+being considerably diminished, she can easily fly, whereas, her belly is
+previously so heavy she can hardly drag it along. Therefore it is
+necessary she should lay in order to be in a condition for undertaking
+her journey, which may sometimes be very long.
+
+But this single condition is not enough. It is also requisite that the
+bees be very numerous: they should even be superabundant, and a person
+might say they are aware of it: for, if the hive is thin, no royal cells
+are constructed when the male eggs are laid, which is solely at the
+period that the queen is able to conduct a colony. This fact was proved
+by the following experiment on a large scale.
+
+On the third of May 1788, we divided eighteen hives into two portions;
+all the queens were about a year old. Thus each portion of the hives
+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 of the hives threw a swarm.--Therefore, if the
+hive containing the old queen is not very populous, she remains in it
+until the subsequent spring; and if the population is then sufficient,
+royal cells will be constructed: the queen 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 very brief abstract of my observations on swarms conducted by
+old queens. You must excuse the long detail on which I am about to
+enter, concerning 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
+hitherto been very obscure. A long course of observations, protracted
+even during several years, was necessary to remove, in some degree, the
+veil that concealed these mysteries. I have been indemnified for the
+trouble, indeed, by the pleasure of seeing my experiments reciprocally
+confirmed; but, considering the assiduity 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 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
+queens; and I shall now extract the chief experiments from my journal.
+
+On the fourteenth of May, we introduced two portions of bees, from the
+straw hives, into a large glass hive very flat; and allowed them only
+one queen of the preceding year, and which had already commenced laying
+in its native hive. We introduced her on the fifteenth. She was very
+fertile. The bees received her well, and she soon began to lay in large
+and small cells alternately.
+
+On the twentieth, we saw the formation of twelve royal cells, all on the
+edges of the communications, or passages through the combs, and shaped
+liked stalactites.
+
+On the twenty-seventh, ten were much but unequally enlarged; 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 became more lively, yet she still continued examining
+the cells as when about to lay; sometimes introducing half her belly,
+but suddenly withdrawing it, without having laid. At other times she
+deposited an egg, which lay in an irregular position, on one side of the
+hexagon, and not fixed by an end to the bottom of the cell. The queen
+produced no distinct sound in her course, and we heard nothing different
+from the ordinary humming of bees. She passed over those in her way;
+sometimes when she stopped, the bees meeting her also stopped; and
+seemed to consider her. They advanced briskly, struck her with their
+antennæ, and mounted on her back. She then went on carrying some of the
+workers on her back. None gave her honey, but she voluntarily took it
+from the cells in her way. The bees no longer inclosed and formed
+regular circles around her. The first, aroused by her motions, followed
+her running in the same manner, and in their passage excited those still
+tranquil on the combs. The way the queen had traversed was evident after
+she left it, by the agitation created, which was never afterwards
+quelled: she had soon visited every part of the hive, and occasioned a
+general agitation; if some places still remained tranquil, the bees in
+agitation arrived, and communicated their motion. The queen no longer
+deposited her eggs in cells; she let them fall fortuitously: nor did the
+bees any longer watch over the young; they ran about in every different
+direction; even those returning from the fields, before the agitation
+came to its height, no sooner entered the hive than they participated in
+these tumultuous motions. They neglected to free themselves of the waxen
+pellets on their limbs, and ran blindly about. At last the whole rushed
+precipitately towards the outlets 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, and, for that reason, as I wished it to continue very populous, I
+removed the queen, at the moment she came out, that the bees might not
+fly too far, and 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 removed its queen also.
+
+All these facts were certain, and appeared susceptible of no error.
+Notwithstanding this, I was particularly earnest to learn whether old
+queens always followed the same course; which induced me, on the
+twenty-ninth, to replace, in the glass hive, the queen a year old, which
+had hitherto been the subject of my experiments, and had just began to
+lay the eggs of males. On the same day, we found one of the royal cells
+left by the preceding queen larger than the rest; and, from its length,
+supposed the included worm two days old: the egg had, therefore, been
+laid on the twenty-fourth by that queen, and the worm was hatched on the
+twenty-seventh. On the thirtieth, the queen laid a great deal in the
+large and small cells alternately. Now, and the two following days, the
+bees enlarged several royal cells, but unequally, which proved that they
+included larvæ of different ages. One was closed on the first of June,
+and on the second another. The bees also commenced some new ones. All
+was perfectly quiet at eleven in the morning; but, at mid-day, the
+queen, from the utmost tranquillity, became evidently agitated; and her
+agitation insensibly communicated to the workers in every part of their
+dwelling. In a few minutes they precipitately crowded to the entrances,
+and, along with the queen, left the hive. After they had settled on the
+branch of a neighbouring tree, I sought for the queen; thinking that, by
+removing her, the bees might return to the hive, which actually ensued.
+Their first care seemed to consist in seeking the female; they were
+still in great agitation, but gradually calmed; and in three hours
+complete tranquillity was restored.
+
+They had resumed their usual occupations on the third: 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, not by applying wax
+cordons, but by removing wax from the surface. Towards the top this
+waved work is almost imperceptible; it becomes deeper above, and the
+workers excavate it still more from thence to the base of the pyramid.
+The cell, when once shut, also becomes thinner; and is so much so,
+immediately preceding the queen's metamorphosis from a nymph, that all
+its motions are perceptible through the thin covering of wax on which
+the waved work is founded. It is a very remarkable circumstance, that in
+making the cells thinner, from the moment they are closed, the bees know
+to regulate their labour so that it terminates only when the nymph is
+ready to undergo its last metamorphosis.
+
+On the seventh day the coccoon is almost completely _unwaxed_, if I may
+use the expression, at the part next to the head and thorax of the
+queen. This operation facilitates her exit; for she has nothing to do
+but cut the silk that forms the coccoon. Most probably the object is, to
+promote evaporation of the superabundant fluids of the nymph. I have
+made some direct experiments to ascertain the fact, but 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 closing the second. The like
+was done to other royal cells successively, during the subsequent days.
+
+Every moment of the seventh, we expected the queen to leave the royal
+cell shut on the thirtieth of May. The seven days had elapsed. 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 being
+unwilling that she should yet quit her cell, they had soldered the
+covering to it with some particles of wax. What seemed most singular
+was, that this female emitted a very distinct sound, or clacking from
+her prison. It was still more audible in the evening, and even consisted
+of several monotonous notes in rapid succession.
+
+The same sound proceeded from the royal cell on the eighth. Several
+bees kept guard round each royal cell.
+
+The first cell opened on the ninth. The young queen was lively, slender,
+and of a brown colour. Now, we understood why bees retain the female
+captive in their cells, after the period for transformation has elapsed;
+it is, that they may be able to fly the instant they are hatched. The
+new queen occupied all our attention. When she approached the other
+royal cells, the bees on guard pulled, bit her, and chased her away;
+they seemed to be greatly irritated against her, and she enjoyed
+tranquillity only when at a good distance from these cells. This
+procedure was frequently repeated through the day. She twice emitted the
+sound; in doing so she stood, her thorax against a comb, and the wings
+crossed on her back; they were in motion but without being unfolded or
+further opened. Whatever might be the cause of her assuming this
+attitude, the bees were affected 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 approached, all the guards became agitated,
+surrounded her on all sides, bit, and commonly drove her away; sometimes
+when in these circumstances, she emitted her sound, assuming the
+position just described, from that moment the bees became motionless.
+
+The queen confined in the second cell had not yet left it, and was heard
+to hum several times. We accidentally discovered how the bees fed her.
+On attentive examination, a small aperture was perceptible in the end of
+the coccoon which she had cut to escape, and which her guards had again
+covered with wax, to confine her still longer. She thrust her trunk
+through the cleft; at first the bees did not observe it alternately
+thrust out and drawn in, but one at length perceiving it, came to apply
+its trunk to that of the captive queen, and then gave way to others that
+also approached her with honey. When satisfied she retracted her trunk,
+and the bees again closed up the opening with wax.
+
+The queen this day between twelve and one became extremely agitated. The
+royal cells had multiplied very much; she could go no where without
+meeting them, and on approaching she was very roughly treated. Then she
+fled, but to obtain no better reception. At last, these things agitated
+the bees; they precipitately rushed through the outlets of the hive, and
+settled on a tree in the garden. It singularly happened that the queen
+was herself unable to follow or conduct the swarm. She had attempted to
+pass between two royal cells before they were abandoned by the bees
+guarding them, and she was so confined and maltreated as to be incapable
+of moving. We then removed her into a separate hive prepared for a
+particular experiment; the bees, which had clustered on a branch, soon
+discovered their queen was not present, and returned of their own accord
+to the hive. Such is an account of the second colony of this hive.
+
+We were extremely solicitous to ascertain what would become of the other
+royal cells. Four of the close ones had attained complete maturity, and
+the queens would have left them had not the bees prevented it. They were
+not open either previous to the agitation of the swarms, or at the
+moment 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 formed the swarm. We could not
+discover what occasioned the difference in their captivity.
+
+On the twelfth, the queen was at last liberated, 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. Returning at mid-day, we were greatly surprised to find it
+almost totally deserted. During our absence, it had thrown a prodigious
+swarm, which still clustered on the branch of a neighbouring tree. We
+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, escaped. Thus, there was no doubt
+of both queens being in the swarm. We found it so; and removed them,
+that the bees might return to the hive, which they did very soon.
+
+While we were occupied in this operation, the fourth captive queen left
+her prison, 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 guard around the royal cells, and took
+great care to remove the queen whenever she attempted to approach.
+Eighteen royal cells now remained to be guarded.
+
+The fifth queen left her cell at ten at night; therefore two queens were
+now in the hive. They immediately began fighting, but came to disengage
+themselves from each other. However they fought several times during the
+night without any thing decisive. Next day, the thirteenth, we witnessed
+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 strikes the bees motionless. For some
+minutes, we conceived, that taking advantage of the dread exhibited by
+the workers on guard, she would open it, and destroy the young female;
+also she prepared to mount the cell; but in doing so she ceased the
+sound, and quitted that attitude which paralyses 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 described. The agitation
+was so considerable, that a sufficient number of bees did not remain to
+guard the royal cells, and several of the imprisoned queens were thus
+enabled to make their escape. Three were in the cluster formed by the
+swarm, and other three remained in the hive. We removed those that had
+led the colony, to force the bees to return. They entered the hive,
+resumed their post around the royal cells, and maltreated the queens
+when approaching.
+
+A duel took place in the night of the fifteenth, in which one queen
+fell. We found her dead next morning before the hive; but three still
+remained, as one had been hatched during night. Next morning we saw a
+duel. Both combatants were extremely agitated, either with the desire of
+fighting, or the treatment of the bees, when they came near the royal
+cells. Their agitation quickly communicated to the rest of the bees, and
+at mid-day they departed impetuously 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, which I shall
+give you no account of, as it shewed nothing new.
+
+Unfortunately we lost this, which was a very strong swarm; the bees flew
+out of sight, and could never be found. The hive was now very thinly
+inhabited. Only the few bees that had not participated in the general
+agitation remained, and those that returned from the fields after the
+swarm had departed. The cells were, therefore, slenderly guarded; the
+queens escaped from them, and engaged in several combats, until the
+throne remained with the most successful.
+
+Notwithstanding the victories of this queen, she was treated with great
+indifference from the sixteenth to the nineteenth, that is, the three
+days that she preserved her virginity. At length, having gone to seek
+the males, she returned with all the external signs of fecundation, and
+was henceforth received with every mark of respect; she laid her first
+eggs 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
+experiments which I made at the same time for 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, 6. September 1791._
+
+_P. S._--In revising this letter, I find I have neglected taking notice
+of an objection that may embarrass my readers, and which ought to be
+answered.
+
+After the first five swarms had thrown, I had always returned the bees
+to the hive: it is not surprising, therefore, that it was continually so
+sufficiently stocked that each colony was numerous. But things are
+otherwise in the natural state: the bees composing a swarm do not return
+to the hive; and it will undoubtedly be asked, What 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 the 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 workers' 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 affording a second colony without
+being too much weakened. The third and fourth swarm weaken it more
+sensibly; but the inhabitants always remain in sufficient numbers to
+preserve the course of their labours uninterrupted; and the losses are
+soon repaired by the great fecundity of the queen, as she lays above an
+hundred eggs a day.
+
+If, in some cases, the agitation of swarming 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, cannot share in the agitation; when returned to the
+hive, they quietly resume their labours; and their number is not small,
+for, when the weather is fine, at least a third of the bees 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 endeavouring to depart
+become members of the new colony. When this agitation or delirium 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 the rest, seem
+drenched in perspiration; their wings grow moist; they are incapable of
+flight; and even when able to escape, they advance no farther than the
+board of the hive, and soon return.
+
+Those that have lately left their cells remain behind the swarm, still
+feeble, they could not support themselves in flight. Here then are also
+many recruits to people what we should have thought a deserted
+habitation.
+
+
+
+
+LETTER X.
+
+_THE SAME SUBJECT CONTINUED._
+
+
+To preserve greater regularity in continuing the history of swarms, I
+think it proper to recapitulate in a few words the principal points of
+the preceding letter, and to expatiate on each, concerning the result of
+new experiments, respecting which I have still been silent.
+
+In the first place. _If at the return of spring, we examine a hive well
+peopled, and governed by a fertile queen, we shall see her lay a
+prodigious number of male eggs in the course of May, and the workers
+will chuse that moment for constructing several royal cells of the kind
+described by M. de Reaumur._ Such is the result of several long
+continued observations, among which there has not been the slightest
+variation, and I cannot hesitate in announcing it as demonstrated.
+However, I should here add the necessary explanation. It is necessary
+that the queen, before commencing her _great_ laying of the eggs of
+males, be eleven months old; when young she lays only those of workers.
+A queen, hatched in spring, will perhaps lay fifty or sixty eggs of
+drones in whole, but before beginning her great laying of them, which
+should be two thousand in a month, she must have completed her eleventh
+month in age. In the course of our experiments, which more or less
+disturbed the natural state of things, it often happened that the queen
+did not attain this age until October, and immediately began laying male
+eggs. The workers, as if induced by some emanation from the eggs, also
+adopted this time for building the royal cells. No swarm resulted
+thence, it is true, because in autumn all the necessary circumstances
+are absolutely wanting, but it is not less evident, that there is a
+secret relation between the production of the eggs of males, and the
+construction of royal cells.
+
+This laying commonly continues thirty days. The bees on the twentieth or
+twenty-first lay the foundation of several royal cells. Sometimes they
+build sixteen or twenty; we have even had twenty-seven. When the cells
+are three or four lines high, the queen lays those eggs from which her
+own species will come, but not the whole in one day. That the hive may
+throw several swarms, it is essential that the young females conducting
+them be not all produced at the same time. One may affirm, that the
+queen anticipates the fact, for she takes care to allow at least the
+interval of a day between every egg deposited in the cells. It is proved
+by the bees knowing to close the cells the moment the worms are ready to
+metamorphose to nymphs. Now, as they close all the royal cells at
+different periods, it is evident the included worms are not all of an
+equal age.
+
+The queen's belly is very turgid before she begins laying the eggs of
+drones; but it sensibly decreases as she advances, and when terminated
+is very small. Thus she finds herself in a condition to undertake a
+journey which circumstances may prolong; thus this condition was
+necessary; 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 larvæ hatched from the eggs laid by the queen, in
+the royal cells, are ready to transform to nymphs, this queen leaves the
+swarm conducting a swarm along with her; and the first swarm that
+proceeds from the hive is uniformly conducted by the old queen._{M} I
+think I can divine the reason of it.
+
+That there may never be a plurality of females in a hive, nature has
+inspired queens with a natural horror against each other; they never
+meet without endeavouring to fight, and to accomplish their mutual
+destruction. Thus, the chance of combat is equal between them, and
+fortune will decide to which the empire shall pertain. But if one
+combatant is older than the rest, she is stronger, and the advantage
+will be with her. She will destroy her rivals successively as produced.
+Thus, if the old queen did not leave the hive, when the young ones
+undergo their last metamorphosis, it could produce no more swarms, and
+the species would perish. Therefore, to preserve the species, it is
+necessary that the old queen 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 old queen; and that may
+happen thus. Nature has not willed that the queen shall quit the first
+hive before her production of male eggs is finished. It is necessary 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 workers. The bees in that case would have no means of replacing
+her, and the colony would go to ruin.
+
+All these things have been with infinite wisdom foreseen. The first
+operation of the bees of a swarm is to construct the cells of workers.
+They labour at them with great ardour, and as the ovaries of the queen
+have been disposed 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. It may be affirmed, that the bees
+know their queen will also lay the eggs of drones; she actually does
+begin to deposit some, though in much smaller number than at first;
+enough however to encourage the bees to construct royal cells. Now, if
+in these circumstances the weather is favourable, it is not impossible
+that a second colony may be formed, and that the queen may depart at the
+head of it within three weeks of conducting the first swarm. But I
+repeat, the fact is rare in our climate. Let me now return to the hives
+from which the queen has led the first colony.
+
+Thirdly. _After the old queen has conducted the first swarm from the
+hive, the remaining bees take particular care of the royal cells, and
+prevent the young queens successively hatched from leaving them, unless
+at an interval of several days between each._
+
+In the preceding letter, I have given you the detail and proof of this
+fact, and I shall here add some reflexions. During the period of
+swarming, the conduct or instinct of bees seems to receive a particular
+modification. At all other times, when they have lost their queen, they
+appropriate workers worms to replace her; they prolong and enlarge the
+cells of these worms; they supply them with aliment more abundantly, and
+of a more pungent taste; and by this alteration, the worms that would
+have changed to common bees are transformed to queens. We have seen
+twenty-seven 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 may perhaps leave her cell, and attack all
+the other royal cells successively, which she will tear open to destroy
+her rivals, without the workers taking any part in their defence. Should
+several queens be hatched at once, they will pursue each other, and
+fight until the throne remain with her that is victorious. Far from
+opposing such duels, the other bees rather seem to excite the
+combatants.
+
+Things are quite reversed during the period of swarming. The royal cells
+then constructed are of a different figure from the former. They
+resemble stalactites, and in the beginning are like the cup of an acorn.
+The bees assiduously guard the cells when the young queens are ready for
+their last metamorphosis. At length the female hatched from the first
+egg laid by the old queen leaves her cell; the workers at first treat
+her with indifference. But she, immediately 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
+drive her away, so that she is forced to remove; but the royal cells
+being numerous, scarce can she find a place of rest. Incessantly
+harassed with the desire of attacking the other queens, and incessantly
+repelled, she becomes agitated, and hastily traverses the different
+groupes of workers, to which she communicates her agitation. At this
+moment numbers of bees rush towards the aperture of the hive, and, with
+the young queen at their head, depart to seek another habitation.
+
+After the departure of the colony, the remaining workers set another
+queen at liberty, and treat her with equal indifference as the first.
+They drive her from the royal cells; being perpetually harrassed, she
+becomes agitated; departs, and carries a new swarm along with her. In a
+populous hive this scene is repeated three or four times during spring.
+As the number of bees is so much reduced, that they are no longer
+capable of preserving a strict watch over the royal cells, several
+females then leave their confinement at once; they seek each other,
+fight, and the queen at last victorious 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 after the first colony is led out by the old queen,
+until the next swarm is conducted by the first young queen set at
+liberty. The interval between the second and third is still shorter; and
+the fourth sometimes departs 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 the day when the sun shines, and the air is calm. Sometimes
+we have observed all the precursors of swarming, disorder and agitation,
+but a cloud passed before the sun, and tranquillity was restored; the
+bees thought no more of swarming. An hour afterwards, the sun having
+again appeared, the tumult was renewed; it rapidly augmented; and the
+swarm departed.
+
+Bees generally seem much alarmed at the prospect of bad weather. While
+ranging in the fields the passage of a cloud before the hive induces
+them precipitately to return. I am induced to think they are disquieted
+by the sudden diminution of light. For if the sky is uniformly obscured,
+and there is no alteration in clearness or in the clouds dispelling,
+they proceed to the fields for their ordinary collections, and the first
+drops of a soft rain does not 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 sometimes seem to use this right in an arbitrary manner. However
+the confinement of the queens is always longer when bad weather lasts
+several days together. Here the final object cannot be mistaken. If the
+young females were at liberty to leave their cradles during these bad
+days, there would be a plurality of queens in the hive, consequently
+combats; and victims would fall. Bad weather might continue so long,
+that all the queens might at once have undergone their last
+metamorphosis, or attained their liberty. One victorious over the whole
+would enjoy the throne, and the hive, which should naturally produce
+several swarms, could give only one. Thus the multiplication of the
+species would have been left to the chance of rain, or fine weather,
+instead of which it is rendered independent of either, by the wise
+dispositions of nature. By allowing only a single female to escape at
+once, the formation of swarms is secured. This explanation appears so
+simple, that it is superfluous to insist farther on it.
+
+But I should mention another important circumstance resulting from the
+captivity of queens; which is, that they are in a condition to fly, when
+the bees have given them liberty, and by this means are capable of
+profiting by the first moment of sunshine to depart at the head of a
+colony.
+
+You well know, Sir, that all drones and workers are not in a condition
+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 hours must elapse before the acquisition of perfect strength, and
+the development of all their faculties. It would be the same with the
+females was not their confinement protracted after the period of
+transformation; 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 used to retain the queens in
+captivity. The bees solder the covering to the sides of the cell by a
+cordon 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 according to their age. Immediately when the royal
+cells were sealed, we marked them all with numbers, and we chose this
+period because it indicated the age of the queens exactly. The oldest
+was first liberated, then the one immediately younger, and so on with
+the rest. None of the younger queens were set at liberty before the
+older ones.
+
+I have a thousand times asked myself how the bees could so accurately
+distinguish the age of their captives. Undoubtedly I should do better to
+answer this question by a simple avowal of my ignorance. At the same
+time, I must be permitted to state a conjecture. You will admit, that I
+have not, as some authors, abused the right of giving myself up to
+hypothesis; may not the humming 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 queens? It is certain that the female, whose
+cell is first sealed, is also the first to emit this sound. That in the
+next emits it sooner than the rest, and so on with those immediately
+subsequent. As their captivity may continue six days, it is possible
+that the bees in this space of time may forget which has emitted it
+first; but it is also possible, that the queens diversify the sounds,
+encreasing the loudness as they become older, and that the bees can
+distinguish these variations. We have even ourselves been able to
+distinguish differences in the sound, either with relation to the
+succession of notes, or their intensity; and probably there are
+gradations still more imperceptible that escape our organs, but may be
+sensible to those of the workers.
+
+What gives weight to this conjecture is, that the queens brought up by
+M. Schirach's method, are perfectly mute; neither do the workers form
+any guard around their cells, nor do they retain them in captivity a
+moment beyond the period of transformation, and, when they have
+undergone it, they are allowed to combat until one has become
+victorious over all the rest. Why? Because the object is only to replace
+the last queen. Now, provided that among the worms reared as queens,
+only one succeeds, the fate of the others is uninteresting to the bees,
+whereas, during the period of swarming, it is necessary to preserve a
+succession of queens, for conducting the different colonies; and to
+ensure the safety of the queens, it is necessary to avert the
+consequences of the mutual horror by which they are animated against
+each other. Behold the evident cause of all the precautions that bees,
+instructed by nature, take during the period of swarming; behold an
+explanation of the captivity of females; and that the duration of their
+captivity 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 have never been able to discover the
+situation of the organ which produces the sound. But I have instituted a
+new course of experiments on the subject, which are still unfinished.
+
+Another problem still remains for solution. Why are the queens reared,
+according to M. Schirach's method, mute, whilst those bred in 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 M. Schirach's method,
+receive it only the second or third day of their existence. It appears
+to me that this circumstance may have an influence on the different
+parts of organisation, and particularly on the organ of voice.
+Experiment has not confirmed this conjecture. I constructed glass cells
+in perfect imitation of royal cells, that the metamorphosis of the worms
+into nymphs, and of the nymphs to queens, might be visible. These
+experiments are related in a preceding letter. Into one of these
+artificial cells we introduced the nymph of a worm, reared according to
+M. Schirach's method, twenty-four hours before it could naturally
+undergo its last metamorphosis; and we replaced the glass cell in the
+hive, that the nymph might have the necessary degree of heat. Next 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 had contrived an aperture for her thrusting out her
+trunk, and that the bees might feed her. I expected that she would have
+been completely mute; but it was otherwise; for she emitted sounds
+similar to those already described, therefore my conjecture was
+erroneous.
+
+I next conceived that the queen being restrained in her motions, and in
+her desire for liberty, was induced to emit certain sounds. All queens,
+in this new point of view, are equally capable of emitting the sound,
+but to induce them to it, they must be in a confined situation. In the
+natural state, the queens that come from workers are not a single
+instant in restraint; 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 are 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 something more probable.
+
+I do not ascribe to myself the credit of having discovered the humming
+of the queen bee. Old authors speak of it. M. de Reaumur cites a Latin
+work published 1671, _Monarchia Femina_, by Charles Butler. He gives a
+very brief abstract of this naturalist's observations, who we easily see
+has exaggerated or rather disguised the truth, by mixing it with the
+most absurd fancies; but it is not the less evident that Butler has
+heard this peculiar humming of queens, and that he did not confound it
+with the confused humming sometimes heard in hives.
+
+Fourthly. _The young queens conducting swarms from their native hive are
+still in a virgin state._ The day after, being settled in their new
+abode, they generally depart in quest of the males; and this is usually
+the fifth day of their existence as queens; for two or three pass in
+captivity, one in their native hive, and a fifth in their new dwelling.
+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 they begin to lay. The old queen,
+which leads the first swarm in spring, requires no farther commerce with
+the males, for preservation of her fecundity. A single copulation is
+sufficient to impregnate all the eggs she will lay for at least two
+years.
+
+  _PREGNY, 8. September 1791._
+
+
+FOOTNOTES:
+
+{M} Schirach seems to have been aware of this fact.--T.
+
+
+
+
+LETTER XI.
+
+_THE SAME SUBJECT CONTINUED._
+
+
+I have collected my chief observations on swarms in the two preceding
+letters; those most frequently repeated, and of which the uniformity of
+result leads me to apprehend no error. I have deduced what seem the most
+direct consequences; and in all the theoretical part, I have sedulously
+avoided going beyond facts. What is yet to be mentioned is more
+hypothetical, but it engrosses several curious experiments.
+
+It has been demonstrated, that the principal motive of the young females
+departing when hives swarm, is their insuperable antipathy to each
+other. I have repeatedly observed that they cannot gratify their
+aversion, because the workers with the utmost care prevent them from
+attacking the royal cells. This perpetual opposition at length creates a
+visible inquietude, and excites a degree of agitation that induces 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 different. Always
+accustomed to respect fertile queens, they do not forget what they owe
+to her; they allow her the most uncontrouled liberty. She is permitted
+to approach the royal cells; and if she even attempts to destroy them,
+no opposition is presented 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. Therefore, I candidly confess
+myself ignorant of the motives of her departure.
+
+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 young ones, have the greatest aversion to 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 departure of
+old queens, seven royal cells opened at one side were destroyed by the
+queen. If rain continues several days, the whole are destroyed; in this
+case, there is no swarm, which too often happens in our climate, where
+spring is generally rainy. Queens never attack cells containing an egg
+or a very young worm; but only when the worm is ready for transforming
+to a nymph, or when it has undergone its last metamorphosis.
+
+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 is 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
+immediate success, her inquietude during this work becomes a terrible
+agitation. If the weather continues favourable, while she remains in
+this condition, she is naturally disposed to depart.
+
+It may easily be understood, that the workers 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
+respect. But you will undoubtedly ask, Sir, What motive can induce the
+workers to follow their queen from the hive, while they treat the young
+queens very ill, and, even in their most amicable moments, testify
+perfect indifference towards them. Probably it is to escape the heat to
+which the hive is then exposed. The extreme agitation of the females
+leads them to traverse the combs in all directions. They pass through
+groupes of bees, injure and derange them; they communicate a kind of
+delirium, and these tumultuous motions raise the temperature to an
+insupportable degree. We have frequently proved it by the thermometer.
+In a populous hive it commonly stands between 92° and 97°, in a fine day
+of spring; but during the tumult which precedes swarming, it rises above
+104°. And this is heat intolerable to bees. When exposed to it, they
+rush impetuously towards the outlets of the hive and depart. In general
+they cannot endure the sudden augmentation of heat, and in that case
+quit their dwelling; neither do those returning from the fields enter
+when the temperature is extraordinary.
+
+I am certain, from direct experiments, that the impetuous courses of the
+queen over the combs, actually throws the workers into agitation; and I
+was able to ascertain it in the following manner. I wished to avoid a
+complication of causes. It was particularly important to learn, whether
+the queen would impart her agitation but not at the time of swarming.
+Therefore I took two females still virgins, but capable of fecundation
+for above five days, and put one in a glass hive sufficiently populous;
+the other I put into a different hive of the same kind. Then I shut the
+hives in such a way that there was no possibility of their escape. The
+air had free circulation. I then prepared to observe the hives every
+moment that the fineness of the weather would invite both males and
+females to go abroad, for the purpose of fecundation. Next morning, the
+weather being gloomy, no male left the hive, and the bees were tranquil;
+but towards eleven of the following day, the sun shining bright, both
+queens began to run about seeking an exit from every part of their
+dwelling; 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 that part of the
+hive where the openings were placed; unable to escape they ascended with
+equal rapidity, and ran heedlessly over the cells until four in the
+afternoon. It is nearly about this time that the sun declining in the
+horizon recalls the males; queens requiring fecundation never remain
+later abroad. The two females became calmer, and tranquillity was in a
+short time restored. This was repeated 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
+always in a tumultuous state when the queen herself is in agitation.
+
+I have but one fact more to mention. It has already been observed, that
+on losing the female, bees give the larvæ of simple workers the royal
+treatment, and, according to M. Schirach, in five or six days they
+repair the loss of their queen. In this case there are no swarms. All
+the females leave their cells almost at the same moment, and after a
+bloody combat the throne remains with the most fortunate.
+
+I can very well comprehend that the object 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 metamorphosis almost at
+the same time? Since they are enabled to retain the young females in
+their cells, why do they allow all the queens, reared according to
+Schirach's method, to escape at once. By prolonging their captivity
+more or less, they would fulfil two most important objects at once, in
+repairing the loss of their females and preserving a succession of
+queens to conduct several swarms.
+
+At first it was my opinion, that this difference of conduct proceeded
+from the difference of circumstances in which they found themselves
+situated. They are induced to make all their dispositions relative to
+swarming only when in great numbers, and when they have a queen occupied
+with her principal laying of male eggs; whereas, having lost their
+female, the eggs of drones are no longer in the combs to influence their
+instinct. They are in a certain degree restless and discouraged.
+
+Therefore, after removing the queen from a hive, I thought of rendering
+all the other circumstances as similar as possible to the situation of
+bees preparing to swarm. By introducing a great many workers, I
+encreased the population to excess, and supplied them with combs of
+male brood in every stage. Their first occupation was to construct royal
+cells after Schirach's method, and to rear 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 this they discontinued, as there was
+no queen to deposit her eggs. Finally, I gave them several close royal
+cells, taken indifferently from hives preparing to swarm. However, all
+these precautions were fruitless; the bees were occupied only with
+replacing their lost queen; they neglected the royal cells entrusted to
+their care; the included queens came out at the ordinary time, without
+being detained prisoners a moment; they engaged in several combats, and
+there were no swarms.
+
+Recurring to subtleties, we may perhaps suggest a cause for this
+apparent contradiction. 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 beautiful system, and the more must we distrust that
+facility of imagination from which we think by embellishment to
+elucidate facts.
+
+In general, Naturalists who have long observed animals, and those in
+particular who have chose insects for their favourite study, have too
+readily ascribed to them our sentiments, our passions, and even our
+intentions and designs. Incited 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 painted different traits of the industry of these
+minute animals, with the colours inspired by an exalted imagination. Nor
+is even the celebrated Reaumur to be acquitted of such a charge. He
+frequently ascribes combined intentions to bees; love, anticipation, and
+other faculties of too elevated a kind. I think I can perceive that
+although he formed very just ideas of their operations, he would be
+well pleased that his reader should admit they were sensible of their
+own interests. He is a painter who by a happy interest 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 association of ideas _simply sensible_. It is
+not my object here to penetrate those depths, or to insist on the
+details.
+
+But, on the whole, facts relative to the formation of swarms perhaps
+present more subjects of admiration than any other part of the history
+of bees. I think it proper to state, in a few words, the simplicity of
+the methods by which the wisdom of nature guides their instinct. It
+cannot allow them the slightest portion of understanding; it leaves them
+no precautions to be taken, no combination to be followed, no foresight
+to exercise, no knowledge to acquire. But having adapted their
+sensorium to the different operations with which they are charged, it is
+the impulse 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 larvæ, and collect
+provisions, we must not seek for method, affection, or foresight. The
+only inducement must be sought for in some pleasing sensation attached
+to each of these operations. I address a philosopher; 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 works which treats of the industry of animals. Let me
+add but another sentence. The inducement of pleasure is not the sole
+agent; there is another principle, the prodigious influence of which, at
+least with regard to bees, has hitherto been unknown, that is the
+sentiment of aversion which all females continually feel against each
+other, a sentiment whose existence is so fully demonstrated by my
+experiments, and which explains many important facts in the theory of
+swarms.
+
+  _PREGNY, 10. September 1791._
+
+
+
+
+LETTER XII.
+
+_ADDITIONAL OBSERVATIONS ON QUEENS THAT LAY ONLY THE EGGS OF DRONES, AND
+ON THOSE DEPRIVED OF THE ANTENNÆ._
+
+
+In relating my first observations on queens that lay male eggs alone, I
+have proved that they lay 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 workers appeared defective. It is
+indeed most singular, that bees which know the worms of males so well
+when the eggs are laid in small cells, and never fail to give them a
+convex 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
+irregularity depends on something I cannot comprehend.
+
+In revising what is said on this subject, I observe still wanting an
+interesting experiment to complete the history of queens that lay only
+the eggs of drones. I had to investigate whether these females could
+themselves distinguish that the eggs they deposit in the royal cells
+would not produce queens. I have already observed that they do not
+endeavour to destroy these cells when close, and I thence concluded,
+that in general the presence of royal cells in their hive does not
+inspire them with the same aversion to females whose fecundation has
+been retarded; but to ascertain the fact more correctly, it was
+essential to examine how the presence of a cell containing a royal nymph
+would affect a queen that had never laid any other than the eggs of
+drones.
+
+This experiment was easy; and I put it in practice on the fourth of
+September, in a hive some time deprived of its queen. The bees had not
+failed to construct several royal cells for replacing their females. I
+chose this opportunity for supplying them with a queen, 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. One remaining was
+enough to shew the impression it would make on the stranger queen
+introduced; had she endeavoured to destroy it; this, in my opinion,
+would have proved that she anticipated the origin of a dangerous rival.
+You must admit the use I make of the word anticipate; it saves a long
+paraphrase; I feel the impropriety of it. If, on the contrary, she did
+not attack the cell I would thence conclude that the delay of
+fecundation, which deprived her of the power of laying workers eggs, had
+also impaired her instinct. This was the fact; the queen passed several
+times over the royal cell, both 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 bestowed 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 honey 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 is no longer preserved against their own
+sex in the nymphine state, nor do they longer 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, should be so well
+treated and become as dear to them as females laying both kinds of eggs.
+But I remember to have observed a fact more astonishing still. I have
+seen workers bestow every attention on a queen though sterile; and after
+her death treat her dead body as they had treated herself when alive,
+and long prefer this inanimate body to the most fertile queens I had
+offered them. This sentiment, which assumes the appearance of so lively
+an affection, is probably the effect of some agreeable sensation
+communicated to bees by their queen, independent of fertility. Those
+laying only the eggs of males probably excite the same sensation in the
+workers.
+
+I now recollect that the celebrated Swammerdam somewhere observes, that
+when a queen is blind, sterile, or mutilated, she ceases to lay, and the
+workers of her hive no longer labour or make any collections, as if
+aware that it was now useless to work. He cites no experiment that led
+him to the discovery. Those made by myself have afforded some very
+singular results.
+
+I frequently amputated the four wings of queens; and not only did they
+continue laying, but the same confederation of them was testified by the
+workers as before. Therefore, Swammerdam has no foundation for
+asserting, that mutilated queens cease to lay. Indeed, from his
+ignorance of fecundation taking place without the hives, it is possible
+he cut the wings off virgin queens, and they, becoming incapable of
+flight, remained sterile from inability to seek the males in the air.
+Thus, amputation of the wings does not produce sterility in queens.
+
+       *       *       *       *       *
+
+I have frequently cut off one antennæ 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 one still remained, the mutilation was imperfect; and the
+experiment decided nothing. But amputation of both antennæ produced most
+singular effects. On the fifth of September, I cut both 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 vivacity. Scarcely
+had the workers time to separate and recede before her; she dropped her
+eggs, without attending to deposit them in any cell. The hive not being
+very populous, part was without comb. Hither she seemed particularly
+earnest to repair, and long remained motionless. She appeared to avoid
+the bees; however, several workers followed her into this solitude, and
+treated her with the most evident respect. She seldom required honey
+from them, but, when that occurred, 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 appeared tormented with the desire of leaving her
+habitation. She rushed towards the opening, and entered the glass tube
+adapted there; but the external orific being too small, after fruitless
+exertion, she returned. Notwithstanding these symptoms of delirium, the
+bees did not cease to render her the same attention as they ever pay to
+their queens, but this one received it with indifference. All that I
+describe appeared to me the consequence of amputating the antennæ.
+However, her organization having already suffered from retarded
+fecundation, and as I had observed her instinct in some degree impaired,
+both causes might possibly concur in producing the same effect. To
+distinguish properly what belonged to the privation of the antennæ, a
+repetition of the experiment was necessary, in a queen otherwise well
+organised, and capable of laying both kinds of eggs.
+
+This I did on the sixth of September. I amputated both the antennæ of a
+female which had been several months the subject of observation, and
+being of great fecundity 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 same marks 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 antennæ, on the industry and
+instinct of bees, I attentively considered the manner in which these two
+mutilated queens treated each other. You cannot have forgot, Sir, the
+animosity with which queens, possessing all their organs, combat, on
+which account it became extremely interesting to learn whether they
+would experience the same reciprocal aversion after losing their
+antennæ. We studied these queens a long time; they met several times in
+their courses, and without exhibiting the smallest resentment. This last
+instance is, in my opinion, the most complete evidence of a change
+operated in their instinct.
+
+Another very remarkable circumstance, which this experiment gave me
+occasion to observe, consists in the good reception given by the bees to
+the stranger queen, while they still preserved the first. Having so
+often seen the symptoms of discontent that a plurality of queens
+occasions, after having witnessed the clusters formed around these
+supernumerary queens to confine them, I could not expect they would pay
+the same homage to a second mutilated one they still testified towards
+the first. Is it because after losing the antennæ, these queens have no
+more any characteristic which distinguishes the one from the other?
+
+I was the more inclined to admit this conjecture from the bad reception
+of a third fertile queen preserving her antennæ, which was introduced
+into the same hive. The bees seized, bit her, and confined her so
+closely, that she could hardly breath or move. Therefore, if they treat
+two females deprived of antennæ in the same hive equally well, it is
+probably because they experience the same sensation from these two
+females, and want the means of longer distinguishing them from each
+other.
+
+From all this, I conclude, that the antennæ are not a frivolous ornament
+to insects, but, according to all 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 organised in such a manner as to fulfil
+both functions at once.
+
+As in the course of this experiment both mutilated females constantly
+endeavoured 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 I removed the first and third queen from the hive,
+leaving the fertile mutilated one, and enlarged the entrance.
+
+The queen left her habitation the same day. At first she tried to fly,
+but, her belly being full of eggs, she fell down and never attempted it
+again. No workers accompanied her. Why, after rendering the queen so
+much 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, carrying 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 I can hazard on this question is a conjecture.
+It appears that bees are induced to quit the hives from the increased
+heat which occasions the agitation of their queen, and the tumultuous
+motion which she communicates to them. A mutilated queen,
+notwithstanding her delirium, does not agitate the workers, because she
+seeks the uninhabited parts of the hive, and the glass panes of it: she
+hurries over clusters of bees, but the shock resembles that of any other
+body, and produces only a local and instantaneous motion. The agitation
+arising from it, is not communicated from one place to another, like
+that produced by a queen, which in the natural state wishes to abandon
+her hive and lead out a swarm; there is no increased heat, consequently
+nothing that renders the hive insupportable to her.
+
+This conjecture, which affords a tolerable explanation why bees persist
+in remaining in the hive, though the mutilated queen has left it, is no
+reason for the motive inducing the queen herself to depart. Her instinct
+is altered; that is the whole that I can perceive. I can discern nothing
+more. It is very fortunate for the hive, that this queen departs, for
+the bees incessantly attend her; nor do they ever think of procuring
+another while she remains; and if she was long of leaving them, it would
+be impossible to replace her; for the workers worms would exceed the
+term at which they are convertible into royal worms, and the hive would
+perish. Observe, that the eggs dropped by the mutilated queen can never
+serve for replacing her, for, not being deposited in cells, they dry and
+produce nothing.
+
+I have yet to say a few words on females laying male eggs only. M.
+Schirach supposes that one branch of their double ovary suffers some
+alteration. He seems to think that one of these branches contains the
+eggs of males, while the other has none but common eggs, and as he
+ascribes the inability of certain queens to lay the latter to some
+disease, his conjecture becomes very plausible. In fact, if the eggs of
+males and workers are indiscriminately mixed in both branches of the
+ovary, it appears at first sight that whatever cause acts on that organ,
+it should equally affect both species of eggs. If on the contrary, one
+branch is occupied by the eggs of drones only, and the other contains
+none but common eggs, we may conceive how disease affects the one, while
+the other remains untouched. Though this conjecture is probable, it is
+confuted by observation. We lately dissected queens, which laid none but
+male eggs, and found both branches of the ovary equally well expanded,
+and equally sound, if I may use the expression. The only difference
+that struck us was that in these two branches, the eggs were apparently
+not so close together as in the ovaries of queens laying both kinds of
+eggs.
+
+  _PREGNY, 12. September 1791._
+
+
+
+
+LETTER XIII.
+
+_ECONOMICAL CONSIDERATIONS ON BEES._
+
+
+In this letter I shall treat of the advantages that may be derived from
+the new invented hives, called _book_ or _leaf_ hives, in promoting the
+_economical knowledge_ of bees.
+
+It is needless to relate the different methods hitherto employed in
+forcing bees to yield up a portion of their honey and wax; all resemble
+each other in being cruel and ill understood.
+
+It is evident, when bees are cultivated for the purpose of sharing the
+produce of their labours, we must endeavour to multiply them as much as
+the nature of the country admits; and consequently to regard their lives
+at the time we plunder them. Therefore it is an absurd custom to
+sacrifice whole hives to get at the riches they contain. The inhabitants
+of this country, who follow no other method, annually lose immense
+numbers of hives; and spring, being generally unfavourable to swarms,
+the loss is irreparable. I well know that at first they will not adopt
+any other method; 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. Though the bees are indifferent as
+to the position of their combs, and as to their greater or lesser size,
+they are obliged to construct them perpendicular to the horizon, and
+parallel to each other. Therefore, if left entirely to themselves, when
+establishing a colony in one of those new hives, they would frequently
+construct several small combs parallel indeed, but perpendicular to the
+plane of the frames or leaves, and by this disposition prevent the
+advantages which I think to derive from the figure of my hives, since
+they could not be opened without breaking the combs. Thus they must
+previously have a guide to follow; the cultivator himself lays the
+foundation of their edifices, and that by a simple method. A portion of
+comb must be solidly fixed in some of the boxes composing the hive; the
+bees will extend it; and, in prosecution of their work, will accurately
+follow the plan already given them. Therefore on opening the hive, no
+obstacle is to be removed, nor stings to be dreaded, for one of the
+most singular and valuable properties attending this construction, 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, and the tranquillity of the bees has given you great
+surprise. I can desire no other evidence of my assertion. It is in the
+facility of opening these hives at pleasure that all the advantages lie,
+which I expect in perfecting the economical knowledge of bees.
+
+I conceive, when I observe bees may be rendered tractable, that it need
+not be added, I do not arrogate to myself the absurd pretence of
+_taming_ them, for this excites a vague idea of tricks; and I would
+willingly avoid the hazard of exposing myself to any such reproach. I
+ascribe their tranquillity on opening the hives, to the manner that the
+sudden introduction of light affects them; then, they seem rather to
+testify fear than anger. Many retire and enter the cells, and appear to
+conceal themselves. What confirms my conjecture is, 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. The divisions must be separated slowly, and
+care taken not to wound any of the bees. If they cluster too much on the
+combs, they must be brushed off with a feather; and breathing on them
+carefully avoided. The air we expire seems to excite their fury; it
+certainly has some irritating quality, for if bellows are used, they are
+rather disposed to escape than to sting.
+
+Respecting the advantages of leaf hives, I shall observe, they are very
+convenient for forming _artificial_ swarms. In the history of natural
+swarms, I have shewn how many favourable circumstances are necessary for
+their success. From experience I know that they very often fail in our
+climate; and even when a hive is disposed to swarm, it frequently
+happens that the swarm is lost either because the moment of its
+departure has not been foreseen, because it rises out of sight, or
+settles on inaccessible places. Instructing the cultivators of bees how
+to make artificial swarms is a real service, and the form of my hives
+renders this an easy operation. But it requires farther illustration.
+
+Since bees, according to M. Schirach's discovery, can procure another
+queen after having lost their own, provided there is workers brood in
+the combs not above three days old, it results that we can at pleasure
+produce queens, by removing the reigning one. Therefore, if a hive
+sufficiently populous is divided in two, one half will retain the old
+queen, and the other will not be long of obtaining 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
+sufficient to admit of division, if the brood is of the proper age, if
+males exist or are ready to be produced for impregnating the young
+queens.
+
+Supposing the union of all these conditions, the following is the method
+to be pursued. The leaf hive may be divided through the middle without
+any shock. Two empty divisions may be insinuated between the halves,
+which, when exactly applied to each other, are close on the outside. The
+queen must be sought in one of the halves, and marked to avoid mistake.
+If she by chance remains in the division with most brood, she is to be
+transferred to the other with less, that the bees may have every
+possible opportunity 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 occupied. The old entrance, now become useless, will be shut
+up; but as each half requires a new one, it ought to be made at the
+bottom of each division, on purpose that they may be as far asunder as
+possible. Both entrances should not be made on the same day. The bees in
+the half deprived of the queen ought to be confined twenty-four hours,
+and no opening made before then except for admission of air. Without
+this precaution, they would soon search for their queen, and infallibly
+find her in the other division. They will then retire in great numbers
+from their own division, until too few remain to perform the necessary
+labours. But this will not ensue if they are confined twenty-four hours,
+provided that interval is sufficient to make them forget the queen. When
+all these circumstances are favourable, the bees, in the division
+wanting the queen, will the same day begin to labour in procuring
+another, and ten or fifteen days after the operation, their loss will be
+repaired. The young female they have reared, soon issues forth to seek
+impregnation, and in two days commences the laying of workers eggs.
+Nothing more is wanting to the bees of this half hive, and the success
+of the artificial swarm is ensured.
+
+It is to M. Schirach that we are indebted for this ingenious method of
+forming swarms. He supposes, by producing young queens early in spring,
+that early swarms might be procured, which would certainly be
+advantageous in favourable circumstances. But unfortunately this is
+impossible. Schirach believed that queens were impregnated 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 become 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. Thus, if a
+swarm were artificially formed before the usual time of the males
+originating, the bees would be discouraged by the sterility of the young
+female. Or should they remain faithful to her, awaiting the period of
+fecundation, as she could not for three or four weeks receive the
+approaches of the male, she would lay eggs producing males only, and the
+hive in this case would perish. Thus the natural order must not be
+deranged, but we must delay the division of hives until males are about
+to originate or actually exist.
+
+Besides, if M. Schirach did succeed in obtaining artificial swarms,
+notwithstanding the great inconvenience of his hives, it was owing to
+his singular address and unremitting assiduity. He had some pupils in
+the art; these communicated the method of forming artificial swarms to
+others, and there are people now in Saxony who traverse the country
+practising this operation. Those versant in the matter can alone venture
+to undertake it with common hives, whereas, every cultivator can do it
+himself with the leaf hives.
+
+In this construction, another very great advantage will also be found.
+Bees can be forced to work in wax. Here I am led to what I believe is a
+new observation. While naturalists have directed our admiration to the
+parallel position of the combs, they have overlooked another trait in
+the industry of bees, namely, the equal distance uniformly between them.
+On measuring the interval separating the combs, it will generally be
+found four lines. Were they too distant, it is very evident the bees
+would be much dispersed and unable to communicate their heat
+reciprocally; whence the brood would not be exposed to sufficient
+warmth. Were the combs too close, on the contrary, the bees could not
+freely traverse the intervals, and the work of the hive would suffer.
+Therefore, a certain distance always uniform is requisite, which
+corresponds equally well with the service of the hive, and the care
+necessary for the worms. Nature, which has taught bees so much, has
+instructed them regularly to preserve this distance. At the approach of
+winter, they sometimes elongate the cells which are to contain the
+honey, and thus contract the intervals between the combs. But this
+operation is a preparation for a season, when it is important to have
+plentiful magazines, and when their activity being very much relaxed, it
+is unnecessary for their communications to be so spacious and free. On
+the return of spring, the bees hasten to contract these elongated cells,
+that they may become fit for receiving the eggs which the queen will
+lay, and thus re-establish the just distance which nature has ordained.
+
+This being admitted, bees may be forced 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
+build others in the interval. Suppose an artificial swarm is lodged in a
+leaf hive, composed of six divisions, each containing a comb, if the
+young queen is as fertile as she ought to be, the bees will be very
+active in their labours, and disposed to make great collections of wax.
+To induce them towards this an empty box or division must be placed
+between two others, each containing a comb. As all the boxes are of
+equal dimensions, and of the necessary width for receiving a comb, the
+bees having sufficient 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 having more than four lines
+between them. Without any guide, this new comb will be parallel to the
+old ones, to preserve that law which establishes an equal distance
+throughout the whole.
+
+If the hive is strong and the weather good, three empty divisions may at
+first be left between the old combs; one between the first and second,
+another between the third and fourth, and the last between the fifth and
+sixth. The bees will fill them in seven or eight days, and the hive then
+contains nine combs. Should the temperature of the weather continue
+favourable, three new leaves or divisions may be introduced;
+consequently 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 where flowers perpetually blow. But in our
+country, I have reason to think that the labour should not be forced
+more during the first year.
+
+From these details, you are sensible, Sir, how preferable _leaf hives_
+are to those of any other construction, and even to those ingenious
+stages described by _M. Palteau_, for the bees cannot by means of them
+be forced to labour more in wax than they would do if left to
+themselves; whereas, they are obliged to do it by inserting empty
+divisions. Next, the combs constructed on those stages cannot be removed
+without destroying considerable portions of brood, deranging the bees,
+and creating real disorder in the hive.
+
+Mine have also this advantage, that what passes within may daily be
+observed, and we may judge of the most favourable moments for depriving
+the bees of part of their stores. With all the combs before us we can
+distinguish those containing brood only, and what it is proper to
+preserve. The scarcity or abundance of provisions is visible, and the
+portion suitable may be taken away.
+
+I should protract this letter too much, if I gave an account of all my
+observations 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 their riches with them. The subject would require a separate
+work; and I may perhaps one day engage in it; but until that arrives I
+shall always feel gratification in communicating to cultivators, who
+wish to follow my method, directions of which long practice has
+demonstrated the utility.
+
+Here I shall only observe, that we hazard absolute ruin of the hives,
+by robbing them of too great a proportion of honey and wax. In my
+opinion, the art of cultivating these animals consists in moderately
+exercising the privilege of sharing their labours; but as a compensation
+for this, every method must be employed which promotes the
+multiplication of bees. Thus, for example, if we desire to procure a
+certain quantity of honey and wax annually, it will be better to seek it
+in a number of hives, managed with discretion, than to plunder a few of
+a great proportion of their treasures.
+
+It is indubitable that the multiplication of these industrious animals
+is much injured by privation of several combs, in a season unfavourable
+to the collection of wax, because the time consumed in replacing them is
+taken from that which should be consecrated to the care of the eggs and
+worms, and by this means the brood suffers. Besides, they must always
+have a sufficient provision of honey left for winter, for although less
+is consumed during this season, they do consume some; because they are
+not torpid, as some authors have conceived.{N} Therefore if they have
+not enough, they must be supplied with it, which requires great
+exactness. I admit that in determining to what extent hives may be
+multiplied in a particular country, it is necessary first to know how
+many the country can support, which is a problem yet unsolved. It also
+depends on another, the solution of which is as little known, namely the
+greatest distance that bees fly in collecting their provender. Different
+authors maintain, they can fly several leagues from the hive. But by the
+few observations I have been able to make, this distance seems greatly
+exaggerated. It appears to me that the radius of the circle they
+traverse does not exceed half a league. As they return to the hive with
+the greatest precipitation whenever a cloud passes before the sun, it is
+probable they do not fly far. Nature which has inspired them with such
+terror for a storm, and even for rain, undoubtedly 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 distances bees with the thorax painted, that they might again
+be recognised. But none ever returned that I had carried for twenty-five
+or thirty minutes from their dwelling, while those at a shorter distance
+have found their way and returned. I do not state this experiment as
+decisive. Though bees do not generally fly above half a league, it is
+very possible they go much farther, when flowers are scarce in their own
+vicinity. A conclusive experiment must be made in vast arid or sandy
+plains, separated by a known distance from a fertile region.
+
+Thus, the question yet remains undecided. But without ascertaining the
+number of hives that any district can maintain, I shall remark that
+certain vegetable productions are much more favourable to bees than
+others. More hives, for example, may be kept in a country abounding
+meadows, and where black grain is cultivated, than in a district of
+vineyards or corn.
+
+       *       *       *       *       *
+
+Here 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. Several problems concerning the history of these
+animals still remain unsolved. The experiments I project may perhaps
+throw some light on them; and I shall be animated 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 HUBER.
+  _PREGNY, 1. October 1791._
+
+
+FOOTNOTES:
+
+{N} So far from being torpid in winter, when the thermometer 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.
+
+Now that I am on the subject of thermometrical observations, I may
+cursorily remark, that M. Dubois of Bourg en Bresse, in a memoir
+otherwise valuable, is of opinion, that the larvæ cannot be hatched
+below (104). I have repeatedly made the experiment with the most
+accurate thermometers, and obtained a very different result. When the
+thermometer rises to (104°), the heat is so much greater than the eggs
+require, that it is intolerable to the bees. M. Dubois has been
+deceived, I imagine, by too suddenly introducing his thermometer into a
+cluster of bees, and putting them in agitation, the mercury has rose
+higher than it should naturally do. Had he delayed introducing the
+thermometer, he would soon have seen it fall to between 95 and 97, which
+is the usual temperature of hives in summer. In August this year, when
+the thermometer in the open air stood at 94, it did not rise above 99 in
+the most populous hives. The bees had little motion, and a great many
+rested on the board of the hive.
+
+
+
+
+APPENDIX.
+
+
+[The following passages are chiefly engrossed in the substance of the
+work, but the Translator, as has already been observed, for various
+reasons, judges it expedient to transfer them to an appendix. In his
+opinion these very minute details rather interrupt the connexion of the
+narrative, however interesting they may be considered, and they pertain
+more to researches purely anatomical.
+
+The Translator has likewise in some instances incorporated several long
+and important notes with the text; because it appears to him that they
+actually belong to the substance of the treatise. These are the only
+variations from the original with respect to arrangement.]
+
+       *       *       *       *       *
+
+Swammerdam has given an imperfect description of the ovary of the queen.
+He observes that he has never been able to find the termination of the
+oviducts in the abdomen, nor any other parts excepting those which he
+has described. "Notwithstanding all my exertions, I never could discover
+the site of the vulva, partly because I had not all my apparatus with me
+in the country, when investigating this subject, and partly from my
+apprehension of injuring other parts by pressure, which I had then
+occasion to examine. However, I have clearly observed a muscular
+swelling of the oviduct, where approaching the last ring of the belly;
+that it then contracts and afterwards dilates in becoming membranaceous.
+As I was desirous of preserving the poison bag, which is situated
+exactly here, along with, the muscles aiding the motion of the sting, I
+could follow the oviduct no farther. However, in another female, it
+appeared that the vulva is in the last ring of the abdomen, and under
+the sting. The parts expanding only while the queen lays, renders it
+extremely difficult to penetrate the aperture."
+
+We have attempted to discover what has escaped the indefatigable
+Swammerdam. But his observation that the research can be made to the
+greatest advantage, at the time of laying, has paved the way to us. We
+have remarked that the oviduct did not issue from the body, but that the
+eggs fall into a kind of cavity, where they are retained several seconds
+before being laid.
+
+On the sixth of August, we took a very fertile queen, and holding her
+gently by the wings in a supine position, the whole belly was exposed.
+She seized the extremity with her second pair of legs, and curved it as
+much as possible. This seeming an unfavourable position for laying, we
+forced her to stretch it out. The queen, oppressed with the necessity of
+laying, could no longer retain her eggs. The lower part of the last ring
+then separated so far from the upper part as to leave some of the inside
+discovered. In this cavity the sting lay above in its sheath. As the
+queen now made new efforts, we saw an egg fall into the cavity from the
+end of the oviduct. The lips then closed for several seconds; they
+opened again, and, in a much shorter time, dropped the egg from the
+cavity.
+
+From our own observations we found that the seminal fluid of drones
+coagulated on exposure to the air, and from several experiments had so
+little doubt on the subject, that whenever the female returned with the
+external marks of fecundation, we thought we recognised it in the
+whitish substance filling the sexual organs. It did not then occur to us
+to dissect the females to ascertain the fact more particularly: but this
+year, whether designing to neglect nothing, or to examine the distension
+of the female organs, we determined to dissect several. To our infinite
+surprise, what we had supposed the residue of the prolific fluid,
+actually proved the genital organs of the male, which separate from his
+body during copulation, and remain in the female.
+
+We procured a number of queens according to Schirach's method for the
+purpose of dissection, and set them at liberty that they might seek the
+males. The first which did so, was seized the instant she returned, and
+without dissection spontaneously exhibited what we were so impatient to
+behold. Examining the under part of the belly, we saw the oval end of a
+white substance which distended the sexual organs. The belly was in
+constant motion, by alternate extension and contraction. Already had we
+prepared to sever the rings, and by dissection to ascertain the cause of
+these motions; when the queen curving her belly very much, and
+endeavouring to reach its extremity with her hind legs, seized the
+distending substance with her claws, and evidently made an effort to
+extract it. She at last succeeded, and it fell before us. We expected a
+shapeless mass of coagulated fluid; what therefore was our surprise to
+find it part of the same male that had rendered this queen a mother. At
+first we could not credit our eyes; but after examining it in every
+position, both with the naked eye, and a powerful magnifier, we
+distinctly recognised it to be that part which M. de Reaumur calls the
+_lenticular_ body, or the _lentil_, in the following description.{O}
+
+    'Opening a drone there appears a portion formed by the assemblage of
+    several parts, often whiter than milk. This on investigation is
+    found to be principally composed of four oblong pieces. The two
+    largest are attached to a kind of twisted cord, fig. 4. r, called by
+    Swammerdam the root of the penis; and he has denominated seminal
+    vessels, s. s. two long bodies that we are about to consider. Other
+    two bodies oblong like the preceding, but shorter and not half the
+    diameter, he calls the _vasa deferentia_, d. d. Each communicates
+    with one of the seminal vessels near, g. g. where they unite to the
+    twisted cord, r. From the other extremity proceeds a very delicate
+    vessel, which, after several involutions, terminates in a body, t. a
+    little larger, but difficult to disengage from the surrounding
+    tracheæ. Swammerdam considers these two bodies, t. t. the testicles.
+    Thus there are two parts of considerable size, communicating with
+    other two still thicker and longer. These four bodies are of a
+    cellular texture, and full of a milky fluid, which may be squeezed
+    out. This long twisted cord, r, to which the largest of the seminal
+    vessels is connected, this cord, I say, is doubtless the channel by
+    which the milky fluid issues. After several plications, it
+    terminates in a kind of bladder or fleshy sac, i. i. In different
+    males this part is of various length and flatness. By calling it the
+    _lenticular_ body, or the lentil, it receives a name descriptive of
+    the figure it presents in all males whose internal parts have
+    acquired consistency in spirit of wine. The body, l. i. is therefore
+    a lentil, a little thickened, of which one half, or nearly so, of
+    the circumference is edged along the outline by two chesnut coloured
+    scaly plates, e. i. A small white cord, the real edge of the lentil,
+    is visible, and separates them. This lentil is a little oblong, and,
+    for convenience, we shall ascribe two extremities to it, the
+    anterior and posterior. The anterior, l, next the head, is where the
+    canal, r, dividing the seminal vessels is inserted, and the opposite
+    part; i. next the anus, the posterior. The two scaly plates, e. i.
+    e. i, proceed from the vicinity of this last part, whence each
+    enlarges to cover part of the lentil. Under the broadest part of
+    each plate, there is a division formed by two soft points of unequal
+    length; the largest of which is on the circumference of the lentil.
+    Besides these two scaly plates, there are two others, n. n. of the
+    same colour, narrower, and fully one half shorter, each of which is
+    situated very near the preceding, and originates close to the origin
+    of that it accompanies, namely, at the posterior part of the lentil.
+    The rest of the lentil is white and membranaceous. From behind
+    proceeds a tube, k. a canal also white and membranaceous, but it is
+    difficult to judge of its diameter, for the membranes, of which it
+    consists, are evidently in folds. To one side of this pipe is
+    attached a fleshy part, p. somewhat pallet shaped, one side is
+    concave, and the edges plaited; the other side is convex. In certain
+    places the plaits rise and project from the rest of the outline, and
+    form a kind of rays; the pallet appears prettily figured. Though
+    lying with the concave side applied to the lentil, it is not fixed
+    to it. Swammerdam seems to consider this pallet as the
+    characteristic part of the male.
+
+    'Though the parts we have described are the most conspicuous in the
+    male, they are neither those which protrude first, nor when
+    protruded are the most remarkable. On viewing from the opposite edge
+    of the lentil, forming the division of the two great scaly plates, a
+    sac or canal, k. proceeding from the posterior part of the lentil,
+    there is distinctly visible the body u, which we call the arc; where
+    there are five transverse hairy bands of a yellow colour, while the
+    rest is white. This arc seems out of the membranaceous canal because
+    it is covered only by a very transparent membrane. One end almost
+    reaches the lenticular body, and the other terminates where the
+    membranaceous canal joins the folded yellow membranes, m. which form
+    a species of sac, that is applied to the sides of the aperture,
+    adapted for the genital organs passing through. These reddish
+    membranes are those that appear first on pressure, and form this
+    elongated portion, at whose end is a kind of hairy mask. Finally,
+    with the sac formed by the reddish membranes, there are connected
+    two appendages, c. c. of reddish yellow, and red at the end, s.
+    These are what appear externally like horns.{P}'
+
+The lenticular substance, l. i. provided with each scaly lamina, are the
+only parts of those described by M. de Reaumur, that we have found
+engaged in the organs of our queens. The canal, r, by Swammerdam
+denominated the root of the penis, breaks in copulation; and we have
+seen its fragments at the place where it unites to the end of the
+lentil, l. towards the anterior extremity; but we have found no traces
+of the canal, k, formed of involuted membranes, which in the body of the
+male proceeds from the posterior end of the lentil, l. i. nor of the
+plaited pallet, p. adhering to this canal, called by Swammerdam the
+penis from its resemblance to that of other animals, though he is not of
+opinion that this point, which is not perforated, can perform the
+functions of a real penis, and hold the principal part in generation.
+The canal, k, therefore, and all appertaining to it, must break at i,
+quite close to the posterior part of the lentil, since we found no
+remains of the lenticular bodies left by the fecundating males, in the
+body of our females. The canal, r, which Swammerdam calls the root of
+the penis, with greater reason than he was himself aware, is not
+extended in the body of the male as represented by the figure here
+engraved, but this long twilled canal consists of several involutions,
+from the seminal vessels whence it proceeds, into the lenticular body
+where it terminates, and where it conveys the fluid. This canal
+therefore can extend during copulation, and allow the lenticular
+substance to protrude out of the body of the males.
+
+It is evident this may be the case during copulation as is seen on
+opening a drone, for, by endeavouring to displace the lenticular body,
+the involutions of the cord disappear, and it extends much more than
+necessary for the lentil to protrude from the body; and if we attempt
+to separate it farther, the canal breaks at l. close to the lentil, and
+at the same place where it breaks in copulation.
+
+By dissection two nerves are discovered, towards the origin of the
+canal, r. inserted into the seminal vessels and distribute in them, and
+towards the root of the penis many ramifications undoubtedly serving for
+the motion of these parts. Two small parts, perceptible near the nerves,
+are two ligaments for retaining the generative organs in their proper
+place, so that except the root of the penis, they cannot be drawn out
+without some exertion; it and the lenticular body however can protrude,
+and actually do so during copulation. A certain degree of pressure
+forces all these parts from the body of the male, but they spontaneously
+return, and appear reversed.
+
+Swammerdam, and after him M. de Reaumur have admired this mechanism;
+they have thought, indeed, that the return should be occasioned by the
+effect of the air inflating the parts, and they supposed that the male
+organs proceeded from the body, and returned during copulation, the same
+as when forced out by pressure. Following their example, we have pressed
+them from the body of many males; we have a thousand times witnessed
+this wonderful return, which they detail with the greatest precision;
+but our males never survived the operation. We have seen, as M. de
+Reaumur, a few males protrude them spontaneously, even some of the parts
+inverted, but at that moment they died, and were unable to retract the
+parts which a pressure, most likely accidental, had forced out. Thus it
+is improbable that the male organs protrude by turning out of themselves
+in copulation; and the details which follow prove incontestibly, that it
+is otherwise. Had not Swammerdam been prejudiced with this opinion, he
+would have seen that the lenticular body can proceed from the body in
+erection without reversing itself; he could have proportioned the
+tortuous canal, which he calls the root of the penis; he would have seen
+that, at certain times, it can be sufficiently extended to allows the
+lenticular substance to protrude; he would have discovered the real use
+of the scaly plates; he would have explained that of the canal k, of the
+plaited pallet q, and the movements of all these parts, more admirable
+perhaps than the inversion which he was the first to observe.
+
+Our observations incontestibly prove copulation. The portion of the
+males found engaged in the body of our queens, hitherto called the
+lenticular substance, may be denominated a penis both from its position
+and use. The same surface is presented by it in the queen as in the body
+of the male, which is proved by the position of the laminæ, e. e.
+attached to the interior of the penis, when found in the queen. It is
+evident, if the supposed inversion took place, the laminæ would be found
+within the posterior part of the penis; and we should see them through
+its membrane, by their concave side, instead of which the convex surface
+is presented when in the vulva of females, the same as in the body of
+the males. But what is the use of these laminæ? From their figure,
+hardness, relative position with respect to each other, and their
+situation at the extremity of the penis, we cannot doubt they are real
+pincers. However, to ascertain the fact, we found it necessary to see
+their position, and that of the penis itself in the females. For this
+purpose, we prevented some of the queens from extracting the parts left
+by the impregnating males, and by dissection we discovered that the
+laminæ were pincers as we had conjectured.
+
+The penis was situated under the sting of the queens, and pressed
+against the upper region of the belly. It was supported by the posterior
+end, against the extremity of the vagina, or excretory canal. There we
+were sensible of the motion and use of the scaly pieces. Their
+extremities were separated a little more than in the male, and pressed
+between them some of the female parts below the excretory canal. The
+extreme minuteness of these parts prevented us from distinguishing them
+clearly, but the effort necessary to separate and remove the penis from
+the female, satisfied us of the use of these laminæ.
+
+Inspecting a male from above, the convex side of the plates, e. e. is
+presented, and the summit of the angle formed by their origin. When in
+the body of the female, they are in the inverse position; what was above
+in the male is now below, and the extremity of the pincers directed
+upwards. This makes us suspect that in copulation the male mounts on the
+back of the female, but we are far from asserting it positively. It may
+be asked whether that part we call the penis, is the sole part
+introduced into the female during copulation? We have carefully
+investigated this, and can affirm, that it is the only one of all those
+described by M. de Reaumur, which has been found in our females. But we
+have discovered a new part that escaped both him and Swammerdam, which
+appears from the following experiment.
+
+Separating the lenticular substance from the excretory canal, where it
+was attached, we drew along with it a white body, adhering by one
+extremity, and having the other engaged in the vagina. Towards the end
+of the lentil, where the substance adhered, it appeared cylindrical,
+then it swelled, and again contracted, to dilate anew in a greater
+degree than at first; afterwards it contracted and terminated in a
+point. A powerful magnifier was required to see all this. When pulled
+from the lenticular body, the part was commonly broke, and also when
+extracted by the queens from themselves. The figure and situation seemed
+to authorise our considering it the penis itself, and the lenticular
+body only an appendage. But the last queen we examined exhibited a
+peculiarity that induced us to doubt the fact, and led us to suspect
+that this body is nothing else than the seminal fluid itself, moulded
+and coagulated in the vagina, and which from its viscosity adheres to
+the lenticular substance, and accompanies it when separated from the
+vagina. In this queen was found a little extravasated white matter, near
+the opening of the vagina. This, though at first liquid, soon coagulated
+in the air as the seminal fluid of drones does. In separating the
+lenticular body from the vagina, we drew along with it a thread which
+broke near the lentil; and seemed of too little consistence for the
+penis of a male. The lenticular bodies, found in our queens, appeared
+larger than in the males we dissected, and we have remarked with M. de
+Reaumur, that these parts are not of equal size in every male.
+
+       *       *       *       *       *
+
+_Experiment 1._--On the tenth of July, we set successively at liberty
+three virgin queens four or five days old. Two flew away several times;
+their absence was short and fruitless. The third profited better by her
+liberty; she departed thrice; the first and second time her absence was
+short; but the third lasted thirty-five minutes. She returned in a very
+different state; and in such as allowed no doubt of her employment, for
+she exhibited the part of a male that had rendered her a mother. We
+seized her wings with one hand, and in the other received the
+lenticular body, of which she had disengaged herself with her claws. The
+posterior part was armed with two pincers, e. e. shelly and elastic,
+which could be drawn asunder, and then resumed their original position.
+Towards the anterior part of the lentil appeared the fragment of the
+root of the penis; this canal had broke half a line from the lenticular
+body. We allowed the queen to enter her habitation, and adapted the
+entrance so that she could not leave it unknown to us.
+
+On the seventeenth we found no eggs in the hive; the queen was as
+slender as the first day; therefore the male, with which she had
+copulated, had not impregnated her eggs. She was again set at liberty;
+after twice departing, she returned with evidence of a second
+copulation. We then confined her, and the eggs she afterwards laid
+proved that the second copulation had been more successful than the
+first and that there are some males more fit for impregnating queens
+than others. However, it is very rare that the first copulation is
+inefficient; we have only seen two that required it twice; all the rest
+were impregnated by the first.
+
+       *       *       *       *       *
+
+_Experiment 2._--On the eighteenth we put at liberty a virgin queen
+twenty-seven days old, she departed twice. Her second absence was
+twenty-eight minutes, and she returned with the proofs of copulation. We
+prevented her from entering, and put her under a glass to see how she
+would disengage the male organs. This she was unable to accomplish,
+having only the table and sides of the glass for support; therefore we
+introduced a bit of comb; thus providing the same conveniences as are in
+a hive. Fixing herself on _it_ by the first four legs, she stretched out
+the two last, and extending them along her belly seemed to press it
+between them. At length introducing her claws between the two parts of
+the last ring, she seized the lenticular body, and dropped it on the
+table. The posterior part was provided with shelly pincers, under which
+and in the same direction was a grey cylindrical body. The end farthest
+from the lentil was sensibly thicker than that adhering to it, and
+terminated in a point. This point was double, and open like the bill of
+a bird, which induces us to think the body was broken, a conjecture
+supported by the following experiment.
+
+       *       *       *       *       *
+
+_Experiment 3._--On the nineteenth we set at liberty a queen four days
+old; she departed twice; her first absence was short; the second lasted
+thirty minutes, and then she returned with the marks of fecundation. As
+we wished to obtain the male organs entire, it was necessary to prevent
+the queen from breaking them by extracting them with her feet; we
+therefore suddenly killed her, and cut off the last rings in order to
+lay the vulva open. But though deprived of animation, so much life
+remained in these parts that the lenticular body was thrown out
+spontaneously. Under the pincers appeared the remnant of a cylindrical
+body which had broken near the origin and remained in the female. This
+body was very small at the origin; it afterwards sensibly enlarged; next
+contracting by degrees, it terminated in a sharp point. We found the
+point engaged up to the gland in the excretory canal, and the rest in
+the vulva.
+
+       *       *       *       *       *
+
+_Experiment 4._--We set two virgin queens at liberty on the twentieth.
+The first had been abroad on the preceding days, but the scarcity of
+males prevented her from being previously fecundated. She returned with
+the organs of a male. We tried to prevent her from extracting them, but
+she did this so expeditiously with her feet, that we could not
+accomplish it. She was then allowed to enter the hive.
+
+The second queen departed twice. Her first absence was short as usual;
+the second lasted about half an hour, and she returned impregnated.
+Having killed her as suddenly as possible, we laid open the vulva. The
+lenticular body was deposited as in every queen hitherto dissected; the
+pincers were situated under the excretory canal. Some parts not easily
+distinguishable were pressed between the laminæ, and their office seemed
+to consist in forcing the extremity of the lentil to approach the
+orifice of the vagina, and apply so forcibly to it that some exertion
+was necessary to separate them. We previously examined them, with a very
+powerful magnifier. Then a peculiarity which had escaped us was
+perceptible. In drawing out the lenticular body, there proceeded from
+the vagina a minute part, v. adhering to the posterior end of the
+lentil, and situated below the plates. It spontaneously retracted into
+the lentil, like the horns of a snail. It appeared white, very short,
+and cylindrical. Under the pincers was a little half coagulated seminal
+fluid at the bottom of the vulva. Though much could be expressed, there
+was none pure; it was almost liquid, but soon coagulated, and formed a
+whitish inorganic mass. This observation carefully made removed all our
+doubts, and demonstrated that what we had taken for the penis of males
+was nothing but the seminal fluid, which had coagulated and assumed the
+interior figure of the vagina. The only hard part introduced by the
+male, was the short cylindrical point which retracted into the lentil,
+when we separated it. Its situation and office prove that it is there we
+must look for the issue of the seminal fluid, if we can hope to find an
+opening, when not engaged in copulation.
+
+We found this new part in the first drone we dissected. By pressing the
+seminal vessels, the white liquid then escaped downwards to the root of
+the penis r. and into the lenticular body, l. i. which became sensibly
+swoln. We prevented the fluid from returning, and by new pressure of the
+lentil forced it to advance. However, none escaped, but we saw at the
+posterior end of the lenticular body, and under the scaly pincers, a
+small white cylindrical substance, the same in appearance as that we had
+found engaged in the vagina of the queen. This part retracted on
+pressure, and then returned.
+
+I request you, Sir, while perusing this letter, to inspect the figure of
+the male sexual organs published by M. de Reaumur, and which are copied
+here. The descriptions are most accurate, and present a just idea of the
+situation of these parts when in the male's body. We readily conceive
+how they appear when left in the female by copulation. This detail will
+sufficiently indicate the situation and figure of the new part I have
+discovered.
+
+I suspect that the males perish after losing their sexual organs. But
+why does nature exact so great a sacrifice? This is a mystery which I
+cannot pretend to unveil. I am unacquainted with any analogous fact in
+natural history, but as there are two species of insects whose
+copulation can take place only in the air, namely, ephemeræ and ants, it
+would be extremely interesting to discover whether their males also
+lose their sexual parts, in the same circumstances, and whether, as with
+drones, enjoyment in their flight is the prelude of death.
+
+FINIS.
+
+
+FOOTNOTES:
+
+{O} Memoires sur les Abeilles, p. 450.
+
+{P} Such long and minute descriptions can be very imperfectly
+translated; indeed they are unintelligible without microscopical
+inspections of the parts themselves.--T.
+
+
+
+
+ANALYTICAL INDEX.
+
+
+  Description of a hive invented by the author                   page 4
+  Swammerdam's opinion on the fecundation of bees                     8
+  Sentiments of M. de Reaumur                                        10
+  Mr Debraw's opinion                                                11
+  Hattorf's opinion                                                  19
+  Difficulty of discovering the mode of impregnation                 22
+  Experiments on the subject                                         23
+  Suggestions by M. Bonnet                                           34
+  The queen is impregnated by copulation, which never takes place
+    within the hive                                                  41
+  Experiments on artificial fecundation have not succeeded           42
+  The male loses the sexual organs in copulation                     43
+  Regarded impregnation affects the ovaries of the queen             45
+  She then lays no eggs but those producing males                    47
+  One copulation impregnates all the eggs the queen will lay in
+    two years                                                        54
+  Fecundity of a queen                                               63
+  Common bees do not transport the queen's eggs                      66
+  They sometimes eat them                                            69
+  Eggs producing males are sometimes laid in royal cells             71
+  Common worms may be converted into queens                          77
+  Operations of the bees when this is done                           78
+  Fertile workers sometimes exist                                    89
+  They lay none but the eggs of males                                96
+  All common bees are originally females                             98
+  Receiving the royal food while larvæ, expands their ovaries       105
+  Mutual enmity of queens                                           110
+  The common bees seem to promote their combats                     117
+  A guard is constantly at the entrance of the hive                 123
+  What ensues when bees lose their queen                            126
+  Effects of introducing a stranger queen                           128
+  Massacre of the males                                             132
+  It never ensues in hives deprived of queens                       135
+  A plurality of queens is never tolerated                          142
+  The queen bee is oviparous                                        149
+  Bees seem occasionally to repose                                  150
+  Interval between production of the egg and the perfect state
+    of bees                                                         151
+  Mode of spinning the coccoon                                      153
+  That of the queen is open at one end                              154
+  The size of the bees is not affected by that of the cells         167
+  The old queen always conducts the first swarm                     173
+  But never before depositing eggs in the royal cells               177
+  Singular effect of a sound emitted by perfect queens              189
+  The instinct of bees is affected during the period of swarming    208
+  Queens are liberated from their cells according to their age      214
+  The bees probably judge of this by the sound emitted              217
+  Young queens conducting swarms are virgins                        221
+  The conduct of bees to old queens is peculiar                     224
+  Retarded impregnation affects the instinct of queens              241
+  Amputation of the antennæ produces singular effects               245
+  Advantages of the leaf hive                                       253
+  It renders the bees tractable                                     256
+  They may there be forced to work in wax                           264
+  Uniform distance between the combs                                265
+  Natural heat of bees                                              269
+  Distance to which they fly                                        271
+  Appendix                                                          273
+  Anatomical observations on the sexual organs of bees              276
+  Experiments proving the copulation of the queen                   290
+
+ALEX. SMELLIE, Printer.
+
+
+
+
+{Transcriber's notes
+
+The spelling in the original is sometimes idiosyncratic. It has not
+been changed, but a few obvious errors have been corrected. The
+corrections are listed below.
+
+Inconsistent spellings include:
+  Lusace/Lusaçe, centre/center, choose/chuse,
+  organisation/organization, recognise/recognize
+
+Unusual spellings (which have not been changed) include:
+  centinels, coccoon, diaphraghm, encreased, encreasing, groupes,
+  harrassed, inaccessible, incontestible, indispensible, moveable,
+  perceptible, susceptible, uncontrouled, unintelligible
+
+Letter I
+
+  "secret distinctive characterestics" changed to
+  "secret distinctive characteristics"
+
+Letter II
+
+  "the copulalation of queens" changed to
+  "the copulation of queens"
+
+Letter IV
+
+  "The worms had spun their silk coccons" changed to
+  "The worms had spun their silk coccoons"
+
+Letter V
+
+  "characteristics of commo nbees" changed to
+  "characteristics of common bees"
+
+Letter VI
+
+  "The result of this rencounter" changed to
+  "The result of this encounter"
+
+  "genius such as your's" unchanged.
+
+  "observing that the antennae" changed to
+  "observing that the antennæ"
+
+  "combats and disastrou scenes" changed to
+  "combats and disastrous scenes"
+
+  "M. de Reamur speaks of these executions" changed to
+  "M. de Reaumur speaks of these executions"
+
+Letter IX
+
+  "Only the few bees that not participated" changed to
+  "Only the few bees that had not participated"
+
+Letter XI
+
+  "these tumultous motions" changed to
+  "these tumultuous motions"
+
+Letter XII
+
+  "one antennæ" unchanged.
+
+  "reside in them," changed to
+  "reside in them."
+
+Appendix
+
+  "the cirumference is edged" changed to
+  "the circumference is edged"
+
+  "he could have proportioned the tortous canal" changed to
+  "he could have proportioned the tortuous canal"
+
+  "pressed between the laminae" changed to
+  "pressed between the laminæ"
+}
+
+
+
+
+
+End of the Project Gutenberg EBook of New observations on the natural
+history of bees, by Francis Huber
+
+*** END OF THIS PROJECT GUTENBERG EBOOK NEW OBSERVATIONS ON BEES ***
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+<pre>
+
+The Project Gutenberg EBook of New observations on the natural history of
+bees, by Francis Huber
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: New observations on the natural history of bees
+
+Author: Francis Huber
+
+Translator: Anonymous
+
+Release Date: August 28, 2008 [EBook #26457]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW OBSERVATIONS ON BEES ***
+
+
+
+
+Produced by Louise Pryor, Steven Giacomelli and the Online
+Distributed Proofreading Team at https://www.pgdp.net (This
+file was produced from images produced by Core Historical
+Literature in Agriculture (CHLA), Cornell University)
+
+
+
+
+
+
+</pre>
+
+
+<div class="transnote">
+<h4>Transcriber's note</h4>
+
+<p>The spelling in the original is sometimes idiosyncratic. It has not
+been changed, but a few obvious errors have been corrected. The
+corrections are listed at the <a href="#tnotes">end of this etext</a>  and marked with a <span class="correction" title="like this">mouse-hover</span>.
+</p>
+
+<p>The four figures appear in a single illustration in the original.
+In this etext they also appear close to the text that refers to them: <a href="#fig1">Fig 1</a>, <a href="#fig2">Fig 2</a>, <a href="#fig3">Fig 3</a>, <a href="#fig4">Fig 4</a>.</p>
+</div>
+
+<div class="figcenter" style="width: 727px;">
+<a  id="figmain" name="figmain"></a>
+<img src="images/main-thumb.png" width="727" height="514" alt="The four figures referred to in the text" />
+<span class="caption">Figures 1 to 4</span>
+</div>
+
+<h1><span class="littler">NEW OBSERVATIONS</span><br />
+<span class="littlest">ON THE</span><br />
+<span class="little">NATURAL HISTORY</span><br />
+<span class="littlest">OF</span><br />
+BEES,<br />
+
+<span class="littlest">BY</span><br />
+
+<span class="little">FRANCIS HUBER.</span><br />
+
+<span class="littlest">TRANSLATED FROM THE ORIGINAL.</span></h1>
+
+<hr class="tiny" />
+
+
+<p class="center">EDINBURGH:<br />
+
+<span class="littler" >PRINTED FOR JOHN ANDERSON,<br />
+AND SOLD BY<br />
+LONGMAN, HURST, REES, AND ORME,<br />
+LONDON.</span><br />
+
+<span class="littlest">ALEX SMELLIE, Printer.</span><br />
+
+<span class="littlest bt">1806.</span></p>
+
+
+
+
+ <p class="center biggap dedication"> To<br />
+
+  <span class="bigger">SIR JOSEPH BANKS, Bart.</span><br />
+
+  KNIGHT OF THE MOST HONOURABLE ORDER<br />
+  OF THE BATH, A PRIVY COUNCILLOR,<br />
+  PRESIDENT OF THE ROYAL<br />
+  SOCIETY OF LONDON,<br />
+  &amp;c. &amp;c.<br />
+
+  THIS TRANSLATION<br />
+  IS INSCRIBED.</p>
+
+
+
+
+
+
+<h2>
+<span class="pagebreak" title="vii">&nbsp;</span><a name="Page_vii" id="Page_vii"></a>
+<a name="CONTENTS" id="CONTENTS"></a>CONTENTS.</h2>
+
+
+
+<div class="center">
+<table width="60%" summary="table of contents">
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;1.&mdash;On the impregnation of the queen bee         </td><td class="toc2">page&nbsp;<a href="#Page_1">1</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;2.&mdash;Sequel of observations on the impregnation
+             of the queen bee                                 </td><td class="toc2"><a href="#Page_41">41</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;3.&mdash;The same subject continued; observations
+             on retarding the fecundation of queens           </td><td class="toc2"><a href="#Page_44">44</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;4.&mdash;On M. Schirach's discovery                      </td><td class="toc2"><a href="#Page_76">76</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;5.&mdash;Experiments proving that there are
+             sometimes common bees which lay fertile eggs     </td><td class="toc2"><a href="#Page_89">89</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;6.&mdash;On the combats of queens; the massacre
+             of the males; and what succeeds in a hive
+             where a stranger queen is substituted for
+             the natural one                                 </td><td class="toc2"><a href="#Page_108">108</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;7.&mdash;Sequel of observations on the reception
+             of a stranger queen; M. de Reaumur's
+             observations on the subject                     </td><td class="toc2"><a href="#Page_137">137</a></td></tr>
+
+
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;8.&mdash;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 coccoons </td><td class="toc2"><a href="#Page_145">145</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;9.&mdash;On the formation of swarms                     </td><td class="toc2"><a href="#Page_171">171</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;10.&mdash;The same subject continued                    </td><td class="toc2"><a href="#Page_201">201</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;11.&mdash;The same subject continued                    </td><td class="toc2"><a href="#Page_223">223</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;12.&mdash;Additional observations on queens
+             that lay only the eggs of drones, and on
+             those deprived of the antenn&aelig;                   </td><td class="toc2"><a href="#Page_237">237</a></td></tr>
+
+<tr><td class="toc1"><span class="little">LETTER</span>&nbsp;13.&mdash;Economical considerations on bees             </td><td class="toc2"><a href="#Page_253">253</a></td></tr>
+
+<tr><td class="toc1 smcap">Appendix                                                  </td><td class="toc2"><a href="#Page_275">275</a></td></tr>
+
+</table>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="ix">&nbsp;</span><a name="Page_ix" id="Page_ix"></a>
+<a name="TRANSLATORS_PREFACE" id="TRANSLATORS_PREFACE"></a>TRANSLATOR'S PREFACE</h2>
+
+
+<p>The facts contained in this volume are deeply interesting to the
+Naturalist. They not only elucidate the history of those industrious
+animals, whose nature is the peculiar subject of investigation, but they
+present some singular features in physiology which have hitherto been
+unknown.</p>
+
+<p>The industry of bees has proved a fertile source of admiration in all
+countries and in every age; and mankind have endeavoured to render it
+subservient to their gratifications or emolument. Hence innumerable
+theories, experiments, and observations have ensued, and uncommon
+patience has been displayed in prosecuting
+<span class="pagebreak" title="x">&nbsp;</span><a name="Page_x" id="Page_x"></a>
+the enquiry. But although
+many interesting peculiarities have been discovered, they are so much
+interwoven with errors, that no subject has given birth to more
+absurdities than investigations into the history of bees: and
+unfortunately those treatises which are most easily attained, and the
+most popular, only serve to give such absurdities a wider range, and
+render it infinitely more difficult to eradicate them. A considerable
+portion of the following work is devoted to this purpose. The reader
+will judge of the success which results from the experiments that have
+been employed.</p>
+
+<p>Perhaps this is not the proper place to bestow an encomium on a treatise
+from which so much entertainment and instruction will be derived.
+However, to testify the estimation in which it is held in other nations,
+the remarks upon it by the French philosopher Sue, may be quoted, 'The
+observations are so consistent, and the consequences seem so just, that
+while perusing this work, it appears as if we
+<span class="pagebreak" title="xi">&nbsp;</span><a name="Page_xi" id="Page_xi"></a>
+had assisted the author
+in each experiment, and pursued it with equal zeal and interest. Let us
+invite the admirers of nature to read these observations; few are equal
+to them in excellence, or so faithfully describe the nature, the habits,
+and inclinations of the insects of which they treat.'</p>
+
+<p>It is a remarkable circumstance that the author laboured under a defect
+in the organs of vision, which obliged him to employ an assistant in his
+experiments. Thus these discoveries may be said to acquire double
+authority. But independent of this the experiments are so judiciously
+adapted to the purposes in view, and the conclusions so strictly
+logical, that there is evidently very little room for error. The talents
+of <i>Francis Burnens</i>, this philosophic assistant, had long been devoted
+to the service of the author, who, after being many successive years in
+this manner aided in his researches, was at last deprived of him by some
+unfortunate accident.</p>
+
+<p>
+<span class="pagebreak" title="xii">&nbsp;</span><a name="Page_xii" id="Page_xii"></a>
+Whether the author has prosecuted his investigation farther does not
+appear, as no other production of his pen is known in this island.</p>
+
+<p>It is vain to attempt a translation of any work without being to a
+certain degree skilled in the subject of which it treats. Some parts of
+the original of the following treatise, it must be acknowledged, are so
+confused, and some so minute, that it is extremely difficult to give an
+exact interpretation. But the general tenor, though not elegant, is
+plain and perspicuous; and such has it been here retained.</p>
+
+
+
+<h2>
+<span class="pagebreak" title="1">&nbsp;</span><a name="Page_1" id="Page_1"></a>
+<a name="LETTER_I" id="LETTER_I"></a>LETTER I.<br /><br />
+
+<span class="little italic">ON THE IMPREGNATION OF THE QUEEN BEE.</span></h2>
+
+
+<p class="salutation">SIR,</p>
+
+<p>When I had the honour at Genthod of giving you an account of my
+principal experiments on bees, you desired me to transmit a written
+detail, that you might consider them with greater attention. I hasten,
+therefore, to extract the following observations from my journal.&mdash;As
+nothing can be more flattering to me than the interest you take in my
+researches, permit me to
+<span class="pagebreak" title="2">&nbsp;</span><a name="Page_2" id="Page_2"></a>
+remind you of your promise to suggest new
+experiments<a name="fnm_A" id="fnm_A"></a><a href="#fn_A" class="fnnum">A</a>.</p>
+
+<p>After having long studied bees in glass hives constructed on M. de
+Reaumur's principle, you have found the form unfavourable to an
+observer. The hives being too wide, two parallel combs were made by the
+bees, consequently whatever passed between them escaped observation.
+From this inconvenience, which I have experienced, you recommended much
+thinner hives to naturalists, where the panes should be so near each
+other, that only a single row of combs could be erected between them. I
+have followed your admonitions, Sir, and provided hives only eighteen
+lines in width, in which I have found no difficulty to establish swarms.
+However, bees must not be entrusted with the charge of constructing a
+single comb: Nature has taught them to make parallel ones, which
+<span class="pagebreak" title="3">&nbsp;</span><a name="Page_3" id="Page_3"></a>
+is a
+law they never derogate from, unless when constrained by some particular
+arrangement. Therefore, if left to themselves in these thin hives, as
+they cannot form two combs parallel to the plane of the hive, they will
+form several small ones perpendicular to it, and, in that case, all is
+equally lost to the observer. Thus it became essential previously to
+arrange the position of the combs. I forced the bees to build them
+perpendicular to the horizon, and so that the lateral surfaces were
+three or four lines from the panes of the hive. This distance allows the
+bees sufficient liberty, but prevents them from collecting in too large
+clusters on the surface of the comb. By such precautions, bees are
+easily established in very thin hives. There they pursue their labours
+with the same assiduity and regularity; and, every cell being exposed,
+none of their motions can be concealed.</p>
+
+<p>It is true, that by compelling these insects to a habitation where they
+could construct
+<span class="pagebreak" title="4">&nbsp;</span><a name="Page_4" id="Page_4"></a>
+only a single row of combs, I had, in a certain
+measure, changed their natural situation, and this circumstance might
+possibly have affected their instinct. Therefore, to obviate every
+objection, I invented a kind of hives, which, without losing the
+advantages of those very thin, at the same time approached the figure of
+common hives where bees form several rows of combs.</p>
+
+<p>I took several small fir boxes, a foot square and fifteen lines wide,
+and joined them together by hinges, so that they could be opened and
+shut like the leaves of a book<a name="fnm_B" id="fnm_B"></a><a href="#fn_B" class="fnnum">B</a>. When using a hive of this
+description,
+<span class="pagebreak" title="5">&nbsp;</span><a name="Page_5" id="Page_5"></a>
+we took care to fix a comb in each frame, and then
+introduced all the
+<span class="pagebreak" title="6">&nbsp;</span><a name="Page_6" id="Page_6"></a>
+bees necessary for each particular experiment. By
+opening the different divisions successively, we daily inspected both
+surfaces of every comb. There was not a single cell where we could not
+distinctly see what passed at all times, nor a single bee, I may almost
+say, with which we were not particularly acquainted. Indeed, this
+construction is nothing more than the union of several very flat hives
+which may be separated. Bees, in such habitations, must not be visited
+before their combs are securely fixed in the frames, otherwise, by
+falling out, they may kill or hurt them, as also irritate them to that
+degree that the observer cannot escape stinging, which is always
+painful, and sometimes dangerous: but they soon become accustomed to
+their situation, and in some measure tamed by it; and, in three days, we
+may begin to operate on the
+<span class="pagebreak" title="7">&nbsp;</span><a name="Page_7" id="Page_7"></a>
+hive, to open it, remove part of the combs,
+and substitute others, without the bees exhibiting too formidable
+symptoms of displeasure. You will remember, Sir, that on visiting my
+retreat, I shewed you a hive of this kind that had been a long time in
+experiment, and how much you were surprised that the bees so quietly
+allowed us to open it.</p>
+
+<p>In these hives, I have repeated all my observations, 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
+inconvenience 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 error; and it also appears, that some
+advantages are found in these which may be called <i>Book</i> or
+<i>Leaf-hives</i>, as they prove extremely useful in the economical treatment
+of bees, which shall afterwards be detailed.</p>
+
+<p>
+<span class="pagebreak" title="8">&nbsp;</span><a name="Page_8" id="Page_8"></a>
+I now come to the particular object of this letter, the fecundation of
+the queen bee; and I shall, in a few words, examine the different
+opinions of naturalists on this singular problem. Next I shall state the
+most remarkable observations which their conjectures have induced me to
+make, and then describe the new experiments by which I think I have
+solved the problem<a name="fnm_C" id="fnm_C"></a><a href="#fn_C" class="fnnum">C</a>.</p>
+
+<p>Swammerdam, who studied bees with unremitting attention, and who never
+could see a real copulation between a drone and a queen, was satisfied
+that copulation was unnecessary for fecundation of the eggs: but having
+remarked that, at certain times, the drones exhaled a very strong odour,
+
+<span class="pagebreak" title="9">&nbsp;</span><a name="Page_9" id="Page_9"></a>
+he thought this odour was an emanation of the <i>aura seminalis</i>, or the
+<i>aura seminalis</i> itself, which operated fecundation by penetrating the
+body of the female. His conjecture was confirmed on dissecting the male
+organs of generation; for he was so much struck with the disproportion
+between them and those of the female, that he did not believe copulation
+possible. His opinion, concerning the influence of the odour, had this
+farther advantage, that it afforded a good reason for the prodigious
+number of the males. There are frequently fifteen hundred or two
+thousand in a hive; and, according to Swammerdam, it is necessary they
+should be numerous, that the emanation proceeding from them may have an
+intensity or energy sufficient to effect impregnation.</p>
+
+<p>Though M. de Reaumur has refuted this hypothesis by just and conclusive
+reasoning, he has failed to make the sole experiment that could support
+or overturn it. This was to confine all the drones of a
+<span class="pagebreak" title="10">&nbsp;</span><a name="Page_10" id="Page_10"></a>
+hive in a tin
+case, perforated with minute holes, which might allow the emanation of
+the odour to escape, but prevent the organs of generation from passing
+through. Then, this case should have been placed in a hive well
+inhabited, but completely deprived of males, both of large and small
+size, and the consequence attended to. It is evident, 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
+confuted had she produced no eggs, or only sterile ones. However the
+experiment has been made by us, and the queen remained barren;
+therefore, it is undoubted, that the emanation of the odour of the males
+does not impregnate bees.</p>
+
+<p>M. de Reaumur was of a different opinion. He thought that the queen's
+fecundation followed actual copulation. He confined several drones in a
+glass vessel along with a virgin queen: he saw the female make many
+advances to the males;
+<span class="pagebreak" title="11">&nbsp;</span><a name="Page_11" id="Page_11"></a>
+but, unable to observe any union so intimate
+that it could be denominated copulation, he leaves the question
+undecided. We have repeated this experiment: we have frequently confined
+virgin queens with drones of all ages: we have done so at every season,
+and witnessed all their advances and solicitations to the males: we have
+even believed we saw a kind of union between them, but so short and
+imperfect that it was unlikely to effect impregnation. Yet, to neglect
+nothing, we confined the virgin queen, that had suffered the approaches
+of the male, to her hive. During a month that her imprisonment
+continued, she did not lay a single egg; therefore, these momentary
+junctions do not accomplish fecundation.</p>
+
+<p>In the <i>Contemplation de la Nature</i>, you have cited the observations of
+the English naturalist Mr Debraw. They appear correct, and at last to
+elucidate the mystery. Favoured by chance, the observer one day
+perceived at the bottom of cells containing
+<span class="pagebreak" title="12">&nbsp;</span><a name="Page_12" id="Page_12"></a>
+eggs, a whitish fluid,
+apparently spermatic, at least, very different from the substance or
+jelly which bees commonly collect around their new hatched worms.
+Solicitous to learn its origin, and conjecturing that it might be the
+male prolific fluid, he began to watch the motions of every drone in the
+hive, on purpose to seize the moment when they would bedew the eggs. He
+assures us, that he saw several insinuate the posterior part of the body
+into the cells, and there deposit the fluid. After frequent repetition
+of the first, he entered on a long series of 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 no brood.
+He saw the queen lay eggs, which were bedewed by the males, and from
+which larv&aelig; were hatched, consequently, he could not hesitate advancing
+as a fact demonstrated, that male bees fecundate the queen's eggs in
+
+<span class="pagebreak" title="13">&nbsp;</span><a name="Page_13" id="Page_13"></a>
+the manner of frogs and fishes, that is, after they are produced.</p>
+
+<p>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 number of males in a hive. At the same time,
+the author had neglected to answer one strong objection. Larv&aelig; appear
+when there are no drones. From the month of September until April, hives
+are generally destitute of males, yet, notwithstanding their absence,
+the queen then lays fertile eggs. Thus, the prolific fluid cannot be
+required to impregnate them, unless we can suppose that it is necessary
+at a certain time of the year, while at every other season it is
+useless.</p>
+
+<p>To discover the truth amidst these facts apparently so contradictory, I
+wished to repeat Mr Debraw's experiments, and to observe more precaution
+than he himself had done. First, I sought for the fluid, which he
+supposes the seminal, in cells
+<span class="pagebreak" title="14">&nbsp;</span><a name="Page_14" id="Page_14"></a>
+containing eggs. Several were actually
+found with that appearance; and, during the first days of observation,
+neither my assistant nor myself doubted the reality of the discovery.
+But we afterwards found it an illusion arising from the reflection of
+the light, for nothing like a fluid was visible, except when the solar
+rays reached the bottom of the cells. Fragments of the coccoons of
+worms, successively hatched, commonly cover the bottom; and, as they are
+shining, it may easily be conceived that, when much illuminated, an
+illusory effect results from the light. We proved it by the strictest
+examination, for no vestiges of a fluid were perceptible when the cells
+were detached and cut asunder.</p>
+
+<p>Though the first observation inspired us with some distrust of Mr
+Debraw's discovery, we repeated his other experiments with the utmost
+care. On the 6. of August 1787, we immersed a hive, and, with scrupulous
+attention, examined the whole bees while in the bath. We ascertained
+
+<span class="pagebreak" title="15">&nbsp;</span><a name="Page_15" id="Page_15"></a>
+that there was no male, either large or small; and having examined all
+the combs, we found neither male nymph, nor worm. When the bees were
+dry, we replaced them all, 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 necessary that no male should enter the hive during the
+experiment, a glass tube was adapted 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 removed any male that appeared,
+that the result of the experiment might be undisturbed, and I can
+positively affirm that not one was seen. However, from the first day,
+which was the sixth of August, the queen deposited fourteen eggs in the
+workers cells; and all these
+<span class="pagebreak" title="16">&nbsp;</span><a name="Page_16" id="Page_16"></a>
+were hatched on the tenth of the same
+month.</p>
+
+<p>This experiment is decisive, since the eggs laid by the queen of a hive
+where there were no males, and where it was impossible one could be
+introduced, since these eggs, I say, were fertile, it becomes
+indubitable that the fluid of the males is not required for their
+exclusion.</p>
+
+<p>Though it did not appear that any reasonable objection could be started
+against this conclusion, yet, as I had been accustomed in all my
+experiments to seek for the most trifling difficulties that could arise,
+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 fluid to their own habitations for depositing it
+on the eggs.</p>
+
+<p>It was easy to appreciate the force of this objection, for all that was
+necessary was a repetition of the former experiments, and to confine the
+bees so closely to
+<span class="pagebreak" title="17">&nbsp;</span><a name="Page_17" id="Page_17"></a>
+their hives that none could possibly escape. You
+very well know, Sir, that these animals can live three or four months
+confined in a hive well stored with honey and wax, and if apertures are
+left for circulation of the air. This experiment 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 larv&aelig;.</p>
+
+<p>I extended the precautions so far as to immerse this hive a second time,
+to assure myself that no male had escaped my researches. 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.</p>
+
+<p>In terminating the confutation of Mr Debraw's opinion, I have only to
+explain what led him into error; and that was, his using queens whose
+history he was unacquainted with from their origin. When
+<span class="pagebreak" title="18">&nbsp;</span><a name="Page_18" id="Page_18"></a>
+he observed
+the eggs produced by a queen, confined along with males, were fertile,
+he thence concluded that they had been bedewed by the prolific fluid in
+the cells: but to render his conclusion just, he should first have
+ascertained that the female never had copulated, and this he neglected.
+The truth is, that, without knowing it, he had used, in his experiments,
+a queen after she had commerce with the male. Had he taken a virgin
+queen the moment she came from the royal cell, and confined her along
+with drones in his vessels, the result would have been opposite; for,
+even amidst a seraglio of males, this young queen would never have laid,
+as I shall afterwards prove.</p>
+
+<p>The Lusatian observers, and M. Hattorf in particular, thought the queen
+was fecundated by herself, without concourse with the males. I shall
+here give an abstract of the experiment on which that opinion is
+founded.<a name="fnm_D" id="fnm_D"></a><a href="#fn_D" class="fnnum">D</a></p>
+
+<p>
+<span class="pagebreak" title="19">&nbsp;</span><a name="Page_19" id="Page_19"></a>
+M. Hattorf took a queen whose virginity he could not doubt. He excluded
+all the males both of the large and small species, and, in several days,
+he 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
+absent, the queen laid eggs, from which came worms: whence he considers
+she is impregnated by herself.</p>
+
+<p>Reflecting on this experiment, I do not find it sufficiently accurate.
+Males pass with great facility from hive to hive; and M. Hattorf took no
+precaution that none was introduced into his. He says, indeed, there was
+no male, but is silent respecting the means he adopted to prove the
+fact. Though he might be satisfied of no large drone being there, still
+a small one might have escaped his vigilance, and fecundated the queen.
+With a view to clear up the
+<span class="pagebreak" title="20">&nbsp;</span><a name="Page_20" id="Page_20"></a>
+doubt, I resolved to repeat his experiment,
+in the manner described, and without greater care or precaution.</p>
+
+<p>I put a virgin queen into a hive, from which all the males were
+excluded, but the bees left at perfect liberty. For several days I
+visited the hive, and found new hatched worms in it. Here then is the
+same result as M. Hattorf obtained? But before deducing the same
+consequence from it, we had to ascertain beyond dispute that no male had
+entered the hive. Thus, it was necessary to immerse the bees, and
+examine each separately. By this operation, we actually found four small
+males. Therefore, to render the experiment decisive, not only was it
+requisite to remove all the drones, but also, by some infallible method,
+to prevent any from being introduced, which the German naturalist had
+neglected.</p>
+
+<p>I prepared to repair this omission, by putting a virgin queen into a
+hive, from which the whole males were carefully removed;
+<span class="pagebreak" title="21">&nbsp;</span><a name="Page_21" id="Page_21"></a>
+and to be
+physically certain that none should enter, a glass tube was adapted at
+the entrance of such dimensions that the working bees could freely pass
+and repass, but too narrow for the smallest male. Matters continued thus
+for thirty days, the workers departing and returning performed their
+usual labours: but the queen remained sterile. At the expiration of this
+time, her belly was equally slender as at the moment of her origin. I
+repeated the experiment several times, and always with the same
+consequence.</p>
+
+<p>Therefore, as a queen, rigorously separated from all commerce with the
+male, remains sterile, it is evident she cannot impregnate herself, and
+M. Hattorf's opinion is ill-founded.</p>
+
+<p>Hitherto, by endeavouring to confute or verify the conjectures of all
+the authors who had preceded me, by new experiments, I acquired the
+knowledge of new facts, but these were apparently so contradictory as to
+render the solution of the problem still
+<span class="pagebreak" title="22">&nbsp;</span><a name="Page_22" id="Page_22"></a>
+more difficult. While
+examining Mr. Debraw's hypothesis, I confined a queen in a hive, from
+which all the drones were removed; the queen nevertheless was fertile.
+When considering the opinion of M. Hattorf on the contrary, I put a
+queen, of whose virginity I was perfectly satisfied, in the same
+situation, she remained sterile.</p>
+
+<p>Embarrassed by so many difficulties, I was on the point of abandoning
+the subject of my researches, when at length by more attentive
+reflection, I thought these contradictions might arise from experiments
+made indifferently on virgin queens, and on those with whose history I
+was not acquainted from the origin, and which had perhaps been
+impregnated unknown to me. Impressed with this idea, I undertook a new
+method of observation not on queens fortuitously taken from the hive,
+but on females decidedly in a virgin state, and whose history I knew
+from the instant they left the cell.</p>
+
+<p>
+<span class="pagebreak" title="23">&nbsp;</span><a name="Page_23" id="Page_23"></a>
+From a very great number of hives, I removed all the virgin females,
+and substituted for each a queen taken at the moment of her birth. The
+hives were then divided into two classes. From the first, I took the
+whole males both large and small, and adapted a glass tube at the
+entrance, so narrow, 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 all the drones belonging to them, and even introduced more; and to
+prevent them from escaping, a glass tube, also too narrow for the males,
+was adapted to the entrance of these hives.</p>
+
+<p>For more than a month, I carefully watched this experiment, made on a
+large scale; but much to my surprise, all the queens remained sterile.
+Thus it was proved, that queens confined in a hive would continue barren
+though amidst a seraglio of males.</p>
+
+<p>This result induced me to suspect that the females could not be
+fecundated in the
+<span class="pagebreak" title="24">&nbsp;</span><a name="Page_24" id="Page_24"></a>
+interior of the hive, and that it was necessary 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 29. June 1788.</p>
+
+<p>Aware, that in summer the males usually leave the hive at the warmest
+time of the day, it was natural for me to conclude that if the queens
+were also obliged to go out for impregnation, instinct would induce them
+to do so at the same time as the males.</p>
+
+<p>At eleven in the forenoon, we placed ourselves opposite a hive
+containing an unimpregnated queen five days old. The sun had shone from
+his rising; the air was very warm; and the males began to leave the
+hives. We then enlarged the entrance of that which we wished to observe,
+and paid great attention to the bees that entered and departed. The
+males appeared, and immediately took flight. Soon afterwards, the
+<span class="pagebreak" title="25">&nbsp;</span><a name="Page_25" id="Page_25"></a>
+young
+queen appeared at the entrance; at first she did not fly, but brushed
+her belly with her hind legs, and traversed the board a little; neither
+workers nor males paid any attention to 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 departure, perhaps judging this precaution
+necessary to recognize it; she then flew away, describing horizontal
+circles twelve or fifteen feet above the earth. We contracted the
+entrance 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 observe all her motions. But she did not remain
+long in a situation favourable for us, and rapidly rose out of sight. We
+resumed our place before the hive; and in seven minutes, the young queen
+returned to the entrance of a habitation which she had left for the
+first time. Having found no external appearance of fecundation, we
+allowed her to enter.
+<span class="pagebreak" title="26">&nbsp;</span><a name="Page_26" id="Page_26"></a>
+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. Her second
+absence was much longer than the first; twenty-seven minutes elapsed
+before she came back. We then found her in a state very different from
+that in which she was after her first excursion. The sexual organs were
+distended by a white substance, thick and hard, very much resembling the
+fluid in the vessels of the male, completely similar to it indeed in
+colour and consistence<a name="fnm_E" id="fnm_E"></a><a href="#fn_E" class="fnnum">E</a>.</p>
+
+<p>
+<span class="pagebreak" title="27">&nbsp;</span><a name="Page_27" id="Page_27"></a>
+But more evidence than mere resemblance was requisite to establish that
+the female had returned with the prolific fluid of the males. We allowed
+this queen to enter the hive, and confined her there. In two days, we
+found her belly swoln; and she had already laid near an hundred eggs in
+the worker's cells.</p>
+
+<p>To confirm our discovery, we made several other experiments, and with
+the same success. I shall continue to transcribe my journal.</p>
+
+<p>On the second of July, the weather being very fine, numbers of males
+left the hives. We set at liberty an unimpregnated young queen, eleven
+days old, whose hive
+<span class="pagebreak" title="28">&nbsp;</span><a name="Page_28" id="Page_28"></a>
+had always been deprived of males. Having quickly
+left the hive, she returned to examine it, and then rose out of sight.
+In a few minutes, she returned without any external marks of
+impregnation. In a quarter of an hour, she departed again, but her
+flight was so rapid that we could scarcely follow her a moment. This
+absence continued thirty minutes. On returning, the last ring of the
+body was open, and the sexual organs full of the whitish substance
+already mentioned. She was then replaced in the hive from which all the
+males were excluded. In two days, we found her impregnated.</p>
+
+<p>These observations at length demonstrate why M. Hattorf obtained results
+so different from ours. His queens, though in hives deprived of males,
+had been fecundated, and he thence concludes that sexual intercourse is
+not requisite for their impregnation. But he did not confine the queens
+to their hives, and they had profited by their liberty to unite with the
+males. We,
+<span class="pagebreak" title="29">&nbsp;</span><a name="Page_29" id="Page_29"></a>
+on the contrary, have surrounded our queens with a number of
+males; but they continued sterile; because the precaution of confining
+the males to their hives had also prevented the queens from departing to
+seek that fecundation without, which they could not obtain within.</p>
+
+<p>These experiments were repeated on queens, twenty, twenty-five, and
+thirty days old. All became fertile after a single impregnation;
+however, we have remarked some essential peculiarities in the fecundity
+of those unimpregnated until the twentieth day of their existence; but
+we shall defer speaking of the fact until we can present naturalists
+with observations sufficiently secure and numerous to merit their
+attention: Yet let me add a few words more. Though neither my assistant
+nor myself have witnessed the copulation of a queen and a drone, we
+think that, after the detail which has just been commenced, no doubt of
+it can remain, or of the necessity of copulation to effect impregnation.
+
+<span class="pagebreak" title="30">&nbsp;</span><a name="Page_30" id="Page_30"></a>
+The sequel of experiments, made with every possible precaution, appears
+demonstrative. The uniform sterility of queens in hives wanting males,
+and in those where they were confined along with them; the departure of
+these queens from the hives; and the very conspicuous evidence of
+impregnation with which they return, 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 copulation.</p>
+
+<p>Naturalists have always been very much embarrassed to account for the
+number of males found 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 that extent. As fecundation
+cannot be accomplished within, and as the queen is obliged to traverse
+the expanse of the atmosphere, it is requisite the males should be
+numerous that she may have the chance of meeting some one of
+<span class="pagebreak" title="31">&nbsp;</span><a name="Page_31" id="Page_31"></a>
+them. Were
+only two or three drones in each hive, there would be little probability
+of their departure at the same instant with the queen, or that they
+would meet in their excursions; and most of the females would thus
+remain sterile.</p>
+
+<p>But why has nature prohibited copulation within the hives? This is a
+secret still unknown to us. It is possible, however, that some
+favourable circumstance may enable us to penetrate it in the course of
+our observations. Various conjectures may be formed; but at this day we
+require facts, and reject gratuitous suppositions. It should be
+remembered, that bees do not form the sole republic among insects
+presenting a similar phenomenon; female ants are also obliged to leave
+the ant-hills previous to fecundation.</p>
+
+<p>I cannot request, Sir, that you will communicate the reflections which
+your genius will excite concerning the facts I have related. This is a
+favour to which I am not yet entitled. But as new experiments
+<span class="pagebreak" title="32">&nbsp;</span><a name="Page_32" id="Page_32"></a>
+will
+unquestionably occur to you, whether on the impregnation 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 encouragement that the
+continuance of my labours can receive.</p>
+
+<p class="indent1 italic">
+Pregny, 13th August 1789.
+</p>
+
+<hr />
+
+<h3>Letter from M. Bonnet to M. Huber.</h3>
+
+<p>You have most agreeably surprised me, Sir, with your interesting
+discovery of the impregnation of the queen bee. It was a fortunate idea,
+that she left the hive to be fecundated, and your method of ascertaining
+the fact was extremely judicious and well adapted to the object in view.</p>
+
+<p>Let me remind you, that male and female ants copulate in the air; and
+that after
+<span class="pagebreak" title="33">&nbsp;</span><a name="Page_33" id="Page_33"></a>
+impregnation the females return to the ant hills to deposit
+their eggs. <i>Contemplation de la Nature, Part II. chap. 22. note 1.</i> 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 a copulation in the air. If you have satisfactory evidence
+that the fluid bedewing the last rings of the female is the same with
+that of the male, it is more than mere presumption in favour of
+copulation. Perhaps it may be necessary that the male should seize the
+female under the belly, which cannot easily be done but in the air. The
+large opening at the extremity of the queen, which you have observed in
+so particular a condition, seems to correspond to the singular size of
+the sexual parts of the male.</p>
+
+<p>You wish, my dear Sir, that I should suggest some new experiments on
+these industrious republicans. In doing so, I shall take the greater
+pleasure and interest, as I know to what extent you possess the valuable
+<span class="pagebreak" title="34">&nbsp;</span><a name="Page_34" id="Page_34"></a>
+art of combining ideas, and of deducing from this combination results
+adapted to the discovery of new facts. A few at this moment occur to me.</p>
+
+<p>It may be proper to attempt the artificial fecundation of a virgin
+queen, by introducing a little of the male's prolific fluid with a
+pencil, and at the same time observing every precaution to avoid error.
+Artificial fecundation, you are aware, has already succeeded in more
+than one animal.</p>
+
+<p>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 paint the thorax with some varnish that resists humidity. It will
+also be right to paint the thorax of a considerable number of workers in
+order to discover the duration of their life. This is a more secure
+method than slight mutilations.</p>
+
+<p>For hatching the worm, the egg must be fixed almost vertically by one
+end near
+<span class="pagebreak" title="35">&nbsp;</span><a name="Page_35" id="Page_35"></a>
+the bottom of the cell. Is it true, that it is unproductive
+unless fixed in this manner? I cannot determine the fact; and therefore
+leave it to the decision of experiment.</p>
+
+<p>I formerly mentioned to you that I had long doubted the real nature of
+the small ovular substances deposited by queens in the cells, and my
+inclination to suppose them minute worms not yet begun to expand. Their
+elongated figure seems to favour my suspicions. It would therefore be
+proper to watch them with the utmost assiduity, from the instant of
+production until the period of exclusion. If the integument bursts,
+there can be no doubt that these minute substances are real eggs.</p>
+
+<p>I return to the mode of operating copulation. The height that the queen
+and the males rise to in the air prevent us from seeing what passes
+between them. On that account, the hive should be put into an apartment
+with a very lofty ceiling. M. de Reaumur's experiment of confining a
+<span class="pagebreak" title="36">&nbsp;</span><a name="Page_36" id="Page_36"></a>
+queen with several males in a glass vessel, merits repetition; and if,
+instead of a vessel, a glass tube, some inches in diameter and several
+feet long, were used, perhaps something satisfactory might be
+discovered.</p>
+
+<p>You have had the fortune to observe the small queens mentioned by the
+Abbe Needham, but which he never saw. It will be of great importance to
+dissect them for the purpose of finding their ovaries. When M. Reims
+informed me that he had confined three hundred workers, along with a
+comb containing no eggs, and afterwards found hundreds in it, I strongly
+recommended that he should dissect the workers. He did so; and informed
+me that eggs were found in three. Probably without being aware of it, he
+has dissected small queens. As small drones exist, it is not surprising
+if small queens are produced also, and undoubtedly by the same external
+causes.</p>
+
+<p>It is of much consequence to be intimately acquainted with this species
+of queens, for they may have great influence
+<span class="pagebreak" title="37">&nbsp;</span><a name="Page_37" id="Page_37"></a>
+on different experiments
+and embarrass the observer: we should ascertain whether they inhabit
+pyramidal cells smaller than the common, or hexagonal ones.</p>
+
+<p>M. Schirach's famous experiment on the supposed conversion of a common
+worm into a royal one, cannot be too often repeated, though the Lusatian
+observers have already done it frequently. I could wish to learn
+whether, as the discoverer maintains, the experiment will succeed only
+with worms, three or four days old, and never with simple eggs.</p>
+
+<p>The Lusatian observers, and those of the Palatinate, affirm, that when
+common bees are confined with combs absolutely void of eggs, they then
+lay none but the eggs of drones. Thus, there must 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 ovaries contain male eggs alone?</p>
+
+<p>
+<span class="pagebreak" title="38">&nbsp;</span><a name="Page_38" id="Page_38"></a>
+According to M. de Reaumur, the life of chrysalids may be prolonged by
+keeping them in a cold situation, such as an ice-house. The same
+experiment should be made on the eggs of a queen; on the nymphs of
+drones and workers.</p>
+
+<p>Another interesting experiment would be to take away all the combs
+composing the common cells, and leave none but those destined for the
+larv&aelig; 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.
+Nevertheless, it is likely, by taking away only part of the common
+cells, the workers may be forced to lay common eggs in the cells of
+drones.</p>
+
+<p>I should also wish to have the young larv&aelig; gently removed from the royal
+cell, and deposited at the bottom of a common one, along with some of
+the royal food.</p>
+
+<p>
+<span class="pagebreak" title="39">&nbsp;</span><a name="Page_39" id="Page_39"></a>
+As the figure of hives has much influence 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
+adopted to instruct us how bees can regulate their labours, and apply
+them to existing circumstances. This may enable us to discover
+particular facts which we cannot foresee.</p>
+
+<p>The royal eggs and those producing drones, have not yet been carefully
+compared with the eggs from which workers come. But they ought to be so,
+that we may ascertain whether these different eggs have secret
+distinctive <a name="corr39" id="corr39"></a><a class="correction" href="#cnote39" title="changed from 'characterestics'">characteristics</a>.</p>
+
+<p>The food supplied by the workers to the royal worm, is not the same with
+that given to the common worm. Could we not endeavour, with the point of
+a pencil, to remove 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
+<span class="pagebreak" title="40">&nbsp;</span><a name="Page_40" id="Page_40"></a>
+cells hanging almost vertically, where the queen had laid; and these I
+should prefer for this experiment.</p>
+
+<p>Various facts, which require corroboration, were collected in my Memoirs
+on Bees; of this number are my own observations. You can select what is
+proper, my dear Sir. 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
+<span class="smcap">Contemplator of Nature</span>. <span class="indent3 italic">Genthod, 18. August 1789.</span></p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_A" id="fn_A"></a><span class="label"><a href="#fnm_A">A</a></span> All these letters are addressed to the celebrated
+naturalist M. Bonnet.&mdash;<i>T.</i></p></div>
+
+<div class="footnote"><p><a name="fn_B" id="fn_B"></a><span class="label"><a href="#fnm_B">B</a></span> The leaf or book hive consists of twelve vertical frames or
+boxes, parallel to each other, and joined together. Fig. 1. the sides, f
+f. f g. should be twelve inches long, and the cross spars, f f. g g.
+nine or ten; the thickness of these spars an inch, and their breadth
+fifteen lines. It is necessary that this last measure should be
+accurate; a a. a piece of comb which guides the bees in their work; d. a
+moveable slider supporting the lower part; b b. pegs to keep the comb
+properly in the frame or box; four are in the opposite side; e e. pegs
+in the sides under the moveable slider to support it.</p>
+
+<div class="figcenter" style="width: 398px;">
+<a id="fig1" name="fig1"></a>
+<img src="images/fig1.png" width="398" height="369" alt="The sides of a leaf or book hive" />
+<span class="caption">Figure 1</span>
+</div>
+
+<p>A book hive, consisting of twelve frames, all numbered, is represented
+fig. 2. Between 6 and 7 are two cases with lids, that divide the hive
+into two equal parts, and should only be used to separate the bees for
+forming an artificial swarm; a a. two frames which shut up the two sides
+of the hive, have sliders, b. b.</p>
+<div class="figcenter" style="width: 661px;">
+<a id="fig2" name="fig2"></a>
+<img src="images/fig2.png" width="661" height="457" alt="A leaf or book hive" />
+<span class="caption">Figure 2</span>
+</div>
+
+<p>The entrance appears at the bottom of each frame. All should be close
+but 1 and 12. However it is necessary that they should open at pleasure.</p>
+
+<p>The hive is partly open, fig. 3. and shews how the component parts may
+be united by hinges, and open as the leaves of a book. The two covers
+closing up the sides, a. a.</p>
+<div class="figcenter" style="width: 748px;">
+<a id="fig3" name="fig3"></a>
+<img src="images/fig3.png" width="748" height="465" alt="A partly open leaf or book hive" />
+<span class="caption">Figure 3</span>
+</div>
+
+
+<p><a name="corrfB" id="corrfB"></a><a class="correction" href="#cnotefB" title="This figure does not appear in the original">Fig. 4.</a> is another view of fig. 1. a a. a piece of comb to guide the
+bees; b b. pegs disposed so as to retain the comb properly in the frame;
+c c. parts of two shelves; the one above is fixed, and keeps the comb in
+a vertical position; the under one, which is moveable, supports it
+below.</p></div>
+
+<div class="footnote"><p><a name="fn_C" id="fn_C"></a><span class="label"><a href="#fnm_C">C</a></span> I cannot insist that my readers, the better to comprehend
+what is here said, shall peruse the Memoirs of M. de Reaumur on Bees,
+and those of the Lusace Society; but I must request them to examine the
+extracts in M. Bonnet's works, tom. 5. 4to edit. and tom. 10. 8vo, where
+they will find a short and distinct abstract of all that naturalists
+have hitherto discovered on the subject.</p></div>
+
+<div class="footnote"><p><a name="fn_D" id="fn_D"></a><span class="label"><a href="#fnm_D">D</a></span> Vide M. Schirach's History of Bees, in a memoir by M.
+Hattorf, entitled, <i>Physical Researches whether the Queen Bee requires
+fecundation by Drones?</i></p></div>
+
+<div class="footnote"><p><a name="fn_E" id="fn_E"></a><span class="label"><a href="#fnm_E">E</a></span> It will afterwards appear that what we took for the
+generative fluid, was the male organs of generation, left by copulation
+in the body of the female. This discovery we owe to a circumstance that
+shall immediately be related. Perhaps I should avoid prolixity, by
+suppressing all my first observations on the impregnation of the queen,
+and by passing directly to the experiments 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 think service is
+done to the reader by a candid avowal of my errors. This is an
+additional proof to so many others, of the absolute necessity that an
+observer should repeat all his experiments a thousand times, to obtain
+the certainty of seeing facts as they really exist.</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="41">&nbsp;</span><a name="Page_41" id="Page_41"></a>
+<a name="LETTER_II" id="LETTER_II"></a>LETTER II.<br /><br />
+
+<span class="little italic">SEQUEL OF OBSERVATIONS ON THE IMPREGNATION OF THE QUEEN BEE.</span></h2>
+
+
+<p class="salutation">
+SIR,</p>
+
+<p>All the experiments, related in my preceding letter, were made in 1787
+and 1788. They seem to establish two facts, which had previously been
+the subject of vague conjecture: 1. The queen bee is not impregnated of
+herself, but is fecundated by copulation with the male. 2. Copulation is
+accomplished without the hive, and in the air.</p>
+
+<p>
+<span class="pagebreak" title="42">&nbsp;</span><a name="Page_42" id="Page_42"></a>
+The latter appeared so extraordinary, that notwithstanding all the
+evidence obtained of it, we eagerly desired to take the queen in the
+fact; but, as she always rises to a great height, we never could see
+what passed. On that account you advised us to cut part off the wings of
+virgin queens. We endeavoured to benefit by your advice, in every
+possible manner; but to our great regret, when the wings lost much, the
+bees could no longer fly; and, by cutting off only an inconsiderable
+portion, we did not diminish the rapidity of their flight. Probably
+there is a medium, but we were unable to attain it. On your suggestion,
+we tried to render their vision less acute, by covering the eyes with an
+opaque varnish, which was an experiment equally fruitless.</p>
+
+<p>We likewise attempted artificial fecundation, and took every possible
+precaution to insure success. Yet the result was always unsatisfactory.
+Several queens were the victims of our curiosity; and those surviving
+<span class="pagebreak" title="43">&nbsp;</span><a name="Page_43" id="Page_43"></a>
+remained sterile. Though these different experiments were unsuccessful,
+it was proved that queens leave their hives to seek the males, and that
+they return with undoubted evidence of fecundation. Satisfied with this,
+we could only trust to time or accident for decisive proof of an actual
+copulation. We were far from suspecting a most singular discovery, which
+we made in July this year, and which affords complete demonstration of
+the supposed event, namely, that the sexual organs of the male remain
+with the female.<a name="fnm_F" id="fnm_F"></a><a href="#fn_F" class="fnnum">F</a></p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_F" id="fn_F"></a><span class="label"><a href="#fnm_F">F</a></span> The remainder of this Chapter chiefly consists of
+anatomical details. These may rather be considered an interruption of
+the narrative; and the Translator has judged it expedient to transfer
+them to an <a href="#APPENDIX">Appendix</a>.</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="44">&nbsp;</span><a name="Page_44" id="Page_44"></a>
+<a name="LETTER_III" id="LETTER_III"></a>LETTER III.<br /><br />
+
+<span class="little italic">THE SAME SUBJECT CONTINUED.&mdash;OBSERVATIONS ON RETARDING THE FECUNDATION
+OF QUEENS.</span></h2>
+
+
+<p>In my first letter, I remarked, that when queens were prevented from
+receiving the approaches of the male until the twenty-fifth or thirtieth
+day of their existence, the result presented very interesting
+peculiarities. My experiments at that time were not sufficiently
+numerous; but they have since been so often repeated, and the result
+<span class="pagebreak" title="45">&nbsp;</span><a name="Page_45" id="Page_45"></a>
+so
+uniform, that I no longer hesitate to announce, as a certain discovery,
+the singularities which retarded fecundation, produces on the ovaries of
+the queen. If she receives the male during the first fifteen days of her
+life, she remains capable of laying both the eggs of workers and of
+drones; but should fecundation be retarded until the twenty-second day,
+her ovaries are vitiated in such a manner that she becomes unfit for
+laying the eggs of workers, and will produce only those of drones.</p>
+
+<p>In June 1787, being occupied in researches 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. When a hive is ready to swarm, I had before
+observed, that the moment of swarming is always preceded by a very
+lively agitation, which first affects the queen, is then communicated to
+the workers, and excites such a tumult among them, that they abandon
+their labours, and rush in disorder to the outlets of the hive.
+<span class="pagebreak" title="46">&nbsp;</span><a name="Page_46" id="Page_46"></a>
+I then
+knew very well the cause of the queen's agitation, and it is described
+in the history of swarms, but I was ignorant how the delirium
+communicated to the workers; and this difficulty interrupted my
+researches. I therefore thought of investigating, by direct experiments,
+whether at all times, when the queen was greatly agitated, even not in
+the time of the hive swarming, her agitation would in like manner be
+communicated to the workers. The moment a queen 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 it
+would produce a kind of delirium. I had the patience to observe this
+queen thirty-four days. Every morning about eleven o'clock, when the
+weather was fine and the sunshine invited the males to leave their
+hives, I saw her impetuously traverse every corner of her habitation,
+<span class="pagebreak" title="47">&nbsp;</span><a name="Page_47" id="Page_47"></a>
+seeking to escape. Her fruitless efforts threw her into an uncommon
+agitation, the symptoms of which I shall elsewhere describe, and all the
+common bees were affected by it. As she never was out all this time, she
+could not be impregnated. At length, on the thirty-sixth day, I set her
+at liberty. She soon took advantage of it; and was not long of returning
+with the most evident marks of fecundation.</p>
+
+<p>Satisfied with the particular object of this experiment, I was far from
+any hopes that it would lead to the knowledge of another very remarkable
+fact; how great was my astonishment, therefore, on finding that this
+female, which, as usual, began to lay forty-six hours after copulation,
+laid the eggs of drones, but none of workers, and that she continued
+ever afterwards to lay those of drones only.</p>
+
+<p>At first, I exhausted myself with conjectures on this singular fact; the
+more I reflected on it, the more did it seem inexplicable. At length, by
+attentively meditating
+<span class="pagebreak" title="48">&nbsp;</span><a name="Page_48" id="Page_48"></a>
+on the circumstances of the experiment it
+appeared there were two principles, the influence of which I should
+first of all endeavour to appreciate separately. On the one hand, this
+queen had suffered long confinement; on the other, her fecundation had
+been extremely retarded. You know, Sir, that queens generally receive
+the males about the fifth or sixth day, and this queen had not copulated
+until the thirty-sixth. Little weight could be given to the supposition,
+that the peculiarity could be occasioned by confinement. Queens, in the
+natural state, leave their hives only once to seek the males. All the
+rest of their life they remain voluntary prisoners. Thus, it was
+improbable that captivity could produce the effect I wished to explain.
+At the same time, as it was essential to neglect nothing in a subject so
+new, I wished to ascertain whether it was owing to the length of
+confinement, or to retarded fecundation.</p>
+
+<p>
+<span class="pagebreak" title="49">&nbsp;</span><a name="Page_49" id="Page_49"></a>
+Investigating this was no easy matter. To discover whether captivity,
+and not retarded fecundation, vitiated the ovaries, it was necessary to
+allow a female to receive the approaches of a male, and also to keep her
+imprisoned. Now this could not be, for bees never copulate in hives. On
+the same account, it was impossible to retard the copulation of a queen
+without keeping her in confinement. I was long embarrassed by the
+difficulty. At length, I contrived an apparatus, which, though
+imperfect, nearly fulfilled my purpose.</p>
+
+<p>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 allowed free passage 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 the queen could at all times come and fly about, enjoying a
+purer air than was to be
+<span class="pagebreak" title="50">&nbsp;</span><a name="Page_50" id="Page_50"></a>
+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 between them. By the
+experiments related in my first letter, copulation takes place high in
+the air only: therefore, in this apparatus, I found the advantage of
+retarding fecundation, while the liberty the queen now had, did not
+render her situation too remote from the natural state. I attended to
+the experiment fifteen days. Every fine morning, the young captive left
+her hive; 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 aloft in the air, and soon returned with full evidence of
+impregnation. In two days, she laid, first the eggs of workers, and
+afterwards as many as the most fertile queens.</p>
+
+<p>
+<span class="pagebreak" title="51">&nbsp;</span><a name="Page_51" id="Page_51"></a>
+It thence followed, 1. That captivity did not alter the organs of
+queens. 2. When fecundation took place within the first sixteen days,
+she produced both species of eggs.</p>
+
+<p>This was an important experiment. It rendered my labours much more
+simple, by clearly pointing out the method to be pursued: it absolutely
+precluded the supposed influence of captivity; and left nothing for
+investigation but the consequences of retarded fecundation.</p>
+
+<p>With this view, I repeated the experiment; but, instead of giving the
+virgin queen liberty on the sixteenth day, I retained her until the
+twenty-first. She departed, rose high in the air, was fecundated, and
+returned. Thirty-six hours afterwards, she began to lay: but it was the
+eggs of males only, and, although very fruitful afterwards, she laid no
+other kind.</p>
+
+<p>I occupied myself the remainder of 1787, and the two subsequent years,
+with experiments on retarded fecundation, and
+<span class="pagebreak" title="52">&nbsp;</span><a name="Page_52" id="Page_52"></a>
+had constantly the same
+results. It is undoubted, therefore, that when the <a name="corr52" id="corr52"></a><a class="correction" href="#cnote52" title="changed from 'copulalation">copulation</a> of queens is retarded beyond the
+twentieth day, only an imperfect impregnation is operated: instead of
+laying the eggs of workers and males equally, they will lay none but
+those of males.</p>
+
+<p>I do not aspire to the honour of explaining this singular fact. When the
+course of my experiments led me to observe 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
+fecundation. My evidence is demonstrative, for I can always prevent
+queens from laying the eggs of workers, by retarding their fecundation
+until the twenty-second or twenty-third day. But, what is the remote
+cause of this peculiarity; or, in other words, why does the delay of
+impregnation render queens incapable of laying the eggs of workers? This
+is a problem on which analogy throws no light: nor in all physiology am
+I
+<span class="pagebreak" title="53">&nbsp;</span><a name="Page_53" id="Page_53"></a>
+acquainted with any fact that bears the smallest similarity.</p>
+
+<p>The problem becomes still more difficult by reflecting on the natural
+state of things, that is when fecundation has not been delayed. The
+queen then lays the eggs of workers forty-six hours after copulation,
+and continues for the subsequent eleven months to lay these alone: and
+it is only after this period that a considerable and uninterrupted
+laying of the eggs of drones commences. When, on the contrary,
+impregnation is retarded after the twentieth day, the queen begins, from
+the forty-sixth hour, to lay the eggs of males, and no other kind during
+her whole life. As, in the natural state, she lays the eggs of workers
+only, during the first eleven months, it is clear that these, and the
+male eggs, are not indiscriminately mixed in the oviducts. Undoubtedly
+they occupy a situation corresponding to the principles that regulate
+laying: the eggs of workers are first, and those of drones behind them.
+Farther, it
+<span class="pagebreak" title="54">&nbsp;</span><a name="Page_54" id="Page_54"></a>
+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 inverted by retarded copulation? How does it happen
+that all the workers eggs which the queen ought to lay, if fecundation
+was in due time, now wither and disappear, yet do not, impede the
+passage of the eggs of drones, which occupy only the second place in the
+ovaries. Nor is this all. I have satisfied myself that a single
+copulation 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 a single copulation will impregnate all the eggs that she will lay
+during her whole life: but I want absolute proof for more than two
+years. This, which is truly a very singular fact in itself, renders the
+influence of retarded fecundation still more difficult to be accounted
+for. Since a single copulation suffices, it is clear that the male fluid
+acts from the first moment on all the eggs
+<span class="pagebreak" title="55">&nbsp;</span><a name="Page_55" id="Page_55"></a>
+that the queen will lay in
+two years. It gives them, according to your principles, that degree of
+<i>animation</i> that afterwards effects their successive expansion. Having
+received the first impressions of life, they grow, they mature, so to
+speak, until the day they are laid: and as the laws of laying are
+constant, because the eggs of the first eleven months are always those
+of workers, it is evident that those which appear first are also the
+eggs that come soonest to maturity. Thus, in the natural state, the
+space of eleven months is necessary for the male eggs to acquire that
+degree of increment they must have attained when laid. This consequence,
+which to me seems immediate, renders the problem insoluble. How can the
+eggs, which should grow slowly for eleven months, suddenly acquire their
+full expansion in forty-eight hours, when fecundation has been retarded
+twenty-one days, and by the effect of this retardation alone? Observe, I
+beseech you, that the hypothesis of successive expansion is
+<span class="pagebreak" title="56">&nbsp;</span><a name="Page_56" id="Page_56"></a>
+not
+gratuitous; it rests on the principles of sound philosophy. Besides, for
+conviction that it is well founded, we have only to look at the figures
+given by Swammerdam of the ovaries of the queen bee. There we see eggs
+in that part of the oviducts contiguous to the vulva, much farther
+advanced, and larger than those contained in the opposite part.
+Therefore the difficulty remains in full force: it is an abyss where I
+am lost.</p>
+
+<p>The only known fact bearing any relation to that now described, is the
+state of certain vegetable seeds, which, although extremely well
+preserved, lose the faculty of germination from age. The eggs of workers
+may also preserve, only for a very short time, the property of being
+fecundated by the seminal fluid; and, after this period, which is about
+fifteen or eighteen days, become disorganised to that degree, that they
+can no longer be animated by it. I am sensible that the comparison is
+very imperfect; besides, it explains nothing,
+<span class="pagebreak" title="57">&nbsp;</span><a name="Page_57" id="Page_57"></a>
+nor does it even put us
+on the way of making any new experiments. I shall add but one reflection
+more.</p>
+
+<p>Hitherto no other effect has been observed from the retarded
+impregnation of animals, but that of rendering them absolutely sterile.
+The first instance of a female still preserving the faculty of
+engendering males, is presented by the queen bee. But as no fact in
+nature is unique, it is most probable that the same peculiarity will
+also be found in other animals. An extremely curious object of research
+would be to consider insects in this new point of view, I say <i>insects</i>,
+for I do not conceive that any thing analogous will be found in other
+species of animals. The experiments now suggested would necessarily
+begin with insects the most analogous to bees; as wasps, humble bees,
+mason bees, all species of flies, and the like. Some experiments might
+also be made on butterflies; and, perhaps, an animal might be found
+whose retarded fecundation would be attended
+<span class="pagebreak" title="58">&nbsp;</span><a name="Page_58" id="Page_58"></a>
+with the same effects as
+that of queen bees. Should the animal be larger, dissection will be more
+easily accomplished; and we may discover what happens to the eggs when
+retarded fecundation prevents their expansion. At least, we might hope
+that some fortunate circumstance would lead to solution of the
+problem<a name="fnm_G" id="fnm_G"></a><a href="#fn_G" class="fnnum">G</a>.</p>
+
+<p>Let us now return to my experiments. In May 1789, I took two queens just
+when they had undergone the last metamorphosis: one was put in a <i>leaf
+hive</i>, well provided with honey and wax, and sufficiently inhabited by
+workers and males. The other was put into a hive exactly similar, from
+which all the drones were removed.
+<span class="pagebreak" title="59">&nbsp;</span><a name="Page_59" id="Page_59"></a>
+The entrances of these hives were
+too confined for the passage of the females and drones, but the common
+bees enjoyed perfect liberty. The queens were imprisoned thirty days;
+and being then set at liberty, they departed, and returned impregnated.
+Visiting the hives in the beginning of July, I found much brood, but
+wholly consisting of the worms and nymphs of males. There actually was
+not a single worker's worm or nymph. Both queens laid uninterruptedly
+until autumn, and constantly the eggs of drones. Their laying ended in
+the first week of November, as that of my other queens.</p>
+
+<p>I was very earnest to learn what would become of them in the subsequent
+spring, whether they would resume laying, or if new fecundation would be
+necessary; and if they did lay, of what species the eggs would be.
+However, the hives being very weak, I dreaded they might perish during
+winter. Fortunately, we were able to preserve them; and from April 1790,
+they recommenced
+<span class="pagebreak" title="60">&nbsp;</span><a name="Page_60" id="Page_60"></a>
+laying. The precautions we had taken prevented them
+from receiving any new approaches of the male. Their eggs were still
+those of males.</p>
+
+<p>It would have been extremely interesting to have followed the history of
+these two females still farther, but, to my great regret, the workers
+abandoned their hives on the fourth of May, and that same day I found
+both queens dead. No weevils were in the hive, which could disturb the
+bees; and the honey was still very plentiful: but as no workers had been
+been produced in the course of the preceding year, and winter had
+destroyed many, they were too few in spring to engage in their wonted
+labours, and, from discouragement, deserted their habitation to occupy
+the neighbouring hives.</p>
+
+<p>In my Journal, I find a detail of many experiments on the retarded
+impregnation of queen bees, so many, that transcribing the whole would
+be tedious. I may repeat, however, that there was not the least
+<span class="pagebreak" title="61">&nbsp;</span><a name="Page_61" id="Page_61"></a>
+variation in the principle, and that whenever the copulation of queens
+was postponed beyond the twenty-first day, the eggs of males only were
+produced. Therefore, I shall limit my narrative to those experiments
+that have taught me some remarkable facts.</p>
+
+<p>A queen being hatched on the fourth of October 1789, we put her into a
+leaf-hive. Though the season was well advanced, a considerable number of
+males was still in the hive; and it here became important to learn,
+whether, at this period of the year, they could equally effect
+fecundation; also, in case it succeeded, whether a laying, begun in the
+middle of autumn, would be interrupted or continued during winter. Thus,
+we allowed the queen to leave the hive. She departed, indeed, but made
+four and twenty fruitless attempts before returning with the evidence of
+fecundation. Finally, on the thirty-first of October, she was more
+fortunate: She departed, and returned with the most undoubted
+<span class="pagebreak" title="62">&nbsp;</span><a name="Page_62" id="Page_62"></a>
+proof of
+the success of her amours: She was now twenty-seven days old,
+consequently fecundation had been retarded. She ought to have begun
+laying within forty-six hours, but the weather was cold, and she did not
+lay; which proves, as we may cursorily remark, that refrigeration of the
+atmosphere is the principal agent that suspends the laying of queens
+during winter. I was excessively impatient to learn whether, on the
+return of spring, she would prove fertile, without a new copulation. The
+means of ascertaining the fact was easy; for the entrances of the hives
+only required contraction, so as to prevent her from escaping. She was
+confined from the end of October until May. In the middle of March, we
+visited the combs, and found a considerable number of eggs, but, none
+being yet hatched, we could not know whether they would produce workers
+or males. On the fourth of April, having again examined the state of the
+hive, we found a prodigious quantity of nymphs
+<span class="pagebreak" title="63">&nbsp;</span><a name="Page_63" id="Page_63"></a>
+and worms, all of
+drones; nor had this queen laid a single worker's egg.</p>
+
+<p>Here, as well as in the preceding experiment, 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
+copulation 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
+begin to lay in spring without requiring new copulation.</p>
+
+<p>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, already flies, two thousand four hundred and
+thirty-eight cells, containing either eggs or nymphs of drones. Thus,
+<span class="pagebreak" title="64">&nbsp;</span><a name="Page_64" id="Page_64"></a>
+she had laid more than three thousand male eggs during March and April,
+which is above fifty each day. Her death soon afterwards unfortunately
+interrupted my observation, I intended to calculate the total number of
+male eggs that she should lay throughout the year, and compare it with
+those 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 considerable, commences in August, also in the
+interval, that they produce the eggs of workers almost solely. But it is
+otherwise with the females whose copulation has been retarded: they
+produce no workers' eggs. For four or five months following, they lay
+the eggs of males without interruption, and in such numbers, that, in
+this short time, I suppose one queen gives birth to more drones than a
+female, whose fecundation has not been retarded, produces in the course
+of two years. It
+<span class="pagebreak" title="65">&nbsp;</span><a name="Page_65" id="Page_65"></a>
+gives me much regret, that I have not been able to
+verify this conjecture.</p>
+
+<p>I should also describe the very remarkable manner in which queens, that
+lay only the eggs of drones, sometimes deposit them in the cells.
+Instead of being placed in the lozenges forming the bottom, they are
+frequently deposited on the lower side of the cells, two lines from the
+mouth. This arises from the body of such queens being shorter than that
+of those whose fecundation has not been retarded. The extremity remains
+slender, while the first two rings next the thorax are uncommonly swoln.
+Thus, in disposing themselves for laying, the extremity cannot reach the
+bottom of the cells on account of the swoln rings; consequently the eggs
+must remain attached to the part that the extremity reaches. The worms
+proceeding from them pass their vermicular state in the same place where
+the eggs were deposited, which proves that bees are not charged with the
+care of transporting the eggs as has been supposed.
+<span class="pagebreak" title="66">&nbsp;</span><a name="Page_66" id="Page_66"></a>
+But here they
+follow another plan. They extend beyond the surface of the comb those
+cells where they observe the eggs deposited, two lines from the mouth.</p>
+
+<p>Permit me, Sir, to digress a moment from the subject, to give the result
+of an experiment which seems interesting. Bees, I say, are not charged
+with the care of transporting into cells, the eggs misplaced by the
+queen: and, judging by the single instance I have related, you will
+think me well entitled to deny this feature of their industry. However,
+as several authors have maintained the reverse, and even demanded our
+admiration of them in conveying the eggs, I should explain clearly that
+they are deceived.</p>
+
+<p>I had a glass hive constructed of two stages; the higher was filled with
+combs of large cells, and the lower with those of common ones. A kind of
+division, or diaphraghm, separated these two stages from
+each other, having at each side an opening for the passage of the
+workers
+<span class="pagebreak" title="67">&nbsp;</span><a name="Page_67" id="Page_67"></a>
+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 had only those of workers to lay, and
+she was obliged to deposit them in the surrounding large cells from the
+want of others. My object in this arrangement will already be
+anticipated. My reasoning was simple. If the queen laid workers' eggs in
+the large cells, and the bees were charged with transporting them if
+misplaced, they would infallibly take advantage of the liberty allowed
+to pass from either stage: they would seek the eggs deposited in the
+large cells, and carry them down to the lower stage containing the cells
+adapted for that species. If, on the contrary, they left the common eggs
+in the large cells, I should obtain certain proof that they had not the
+charge of transporting them.</p>
+
+<p>
+<span class="pagebreak" title="68">&nbsp;</span><a name="Page_68" id="Page_68"></a>
+The result of this experiment excited my curiosity extremely. We
+observed the queen several days without intermission. During the first
+twenty-four hours, she persisted in not laying a single egg in the
+surrounding cells; she examined them one after another, but passed on
+without insinuating her belly into one. She was restless, and traversed
+the combs in all directions: her eggs appeared an oppressive burden, but
+she persisted 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 mother. I was amused to observe, when she
+approached the edges of the division separating the two stages, that she
+gnawed at them to enlarge the passage: the workers approached her, and
+also laboured with their teeth, and made every exertion to enlarge the
+entrance to her prison, but ineffectually. On the second day, the queen
+could no longer retain her eggs: they escaped in spite of her,
+<span class="pagebreak" title="69">&nbsp;</span><a name="Page_69" id="Page_69"></a>
+and fell
+at random. Then we conceived that the bees would convey them into the
+small cells of the lower stage, and we sought them there with the utmost
+assiduity; but I can safely affirm there was not one. The eggs that the
+queen still laid the third day disappeared as the first. We again sought
+them in the small cells, but none were there. The fact is, they are ate
+by the workers; and this is what has deceived the naturalists, who
+supposed them carried away. They have observed the misplaced eggs
+disappear, and, without farther investigation, have asserted that the
+bees convey them elsewhere: they take them, indeed, not to convey them
+any where, but to devour them. Thus nature has not charged bees with the
+care of placing the eggs in the cells appropriated for them, but she has
+inspired females themselves with sufficient instinct to know the species
+of eggs they are about to lay, and to deposit them in suitable cells.
+This has already been observed by M. de
+<span class="pagebreak" title="70">&nbsp;</span><a name="Page_70" id="Page_70"></a>
+Reaumur, and here my
+observations correspond with his. Thus it is certain that in the natural
+state, when fecundation takes place at the proper time, and the queen
+has suffered from nothing, she is never deceived in the choice of the
+cells where her eggs are to be deposited; she never fails to lay those
+of workers in small cells, and those of males in large ones. The
+distinction is important, for the same certainty of instinct is no
+longer conspicuous in the conduct of those females whose impregnation
+has been deferred. I was oftener than once deceived respecting the eggs
+that such queens laid, for they were deposited indiscriminately in small
+cells and those of drones; and not aware of their instinct having
+suffered, I conceived that the eggs in small cells would produce
+workers; therefore I was very much surprised, when, at the moment they
+should have been hatched, the bees closed up the cells, and
+demonstrated, by anticipation, that the included worms would change into
+drones; they actually became males;
+<span class="pagebreak" title="71">&nbsp;</span><a name="Page_71" id="Page_71"></a>
+those produced in small cells were
+small, those in large cells large. Thus I must warn observers, who would
+repeat my experiments on queens that lay only the eggs of males, not to
+be deceived by these circumstances, and expect that eggs of males will
+be deposited in the workers cells.</p>
+
+<p>It is a singular fact, that the females, whose fecundation has been
+retarded, sometimes lay the eggs of males in royal cells. I shall prove,
+in the history of swarms, that immediately when queens, in the natural
+state, begin their great laying of male eggs, the workers construct
+numerous royal cells. Undoubtedly, there is some secret relation between
+the appearance of male eggs and the construction of these cells; for it
+is a law of nature from which bees never derogate. It is not surprising,
+therefore, that such cells are constructed in hives governed by queens
+laying the eggs of males only. It is no longer extraordinary that these
+queens deposit in the royal cells, eggs of the only species they can
+lay,
+<span class="pagebreak" title="72">&nbsp;</span><a name="Page_72" id="Page_72"></a>
+for in general their instinct seems affected. But what I cannot
+comprehend is, why the bees take exactly the same care of the male eggs
+deposited in royal cells, as of those that should become queens. They
+provide them more plentifully with food, they build up the cells as if
+containing a royal worm; in a word, they labour with such regularity
+that we have frequently been deceived. More than once, in the firm
+persuasion of finding royal nymphs, we have opened the cells after they
+were sealed, yet the nymph of a drone always appeared. Here the instinct
+of the workers seemed defective. In the natural state, they can
+accurately distinguish the male worms from those of common bees, as they
+never fail giving 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 to investigate the instinct of
+<span class="pagebreak" title="73">&nbsp;</span><a name="Page_73" id="Page_73"></a>
+animals, we must
+carefully observe where it appears to err.</p>
+
+<p>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.</p>
+
+<p>In a work, <i>Histoire de la Reine des Abeilles</i>, translated from the
+German by <i>Blassiere</i>, there is printed a letter from M. Schirach to
+you, dated 15 April 1771, where he speaks of some hives, in which the
+whole brood changed into drones. You will remember that he ascribes this
+circumstance to some unknown vice in the ovaries of the queen; but he
+was far from suspecting that retarded fecundation had been the cause of
+vitiation. He justly felicitated himself on discovering a method to
+prevent the destruction of hives in this situation, which was simple,
+for it consisted in removing the queen that laid the eggs of males only,
+and substituting one for her whose ovaries were not impaired.
+<span class="pagebreak" title="74">&nbsp;</span><a name="Page_74" id="Page_74"></a>
+But to
+make the substitution effectual, it was necessary to procure queens at
+pleasure; a secret reserved for M. Schirach, and of which I shall speak
+in the following letter. You observe that the whole experiments of the
+German naturalist tended to the preservation of the hives whose queens
+laid none except male eggs; and that he did not attempt to discover the
+cause of the vice evident in their ovaries.</p>
+
+<p>M. de Reaumur also says a few words, somewhere, of a hive containing
+many more drones than workers, but advances no conjectures on the cause.
+However, he adds, as a remarkable circumstance, that the males were
+tolerated in this hive until the subsequent spring. It is true that bees
+governed 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. I can ascribe no reason for it, but it is a fact I have
+several times witnessed during my long course of observations on
+retarded impregnation. In
+<span class="pagebreak" title="75">&nbsp;</span><a name="Page_75" id="Page_75"></a>
+general it has appeared that while the queen
+lays male eggs, bees do not massacre the males already perfect in the
+hive. <span class="smcap indent3">Pregny</span>, <i>21. August 1791</i>.</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_G" id="fn_G"></a><span class="label"><a href="#fnm_G">G</a></span> The experiments suggested in this paragraph, recall a
+singular reflection of M. de Reaumur. Where treating of oviparous flies,
+he says, it would not be impossible for a hen to produce a living
+chicken, if, after fecundation, the eggs she should first lay could by
+any means be retained twenty-one days in the oviducts. <i>Mem. sur. les
+Insect. tom. 4. mem. 10.</i></p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="76">&nbsp;</span><a name="Page_76" id="Page_76"></a>
+<a name="LETTER_IV" id="LETTER_IV"></a>LETTER IV.<br /><br />
+
+<span class="little italic">ON M. SCHIRACH'S DISCOVERY.</span></h2>
+
+
+<p>When you found it necessary, Sir, in the new edition of your works, to
+give an account of M. Schirach'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
+testimonies. For this reason, I hasten to inform you that all my
+researches establish the reality of the discovery. During
+<span class="pagebreak" title="77">&nbsp;</span><a name="Page_77" id="Page_77"></a>
+ten years
+that I have studied bees, I have repeated M. Schirach's experiment so
+often, and with such uniform success, that I can no longer have the
+least doubt on the subject. Therefore, I consider it an established
+fact, when bees lose their queen, and 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, but a greater quantity of it,
+and 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 a fact, and the philosophical consequences
+you have deduced from it. <i>Contemplation de la Nature, part. II, chap.
+27.</i></p>
+
+<p>In this letter I shall content myself with some account of the figure of
+the royal cells constructed by bees around those worms that are destined
+for the royal state, and terminate with discussing some points
+<span class="pagebreak" title="78">&nbsp;</span><a name="Page_78" id="Page_78"></a>
+wherein
+my observations differ from those of M. Schirach.</p>
+
+<p>Bees soon become sensible of having lost their queen, and in a few hours
+commence the labour necessary to repair their loss. First, they select
+the young common worms, which the requisite treatment is to convert into
+queens, and immediately begin with enlarging the cells where they are
+deposited. Their mode of proceeding is curious; and the 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.
+Having chosen a worm, they sacrifice three of the contiguous cells:
+next, they supply it with food, and raise a cylindrical inclosure
+around, by which the cell becomes a perfect tube, with a rhomboidal
+bottom; for the parts forming the bottom are left untouched. If the bees
+damaged it, they would lay open three corresponding cells on the
+opposite surface of the comb, and, consequently, destroy their worms,
+which
+<span class="pagebreak" title="79">&nbsp;</span><a name="Page_79" id="Page_79"></a>
+would be an unnecessary sacrifice, and Nature has opposed it.
+Therefore, leaving the bottom rhomboidal, they are satisfied with
+raising a cylindrical tube around the worm, which, like the other cells
+in the comb, is horizontal. But this habitation remains suitable to the
+worm called to the royal state only during the first three days of its
+existence: another situation is requisite for the other two days it is a
+worm. Then, which is so small a portion of its life, it must inhabit a
+cell nearly of a pyramidal figure, and hanging perpendicularly; we may
+say the workers know it; for, after the worm has completed the third
+day, they prepare the place to be occupied by its new lodging. They gnaw
+away the cells surrounding the cylindrical tube, mercilessly sacrifice
+their worms, and use the wax in constructing a new pyramidal tube, which
+they solder at right angles to the first, and work it downwards. The
+diameter of this pyramid decreases insensibly from the base, which is
+very wide, to the
+<span class="pagebreak" title="80">&nbsp;</span><a name="Page_80" id="Page_80"></a>
+point. During the two days that it is inhabited by
+the worm, a bee constantly keeps its head more or less inserted into the
+cell, and, when this worker quits it, another comes to occupy its place.
+In proportion as the worm grows, the bees labour in extending the cell,
+and bring food, which they place before its mouth, and around its body,
+forming a kind of cord around it. The worm, which can move only in a
+spiral direction, turns incessantly to take the food before its head: it
+insensibly descends, and at length arrives at the orifice of the cell.
+Now is the time of transformation to a nymph. As any farther care is
+unnecessary, the bees close the cell with a peculiar substance
+appropriated for it, and there the worm undergoes both its
+metamorphoses.</p>
+
+<p>Though M. Schirach supposes that none but worms three days old are
+selected for the royal treatment, I am certain of the contrary; and that
+the operation succeeds equally well on those of two days only.
+<span class="pagebreak" title="81">&nbsp;</span><a name="Page_81" id="Page_81"></a>
+I must
+be permitted to relate at length the evidence I have of the fact, which
+will both demonstrate the reality of common worms being converted into
+queens, and the little influence which their age has on the effect of
+the operation.</p>
+
+<p>I put some pieces of comb, with some workers eggs, in the cells, and of
+the same kind as those already hatched, into a hive deprived of the
+queen. The same day several cells were enlarged by the bees, and
+converted into royal cells, and the worms supplied with a thick bed of
+jelly. Five were then removed from those cells, and five common worms,
+which, forty-eight hours before we had seen come from the egg
+substituted for them. The bees did not seem aware of the change; they
+watched over the new worms the same as over those chosen by themselves;
+they continued enlarging the cells, and closed them at the usual time.
+When they had hatched on them seven days<a name="fnm_H" id="fnm_H"></a><a href="#fn_H" class="fnnum">H</a>, we removed the
+<span class="pagebreak" title="82">&nbsp;</span><a name="Page_82" id="Page_82"></a>
+cells to
+see the queens that were to 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 having elapsed, and no queen appearing, we
+opened them. In one, was a dead queen, but still a nymph; the other two
+were empty. The worms had spun their silk <a name="corr82" id="corr82"></a><a class="correction" href="#cnote82" title="changed from 'coccons'">coccoons</a>, but died
+before passing into their nymphine state, and presented only a dry skin.
+I can conceive nothing more conclusive than this experiment. It
+demonstrates that bees have the power of converting the worms of workers
+into queens; since they succeeded in procuring queens, by operating on
+the worms which we ourselves had selected. It is equally demonstrated,
+that the success of the operation does not depend on the worms being
+three days old, as those entrusted to the bees were only two. Nor is
+this all; bees can convert worms still younger into queens. The
+following experiment showed, that
+<span class="pagebreak" title="83">&nbsp;</span><a name="Page_83" id="Page_83"></a>
+when the queen is lost, they destine
+worms only a few hours old to replace her.</p>
+
+<p>I was in possession of a hive, which being long deprived of the female,
+had neither egg nor worm. I provided a queen of the greatest fertility;
+and she immediately began laying in the cells of workers. I removed this
+female before being quite three days in the hive, and before any of her
+eggs were hatched. The following morning, that is, 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 depositing around them a much more abundant provision
+of food than is supplied to common worms. Next day, the worms were near
+forty hours old: the bees had enlarged and converted their hexagonal
+cells into cylindrical ones of the greatest capacity. During the
+subsequent days, they still laboured at them, and closed them on the
+fifth from the origin of the worms.
+<span class="pagebreak" title="84">&nbsp;</span><a name="Page_84" id="Page_84"></a>
+Seven days after sealing of the
+first of these royal cells, a queen of the largest size proceeded from
+it. She immediately rushed towards the other royal cells, and
+endeavoured to destroy their nymphs and worms. In another letter, I
+shall recount the effects of her fury.</p>
+
+<p>From these details, you will observe, Sir, that M. Schirach's
+experiments had not been sufficiently diversified when he affirmed that
+it was essential for the conversion of common 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 existence.</p>
+
+<p>After my researches to corroborate M. Schirach's discovery, I was
+desirous of learning whether, as this observer conceives, the only means
+which the bees have of procuring a queen, is giving the common worms a
+certain kind of aliment, and rearing them in the largest cells. You
+
+<span class="pagebreak" title="85">&nbsp;</span><a name="Page_85" id="Page_85"></a>
+will remember, that M. de Reaumur's sentiments are very different: "The
+mother should lay, and she does lay, eggs from which flies fit for being
+mothers must in their turn proceed. She does 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 peculiar interest in the eggs that one is to
+proceed from, and to consider them of the greatest value. They construct
+particular cells where they are to be deposited.&mdash;The figure of a royal
+cell only begun, very much resembles a cup, or, more correctly speaking,
+the cup that has lost its acorn."</p>
+
+<p>M. de Reaumur, though he did not suspect 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 M. Schirach, on the other hand,
+bees always having the power of procuring
+<span class="pagebreak" title="86">&nbsp;</span><a name="Page_86" id="Page_86"></a>
+a queen by bringing up worms
+three days old in a particular manner, it would be needless for nature
+to grant females the faculty of laying royal eggs. Such prodigality is,
+in his eyes, inconsistent with the ordinary laws of nature. Therefore he
+maintains, in direct terms, that she 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 would always be
+too long for the belly of the mother to reach the bottom.</p>
+
+<p>I admit that M. de Reaumur no where says he has seen the queen lay in
+the royal cell. However he did not doubt the fact; and, after all my
+observations, I must esteem his opinion just. It is quite certain that,
+at particular periods of the year, the bees prepare royal cells; that
+the females deposit their eggs in them; and that worms,
+<span class="pagebreak" title="87">&nbsp;</span><a name="Page_87" id="Page_87"></a>
+which shall
+became queens, proceed from these eggs.</p>
+
+<p>M. Schirach's objection, concerning the length of the cells, proves
+nothing; for the queen does not delay depositing her egg till they are
+finished. While only sketched and shaped like the cup of an acorn, she
+lays it. This naturalist, dazzled by the brilliancy of his discovery,
+saw only part of the truth. He was the first to find out the resource
+granted to bees by nature, for repairing the loss of their queen; and
+too soon persuaded himself that she had provided no other resource for
+the production of females. This error arose from not observing bees in
+very flat hives: had he used such as mine, he would have found, on
+opening them in spring, a confirmation of M. de Reaumur's opinion. Then,
+which is the season of swarming, hives in good condition are governed by
+a very fruitful queen: there are royal cells of a figure widely
+different from those constructed around the worms destined by the bees
+
+<span class="pagebreak" title="88">&nbsp;</span><a name="Page_88" id="Page_88"></a>
+for queens. They are large, attached to the comb by a stalk, and
+hanging vertically like stalactites, such, in short, as M. de Reaumur
+has described them. The females lay in them before completion. 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 the egg has been
+laid. In proportion as the worm grows, they are enlarged, and closed by
+the bees when the first transformation approaches. Thus it is true,
+that, in spring, the queen deposits in royal cells, previously prepared,
+eggs from which flies of her own species are to come. Nature has,
+therefore, provided a double means for the multiplication and
+conservation of their race.</p>
+
+<p class="indent1 italic">
+<span class="smcap">Pregny</span>, 24. August 1791.
+</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_H" id="fn_H"></a><span class="label"><a href="#fnm_H">H</a></span> The author's meaning here is obscure.&mdash;T.</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="89">&nbsp;</span><a name="Page_89" id="Page_89"></a>
+<a name="LETTER_V" id="LETTER_V"></a>LETTER V.<br /><br />
+
+<span class="little italic">EXPERIMENTS PROVING THAT THERE ARE SOMETIMES COMMON BEES WHICH LAY
+FERTILE EGGS.</span></h2>
+
+
+<p>The singular discovery of M. Riems, concerning the existence of fertile
+workers, has appeared very doubtful to you, Sir. You have suspected that
+the eggs ascribed to workers by this naturalist had actually been
+produced by small queens, which, on account of their size, were
+confounded with common bees. But you do not positively insist that M.
+Riems is deceived; and, in the letter which you did me the honour to
+address to me, you requested
+<span class="pagebreak" title="90">&nbsp;</span><a name="Page_90" id="Page_90"></a>
+me to investigate, by new experiments,
+whether there are actually working bees capable of laying fertile eggs.
+I have made these experiments with great care: and it is for you to
+judge of the confidence they merit.</p>
+
+<p>On the fifth of August 1788, we found the eggs and worms of large drones
+in two hives, which had both been some time deprived of queens. We also
+observed the rudiments of some royal cells appended like stalactites to
+the edges of the combs. The eggs of males were in them. Being perfectly
+secure that there was no queen of large size among the bees of these two
+hives, the eggs, which daily became more numerous, were evidently laid
+either by queens of small size or by fertile workers. I had reason to
+believe it was actually by common bees, for we had frequently observed
+them inserting the posterior part into the cells; and assuming the same
+attitude as the queen when laying. But, not withstanding every exertion,
+we had never
+<span class="pagebreak" title="91">&nbsp;</span><a name="Page_91" id="Page_91"></a>
+been able to seize one in this situation, 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 observations were continued with equal assiduity, in hopes that, by
+some fortunate chance, or in a moment of address, we could secure one of
+them. More than a month all our endeavours were abortive.</p>
+
+<p>My assistant then offered to perform an operation that required both
+courage and patience, and which I could not resolve to suggest, though
+the same expedient had occurred to myself. He proposed to examine each
+bee in the hive separately, to discover whether some small queen had not
+insinuated herself among them, and escaped our first researches. This
+was an important experiment; for, should no small queen be found, it
+would be demonstrative evidence that the eggs had been laid by simple
+workers.</p>
+
+<p>
+<span class="pagebreak" title="92">&nbsp;</span><a name="Page_92" id="Page_92"></a>
+To perform this operation with all possible exactness, immersing the
+bees was not enough. You know, Sir, that the contact of water stiffens
+their organs, that it produces a certain alteration of their external
+figure: and, from the resemblance of small queens to workers, the
+slightest alteration of shape would prevent us from distinguishing with
+sufficient accuracy to what species those immersed might belong.
+Therefore it was necessary to seize the whole bees of both hives,
+notwithstanding their irritation, and examine their specific character
+with the utmost 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, but what the relief of
+his eyes required. He took every bee in his hand; he attentively
+examined the trunk, the hind limbs, and the sting: there was not one
+without the characteristics of the common bee, that
+<span class="pagebreak" title="93">&nbsp;</span><a name="Page_93" id="Page_93"></a>
+is, the little
+basket on the hind legs, the long trunk, and the straight sting. He had
+previously prepared glass cases containing combs. Into these, he put
+each bee after examination. It is superfluous to observe they were
+confined, which was a precaution indispensible until termination of the
+experiment. Neither was it enough to establish that the whole were
+workers; we had also to continue the experiment, and observe whether any
+would produce eggs. Thus we examined the cells for several days, and
+soon observed new laid eggs, from which the worms of drones came at the
+proper time. My assistant held in his hands the bees that produced them;
+and as he was perfectly certain they were common ones, it is proved that
+there are sometimes fertile workers in hives.</p>
+
+<p>Having ascertained M. Schirach's discovery, by so decisive an
+experiment, we replaced all the bees examined, in very thin glass hives,
+being only eighteen lines thick, and capable of containing but a single
+row
+<span class="pagebreak" title="94">&nbsp;</span><a name="Page_94" id="Page_94"></a>
+of combs, and thus were extremely favourable to the observer. We
+thought, by strictly persisting to watch the bees, we might surprise a
+fertile one in the act of laying, seize and dissect her. This we were
+desirous of doing, for the purpose of comparing her ovaries with those
+of queens, and to ascertain the difference. At length, on the eighth of
+September, we had the good fortune to succeed.</p>
+
+<p>A bee appeared in the position of a female laying. Before she had time
+to leave the cell, we suddenly opened the hive and seized her. She
+presented all the external characteristics of <a name="corr94" id="corr94"></a><a class="correction" href="#cnote94" title="changed from 'commo nbees'">common bees</a>; the only difference we could recognise, and that was a very
+slight one, consisted in the belly seeming less and more slender than
+that of workers. On dissection, her ovaries were found more fragile,
+smaller and composed of fewer oviducts than the ovaries of queens. The
+filaments containing the eggs were extremely fine, and exhibited
+swellings at equal distances. We counted
+<span class="pagebreak" title="95">&nbsp;</span><a name="Page_95" id="Page_95"></a>
+eleven eggs of sensible size,
+some of which appeared ripe for laying. This ovary was double like that
+of queens.</p>
+
+<p>On the ninth of September, we seized another fertile worker the instant
+she laid, and dissected her. The ovary was still less expanded than that
+of the preceding bee, and only four eggs had attained maturity. My
+assistant extracted one from the oviducts, and succeeded in fixing it by
+an end on a glass slider. We may take this opportunity of remarking,
+that it is in the oviducts themselves the eggs are imbued with the
+viscous liquid, with which they are produced, and not in passing through
+the spherical sac as Swammerdam believed. During the remainder of this
+month, we found ten fertile workers in the same hives, and dissected
+them all. In most, the ovaries were easily distinguished, but in some we
+could not discern the faintest traces of them. In these last, the
+oviducts to all appearance were but imperfectly developed, and more
+address than we had acquired in
+<span class="pagebreak" title="96">&nbsp;</span><a name="Page_96" id="Page_96"></a>
+dissection was necessary to distinguish
+them.</p>
+
+<p>Fertile workers never lay the eggs of common bees; they produce none but
+those of males. M. Riems had already observed this singular fact; and
+here all my observations correspond with his. I shall only add to what
+he says, that fertile workers are not absolutely indifferent in the
+choice of cells for depositing their eggs. They always prefer large
+ones; and only use small cells when unable to find those of larger
+diameter. But they so far correspond with queens whose impregnation has
+been retarded, that they sometimes lay in royal cells.</p>
+
+<p>Speaking of females laying male eggs alone, I have already expressed my
+surprise that bees bestow, on those deposited in royal cells, such care
+and attention as to feed the worms proceeding from them, and, at the
+period of transformation, to close them up. But I know not, Sir, why I
+omitted to observe that, after sealing the royal cells,
+<span class="pagebreak" title="97">&nbsp;</span><a name="Page_97" id="Page_97"></a>
+the workers
+build them up, and sit on them until the last metamorphosis of the
+included male<a name="fnm_I" id="fnm_I"></a><a href="#fn_I" class="fnnum">I</a>. The treatment of the royal cells where fertile workers
+lay the eggs of drones is very different. They begin indeed with
+bestowing every care on their eggs and worms; they close the cells at a
+suitable time, but never fail to destroy them three days afterwards.</p>
+
+<p>Having finished these first experiments with success, I had still to
+discover the cause of the expansion of the sexual organs of fertile
+workers. M. Riems had not engaged in this interesting problem; and at
+first I dreaded that I should have no other guide towards its solution
+than conjecture. Yet from serious reflection, it appeared, that, by
+connecting the facts contained in this letter, there was some light that
+might elucidate my procedure in this new research.</p>
+
+<p>
+<span class="pagebreak" title="98">&nbsp;</span><a name="Page_98" id="Page_98"></a>
+From M. Schirach's elegant discoveries, it is beyond all doubt that
+common bees are originally of the female sex. They have received from
+nature the germs of an ovary, but she has allowed 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 get that aliment while worms.</p>
+
+<p>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 larv&aelig; destined to replace her.
+Therefore, if fertile workers are produced in this situation alone, it
+is evident their origin is only in those hives where bees prepare the
+royal jelly. Towards this circumstance, I bent all my attention. It
+induced me to suspect that when bees give the <i>royal treatment</i> to
+certain worms, they either by accident or a particular instinct, the
+principle of which
+<span class="pagebreak" title="99">&nbsp;</span><a name="Page_99" id="Page_99"></a>
+is unknown to me, drop some particles of royal jelly
+into cells contiguous to those containing the worms destined for queens.
+The larv&aelig; of workers that have accidentally received portions of so
+active an aliment, must be more or less affected by it; and their
+ovaries should acquire a degree of expansion. But this expansion will be
+imperfect; why? because the royal food has been administered only in
+small portions, and, besides, the larv&aelig; having lived in cells of the
+smallest dimensions, their parts cannot extend beyond the ordinary
+proportions. Thus, the bees produced by them will resemble common
+workers in size and all the external characteristics. Added to that,
+they will have the faculty of laying some eggs, solely from the effect
+of the trifling portion of royal jelly mixed with their aliment.</p>
+
+<p>That we may judge of the justness of this explanation, it is necessary
+to consider fertile workers from their origin; to investigate whether
+the cells, where they are
+<span class="pagebreak" title="100">&nbsp;</span><a name="Page_100" id="Page_100"></a>
+brought up, are constantly in the vicinity of
+the royal cells, and if their food is mixed with particles of the royal
+jelly. Unfortunately, the execution of these experiments is very
+difficult. When pure, the royal jelly is recognised by its sharp and
+pungent taste; but, when mixed with other substances, the peculiar
+savour is very 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<a name="fnm_J" id="fnm_J"></a><a href="#fn_J" class="fnnum">J</a>.</p>
+
+<p>In June 1790, I observed that one of my thinnest hives had wanted the
+queen several days, and that the bees had no mean 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 worms destined for
+
+<span class="pagebreak" title="101">&nbsp;</span><a name="Page_101" id="Page_101"></a>
+queens, in 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
+destined for replacing the queens, it was very interesting to follow
+their history, and necessary to watch the moment of their last
+transformation. I removed the nineteen cells into a grated box, which
+was introduced 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 experiment. But here another
+precaution was also requisite. It was to be feared, that the bees being
+deprived of the produce of their labour, and the object of their hope
+might be totally discouraged; therefore, I supplied them with another
+piece of comb, containing
+<span class="pagebreak" title="102">&nbsp;</span><a name="Page_102" id="Page_102"></a>
+the brood of workers, reserving power to
+destroy the young brood when necessary. This plan succeeded admirably.
+The bees, in bestowing all their attention on these last worms, forgot
+those that had been removed.</p>
+
+<p>When the moment of transformation of the nymphs in the nineteen cells
+arrived, I examined the grated box frequently every day, and at length
+found six bees exactly similar to <i>common bees</i>. The worms of the
+remaining thirteen had perished without changing.</p>
+
+<p>The portion of brood comb that had been put into the hive to prevent the
+discouragement of the bees was then removed. I put 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 six
+into the hive, where they were well received.</p>
+
+<p>You easily conceive my object, Sir, in this course of observations. I
+knew there
+<span class="pagebreak" title="103">&nbsp;</span><a name="Page_103" id="Page_103"></a>
+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 produced by some of the six bees?
+But, to attain absolute certainty, it was necessary to take them in the
+act of laying. Some ineffaceable mark was also required for
+distinguishing them in particular.</p>
+
+<p>This proceeding was attended with the most ample success. We 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
+perceived one introducing the posterior part into 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 privation of the right antenna, instantly
+recognised that it was one of the six that had passed to the vermicular
+state in the vicinity of the royal cells.</p>
+
+<p>
+<span class="pagebreak" title="104">&nbsp;</span><a name="Page_104" id="Page_104"></a>
+I could no longer doubt the truth of my conjecture; at the same time, I
+know not whether the truth will appear as rigorous to you, Sir, as it
+does to myself. But I reason in the following manner: If it is certain
+that fertile workers are always produced in the vicinity of royal cells,
+it is no less true, that in itself, the vicinity is indifferent; for the
+size and figure of these cells can produce no effect on the worms in
+those surrounding them; there must be something more; we 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 the worms in the
+adjacent cells have had a portion of the same food. This is what they
+gain, therefore, by vicinity to the royal cells. The bees, in their
+course thither, will pass in numbers over them, stop and drop some
+portion of the jelly destined for the royal larv&aelig;. This
+<span class="pagebreak" title="105">&nbsp;</span><a name="Page_105" id="Page_105"></a>
+reasoning, I
+presume, is consistent with the principles of sound logic.</p>
+
+<p>I have repeated the experiment now described so often, and weighed all
+the concomitant circumstances with so much care, that whenever I please,
+I can produce fertile workers in my hives. The method is simple. 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
+supplying the included worms with the royal jelly. Portions of this
+aliment also fall on the young larv&aelig; deposited in the adjacent cells,
+and it unfolds the ovaries to a certain degree. 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. Therefore, to
+save them, all their enemies must be removed, and the larv&aelig; of the royal
+cells taken away before undergoing their last metamorphoses. Then the
+fertile
+<span class="pagebreak" title="106">&nbsp;</span><a name="Page_106" id="Page_106"></a>
+workers, being without rivals at the time of their origin, will
+be well received, and, by taking the precaution to mark them, it will be
+seen, in a few days, that they produce the eggs of males. Thus, the
+whole secret of this proceeding consists in removing the royal cells at
+the proper time; that is, after being sealed, and previous to the young
+queens leaving them<a name="fnm_K" id="fnm_K"></a><a href="#fn_K" class="fnnum">K</a>.</p>
+
+<p>I shall add but a few words to this long letter. There is nothing so
+very surprising in the production of fertile workers, when we consider
+the consequences of M. Shirach's
+<span class="pagebreak" title="107">&nbsp;</span><a name="Page_107" id="Page_107"></a>
+beautiful discovery. But why do they
+lay male eggs only? I can conceive, indeed, that the reason of their
+laying few is from their ovaries being but imperfectly expanded, but I
+can form no idea why all the eggs should be those of males, neither can
+I any better account for their use in hives; and hitherto, I have made
+no experiments on their mode of fecundation.</p>
+
+<p class="indent1 italic">
+<span class="little">PREGNY</span>, 25. August 1791.
+</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_I" id="fn_I"></a><span class="label"><a href="#fnm_I">I</a></span> It is difficult to discover whether the author thinks, as
+some naturalists, that bees are instrumental in hatching the eggs.&mdash;T.</p></div>
+
+<div class="footnote"><p><a name="fn_J" id="fn_J"></a><span class="label"><a href="#fnm_J">J</a></span> The original is extremely confused in the preceding
+passages.&mdash;T.</p></div>
+
+<div class="footnote"><p><a name="fn_K" id="fn_K"></a><span class="label"><a href="#fnm_K">K</a></span> I have frequently seen queens, at the moment of production,
+begin first by attacking the royal cells and then the common ones beside
+them. As I had not seen fertile workers when I first observed this fact,
+I could not conceive from what motive the fury of the queen was thus
+directed towards the common cells. But now I know they can distinguish
+the species included, and have the same instinctive jealousy or aversion
+towards them as against the nymphs of queens properly so denominated.</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="108">&nbsp;</span><a name="Page_108" id="Page_108"></a>
+<a name="LETTER_VI" id="LETTER_VI"></a>LETTER VI.<br /><br />
+
+<span class="little italic">ON THE COMBATS OF QUEENS: THE MASSACRE OF THE MALES: AND WHAT SUCCEEDS
+IN A HIVE WHERE A STRANGER QUEEN IS SUBSTITUTED FOR THE NATURAL ONE.</span></h2>
+
+
+<p>M. de Reaumur had not witnessed every thing relative to bees when he
+composed his history of these industrious animals. Several observers,
+and those of Lusa&ccedil;e in particular, have discovered many important facts
+that escaped him; and I, in my turn, have made various observations of
+which he had no suspicion: at the same
+<span class="pagebreak" title="109">&nbsp;</span><a name="Page_109" id="Page_109"></a>
+time, and this is a very
+remarkable circumstance, not only has all that he expressly declares he
+saw been verified by succeeding naturalists, but all his conjectures are
+found just. The German naturalists, Schirach, Hattorf, and Riems
+sometimes contradict him, indeed, in their memoirs; but I can maintain
+that, while combating the opinion of M. de Reaumur, it is they who are
+almost always wrong; of which several instances might be adduced.</p>
+
+<p>What I shall now proceed to say will give me an opportunity of detailing
+some interesting facts.</p>
+
+<p>It was observed by M. de Reaumur, that when any supernumerary queen is
+either produced in a hive, or comes into it, one of the two soon
+perishes. He has not actually witnessed 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. M. Schirach, on the other
+hand, and, after him,
+<span class="pagebreak" title="110">&nbsp;</span><a name="Page_110" id="Page_110"></a>
+M. Riems, thinks that the working bees assail the
+stranger, and sting her to death. I cannot comprehend by what means they
+have been able to make this observation: as they used very thick hives
+only, with several rows of combs, they could at most but observe the
+commencement of hostilities. While the combat lasts, the bees move with
+great rapidity; they fly on all sides; and, gliding between 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 queens and
+workers, but I have very often beheld one between the queens themselves.</p>
+
+<p>In one of my hives in particular, there were five or six royal cells,
+each including a nymph. The eldest first underwent its transformation.
+Scarcely did ten minutes elapse from the time of this young queen
+leaving her cradle, when she visited the other royal cells still close.
+She furiously attacked the nearest; and, by dint of labour,
+<span class="pagebreak" title="111">&nbsp;</span><a name="Page_111" id="Page_111"></a>
+succeeded
+in opening the top: we saw her tearing the silk of the coccoon with her
+teeth; but her efforts were probably inadequate to the object, for she
+abandoned this end of the cell, and began at the other, where she
+accomplished a larger aperture. When it was sufficiently enlarged, she
+endeavoured to introduce her belly, and made many exertions until she
+succeeded in giving her rival a deadly wound with her sting. Then having
+left the cell, all the bees that had hitherto been spectators of her
+labour, began to increase the opening, and drew out the dead body of a
+queen scarcely come from its envelope of a nymph.</p>
+
+<p>Meanwhile, the victorious young queen attacked another royal cell, but
+did not endeavour to introduce her extremity into it. There was only a
+royal nymph, and no queen, come to maturity, as in the first cell. In
+all probability, nymphs of queens inspire their rivals with less
+animosity; still they do not escape destruction: because,
+<span class="pagebreak" title="112">&nbsp;</span><a name="Page_112" id="Page_112"></a>
+whenever a
+royal cell has been opened before the proper time, the bees extract the
+contents in whatever form they may be, whether worm, nymph, or queen.
+Lastly, the young queen attacked the third cell, but could not succeed
+in penetrating it. She laboured languidly, and appeared as if exhausted
+by her first exertions. As we now required queens for some particular
+experiments, we resolved to remove the other royal cells, yet in safety,
+to secure them from her fury.</p>
+
+<p>After this observation, we wished to see what ensued on two queens
+leaving their cells at the same time, and in what manner one perished. I
+find an observation on this head in my Journal, 15. May 1790.</p>
+
+<p>In one of our thinnest hives, two queens left their cells almost at the
+same moment. Whenever they observed each other, they rushed together,
+apparently with great fury, and were in such a position that the antenn&aelig;
+of each was seized by the teeth of the other: the head, breast, and
+belly of
+<span class="pagebreak" title="113">&nbsp;</span><a name="Page_113" id="Page_113"></a>
+the one were exposed to the head, breast, and belly of the
+other: the extremity of their bodies were curved; they were reciprocally
+pierced with the stings; and both fell dead at the same instant. But it
+seems as if nature has not ordained that both combatants should perish
+in the duel; but rather that, when finding themselves in the situation
+described, namely, opposite, and belly to belly, they fly at that moment
+with the utmost precipitation. Thus, when these two rivals felt the
+extremities about to meet, they disengaged 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 think that we may here
+penetrate the intention of nature.</p>
+
+<p>There ought to be none but one queen in a hive: therefore it is
+necessary, if by chance a second is either produced or comes into the
+hive, that one of the two must be destroyed. This cannot be committed to
+the working bees, because, in a
+<span class="pagebreak" title="114">&nbsp;</span><a name="Page_114" id="Page_114"></a>
+republic composed of so many
+individuals, an equal consent cannot be supposed always to exist; it
+might frequently happen that one group of bees destroyed one of the
+queens, while a second would massacre the other; and the hive thus be
+deprived of queens. Therefore it was necessary that the queens
+themselves should be entrusted with the destruction of their rivals: but
+as, in these combats, nature demands but 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.</p>
+
+<p>I am well aware of the hazard of error in minute researches into the
+causes of the most trifling facts. But here the object and the means
+seem so plain, that I have ventured to advance my conjectures. You will
+judge better than I can, whether they are well founded.&mdash;Let me now
+return from this digression.</p>
+
+<p>
+<span class="pagebreak" title="115">&nbsp;</span><a name="Page_115" id="Page_115"></a>
+A few minutes after the two queens separated, their terror ceased, and
+they again began to seek each other. Immediately on coming in sight,
+they rushed together, seized one another, and resumed exactly their
+former position. The result of this <a name="corr115" id="corr115"></a><a class="correction" href="#cnote115" title="changed from 'rencounter'">encounter</a> was the
+same. When their bellies approached, they hastily disengaged themselves,
+and fled with precipitation. During all this time, the workers seemed in
+great agitation; and the tumult appeared to increase when the
+adversaries separated. Two different times, we observed them stop the
+flight of the queens, seize their limbs, and retain them prisoners above
+a minute. At last, the queen, which was either the strongest 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, brought the
+extremity of her own body under the belly of the other; and, by this
+means, easily pierced her with the sting. Then she withdrew her sting
+after losing hold of the
+<span class="pagebreak" title="116">&nbsp;</span><a name="Page_116" id="Page_116"></a>
+wing. The vanquished queen fell down, dragged
+herself languidly along, and, her strength failing, she soon expired.</p>
+
+<p>This observation proved that virgin queens engage in single combats; but
+we wished to discover whether those fecundated, and mothers, had the
+same animosity.</p>
+
+<p>On the 22. of July, we selected a flat hive, containing a very fertile
+queen: and being curious to learn whether, as virgin queens, she would
+destroy the royal cells, three were introduced into the middle of the
+comb. Whenever she observed this, <i>she</i> sprung forward on the whole, and
+pierced them towards the bottom; nor did she desist until the included
+nymphs were exposed. The workers which had hitherto been spectators of
+this destruction, now came to carry the nymphs away. They greedily
+devoured the food remaining at the bottom of the cells, and also sucked
+the fluid from the abdomen of the nymphs: and then terminated with
+destroying the cells from which they had been drawn.</p>
+
+<p>
+<span class="pagebreak" title="117">&nbsp;</span><a name="Page_117" id="Page_117"></a>
+In the next place, we introduced a very fertile queen into this hive;
+after painting the thorax to distinguish her from the reigning queen. A
+circle of bees quickly formed around the stranger, but their intention
+was not to caress and receive her well; for they insensibly accumulated
+so much, and surrounded her so closely, that in scarcely a minute she
+lost her liberty and became a prisoner. It is a remarkable circumstance,
+that other workers at the same time collected round the reigning queen
+and restrained all her motions; we instantly saw her confined like the
+stranger. Perhaps it may be said, 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 way to allow her full liberty for the attack; then if the queens
+testified a disposition
+<span class="pagebreak" title="118">&nbsp;</span><a name="Page_118" id="Page_118"></a>
+to fly, they returned to enclose them.</p>
+
+<p>We have repeatedly witnessed this fact, but it presents so new and
+singular a characteristic in the policy of bees, that it must be seen
+again a thousand times before any positive assertion can be made on the
+subject. I would therefore recommend that naturalists should attentively
+examine the combat of queens, and particularly ascertain what part is
+taken by the workers. Is their object to accelerate the combat? Do they
+by any secret means excite the fury of the combatants? Whence does it
+happen that accustomed to bestow every care on their queen, in certain
+circumstances, they oppose her preparations to avoid impending danger?</p>
+
+<p>A long series of observations are necessary to solve these problems. It
+is an immense field for experiment, which will afford infinitely curious
+results. I intreat you to pardon my frequent digressions. The subject is
+deeply philosophical, genius
+<span class="pagebreak" title="119">&nbsp;</span><a name="Page_119" id="Page_119"></a>
+such as <a name="corr119" id="corr119"></a><a class="correction" href="#cnote119" title="unchanged">your's</a> is required to
+treat it properly; and I shall now be satisfied with proceeding in the
+description of the combat.</p>
+
+<p>The cluster of bees that surrounded the reigning queen having allowed
+her some freedom, she seemed to advance towards that part of the comb
+where her rival stood; then, all the bees receded before her, the
+multitude of workers, separating the two adversaries, gradually
+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.</p>
+
+<p>In the last place, to exhaust every combination, we had still to examine
+whether a combat would ensue between two queens,
+<span class="pagebreak" title="120">&nbsp;</span><a name="Page_120" id="Page_120"></a>
+one impregnated, and
+the other a virgin; and what circumstances attended it.</p>
+
+<p>On the 18. of September, we introduced a very fruitful queen into a
+glass hive, already containing a virgin queen, and put her on the
+opposite side of the comb, that we might have time to see how the
+workers would receive her. She was immediately surrounded, but they
+confined her only a moment. Being oppressed with the necessity of
+laying, she dropped some eggs; however, 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 advanced towards the edge of the
+comb, and soon approached very near the virgin queen. When in sight,
+they rushed together; the virgin queen got on the back of the other, and
+gave her several stings in the belly, but, having aimed at the scaly
+part, they did not injure her, and the combatants separated. In a few
+minutes, they
+<span class="pagebreak" title="121">&nbsp;</span><a name="Page_121" id="Page_121"></a>
+returned to the charge; but this time the impregnated
+queen mounted on her rival; however, she sought in vain to pierce her,
+for the sting did not enter; the virgin queen then disengaged herself
+and fled; she also succeeded in escaping another attack, where her
+adversary 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 successful exertion, the virgin queen
+mortally wounded the stranger, and she expired in a moment. The sting
+had penetrated so far that the victorious queen was unable to extract
+it, and she was overthrown by the fall of her enemy. She made great
+exertions to disengage the sting: but could succeed by no other means
+than turning on the extremity of the belly, as on a pivot. Probably the
+barbs of the sting fell by this motion, and, closing like a spiral
+around the stem, came more easily from the wound.</p>
+
+<p>
+<span class="pagebreak" title="122">&nbsp;</span><a name="Page_122" id="Page_122"></a>
+These observations, Sir, I think will satisfy you, respecting the
+conjecture of our celebrated 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 will
+at no time attempt to employ their stings against the stranger queen. I
+can conceive what has misled M. Riems and Schirach; but it is necessary
+for explaining it that I should relate a new feature in the policy of
+bees, at considerable length.</p>
+
+<p>In the natural state of hives, several queens 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 supernumerary queens; and if an observer wishes to introduce one,
+he can accomplish it only by force, that is by opening the hive. In a
+word, no queen can insinuate
+<span class="pagebreak" title="123">&nbsp;</span><a name="Page_123" id="Page_123"></a>
+herself into a hive in a natural state,
+for the following reasons.</p>
+
+<p>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
+antenn&aelig; every individual endeavouring to penetrate the hive, and also
+the various substances put within their reach; which affords us an
+opportunity of observing that the <a name="corr123" id="corr123"></a><a class="correction" href="#cnote123" title="changed from 'antennae'">antenn&aelig;</a> are certainly the
+organs of feeling. If a stranger queen appears, she is instantly seized
+by the bees on guard, which prevent her entry by laying hold of her legs
+or wings with their teeth, and crowd so closely around her, that she
+cannot move. Other bees, from the interior of the hive, gradually come
+to their assistance, and confine her still more narrowly, all having
+their heads towards the centre where the queen is inclosed; and they
+remain with such evident anxiety, eagerness, and attention, that the
+cluster they form may be carried about for some time,
+<span class="pagebreak" title="124">&nbsp;</span><a name="Page_124" id="Page_124"></a>
+without their
+being sensible of it. A stranger queen, so closely confined and hemmed
+in, cannot possibly penetrate the hive. If the bees retain her too long
+imprisoned, she perishes. Her death probably ensues from hunger, or the
+privation of air; it is undoubted, at least, that she is never stung. We
+never saw the bees direct their stings against her, except a single
+time, and then it was owing to ourselves. We endeavoured, from
+compassion for a queen's situation, to remove her from the center 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 several of the workers were
+themselves killed; and surely they could not intend destroying one
+another. Had we not interfered, they would have been content with
+confining the queen, and would not have massacred her.</p>
+
+<p>It was in similar circumstances that M. Riems saw the workers anxiously
+pursue a
+<span class="pagebreak" title="125">&nbsp;</span><a name="Page_125" id="Page_125"></a>
+queen. He thought they designed to sting her, and thence
+concluded, that the office of the common bees is to kill supernumerary
+queens. You have quoted his observations in the <i>Contemplation de la
+Nature, part II, chap. 27, note 7</i>. But you are sensible, Sir, from
+these details, that he has been mistaken. He did not know the attention
+that bees bestow on what passes at the entrance of their hive, and he
+was entirely ignorant of the means they take to prevent supernumerary
+queens from penetrating it.</p>
+
+<hr />
+
+<p>After ascertaining that the workers in no situation sting the
+supernumerary queens, we were curious to learn how a stranger queen
+would be received in a hive wanting a 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.</p>
+
+<p>
+<span class="pagebreak" title="126">&nbsp;</span><a name="Page_126" id="Page_126"></a>
+Bees do not immediately observe the removal of their queen; their
+labours are uninterrupted; they watch over the young, and perform all
+their ordinary occupations. 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 a
+delirious impetuosity. Then they discover their queen is no longer among
+them. But how do they become sensible of it? How do the bees on the
+surface of the comb discover that the queen is not on the next comb? In
+treating of another characteristic of these animals, you have yourself,
+Sir, proposed the same question; I am incapable of answering it indeed,
+but I have collected some facts, that may perhaps facilitate the
+elucidation of this mystery.</p>
+
+<p>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
+<span class="pagebreak" title="127">&nbsp;</span><a name="Page_127" id="Page_127"></a>
+singular, they <i>recognise</i> her: you must
+interpret this expression strictly. Substitution of another queen is not
+attended with the same effect, if she is introduced into the hive within
+the first twelve hours after removal of the reigning one. Here the
+agitation continues; and the bees treat the stranger the same as when
+the presence of their own leaves them nothing to desire. They surround,
+seize, and keep her captive, a very long time, in an impenetrable
+cluster; and she commonly dies either from hunger or privation of air.</p>
+
+<p>If eighteen hours elapse before substitution of a stranger queen for the
+native one removed, she is at first treated 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 moves
+languidly; and sometimes 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
+<span class="pagebreak" title="128">&nbsp;</span><a name="Page_128" id="Page_128"></a>
+where they had originally been ill
+received.</p>
+
+<p>If, before substituting the stranger queen, twenty-four hours elapse,
+she 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 word reception is very
+indefinite, it is proper to enter into some detail for explaining the
+exact sense in which I use it. On the 15. of August, I introduced a
+fertile queen, eleven months old, into a glass hive. The bees were
+twenty-four hours deprived of their queen, and had already begun the
+construction of twelve royal-cells, such as described in the preceding
+chapter. Immediately on placing this female stranger on the comb, the
+workers near her touched her with their antenn&aelig;, and, passing their
+trunks over every part of her body, they gave her honey. Then these gave
+place to others that treated her exactly in the same manner.
+<span class="pagebreak" title="129">&nbsp;</span><a name="Page_129" id="Page_129"></a>
+All
+vibrated their wings at once, and ranged themselves in a circle around
+their sovereign. Hence resulted a kind of agitation which gradually
+communicated to the workers situated on the same surface of the comb,
+and induced them to come and reconnoitre, in their turn, what was going
+on. They soon arrived; and, having broke through the circle formed by
+the first, approached the queen, touched her with the antenn&aelig;, and gave
+her honey. After this little ceremony they retired; and, placing
+themselves behind the others, enlarged the circle. There they vibrated
+their wings, and buzzed without tumult or disorder, and as if
+experiencing some very agreeable sensation. The queen had not yet moved
+from the place where I had put her, but in a quarter of an hour she
+began to move. The bees, far from opposing her, opened the circle at
+that part to which she turned, followed her, and formed a guard around.
+She was oppressed with the necessity of laying, and dropped
+<span class="pagebreak" title="130">&nbsp;</span><a name="Page_130" id="Page_130"></a>
+eggs.
+Finally, after four hours abode, she began to deposit male eggs in the
+cells she met.</p>
+
+<p>While these events passed on the surface of the comb where the queen
+stood, all was quiet on the other side. Here the workers were apparently
+ignorant of a queen's arrival in the hive. They laboured with great
+activity at the royal cells, as if ignorant that they no longer stood in
+need of them: they watched over the royal worms, supplied them with
+jelly and the like. But the queen having at length come to this side,
+she was received with the same respect that she had experienced from
+their companions on the other side of the comb. They encompassed her;
+gave her honey; and touched her with their antenn&aelig;: and what proved
+better that they treated her as a mother, was their immediately
+desisting from work at the royal cells; they removed the worms, and
+devoured the food collected around them. From this moment the queen was
+recognised
+<span class="pagebreak" title="131">&nbsp;</span><a name="Page_131" id="Page_131"></a>
+by all her people, and conducted herself in this new
+habitation as if it had been her native hive.</p>
+
+<p>These particulars will give a just idea of the manner that bees receive
+a stranger queen; when they have time to forget their own, she is
+treated exactly as if she was their natural one, except that there is
+perhaps at first greater interest testified in her, or more conspicuous
+demonstrations of it. I am sensible of the impropriety of these
+expressions, but M. de Reaumur in some respect authorises them. He does
+not scruple to say, that bees pay <i>attention</i>, <i>homage</i>, and <i>respect</i>,
+to their queen, and from his example the like expressions have escaped
+most authors that treat on bees.</p>
+
+<p>Twenty-four or thirty hours absence is sufficient to make them forget
+their first queen, but I can hazard no conjecture on the cause.</p>
+
+<hr />
+
+<p>Before terminating this letter, which is full of combats and
+<a name="corr131" id="corr131"></a><a class="correction" href="#cnote131" title="changed from 'disastrou'">disastrous</a> scenes, I
+<span class="pagebreak" title="132">&nbsp;</span><a name="Page_132" id="Page_132"></a>
+should, perhaps, give you an account
+of some more pleasing and interesting facts relative to their industry.
+However, to avoid returning to duels and massacres, I shall here subjoin
+my observations on the massacre of the males.</p>
+
+<p>You will remember, Sir, it is agreed by all observers, that at a certain
+period of the year, the workers kill and expel the drones. M. de
+<a name="corr132" id="corr132"></a><a class="correction" href="#cnote132" title="changed from 'Reamur'">Reaumur</a> speaks of these executions as a horrible massacre. He
+does not expressly affirm, indeed, that he has himself witnessed it, but
+what we have seen corresponds so well with his account, that there can
+be no doubt he has beheld the peculiarities of the massacre.</p>
+
+<p>It is usually in the months of July and August, that the bees free
+themselves of the males. Then they are drove away and pursued to the
+inmost parts of the hive, where they collect in numbers; and as at the
+same time we find many dead drones on the ground before the hives, it is
+indubitable that after being expelled, the bees sting
+<span class="pagebreak" title="133">&nbsp;</span><a name="Page_133" id="Page_133"></a>
+them to death.
+Yet on the surface of the comb, we do not see the sting used against
+them; there the bees are content to pursue and drive them away. You
+observe this, Sir, yourself, in the new notes added to <i>la Contemplation
+de la Nature</i>; and you seem disposed to think, that the drones forced to
+retire to the extremity of the hive, perish from hunger. Your conjecture
+was extremely probable. Still it was possible the carnage might take
+place in the bottom of the hive, and had been unobserved, because that
+part is dark, and escapes the observer's eye.</p>
+
+<p>To appreciate the justice of this suspicion, we thought of making the
+support of the hive of glass, and of placing ourselves below to see what
+passed in the scene of action. Therefore, a glass table was constructed,
+on which were put six hives with swarms of the same year; and, lying
+under it, we endeavoured to discover how the drones were destroyed. The
+invention succeeded to admiration. On the 4 of
+<span class="pagebreak" title="134">&nbsp;</span><a name="Page_134" id="Page_134"></a>
+July, we saw the workers
+actually massacre the males, in the whole six swarms, at the same hour,
+and with the same peculiarities.</p>
+
+<p>The glass table was covered with bees full of animation, which flew upon
+the drones, as they came from the bottom of the hive; seized them by the
+antenn&aelig;, the limbs, and the wings, and after having dragged them about,
+or, so to speak, after quartering them, they killed them by repeated
+stings directed between the rings of the belly. The moment that this
+formidable weapon reached them, was the last of their existence; they
+stretched 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 stuck
+the sting so deep, that it could hardly be withdrawn, and these bees
+were obliged to turn upon themselves before the stings could be
+disengaged.</p>
+
+<p>Next day, having resumed our former position, we witnessed new scenes of
+carnage. During three hours, the bees furiously destroyed the males.
+They had
+<span class="pagebreak" title="135">&nbsp;</span><a name="Page_135" id="Page_135"></a>
+massacred all their own on the preceding evening, but now
+attacked those which, driven from the neighbouring hives, had taken
+refuge amongst them. We saw them also tear some remaining nymphs from
+the combs; they greedily sucked all the fluid from the abdomen, and then
+carried them away. The following days no drones remained in the hives.</p>
+
+<p>These two observations seem to me decisive. It is incontestible that
+nature has charged the workers with the destruction of the males at
+certain seasons of the year. But what means does she use to excite their
+fury against them? This is a question that I cannot pretend to answer.
+However, an observation I have made may one day lead to solution of the
+problem. The males are never destroyed in hives deprived of queens, on
+the contrary, while a savage massacre prevails in other places, they
+there find an asylum. They are tolerated and fed, and many are seen even
+in the middle of January. They are also preserved in hives,
+<span class="pagebreak" title="136">&nbsp;</span><a name="Page_136" id="Page_136"></a>
+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 but in hives where the queens are completely
+fertile, and it never begins until the season of swarming is past.</p>
+
+<p class="toright italic">
+<span class="smcap">Pregny</span>, 28 August 1791.
+</p>
+
+
+
+<h2>
+<span class="pagebreak" title="137">&nbsp;</span><a name="Page_137" id="Page_137"></a>
+<a name="LETTER_VII" id="LETTER_VII"></a>LETTER VII.<br /><br />
+
+<span class="little italic">SEQUEL OF EXPERIMENTS ON THE RECEPTION OF A STRANGER QUEEN. M. DE
+REAUMUR'S OBSERVATIONS ON THE SUBJECT.</span></h2>
+
+
+<p>I have frequently testified my admiration of M. de Reaumur's
+observations on bees. I feel a sensible pleasure in acknowledging that
+if I have made any progress in the art of observation, I am indebted for
+it to profound study of the works of this
+<span class="pagebreak" title="138">&nbsp;</span><a name="Page_138" id="Page_138"></a>
+naturalist. In general his
+authority has such weight, that I can scarcely trust my own experiments
+when the results are different from his. Likewise, on finding myself in
+opposition to the <i>historian of bees</i>, I repeat my experiments. I vary
+the mode of conducting them; I examine with the utmost caution all the
+circumstances that might mislead me, and never are my labours
+interrupted before acquiring the moral certainty of avoiding error. With
+the aid of these precautions, I have discovered the justice of M. de
+Reaumur's suggestions, and I have a thousand times seen, if certain
+experiments seemed to combat them, it was from incorrectness of
+execution. Yet I must except some facts where my results have constantly
+been different from his. Those respecting the reception of a stranger
+queen substituted for the natural one, are of the number.</p>
+
+<p>If, after removing the natural queen, a stranger is immediately
+substituted, the usurper is ill received. I never could succeed
+<span class="pagebreak" title="139">&nbsp;</span><a name="Page_139" id="Page_139"></a>
+in
+making them adopt her, but by allowing an interval of twenty or
+twenty-four hours to elapse. Then they seemed to have forgot their own
+queen; and respectfully received any female put in her place. M. de
+Reaumur, on the contrary, asserts, that should the original queen be
+removed, and another presented, this new one will be perfectly well
+received from the beginning. As evidence of this assertion, he gives the
+detail of an experiment which must be read in his work, for I shall here
+give only an extract of it<a name="fnm_L" id="fnm_L"></a><a href="#fn_L" class="fnnum">L</a>. 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 this moment, the
+tumult ceased, and the stranger queen was received with all respect.</p>
+
+<p>
+<span class="pagebreak" title="140">&nbsp;</span><a name="Page_140" id="Page_140"></a>
+I do not dispute the truth of this experiment; but, in my opinion, it
+does not warrant the conclusion that M. de Reaumur deduces from it. His
+apparatus removed the bees too much from their natural condition, to
+allow him to judge of their instinct and dispositions. In other
+situations, he has himself observed, that these animals, reduced to
+small numbers, lost their industry and activity, and feebly continued
+their ordinary labours. Thus their instinct is affected by every
+operation that too much diminishes their number. To render such an
+experiment truly conclusive, it must be made in a populous hive; and on
+removing the native queen, a stranger must immediately be substituted in
+her place. Had this been done, I am fully persuaded, that M. de Reaumur
+would have seen the bees imprison the usurper, confine her at least
+twelve or fifteen hours among them, and frequently suffocate her: nor
+would he have witnessed any favourable reception before an interval of
+twenty-four
+<span class="pagebreak" title="141">&nbsp;</span><a name="Page_141" id="Page_141"></a>
+hours after removal of the original queen. No variation has
+occurred in my experiments regarding this fact. Their number, and the
+attention bestowed on them, make me presume they merit your confidence.</p>
+
+<p>M. de Reaumur, in another passage of the same Memoir, affirms, that
+<i>bees, which have a queen they are satisfied with, are nevertheless
+disposed to give the best possible reception to any female that seeks
+refuge among them</i>. In the preceding letter, I have related my
+experiments on this head: their success has been very different from
+that of M. de Reaumur's. I have proved that the workers never employ
+their stings against the queen; but this cannot be called the welcome
+reception of a stranger. They retain her within their ranks, and seem to
+allow her liberty only when she prepares to combat the reigning queen.
+This observation cannot be made except in the thinnest hives. Those used
+by M. de Reaumur had always two parallel combs
+<span class="pagebreak" title="142">&nbsp;</span><a name="Page_142" id="Page_142"></a>
+at least, which must
+have prevented him from observing some very important circumstances that
+influence the conduct of workers when supplied with several females. The
+first circles formed around a stranger queen he has taken for caresses;
+and, from the little that this queen could advance between the combs, it
+must have been impossible for him to observe that the circles, which
+always continued contracting, ended in restraint of the females there
+inclosed. Had he used thinner hives, he would have discovered that what
+he supposed indication of a favourable reception was the prelude of
+actual imprisonment.</p>
+
+<p>I feel reluctant to assert that M. de Reaumur was deceived. Yet I cannot
+admit that, on certain occasions, bees tolerate a plurality of females
+in their hives. The experiment on which this affirmation rests will not
+be considered decisive. In the month of December, he introduced a
+stranger queen into a glass hive, in his cabinet,
+<span class="pagebreak" title="143">&nbsp;</span><a name="Page_143" id="Page_143"></a>
+and confined her
+there. The bees had no opportunity of going out. This stranger was well
+received; her presence awakened the workers from their lethargic state,
+into which they did not relapse; she excited no carnage; the number of
+dead bees on the board of the hive did not sensibly increase; and no
+dead queens were found.</p>
+
+<p>Before concluding any thing favourable to the plurality of queens, it
+was necessary to ascertain whether the native queen was living when the
+new one was introduced into the hive: however the author neglected this;
+and it is very probable the hive had lost its queen, since the bees were
+languid, and the presence of a stranger restored their activity.</p>
+
+<p>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 observations coincide with his, and still
+more, when my experiments justify his conjectures.
+<span class="pagebreak" title="144">&nbsp;</span><a name="Page_144" id="Page_144"></a>
+But I think it
+proper to point out those cases where the imperfections of his hives
+have led him into error, and to explain from what causes I have not seen
+certain facts in the same manner he did. I feel particular anxiety to
+merit your confidence, and I am aware that the greatest exertions are
+necessary, when I have to combat the historian of bees. I confide in
+your judgment; and pray you to be assured of my respect.</p>
+
+<p class="indent1 italic">
+<span class="little">PREGNY</span>, 30. August 1791.
+</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_L" id="fn_L"></a><span class="label"><a href="#fnm_L">L</a></span> Edit. 4to, Tom. V. p. 258.</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="145">&nbsp;</span><a name="Page_145" id="Page_145"></a>
+<a name="LETTER_VIII" id="LETTER_VIII"></a>LETTER VIII.<br /><br />
+
+<span class="little italic">IS THE QUEEN OVIPAROUS? WHAT INFLUENCE HAS THE SIZE OF THE CELLS, WHERE
+THE EGGS ARE DEPOSITED, ON THE BEES PRODUCED?&mdash;RESEARCHES ON THE MODE OF
+SPINNING THE COCCOONS.</span></h2>
+
+
+<p>In this letter I shall collect some isolated observations relative to
+various points in the history of bees, concerning which you wished me to
+engage.</p>
+
+<p>You desired me to investigate whether the queen is really <i>oviparous</i>.
+M. de Reaumur leaves this question undecided. He
+<span class="pagebreak" title="146">&nbsp;</span><a name="Page_146" id="Page_146"></a>
+observes, that he has
+never seen the worm hatched; and he only asserts that worms are found in
+those cells where eggs have been deposited three days preceding. If we
+attempt to catch the moment when the worm leaves the egg, we must extend
+our observations beyond the interior of the hive; for there the
+continual motion of the bees obscures what passes at the bottom of
+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 to hatch the worms, should the eggs
+be too soon deprived of it they wither and perish. The sole method of
+succeeding in seeing the worm come out, consists in watching the queen
+while she lays, in marking the egg so as to be recognised, and removing
+it from the hive to the microscope only an hour or two before the three
+days elapse. The worm will certainly be hatched, provided it has been
+exposed as long as possible to the full degree of
+<span class="pagebreak" title="147">&nbsp;</span><a name="Page_147" id="Page_147"></a>
+heat. Such is the
+course I have pursued; and the following are the results obtained.</p>
+
+<p>In the month of August, we removed several cells containing eggs that
+had been three days deposited: we cut off the top of the cell, and put
+the pyramidal bottom, where the egg was fixed, on a glass slider. Slight
+motions were soon perceptible in the eggs. At first, we could observe no
+external organization: the worm was entirely concealed from us by its
+pellicle. We then prepared to examine the egg with a powerful magnifier;
+however, during the interval, the worm burst its surrounding membrane,
+and cast off part of the envelope, which was torn and ragged on
+different parts of the body, and more evidently so towards the last
+rings. The worm alternately curved and stretched itself, with very
+lively action. Twenty minutes were occupied in casting off the spoil;
+when this exertion ceased: the worm lay down, curved, and seemed to take
+that rest which it required. An egg laid in a worker's cell
+<span class="pagebreak" title="148">&nbsp;</span><a name="Page_148" id="Page_148"></a>
+produced
+this animal, which would have become a worker itself.</p>
+
+<p>We next directed our attention to the moment when a male worm would be
+hatched. An egg was exposed to the sun on a glass slider; and, with a
+good magnifier, nine rings of the worm were perceptible within the
+transparent pellicle. This membrane was still entire, and the worm
+perfectly motionless. The two longitudinal lines of trache&aelig; were visible
+on the surface, and many ramifications. We never lost sight of the egg a
+single instant, and now succeeded in observing the first motions of the
+worm. The thick end alternately straightened and curved, and almost
+reached the part where the sharp extremity was fixed. These exertions
+burst the membrane, first on the upper part, towards the head, then on
+the back, and afterwards on all the rest successively. The ragged
+pellicle remained in folds on different parts of the body, and then fell
+off.
+<span class="pagebreak" title="149">&nbsp;</span><a name="Page_149" id="Page_149"></a>
+Thus it is beyond dispute, that the queen is oviparous.</p>
+
+<p>Some observers affirm, that the workers attend to the eggs before the
+worms are hatched; and it is certain that, at whatever time a hive is
+examined, we always see some workers with the head and thorax inserted
+into cells containing eggs, and remaining motionless several minutes in
+this position. It is impossible to discover what they do, for the
+interior of the cell is concealed by their bodies; but it is very easily
+ascertained that, in this attitude, they are doing nothing to the eggs.</p>
+
+<p>If, at the moment the queen lays, her eggs are put into a grated box,
+and deposited in a strange hive, where there is the necessary degree of
+heat, the worms come out at the usual time, just as if they had been
+left in the cells. Thus no extraordinary aid or attention is required
+for their exclusion.</p>
+
+<p>When the workers penetrate the cells, and remain fifteen or twenty
+minutes motionless,
+<span class="pagebreak" title="150">&nbsp;</span><a name="Page_150" id="Page_150"></a>
+I have reason to believe, it is only to repose from
+their labours. My observations on the subject seem correct. You know,
+Sir, that a kind of irregular shaped cells, are frequently constructed
+on the panes of the hive. These, being glass on one side, are
+exceedingly convenient to the observer, since all that passes within is
+exposed. I have often seen bees enter these cells when nothing could
+attract them. The cells contained neither eggs nor honey, nor did they
+need further completion. Therefore the workers repaired thither only to
+enjoy some moments of repose. Indeed, they were fifteen or twenty
+minutes so perfectly motionless, that had not the dilatation of the
+rings shewed their respiration, we might have concluded them dead. The
+queen also sometimes penetrates the large cells of the males, and
+continues very long motionless in them. Her position prevents the bees
+from paying their full homage to her, yet even then the workers do not
+fail to form a circle around her,
+<span class="pagebreak" title="151">&nbsp;</span><a name="Page_151" id="Page_151"></a>
+and brush the part of her belly that
+remains exposed.</p>
+
+<p>The drones do not enter the cells while reposing, but cluster together
+on the combs; and sometimes retain this position eighteen or twenty
+hours without the slightest motion.</p>
+
+<p>As it is important, in many experiments, to know the exact time that the
+three species of bees exist before assuming their ultimate form, I shall
+here subjoin my own observations on the point.</p>
+
+<p>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. The worm
+now begins spinning its coccoon, in which operation thirty-six hours are
+consumed. In three days, it changes to a nymph, and passes six days in
+this form. It is only on the twentieth day of its existence, counting
+from the moment the egg is laid, that it attains the fly state.</p>
+
+<p>
+<span class="pagebreak" title="152">&nbsp;</span><a name="Page_152" id="Page_152"></a>
+The royal worm also passes three days in the egg, and is five a worm;
+the bees then close its cell; and it immediately begins spinning the
+coccoon, which occupies twenty-four hours. The tenth and eleventh day it
+remains in complete repose, and even sixteen hours of the twelfth. Then
+the transformation to a nymph takes place, in which state four days and
+a third are passed. Thus it is not before the sixteenth day that the
+perfect state of queen is attained.</p>
+
+<p>The male worm passes three days in the egg, six and a half as a worm,
+and metamorphoses into a fly on the twenty-fourth day after the egg is
+laid.</p>
+
+<p>Though the larv&aelig; of bees are apodal, they are not condemned to absolute
+immobility in their cells; for they can move by a spiral motion. During
+the first three days, this motion is so slow as scarcely to be
+perceptible, but it afterwards becomes more evident. I have then
+observed them perform two complete revolutions in an hour
+<span class="pagebreak" title="153">&nbsp;</span><a name="Page_153" id="Page_153"></a>
+and three
+quarters. When the period of transformation arrives, they are only two
+lines from the orifice of the cells. As their position is constantly the
+same, bent in an arc, those in the workers' and drones' cells are
+perpendicular to the horizon, while those in the royal cells lie
+horizontally. It might be thought, that the difference of position has
+much influence on the increment of the different larv&aelig;; yet it has none.
+By reversing combs containing common cells full of brood, I have put the
+worms in a horizontal position; but they were not injured. I have also
+turned the royal cells, so that the worms came into a horizontal
+direction; however their increment was neither slower nor less perfect.</p>
+
+<hr />
+
+<p>I have been much surprised at the mode of bees spinning their coccoons,
+and I have witnessed many new and interesting facts. The worms both of
+workers and males fabricate <i>complete</i> coccoons in their cells;
+<span class="pagebreak" title="154">&nbsp;</span><a name="Page_154" id="Page_154"></a>
+that
+is, close at both ends, and surrounding the whole body. The royal larv&aelig;,
+on the other hand, spin imperfect coccoons, open behind, and enveloping
+only the head, thorax, and first ring of the abdomen. The discovery of
+this difference, which at first may seem trifling, has given me extreme
+pleasure, for it evidently demonstrates the admirable art with which
+nature connects the various characteristics in the industry of bees.</p>
+
+<p>You will remember, Sir, the evidence I gave you of the mutual aversion
+of queens, of the combats in which they engage, 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 a complete coccoon, she 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 retract
+<span class="pagebreak" title="155">&nbsp;</span><a name="Page_155" id="Page_155"></a>
+it, would
+become the victim of her own fury. Thus, that the queen might destroy
+her rivals, it was necessary the last rings of the body should remain
+uncovered, therefore the royal nymphs must only form imperfect 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.</p>
+
+<p>Hitherto, philosophers have claimed our admiration of nature in her care
+of preserving and multiplying the species. But from the facts I relate,
+we must admire her precautions in exposing certain individuals to a
+mortal danger.</p>
+
+<p>The detail on which I have just entered clearly indicates the final
+cause of the opening left by the royal worms in their coccoons; but it
+does not shew whether it is in consequence of a particular instinct that
+they leave this opening, or whether the wideness of their cells prevents
+them from stretching the thread up to the top. This
+<span class="pagebreak" title="156">&nbsp;</span><a name="Page_156" id="Page_156"></a>
+question interested
+me very much; the only method of deciding it was to observe the worms
+while spinning, which cannot be done in their opaque cells. 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 operation
+consisted in extracting worms and introducing them here; but my
+assistant accomplished it with much address. He opened several sealed
+royal cells, where we knew the larv&aelig; were about to begin their coccoons,
+and, taking them gently out, introduced one into each of my glass cells
+without the smallest injury.</p>
+
+<p>They soon prepared to work; and commenced by stretching the anterior
+part of the body in a straight line, while the other was bent in a
+curve. This formed a curve of which the longitudinal sides of the cells
+were tangents, and afforded two points of support. The head was next
+conducted to the different parts of the cell which it could
+<span class="pagebreak" title="157">&nbsp;</span><a name="Page_157" id="Page_157"></a>
+reach, and
+it carpeted the surface with a thick bed of silk. We remarked that the
+threads were not carried from one side to another, and that this would
+have been impracticable, for the worms being obliged to support
+themselves, and to keep the posterior rings curved, the free and
+moveable part of the body was not long enough for the mouth to reach the
+sides diametrically opposite, and fix the threads to them. You will
+remember, Sir, that the royal cells are of a pyramidal form, with a wide
+base, and a long contracted top. These cells hang perpendicularly in the
+hive, the point downwards, from which position the royal worm can be
+supported in the cell, only when the curvature of the posterior part
+forms two points of support; and that it cannot obtain this support
+without resting on the lower part, or towards the extremity. Therefore
+if it attempted to stretch out and spin towards the wide end of the
+cell, it could not reach both sides from being too distant. One part
+would be touched by its extremity,
+<span class="pagebreak" title="158">&nbsp;</span><a name="Page_158" id="Page_158"></a>
+the other by its back, and it would
+consequently 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
+support themselves.</p>
+
+<p>These first experiments obviated the suspicion of any particular
+instinct in the royal worms. They proved, if the worms spun incomplete
+coccoons, it was because they were forced to do so by the figure of
+their cells. However, I wished to have evidence still more direct. I put
+them into cylindrical glass cells, or portions of glass tubes resembling
+common cells, and I had the satisfaction of seeing them spin complete
+coccoons, as the worms of workers do. Lastly, I put common worms in very
+wide cells, and they left the coccoon open. Thus it is demonstrated,
+that the royal worms, and those of workers, have the same instinct and
+the same industry, or in other words, when situated
+<span class="pagebreak" title="159">&nbsp;</span><a name="Page_159" id="Page_159"></a>
+in the same
+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 coccoons, undergo all their metamorphoses equally well. Thus
+the necessity imposed on them by nature, of having the coccoons open, is
+not necessary for their increment; nor has it any other object than that
+of exposing them to the certainty of perishing by the wounds of their
+natural enemy; an observation new and truly singular.</p>
+
+<hr />
+
+<p>I ought to 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.</p>
+
+<p>As we sometimes find 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, influence their size. With this view, you have
+advised me to
+<span class="pagebreak" title="160">&nbsp;</span><a name="Page_160" id="Page_160"></a>
+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 large cells produced workers of
+larger size, we were bound to conclude that the size of the cells had a
+sensible influence on the size of the bees. The first time I made this
+experiment, it did not succeed, because weevils lodged in the hive
+discouraged the bees. But I repeated it afterwards, and the result was
+very remarkable.</p>
+
+<p>I removed the whole comb, consisting of common cells, from one of my
+best glass hives, and left that composed of males' cells alone: and to
+avoid vacuities, I supplied others of the same kind. This was in June,
+the season most favourable to bees. I expected that the bees would
+quickly have repaired the ravages produced by this operation 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
+<span class="pagebreak" title="161">&nbsp;</span><a name="Page_161" id="Page_161"></a>
+work. Expecting they would resume their activity, I continued
+observing them several days; however, my hopes were disappointed. Their
+homage to the queen was not interrupted indeed; but except in this,
+their conduct to the queen was quite different from what it usually is;
+they clustered on the combs without exciting any sensible heat. A
+thermometer among them rose only to 81&deg;, though standing at 77&deg; in the
+open air. In a word, they appeared in a state of the greatest
+despondency.</p>
+
+<p>The queen herself, though very fertile, and though she must have been
+oppressed by her eggs, hesitated long before depositing them in the
+large cells; she chose rather to drop them at random than lay in cells
+unsuitable. However, on the second day, we found six that had been
+deposited there with all regularity. The worms were hatched three days
+afterwards, and then we began to study their history. Though the bees
+provided them with food, they did not carefully attend to it; yet I was
+in hopes they
+<span class="pagebreak" title="162">&nbsp;</span><a name="Page_162" id="Page_162"></a>
+might be reared. I was again disappointed; for next
+morning all the worms had disappeared, and their cells were left empty.
+Profound silence reigned in the hive; few bees left it, and these
+returned without pellets of wax on the limbs; all was cold and
+inanimate. To promote a little motion, I thought of supplying the hive
+with a comb, composed of large cells, full of male brood of all ages.
+The bees, which had twelve days obstinately refused working in wax, did
+not unite this comb to their own. However, their industry was awakened
+in a way that I had not anticipated. They removed all the brood from
+this comb, cleaned out the whole cells, and prepared them for receiving
+new eggs. I cannot determine whether they expected the queen 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 removed all
+the combs
+<span class="pagebreak" title="163">&nbsp;</span><a name="Page_163" id="Page_163"></a>
+composed of large cells to substitute small cells in their
+place, an operation which restored complete activity among the bees.</p>
+
+<p>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, and
+this order cannot be inverted. We have seen that another fact led me to
+the same consequence; and as that was extremely important, I am
+delighted to have it confirmed by a new observation. 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 certain 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 large cells; for, by the
+experiment just described, we learn, that she will
+<span class="pagebreak" title="164">&nbsp;</span><a name="Page_164" id="Page_164"></a>
+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 has instructed her that their size is
+neither proportioned to the size nor necessities of common worms, would
+not she also have been instructed not to lay 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 bees is not very conspicuous. Here the most prominent
+feature of industry appears in the common bees. When I supplied them
+with a comb of small cells, full of male brood, their activity was
+awakened; but instead of bestowing the necessary care on this brood, as
+they would have done in every other situation, they destroyed the
+<span class="pagebreak" title="165">&nbsp;</span><a name="Page_165" id="Page_165"></a>
+whole
+nymphs and larv&aelig;, and cleaned out their cells, that the queen, now
+oppressed with the necessity of laying, might suffer no delay in
+depositing her eggs. Could we allow them either reason or reflection,
+this would be an interesting proof of their affection for her.</p>
+
+<p>The experiment, now detailed at length, not having fulfilled my object
+in determining the influence of the size of the cells on that of the
+worms, I invented another which proved more successful.</p>
+
+<p>Having selected a comb of large cells, containing the eggs and worms of
+males, I removed all the worms from their farina, and my assistant
+substituted those of workers a day old in their place. Then he
+introduced this comb into a hive that had the queen. The bees did not
+abandon these substituted worms; they covered their cells with a top
+almost flat, a kind quite different from what is put on the cells of
+males; which proves, that they were well aware that these, though
+inhabiting large
+<span class="pagebreak" title="166">&nbsp;</span><a name="Page_166" id="Page_166"></a>
+cells, were not males. This comb remained eight days
+in the hive, counting from the time the cells were sealed. I then
+removed it to examine the included nymphs, which proved those of workers
+in different stages of advancement; but, as to size and figure, they
+perfectly resembled what had grown in the smallest cells. I thence
+concluded, that the larv&aelig; of workers do not acquire greater size in
+large than in small cells. Although this 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; undoubtedly
+she has ordained that their organs should be fully expanded, and there
+is sufficient space for that purpose; therefore more would be useless.
+Their expansion ought to be no greater in the most spacious cells than
+in those appropriated 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 will probably
+<span class="pagebreak" title="167">&nbsp;</span><a name="Page_167" id="Page_167"></a>
+be less than that of common workers,
+because they have been cramped in the cells; but it does not thence
+ensue, that a larger cell will admit of them growing to a greater size.</p>
+
+<p>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 larv&aelig;
+of workers in the same circumstances. The large cells of males are
+sufficiently capacious for the perfect expansion of their organs. Thus,
+although reared in cells of still greater capacity, they will grow no
+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 them, and reared 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
+<span class="pagebreak" title="168">&nbsp;</span><a name="Page_168" id="Page_168"></a>
+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
+will be checked, they will not attain the usual size, of which there is
+proof in the following experiment. I had a comb consisting of the cell
+of large drones, and one with those of workers, which also served for
+the male worms. Of these, my assistant took a certain number from the
+smallest cells, and deposited them on a quantity of food purposely
+prepared in the large ones; and in return he introduced into the small
+cells the worms that had been hatched in the other, and then committed
+both to the care of the workers 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 metamorphosis arrived,
+gave both kinds that convex covering usually put on those of the males.
+Eight days afterwards,
+<span class="pagebreak" title="169">&nbsp;</span><a name="Page_169" id="Page_169"></a>
+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.</p>
+
+<p>You suggested another experiment which I carefully made, but it met with
+an unforeseen obstacle. To appreciate the influence of the royal food on
+the expansion 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
+without success. Nor do I think it can ever succeed. 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 deprived of a queen,
+they change the cells containing worms into cells of the largest kind.
+Then the worms will infallibly be converted to queens.</p>
+
+<p>But there is another situation where we can judge of the influence of
+the royal
+<span class="pagebreak" title="170">&nbsp;</span><a name="Page_170" id="Page_170"></a>
+food administered to worms in common cells. I have spoken at
+great length in my letter on the existence of fertile workers. You
+cannot forget, Sir, that the expansion of their sexual organs is owing
+to the reception of some particles of royal jelly, while in the
+vermicular form. For want of new observations, I must refer you to what
+is previously said on the subject.</p>
+
+<p class="toright italic">
+<span class="smcap">Pregny</span>, 4 September 1791.
+</p>
+
+
+
+<h2>
+<span class="pagebreak" title="171">&nbsp;</span><a name="Page_171" id="Page_171"></a>
+<a name="LETTER_IX" id="LETTER_IX"></a>LETTER IX.<br /><br />
+
+<span class="little italic">ON THE FORMATION OF SWARMS.</span></h2>
+
+
+<p>I can add but a few facts to the information M. de Reaumur has
+communicated relative to swarms.</p>
+
+<p>A young queen, according to this celebrated naturalist, is always or
+almost always at the head of a swarm; but he does not assert the fact
+positively, and had some doubts on the subject. "Is it certain,"
+<span class="pagebreak" title="172">&nbsp;</span><a name="Page_172" id="Page_172"></a>
+says
+he "as we have hitherto supposed, in coincidence with all who have
+treated of bees, that the new colony is always conducted by a young
+mother? May not the old mother be disgusted with her habitation? or may
+she not be influenced by some particular 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."</p>
+
+<p>These expressions seem to indicate, that M. de Reaumur suspected that
+the old queens sometimes conducted the young swarms. By the following
+details, you will observe, that his suspicions are fully justified.</p>
+
+<p>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
+
+<span class="pagebreak" title="173">&nbsp;</span><a name="Page_173" id="Page_173"></a>
+prove; and the peculiarities attending it shall be related.</p>
+
+<p>But previous to entering on this subject, I should repeat what has
+already been frequently observed, that the <i>leaf</i> or flat hives are
+indispensible in studying the industry and instinct of bees. When they
+are left at liberty to conduct several rows of parallel combs, we can no
+longer observe what is continually passing between them, or they must be
+dislodged by water or smoke, for examining what has been constructed; a
+violent proceeding, which has a material influence on their instinct,
+and consequently exposes an observer to the risk of supposing simple
+accidents permanent laws.</p>
+
+<p>I now proceed to experiments proving that an old queen always conducts
+the first swarm.</p>
+
+<p>One of my glass hives consisted of three parallel combs, placed in
+squares that opened like the leaves of a book. It was well peopled and
+abundantly provided with
+<span class="pagebreak" title="174">&nbsp;</span><a name="Page_174" id="Page_174"></a>
+honey, wax, and brood, of every age. On the
+fifth of May 1788, I removed the queen, and on the sixth, transferred
+all the bees into another hive, with a fertile queen at least a year
+old. They entered easily and without fighting, and were in general 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 formed regular
+circles around her. However, there was a little agitation in the
+evening, but confined to the surface of the comb where we had put the
+queen, and which she had not quitted. All was perfectly quiet on the
+other side of this comb.</p>
+
+<p>In the morning of the seventh, the bees had destroyed the twelve royal
+cells, but, independent of that, order continued prevalent in the hive;
+the queen laid the eggs of males in the large cells, and those of
+workers in the small ones, respectively.</p>
+
+<p>
+<span class="pagebreak" title="175">&nbsp;</span><a name="Page_175" id="Page_175"></a>
+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
+perpendicularly 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, and
+the longest were only about two lines and a half in depth from the
+bottom to the orifice.</p>
+
+<p>On the thirteenth, the queen seemed already more slender than when
+introduced into the hive; however she still laid some eggs, both in
+common cells and those of males. We also surprised her this day laying
+in a royal cell: she first dislodged the worker there employed, by
+pushing it away with her head, and then supported herself by the
+adjoining cells while depositing the egg.</p>
+
+<p>
+<span class="pagebreak" title="176">&nbsp;</span><a name="Page_176" id="Page_176"></a>
+On the fifteenth, the queen was still more slender: the bees continued
+their attention to the royal cells, which were all unequally advanced;
+some to three or four lines in height, while others were already an inch
+long; which proved that the queen had not laid in the whole at the same
+time.</p>
+
+<p>At the moment when least expected, the hive swarmed on the nineteenth;
+we were warned of it by the noise in the air; and hastened to collect
+and put the bees into a hive purposely prepared. Though we had
+overlooked the facts attending the departure of the swarm, the object of
+this experiment was fulfilled; for, on examination of all the bees, we
+were convinced they had been conducted by the old queen; by that we
+introduced on the sixth of the month, and which had been deprived of one
+of the antenn&aelig;. Observe, there was no other queen in this colony. In the
+hive she had left, we found seven royal cells close at the top, but open
+at the side, and quite empty.
+<span class="pagebreak" title="177">&nbsp;</span><a name="Page_177" id="Page_177"></a>
+Eleven more were sealed; and some others
+newly begun; no queen remained in the hive.</p>
+
+<p>The new swarm next became the object of our attention: we observed it
+during the rest of the year, during winter and the subsequent spring;
+and, in April, we had the satisfaction of seeing a new swarm depart with
+the same queen at its head that had conducted the former swarm in May
+the preceding year.</p>
+
+<p>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 royal cells; she laid in
+one of them before us; and in the last place led forth the swarm.</p>
+
+<p>We have several times repeated the same experiment with equal success.
+Thus it appears incontestible, that the old queen always conducts the
+first swarm; but never quits the hive before depositing eggs in the
+royal cells, from which other queens will
+<span class="pagebreak" title="178">&nbsp;</span><a name="Page_178" id="Page_178"></a>
+proceed after her departure.
+The bees prepare these cells only while the queen lays male eggs; and a
+remarkable fact attends it, that after this laying terminates, her belly
+being considerably diminished, she can easily fly, whereas, her belly is
+previously so heavy she can hardly drag it along. Therefore it is
+necessary she should lay in order to be in a condition for undertaking
+her journey, which may sometimes be very long.</p>
+
+<p>But this single condition is not enough. It is also requisite that the
+bees be very numerous: they should even be superabundant, and a person
+might say they are aware of it: for, if the hive is thin, no royal cells
+are constructed when the male eggs are laid, which is solely at the
+period that the queen is able to conduct a colony. This fact was proved
+by the following experiment on a large scale.</p>
+
+<p>On the third of May 1788, we divided eighteen hives into two portions;
+all the queens were about a year old. Thus each
+<span class="pagebreak" title="179">&nbsp;</span><a name="Page_179" id="Page_179"></a>
+portion of the hives
+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 of the hives threw a swarm.&mdash;Therefore, if the
+hive containing the old queen is not very populous, she remains in it
+until the subsequent spring; and if the population is then sufficient,
+royal cells will be constructed: the queen 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.</p>
+
+<p>Such is a very brief abstract of my observations on swarms conducted by
+old queens. You must excuse the long detail on which I am about to
+enter, concerning 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
+hitherto been very obscure. A long course of observations, protracted
+even during several
+<span class="pagebreak" title="180">&nbsp;</span><a name="Page_180" id="Page_180"></a>
+years, was necessary to remove, in some degree, the
+veil that concealed these mysteries. I have been indemnified for the
+trouble, indeed, by the pleasure of seeing my experiments reciprocally
+confirmed; but, considering the assiduity required in these researches,
+they were truly very laborious.</p>
+
+<p>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 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
+queens; and I shall now extract the chief experiments from my journal.</p>
+
+<p>On the fourteenth of May, we introduced two portions of bees, from the
+straw hives, into a large glass hive very flat; and allowed them only
+one queen of the preceding year, and which had already commenced laying
+in its native hive. We introduced her on the fifteenth. She was
+<span class="pagebreak" title="181">&nbsp;</span><a name="Page_181" id="Page_181"></a>
+very
+fertile. The bees received her well, and she soon began to lay in large
+and small cells alternately.</p>
+
+<p>On the twentieth, we saw the formation of twelve royal cells, all on the
+edges of the communications, or passages through the combs, and shaped
+liked stalactites.</p>
+
+<p>On the twenty-seventh, ten were much but unequally enlarged; but none so
+long as when the worms are hatched.</p>
+
+<p>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 became more lively, yet she still continued examining
+the cells as when about to lay; sometimes introducing half her belly,
+but suddenly withdrawing it, without having laid. At other times she
+deposited an egg, which lay in an irregular position, on one side of the
+hexagon, and not fixed by an end to the bottom of the cell. The queen
+produced no distinct sound in her course, and we heard nothing different
+from the ordinary humming of bees.
+<span class="pagebreak" title="182">&nbsp;</span><a name="Page_182" id="Page_182"></a>
+She passed over those in her way;
+sometimes when she stopped, the bees meeting her also stopped; and
+seemed to consider her. They advanced briskly, struck her with their
+antenn&aelig;, and mounted on her back. She then went on carrying some of the
+workers on her back. None gave her honey, but she voluntarily took it
+from the cells in her way. The bees no longer inclosed and formed
+regular circles around her. The first, aroused by her motions, followed
+her running in the same manner, and in their passage excited those still
+tranquil on the combs. The way the queen had traversed was evident after
+she left it, by the agitation created, which was never afterwards
+quelled: she had soon visited every part of the hive, and occasioned a
+general agitation; if some places still remained tranquil, the bees in
+agitation arrived, and communicated their motion. The queen no longer
+deposited her eggs in cells; she let them fall fortuitously: nor did the
+bees any longer watch over the young; they
+<span class="pagebreak" title="183">&nbsp;</span><a name="Page_183" id="Page_183"></a>
+ran about in every different
+direction; even those returning from the fields, before the agitation
+came to its height, no sooner entered the hive than they participated in
+these tumultuous motions. They neglected to free themselves of the waxen
+pellets on their limbs, and ran blindly about. At last the whole rushed
+precipitately towards the outlets of the hive, and the queen along with
+them.</p>
+
+<p>As it was of much consequence to see the formation of new swarms in this
+hive, and, for that reason, as I wished it to continue very populous, I
+removed the queen, at the moment she came out, that the bees might not
+fly too far, and 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 removed its queen also.</p>
+
+<p>All these facts were certain, and appeared susceptible of no error.
+Notwithstanding
+<span class="pagebreak" title="184">&nbsp;</span><a name="Page_184" id="Page_184"></a>
+this, I was particularly earnest to learn whether old
+queens always followed the same course; which induced me, on the
+twenty-ninth, to replace, in the glass hive, the queen a year old, which
+had hitherto been the subject of my experiments, and had just began to
+lay the eggs of males. On the same day, we found one of the royal cells
+left by the preceding queen larger than the rest; and, from its length,
+supposed the included worm two days old: the egg had, therefore, been
+laid on the twenty-fourth by that queen, and the worm was hatched on the
+twenty-seventh. On the thirtieth, the queen laid a great deal in the
+large and small cells alternately. Now, and the two following days, the
+bees enlarged several royal cells, but unequally, which proved that they
+included larv&aelig; of different ages. One was closed on the first of June,
+and on the second another. The bees also commenced some new ones. All
+was perfectly quiet at eleven in the morning; but, at
+<span class="pagebreak" title="185">&nbsp;</span><a name="Page_185" id="Page_185"></a>
+mid-day, the
+queen, from the utmost tranquillity, became evidently agitated; and her
+agitation insensibly communicated to the workers in every part of their
+dwelling. In a few minutes they precipitately crowded to the entrances,
+and, along with the queen, left the hive. After they had settled on the
+branch of a neighbouring tree, I sought for the queen; thinking that, by
+removing her, the bees might return to the hive, which actually ensued.
+Their first care seemed to consist in seeking the female; they were
+still in great agitation, but gradually calmed; and in three hours
+complete tranquillity was restored.</p>
+
+<p>They had resumed their usual occupations on the third: 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, not by applying wax
+cordons, but by removing wax from the surface. Towards the top this
+waved work is almost imperceptible; it becomes deeper above, and the
+
+<span class="pagebreak" title="186">&nbsp;</span><a name="Page_186" id="Page_186"></a>
+workers excavate it still more from thence to the base of the pyramid.
+The cell, when once shut, also becomes thinner; and is so much so,
+immediately preceding the queen's metamorphosis from a nymph, that all
+its motions are perceptible through the thin covering of wax on which
+the waved work is founded. It is a very remarkable circumstance, that in
+making the cells thinner, from the moment they are closed, the bees know
+to regulate their labour so that it terminates only when the nymph is
+ready to undergo its last metamorphosis.</p>
+
+<p>On the seventh day the coccoon is almost completely <i>unwaxed</i>, if I may
+use the expression, at the part next to the head and thorax of the
+queen. This operation facilitates her exit; for she has nothing to do
+but cut the silk that forms the coccoon. Most probably the object is, to
+promote evaporation of the superabundant fluids of the nymph. I have
+made some direct experiments to ascertain the fact, but they
+<span class="pagebreak" title="187">&nbsp;</span><a name="Page_187" id="Page_187"></a>
+are yet
+unfinished. A third royal cell was closed by the bees on the same day,
+the third of June, twenty-four hours after closing the second. The like
+was done to other royal cells successively, during the subsequent days.</p>
+
+<p>Every moment of the seventh, we expected the queen to leave the royal
+cell shut on the thirtieth of May. The seven days had elapsed. 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 being
+unwilling that she should yet quit her cell, they had soldered the
+covering to it with some particles of wax. What seemed most singular
+was, that this female emitted a very distinct sound, or clacking from
+her prison. It was still more audible in the evening, and even consisted
+of several monotonous notes in rapid succession.</p>
+
+<p>
+<span class="pagebreak" title="188">&nbsp;</span><a name="Page_188" id="Page_188"></a>
+The same sound proceeded from the royal cell on the eighth. Several
+bees kept guard round each royal cell.</p>
+
+<p>The first cell opened on the ninth. The young queen was lively, slender,
+and of a brown colour. Now, we understood why bees retain the female
+captive in their cells, after the period for transformation has elapsed;
+it is, that they may be able to fly the instant they are hatched. The
+new queen occupied all our attention. When she approached the other
+royal cells, the bees on guard pulled, bit her, and chased her away;
+they seemed to be greatly irritated against her, and she enjoyed
+tranquillity only when at a good distance from these cells. This
+procedure was frequently repeated through the day. She twice emitted the
+sound; in doing so she stood, her thorax against a comb, and the wings
+crossed on her back; they were in motion but without being unfolded or
+further opened. Whatever might be the cause of her assuming this
+attitude, the bees were
+<span class="pagebreak" title="189">&nbsp;</span><a name="Page_189" id="Page_189"></a>
+affected by it; all hung down their heads, and
+remained motionless.</p>
+
+<p>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 approached, all the guards became agitated,
+surrounded her on all sides, bit, and commonly drove her away; sometimes
+when in these circumstances, she emitted her sound, assuming the
+position just described, from that moment the bees became motionless.</p>
+
+<p>The queen confined in the second cell had not yet left it, and was heard
+to hum several times. We accidentally discovered how the bees fed her.
+On attentive examination, a small aperture was perceptible in the end of
+the coccoon which she had cut to escape, and which her guards had again
+covered with wax, to confine her still longer. She thrust her trunk
+through the cleft; at first the bees did not observe it alternately
+thrust out and drawn
+<span class="pagebreak" title="190">&nbsp;</span><a name="Page_190" id="Page_190"></a>
+in, but one at length perceiving it, came to apply
+its trunk to that of the captive queen, and then gave way to others that
+also approached her with honey. When satisfied she retracted her trunk,
+and the bees again closed up the opening with wax.</p>
+
+<p>The queen this day between twelve and one became extremely agitated. The
+royal cells had multiplied very much; she could go no where without
+meeting them, and on approaching she was very roughly treated. Then she
+fled, but to obtain no better reception. At last, these things agitated
+the bees; they precipitately rushed through the outlets of the hive, and
+settled on a tree in the garden. It singularly happened that the queen
+was herself unable to follow or conduct the swarm. She had attempted to
+pass between two royal cells before they were abandoned by the bees
+guarding them, and she was so confined and maltreated as to be incapable
+of moving. We then removed her into a separate hive prepared for a
+particular experiment;
+<span class="pagebreak" title="191">&nbsp;</span><a name="Page_191" id="Page_191"></a>
+the bees, which had clustered on a branch, soon
+discovered their queen was not present, and returned of their own accord
+to the hive. Such is an account of the second colony of this hive.</p>
+
+<p>We were extremely solicitous to ascertain what would become of the other
+royal cells. Four of the close ones had attained complete maturity, and
+the queens would have left them had not the bees prevented it. They were
+not open either previous to the agitation of the swarms, or at the
+moment of swarming.</p>
+
+<p>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 formed the swarm. We could not
+discover what occasioned the difference in their captivity.</p>
+
+<p>On the twelfth, the queen was at last liberated, as we found her in the
+hive. She had been treated exactly as her predecessor; the bees allowed
+her to rest in quiet, when
+<span class="pagebreak" title="192">&nbsp;</span><a name="Page_192" id="Page_192"></a>
+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. Returning at mid-day, we were greatly surprised to find it
+almost totally deserted. During our absence, it had thrown a prodigious
+swarm, which still clustered on the branch of a neighbouring tree. We
+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, escaped. Thus, there was no doubt
+of both queens being in the swarm. We found it so; and removed them,
+that the bees might return to the hive, which they did very soon.</p>
+
+<p>While we were occupied in this operation, the fourth captive queen left
+her prison, and the bees found her on returning. At first they were very
+much agitated, but calmed towards the evening, and resumed
+<span class="pagebreak" title="193">&nbsp;</span><a name="Page_193" id="Page_193"></a>
+their wonted
+labours. They formed a strict guard around the royal cells, and took
+great care to remove the queen whenever she attempted to approach.
+Eighteen royal cells now remained to be guarded.</p>
+
+<p>The fifth queen left her cell at ten at night; therefore two queens were
+now in the hive. They immediately began fighting, but came to disengage
+themselves from each other. However they fought several times during the
+night without any thing decisive. Next day, the thirteenth, we witnessed
+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.</p>
+
+<p>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 strikes the bees motionless. For some
+minutes, we conceived, that taking advantage of the dread exhibited by
+the
+<span class="pagebreak" title="194">&nbsp;</span><a name="Page_194" id="Page_194"></a>
+workers on guard, she would open it, and destroy the young female;
+also she prepared to mount the cell; but in doing so she ceased the
+sound, and quitted that attitude which paralyses the bees. The guardians
+of the cell instantly took courage; and, by means of tormenting and
+biting the queen, drove her away.</p>
+
+<p>On the fourteenth, the sixth young queen appeared, and the hive threw a
+swarm, with all the concomitant disorder before described. The agitation
+was so considerable, that a sufficient number of bees did not remain to
+guard the royal cells, and several of the imprisoned queens were thus
+enabled to make their escape. Three were in the cluster formed by the
+swarm, and other three remained in the hive. We removed those that had
+led the colony, to force the bees to return. They entered the hive,
+resumed their post around the royal cells, and maltreated the queens
+when approaching.</p>
+
+<p>
+<span class="pagebreak" title="195">&nbsp;</span><a name="Page_195" id="Page_195"></a>
+A duel took place in the night of the fifteenth, in which one queen
+fell. We found her dead next morning before the hive; but three still
+remained, as one had been hatched during night. Next morning we saw a
+duel. Both combatants were extremely agitated, either with the desire of
+fighting, or the treatment of the bees, when they came near the royal
+cells. Their agitation quickly communicated to the rest of the bees, and
+at mid-day they departed impetuously 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, which I shall
+give you no account of, as it shewed nothing new.</p>
+
+<p>Unfortunately we lost this, which was a very strong swarm; the bees flew
+out of sight, and could never be found. The hive was now very thinly
+inhabited. Only the few bees that <a name="corr195" id="corr195"></a><a class="correction" href="#cnote195" title="missing word inserted">had</a> not
+participated in the general agitation remained, and those that returned
+from the fields after the swarm had
+<span class="pagebreak" title="196">&nbsp;</span><a name="Page_196" id="Page_196"></a>
+departed. The cells were,
+therefore, slenderly guarded; the queens escaped from them, and engaged
+in several combats, until the throne remained with the most successful.</p>
+
+<p>Notwithstanding the victories of this queen, she was treated with great
+indifference from the sixteenth to the nineteenth, that is, the three
+days that she preserved her virginity. At length, having gone to seek
+the males, she returned with all the external signs of fecundation, and
+was henceforth received with every mark of respect; she laid her first
+eggs forty-six hours after fecundation.</p>
+
+<p>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
+experiments which I made at the same time for elucidating various
+obscure points of their history. These shall be the subject of future
+letters. For, although I
+<span class="pagebreak" title="197">&nbsp;</span><a name="Page_197" id="Page_197"></a>
+have said so much, I hope still to interest
+you.</p>
+
+<p class="toright italic">
+<span class="smcap">Pregny</span>, 6. September 1791.
+</p>
+
+<p><i>P. S.</i>&mdash;In revising this letter, I find I have neglected taking notice
+of an objection that may embarrass my readers, and which ought to be
+answered.</p>
+
+<p>After the first five swarms had thrown, I had always returned the bees
+to the hive: it is not surprising, therefore, that it was continually so
+sufficiently stocked that each colony was numerous. But things are
+otherwise in the natural state: the bees composing a swarm do not return
+to the hive; and it will undoubtedly be asked, What resource enables a
+common hive to swarm three or four times without being too much
+weakened?</p>
+
+<p>I cannot lessen the difficulty. I have observed that the agitation,
+which precedes the swarming, is often so considerable, that most of the
+bees quit the hive, and in that
+<span class="pagebreak" title="198">&nbsp;</span><a name="Page_198" id="Page_198"></a>
+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.</p>
+
+<p>But remark, in the first place, that the queen leaves a prodigious
+quantity of workers' brood, which soon transforms to bees; and in this
+way the population sometimes becomes almost as great after swarming as
+before it.</p>
+
+<p>Thus the hive is perfectly capable of affording a second colony without
+being too much weakened. The third and fourth swarm weaken it more
+sensibly; but the inhabitants always remain in sufficient numbers to
+preserve the course of their labours uninterrupted; and the losses are
+soon repaired by the great fecundity of the queen, as she lays above an
+hundred eggs a day.</p>
+
+<p>If, in some cases, the agitation of swarming 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
+<span class="pagebreak" title="199">&nbsp;</span><a name="Page_199" id="Page_199"></a>
+then that the bees are ranging through the country. Those that are
+out, therefore, cannot share in the agitation; when returned to the
+hive, they quietly resume their labours; and their number is not small,
+for, when the weather is fine, at least a third of the bees are employed
+in the fields at once.</p>
+
+<p>Even in the most embarrassing case, namely, where the whole bees desert
+the hive, it does not follow, that all those endeavouring to depart
+become members of the new colony. When this agitation or delirium 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 the rest, seem
+drenched in perspiration; their wings grow moist; they are incapable of
+flight; and even when able to escape, they advance no farther than the
+board of the hive, and soon return.</p>
+
+<p>
+<span class="pagebreak" title="200">&nbsp;</span><a name="Page_200" id="Page_200"></a>
+Those that have lately left their cells remain behind the swarm, still
+feeble, they could not support themselves in flight. Here then are also
+many recruits to people what we should have thought a deserted
+habitation.</p>
+
+
+
+<h2>
+<span class="pagebreak" title="201">&nbsp;</span><a name="Page_201" id="Page_201"></a>
+<a name="LETTER_X" id="LETTER_X"></a>LETTER X.<br /><br />
+
+<span class="little italic">THE SAME SUBJECT CONTINUED.</span></h2>
+
+
+<p>To preserve greater regularity in continuing the history of swarms, I
+think it proper to recapitulate in a few words the principal points of
+the preceding letter, and to expatiate on each, concerning the result of
+new experiments, respecting which I have still been silent.</p>
+
+<p>
+<span class="pagebreak" title="202">&nbsp;</span><a name="Page_202" id="Page_202"></a>
+In the first place. <i>If at the return of spring, we examine a hive well
+peopled, and governed by a fertile queen, we shall see her lay a
+prodigious number of male eggs in the course of May, and the workers
+will chuse that moment for constructing several royal cells of the kind
+described by M. de Reaumur.</i> Such is the result of several long
+continued observations, among which there has not been the slightest
+variation, and I cannot hesitate in announcing it as demonstrated.
+However, I should here add the necessary explanation. It is necessary
+that the queen, before commencing her <i>great</i> laying of the eggs of
+males, be eleven months old; when young she lays only those of workers.
+A queen, hatched in spring, will perhaps lay fifty or sixty eggs of
+drones in whole, but before beginning her great laying of them, which
+should be two thousand in a month, she must have completed her eleventh
+month in age. In the course of our experiments, which more or less
+disturbed the natural state of things, it often happened that the
+<span class="pagebreak" title="203">&nbsp;</span><a name="Page_203" id="Page_203"></a>
+queen
+did not attain this age until October, and immediately began laying male
+eggs. The workers, as if induced by some emanation from the eggs, also
+adopted this time for building the royal cells. No swarm resulted
+thence, it is true, because in autumn all the necessary circumstances
+are absolutely wanting, but it is not less evident, that there is a
+secret relation between the production of the eggs of males, and the
+construction of royal cells.</p>
+
+<p>This laying commonly continues thirty days. The bees on the twentieth or
+twenty-first lay the foundation of several royal cells. Sometimes they
+build sixteen or twenty; we have even had twenty-seven. When the cells
+are three or four lines high, the queen lays those eggs from which her
+own species will come, but not the whole in one day. That the hive may
+throw several swarms, it is essential that the young females conducting
+them be not all produced at the same time. One may affirm, that the
+queen anticipates the
+<span class="pagebreak" title="204">&nbsp;</span><a name="Page_204" id="Page_204"></a>
+fact, for she takes care to allow at least the
+interval of a day between every egg deposited in the cells. It is proved
+by the bees knowing to close the cells the moment the worms are ready to
+metamorphose to nymphs. Now, as they close all the royal cells at
+different periods, it is evident the included worms are not all of an
+equal age.</p>
+
+<p>The queen's belly is very turgid before she begins laying the eggs of
+drones; but it sensibly decreases as she advances, and when terminated
+is very small. Thus she finds herself in a condition to undertake a
+journey which circumstances may prolong; thus this condition was
+necessary; 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.</p>
+
+<p>Secondly. <i>When the larv&aelig; hatched from the eggs laid by the queen, in
+the royal cells, are ready to transform to nymphs, this queen leaves the
+swarm conducting a swarm along
+<span class="pagebreak" title="205">&nbsp;</span><a name="Page_205" id="Page_205"></a>
+with her; and the first swarm that
+proceeds from the hive is uniformly conducted by the old queen.</i><a name="fnm_M" id="fnm_M"></a><a href="#fn_M" class="fnnum">M</a> I
+think I can divine the reason of it.</p>
+
+<p>That there may never be a plurality of females in a hive, nature has
+inspired queens with a natural horror against each other; they never
+meet without endeavouring to fight, and to accomplish their mutual
+destruction. Thus, the chance of combat is equal between them, and
+fortune will decide to which the empire shall pertain. But if one
+combatant is older than the rest, she is stronger, and the advantage
+will be with her. She will destroy her rivals successively as produced.
+Thus, if the old queen did not leave the hive, when the young ones
+undergo their last metamorphosis, it could produce no more swarms, and
+the species would perish. Therefore, to preserve the species, it is
+necessary that the old queen conduct the first swarm. But what is the
+secret means employed by nature to induce her departure? I am ignorant
+of it.</p>
+
+<p>
+<span class="pagebreak" title="206">&nbsp;</span><a name="Page_206" id="Page_206"></a>
+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 old queen; and that may
+happen thus. Nature has not willed that the queen shall quit the first
+hive before her production of male eggs is finished. It is necessary 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 workers. The bees in that case would have no means of replacing
+her, and the colony would go to ruin.</p>
+
+<p>All these things have been with infinite wisdom foreseen. The first
+operation of the bees of a swarm is to construct the cells of workers.
+They labour at them with great ardour, and as the ovaries of the queen
+have been disposed with admirable foresight, the first eggs she has to
+lay in
+<span class="pagebreak" title="207">&nbsp;</span><a name="Page_207" id="Page_207"></a>
+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. It may be affirmed, that the bees
+know their queen will also lay the eggs of drones; she actually does
+begin to deposit some, though in much smaller number than at first;
+enough however to encourage the bees to construct royal cells. Now, if
+in these circumstances the weather is favourable, it is not impossible
+that a second colony may be formed, and that the queen may depart at the
+head of it within three weeks of conducting the first swarm. But I
+repeat, the fact is rare in our climate. Let me now return to the hives
+from which the queen has led the first colony.</p>
+
+<p>Thirdly. <i>After the old queen has conducted the first swarm from the
+hive, the remaining bees take particular care of the royal cells, and
+prevent the young queens successively hatched from leaving them, unless
+at an interval of several days between each.</i></p>
+
+<p>
+<span class="pagebreak" title="208">&nbsp;</span><a name="Page_208" id="Page_208"></a>
+In the preceding letter, I have given you the detail and proof of this
+fact, and I shall here add some reflexions. During the period of
+swarming, the conduct or instinct of bees seems to receive a particular
+modification. At all other times, when they have lost their queen, they
+appropriate workers worms to replace her; they prolong and enlarge the
+cells of these worms; they supply them with aliment more abundantly, and
+of a more pungent taste; and by this alteration, the worms that would
+have changed to common bees are transformed to queens. We have seen
+twenty-seven 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 may perhaps leave her cell, and attack all
+the other royal cells successively, which she will tear open to destroy
+her rivals, without the workers taking any part in their defence. Should
+several queens be hatched at once, they will pursue each other, and
+fight until
+<span class="pagebreak" title="209">&nbsp;</span><a name="Page_209" id="Page_209"></a>
+the throne remain with her that is victorious. Far from
+opposing such duels, the other bees rather seem to excite the
+combatants.</p>
+
+<p>Things are quite reversed during the period of swarming. The royal cells
+then constructed are of a different figure from the former. They
+resemble stalactites, and in the beginning are like the cup of an acorn.
+The bees assiduously guard the cells when the young queens are ready for
+their last metamorphosis. At length the female hatched from the first
+egg laid by the old queen leaves her cell; the workers at first treat
+her with indifference. But she, immediately 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
+drive her away, so that she is forced to remove; but the royal cells
+being numerous, scarce can she find a place of rest. Incessantly
+harassed with the desire of attacking the other queens, and
+<span class="pagebreak" title="210">&nbsp;</span><a name="Page_210" id="Page_210"></a>
+incessantly
+repelled, she becomes agitated, and hastily traverses the different
+groupes of workers, to which she communicates her agitation. At this
+moment numbers of bees rush towards the aperture of the hive, and, with
+the young queen at their head, depart to seek another habitation.</p>
+
+<p>After the departure of the colony, the remaining workers set another
+queen at liberty, and treat her with equal indifference as the first.
+They drive her from the royal cells; being perpetually harrassed, she
+becomes agitated; departs, and carries a new swarm along with her. In a
+populous hive this scene is repeated three or four times during spring.
+As the number of bees is so much reduced, that they are no longer
+capable of preserving a strict watch over the royal cells, several
+females then leave their confinement at once; they seek each other,
+fight, and the queen at last victorious reigns peaceably over the
+republic.</p>
+
+<p>
+<span class="pagebreak" title="211">&nbsp;</span><a name="Page_211" id="Page_211"></a>
+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 after the first colony is led out by the old queen,
+until the next swarm is conducted by the first young queen set at
+liberty. The interval between the second and third is still shorter; and
+the fourth sometimes departs 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.</p>
+
+<p>A swarm is never seen except in a fine day, or, to speak more correctly,
+at a time of the day when the sun shines, and the air is calm. Sometimes
+we have observed all the precursors of swarming, disorder and agitation,
+but a cloud passed before the sun, and tranquillity was restored; the
+bees thought no more of swarming. An hour afterwards, the sun having
+again appeared,
+<span class="pagebreak" title="212">&nbsp;</span><a name="Page_212" id="Page_212"></a>
+the tumult was renewed; it rapidly augmented; and the
+swarm departed.</p>
+
+<p>Bees generally seem much alarmed at the prospect of bad weather. While
+ranging in the fields the passage of a cloud before the hive induces
+them precipitately to return. I am induced to think they are disquieted
+by the sudden diminution of light. For if the sky is uniformly obscured,
+and there is no alteration in clearness or in the clouds dispelling,
+they proceed to the fields for their ordinary collections, and the first
+drops of a soft rain does not make them return with much precipitation.</p>
+
+<p>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 sometimes seem to use this right in an arbitrary manner. However
+the confinement of the queens is always longer when bad weather lasts
+several days together. Here the final object cannot be mistaken.
+<span class="pagebreak" title="213">&nbsp;</span><a name="Page_213" id="Page_213"></a>
+If the
+young females were at liberty to leave their cradles during these bad
+days, there would be a plurality of queens in the hive, consequently
+combats; and victims would fall. Bad weather might continue so long,
+that all the queens might at once have undergone their last
+metamorphosis, or attained their liberty. One victorious over the whole
+would enjoy the throne, and the hive, which should naturally produce
+several swarms, could give only one. Thus the multiplication of the
+species would have been left to the chance of rain, or fine weather,
+instead of which it is rendered independent of either, by the wise
+dispositions of nature. By allowing only a single female to escape at
+once, the formation of swarms is secured. This explanation appears so
+simple, that it is superfluous to insist farther on it.</p>
+
+<p>But I should mention another important circumstance resulting from the
+captivity of queens; which is, that they are in a condition to fly, when
+the bees have given
+<span class="pagebreak" title="214">&nbsp;</span><a name="Page_214" id="Page_214"></a>
+them liberty, and by this means are capable of
+profiting by the first moment of sunshine to depart at the head of a
+colony.</p>
+
+<p>You well know, Sir, that all drones and workers are not in a condition
+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 hours must elapse before the acquisition of perfect strength, and
+the development of all their faculties. It would be the same with the
+females was not their confinement protracted after the period of
+transformation; 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 used to retain the queens in
+captivity. The bees solder the covering to the sides of the cell by a
+cordon of wax. As I have also explained how they are fed, it need not be
+repeated here.</p>
+
+<p>It is likewise a very remarkable fact, that queens are set at liberty
+earlier or later according
+<span class="pagebreak" title="215">&nbsp;</span><a name="Page_215" id="Page_215"></a>
+to their age. Immediately when the royal
+cells were sealed, we marked them all with numbers, and we chose this
+period because it indicated the age of the queens exactly. The oldest
+was first liberated, then the one immediately younger, and so on with
+the rest. None of the younger queens were set at liberty before the
+older ones.</p>
+
+<p>I have a thousand times asked myself how the bees could so accurately
+distinguish the age of their captives. Undoubtedly I should do better to
+answer this question by a simple avowal of my ignorance. At the same
+time, I must be permitted to state a conjecture. You will admit, that I
+have not, as some authors, abused the right of giving myself up to
+hypothesis; may not the humming 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 queens? It is certain that the female, whose
+cell is first sealed, is also the first to emit this sound.
+<span class="pagebreak" title="216">&nbsp;</span><a name="Page_216" id="Page_216"></a>
+That in the
+next emits it sooner than the rest, and so on with those immediately
+subsequent. As their captivity may continue six days, it is possible
+that the bees in this space of time may forget which has emitted it
+first; but it is also possible, that the queens diversify the sounds,
+encreasing the loudness as they become older, and that the bees can
+distinguish these variations. We have even ourselves been able to
+distinguish differences in the sound, either with relation to the
+succession of notes, or their intensity; and probably there are
+gradations still more imperceptible that escape our organs, but may be
+sensible to those of the workers.</p>
+
+<p>What gives weight to this conjecture is, that the queens brought up by
+M. Schirach's method, are perfectly mute; neither do the workers form
+any guard around their cells, nor do they retain them in captivity a
+moment beyond the period of transformation, and, when they have
+undergone it, they are allowed to combat until one
+<span class="pagebreak" title="217">&nbsp;</span><a name="Page_217" id="Page_217"></a>
+has become
+victorious over all the rest. Why? Because the object is only to replace
+the last queen. Now, provided that among the worms reared as queens,
+only one succeeds, the fate of the others is uninteresting to the bees,
+whereas, during the period of swarming, it is necessary to preserve a
+succession of queens, for conducting the different colonies; and to
+ensure the safety of the queens, it is necessary to avert the
+consequences of the mutual horror by which they are animated against
+each other. Behold the evident cause of all the precautions that bees,
+instructed by nature, take during the period of swarming; behold an
+explanation of the captivity of females; and that the duration of their
+captivity 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.</p>
+
+<p>
+<span class="pagebreak" title="218">&nbsp;</span><a name="Page_218" id="Page_218"></a>
+In spite of all my researches, I have never been able to discover the
+situation of the organ which produces the sound. But I have instituted a
+new course of experiments on the subject, which are still unfinished.</p>
+
+<p>Another problem still remains for solution. Why are the queens reared,
+according to M. Schirach's method, mute, whilst those bred in 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 M. Schirach's method,
+receive it only the second or third day of their existence. It appears
+to me that this circumstance may have an influence on the different
+parts of organisation, and particularly on the organ of voice.
+
+<span class="pagebreak" title="219">&nbsp;</span><a name="Page_219" id="Page_219"></a>
+Experiment has not confirmed this conjecture. I constructed glass cells
+in perfect imitation of royal cells, that the metamorphosis of the worms
+into nymphs, and of the nymphs to queens, might be visible. These
+experiments are related in a preceding letter. Into one of these
+artificial cells we introduced the nymph of a worm, reared according to
+M. Schirach's method, twenty-four hours before it could naturally
+undergo its last metamorphosis; and we replaced the glass cell in the
+hive, that the nymph might have the necessary degree of heat. Next 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 had contrived an aperture for her thrusting out her
+trunk, and that the bees might feed her. I expected that she would have
+been completely mute; but it was otherwise; for she emitted sounds
+similar to those already
+<span class="pagebreak" title="220">&nbsp;</span><a name="Page_220" id="Page_220"></a>
+described, therefore my conjecture was
+erroneous.</p>
+
+<p>I next conceived that the queen being restrained in her motions, and in
+her desire for liberty, was induced to emit certain sounds. All queens,
+in this new point of view, are equally capable of emitting the sound,
+but to induce them to it, they must be in a confined situation. In the
+natural state, the queens that come from workers are not a single
+instant in restraint; 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 are 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 something more probable.</p>
+
+<p>I do not ascribe to myself the credit of having discovered the humming
+of the queen bee. Old authors speak of it. M.
+<span class="pagebreak" title="221">&nbsp;</span><a name="Page_221" id="Page_221"></a>
+de Reaumur cites a Latin
+work published 1671, <i>Monarchia Femina</i>, by Charles Butler. He gives a
+very brief abstract of this naturalist's observations, who we easily see
+has exaggerated or rather disguised the truth, by mixing it with the
+most absurd fancies; but it is not the less evident that Butler has
+heard this peculiar humming of queens, and that he did not confound it
+with the confused humming sometimes heard in hives.</p>
+
+<p>Fourthly. <i>The young queens conducting swarms from their native hive are
+still in a virgin state.</i> The day after, being settled in their new
+abode, they generally depart in quest of the males; and this is usually
+the fifth day of their existence as queens; for two or three pass in
+captivity, one in their native hive, and a fifth in their new dwelling.
+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
+<span class="pagebreak" title="222">&nbsp;</span><a name="Page_222" id="Page_222"></a>
+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 they begin to lay. The old queen,
+which leads the first swarm in spring, requires no farther commerce with
+the males, for preservation of her fecundity. A single copulation is
+sufficient to impregnate all the eggs she will lay for at least two
+years.</p>
+
+<p class="toright italic">
+<span class="smcap">Pregny</span>, 8. September 1791.
+</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_M" id="fn_M"></a><span class="label"><a href="#fnm_M">M</a></span> Schirach seems to have been aware of this fact.&mdash;T.</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="223">&nbsp;</span><a name="Page_223" id="Page_223"></a>
+<a name="LETTER_XI" id="LETTER_XI"></a>LETTER XI.<br /><br />
+
+<span class="little italic">THE SAME SUBJECT CONTINUED.</span></h2>
+
+
+<p>I have collected my chief observations on swarms in the two preceding
+letters; those most frequently repeated, and of which the uniformity of
+result leads me to apprehend no error. I have deduced what seem the most
+direct consequences; and in all the theoretical part, I have sedulously
+
+<span class="pagebreak" title="224">&nbsp;</span><a name="Page_224" id="Page_224"></a>
+avoided going beyond facts. What is yet to be mentioned is more
+hypothetical, but it engrosses several curious experiments.</p>
+
+<p>It has been demonstrated, that the principal motive of the young females
+departing when hives swarm, is their insuperable antipathy to each
+other. I have repeatedly observed that they cannot gratify their
+aversion, because the workers with the utmost care prevent them from
+attacking the royal cells. This perpetual opposition at length creates a
+visible inquietude, and excites a degree of agitation that induces 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 different. Always
+accustomed to respect fertile queens, they do not forget what they owe
+to her; they allow her the most uncontrouled liberty. She is permitted
+to approach the royal cells; and if she even attempts to destroy them,
+no opposition is presented by the bees.
+<span class="pagebreak" title="225">&nbsp;</span><a name="Page_225" id="Page_225"></a>
+Thus her inclinations are not
+obstructed, and we cannot ascribe her flight, as that of the young
+queens, to the opposition she suffers. Therefore, I candidly confess
+myself ignorant of the motives of her departure.</p>
+
+<p>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 young ones, have the greatest aversion to 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 departure of
+old queens, seven royal cells opened at one side were destroyed by the
+queen. If rain continues several days, the whole are destroyed; in this
+case, there is no swarm, which too often happens in our climate, where
+spring is generally rainy. Queens never attack cells containing an egg
+or a very young worm; but only when the worm
+<span class="pagebreak" title="226">&nbsp;</span><a name="Page_226" id="Page_226"></a>
+is ready for transforming
+to a nymph, or when it has undergone its last metamorphosis.</p>
+
+<p>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 is 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
+immediate success, her inquietude during this work becomes a terrible
+agitation. If the weather continues favourable, while she remains in
+this condition, she is naturally disposed to depart.</p>
+
+<p>It may easily be understood, that the workers accustomed to respect
+their queen, whose presence is a real necessity to them, crowd after
+her; and the formation of the
+<span class="pagebreak" title="227">&nbsp;</span><a name="Page_227" id="Page_227"></a>
+first swarm creates no difficulty in this
+respect. But you will undoubtedly ask, Sir, What motive can induce the
+workers to follow their queen from the hive, while they treat the young
+queens very ill, and, even in their most amicable moments, testify
+perfect indifference towards them. Probably it is to escape the heat to
+which the hive is then exposed. The extreme agitation of the females
+leads them to traverse the combs in all directions. They pass through
+groupes of bees, injure and derange them; they communicate a kind of
+delirium, and these <a name="corr227" id="corr227"></a><a class="correction" href="#cnote227" title="changed from 'tumultous'">tumultuous</a> motions raise the
+temperature to an insupportable degree. We have frequently proved it by
+the thermometer. In a populous hive it commonly stands between 92&deg; and
+97&deg;, in a fine day of spring; but during the tumult which precedes
+swarming, it rises above 104&deg;. And this is heat intolerable to bees.
+When exposed to it, they rush impetuously towards the outlets of the
+hive and depart. In general they cannot endure the sudden
+<span class="pagebreak" title="228">&nbsp;</span><a name="Page_228" id="Page_228"></a>
+augmentation
+of heat, and in that case quit their dwelling; neither do those
+returning from the fields enter when the temperature is extraordinary.</p>
+
+<p>I am certain, from direct experiments, that the impetuous courses of the
+queen over the combs, actually throws the workers into agitation; and I
+was able to ascertain it in the following manner. I wished to avoid a
+complication of causes. It was particularly important to learn, whether
+the queen would impart her agitation but not at the time of swarming.
+Therefore I took two females still virgins, but capable of fecundation
+for above five days, and put one in a glass hive sufficiently populous;
+the other I put into a different hive of the same kind. Then I shut the
+hives in such a way that there was no possibility of their escape. The
+air had free circulation. I then prepared to observe the hives every
+moment that the fineness of the weather would invite both males and
+females to go abroad, for the purpose of
+<span class="pagebreak" title="229">&nbsp;</span><a name="Page_229" id="Page_229"></a>
+fecundation. Next morning, the
+weather being gloomy, no male left the hive, and the bees were tranquil;
+but towards eleven of the following day, the sun shining bright, both
+queens began to run about seeking an exit from every part of their
+dwelling; 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 that part of the
+hive where the openings were placed; unable to escape they ascended with
+equal rapidity, and ran heedlessly over the cells until four in the
+afternoon. It is nearly about this time that the sun declining in the
+horizon recalls the males; queens requiring fecundation never remain
+later abroad. The two females became calmer, and tranquillity was in a
+short time restored. This was repeated 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
+<span class="pagebreak" title="230">&nbsp;</span><a name="Page_230" id="Page_230"></a>
+are
+always in a tumultuous state when the queen herself is in agitation.</p>
+
+<p>I have but one fact more to mention. It has already been observed, that
+on losing the female, bees give the larv&aelig; of simple workers the royal
+treatment, and, according to M. Schirach, in five or six days they
+repair the loss of their queen. In this case there are no swarms. All
+the females leave their cells almost at the same moment, and after a
+bloody combat the throne remains with the most fortunate.</p>
+
+<p>I can very well comprehend that the object 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 metamorphosis almost at
+the same time? Since they are enabled to retain the young females in
+their cells, why do they allow all the queens, reared according to
+Schirach's method,
+<span class="pagebreak" title="231">&nbsp;</span><a name="Page_231" id="Page_231"></a>
+to escape at once. By prolonging their captivity
+more or less, they would fulfil two most important objects at once, in
+repairing the loss of their females and preserving a succession of
+queens to conduct several swarms.</p>
+
+<p>At first it was my opinion, that this difference of conduct proceeded
+from the difference of circumstances in which they found themselves
+situated. They are induced to make all their dispositions relative to
+swarming only when in great numbers, and when they have a queen occupied
+with her principal laying of male eggs; whereas, having lost their
+female, the eggs of drones are no longer in the combs to influence their
+instinct. They are in a certain degree restless and discouraged.</p>
+
+<p>Therefore, after removing the queen from a hive, I thought of rendering
+all the other circumstances as similar as possible to the situation of
+bees preparing to swarm. By introducing a great many workers, I
+encreased the population to excess, and supplied
+<span class="pagebreak" title="232">&nbsp;</span><a name="Page_232" id="Page_232"></a>
+them with combs of
+male brood in every stage. Their first occupation was to construct royal
+cells after Schirach's method, and to rear 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 this they discontinued, as there was
+no queen to deposit her eggs. Finally, I gave them several close royal
+cells, taken indifferently from hives preparing to swarm. However, all
+these precautions were fruitless; the bees were occupied only with
+replacing their lost queen; they neglected the royal cells entrusted to
+their care; the included queens came out at the ordinary time, without
+being detained prisoners a moment; they engaged in several combats, and
+there were no swarms.</p>
+
+<p>Recurring to subtleties, we may perhaps suggest a cause for this
+apparent contradiction. 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
+<span class="pagebreak" title="233">&nbsp;</span><a name="Page_233" id="Page_233"></a>
+admitting any theory
+adverse to this beautiful system, and the more must we distrust that
+facility of imagination from which we think by embellishment to
+elucidate facts.</p>
+
+<p>In general, Naturalists who have long observed animals, and those in
+particular who have chose insects for their favourite study, have too
+readily ascribed to them our sentiments, our passions, and even our
+intentions and designs. Incited 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 painted different traits of the industry of these
+minute animals, with the colours inspired by an exalted imagination. Nor
+is even the celebrated Reaumur to be acquitted of such a charge. He
+frequently ascribes combined intentions to bees; love, anticipation, and
+other faculties of too elevated a kind. I think I can perceive that
+although he formed very just ideas of their operations, he
+<span class="pagebreak" title="234">&nbsp;</span><a name="Page_234" id="Page_234"></a>
+would be
+well pleased that his reader should admit they were sensible of their
+own interests. He is a painter who by a happy interest 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 association of ideas <i>simply sensible</i>. It is
+not my object here to penetrate those depths, or to insist on the
+details.</p>
+
+<p>But, on the whole, facts relative to the formation of swarms perhaps
+present more subjects of admiration than any other part of the history
+of bees. I think it proper to state, in a few words, the simplicity of
+the methods by which the wisdom of nature guides their instinct. It
+cannot allow them the slightest portion of understanding; it leaves them
+no precautions to be taken, no combination to be followed, no foresight
+<span class="pagebreak" title="235">&nbsp;</span><a name="Page_235" id="Page_235"></a>
+to exercise, no knowledge to acquire. But having adapted their
+sensorium to the different operations with which they are charged, it is
+the impulse 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 larv&aelig;, and collect
+provisions, we must not seek for method, affection, or foresight. The
+only inducement must be sought for in some pleasing sensation attached
+to each of these operations. I address a philosopher; 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 works which treats of the industry of animals. Let me
+add but another sentence. The inducement of pleasure is not the sole
+agent; there is another principle, the prodigious influence of which, at
+least with regard
+<span class="pagebreak" title="236">&nbsp;</span><a name="Page_236" id="Page_236"></a>
+to bees, has hitherto been unknown, that is the
+sentiment of aversion which all females continually feel against each
+other, a sentiment whose existence is so fully demonstrated by my
+experiments, and which explains many important facts in the theory of
+swarms.</p>
+
+<p class="toright italic">
+<span class="smcap">Pregny</span>, 10. September 1791.
+</p>
+
+
+
+<h2>
+<span class="pagebreak" title="237">&nbsp;</span><a name="Page_237" id="Page_237"></a>
+<a name="LETTER_XII" id="LETTER_XII"></a>LETTER XII.<br /><br />
+
+<span class="little italic">ADDITIONAL OBSERVATIONS ON QUEENS THAT LAY ONLY THE EGGS OF DRONES, AND
+ON THOSE DEPRIVED OF THE ANTENN&AElig;.</span></h2>
+
+
+<p>In relating my first observations on queens that lay male eggs alone, I
+have proved that they lay them in cells of all dimensions indifferently,
+and even in royal cells. It is also proved that the same treatment
+<span class="pagebreak" title="238">&nbsp;</span><a name="Page_238" id="Page_238"></a>
+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 workers appeared defective. It is
+indeed most singular, that bees which know the worms of males so well
+when the eggs are laid in small cells, and never fail to give them a
+convex 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
+irregularity depends on something I cannot comprehend.</p>
+
+<p>In revising what is said on this subject, I observe still wanting an
+interesting experiment to complete the history of queens that lay only
+the eggs of drones. I had to investigate whether these females could
+themselves distinguish that the eggs they deposit in the royal cells
+would not produce queens. I have already observed that they do not
+endeavour to destroy these
+<span class="pagebreak" title="239">&nbsp;</span><a name="Page_239" id="Page_239"></a>
+cells when close, and I thence concluded,
+that in general the presence of royal cells in their hive does not
+inspire them with the same aversion to females whose fecundation has
+been retarded; but to ascertain the fact more correctly, it was
+essential to examine how the presence of a cell containing a royal nymph
+would affect a queen that had never laid any other than the eggs of
+drones.</p>
+
+<p>This experiment was easy; and I put it in practice on the fourth of
+September, in a hive some time deprived of its queen. The bees had not
+failed to construct several royal cells for replacing their females. I
+chose this opportunity for supplying them with a queen, 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. One remaining was
+enough to shew the impression it would make on the stranger queen
+introduced; had she endeavoured
+<span class="pagebreak" title="240">&nbsp;</span><a name="Page_240" id="Page_240"></a>
+to destroy it; this, in my opinion,
+would have proved that she anticipated the origin of a dangerous rival.
+You must admit the use I make of the word anticipate; it saves a long
+paraphrase; I feel the impropriety of it. If, on the contrary, she did
+not attack the cell I would thence conclude that the delay of
+fecundation, which deprived her of the power of laying workers eggs, had
+also impaired her instinct. This was the fact; the queen passed several
+times over the royal cell, both 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 bestowed 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 honey and respect, and formed those regular
+circles around her that seem an expression of homage.</p>
+
+<p>
+<span class="pagebreak" title="241">&nbsp;</span><a name="Page_241" id="Page_241"></a>
+Thus, independent of the derangement occasioned by retarded
+impregnation, in the sexual organs of queens, it certainly impairs their
+instinct. Aversion or jealousy is no longer preserved against their own
+sex in the nymphine state, nor do they longer endeavour to destroy them
+in their cradles.</p>
+
+<p>My readers will be surprised that queens whose fecundation has been
+retarded, and whose fecundity is so useless to bees, should be so well
+treated and become as dear to them as females laying both kinds of eggs.
+But I remember to have observed a fact more astonishing still. I have
+seen workers bestow every attention on a queen though sterile; and after
+her death treat her dead body as they had treated herself when alive,
+and long prefer this inanimate body to the most fertile queens I had
+offered them. This sentiment, which assumes the appearance of so lively
+an affection, is probably the effect of some agreeable sensation
+communicated to bees by their
+<span class="pagebreak" title="242">&nbsp;</span><a name="Page_242" id="Page_242"></a>
+queen, independent of fertility. Those
+laying only the eggs of males probably excite the same sensation in the
+workers.</p>
+
+<p>I now recollect that the celebrated Swammerdam somewhere observes, that
+when a queen is blind, sterile, or mutilated, she ceases to lay, and the
+workers of her hive no longer labour or make any collections, as if
+aware that it was now useless to work. He cites no experiment that led
+him to the discovery. Those made by myself have afforded some very
+singular results.</p>
+
+<p>I frequently amputated the four wings of queens; and not only did they
+continue laying, but the same confederation of them was testified by the
+workers as before. Therefore, Swammerdam has no foundation for
+asserting, that mutilated queens cease to lay. Indeed, from his
+ignorance of fecundation taking place without the hives, it is possible
+he cut the wings off virgin queens, and they, becoming incapable of
+flight, remained sterile from inability to seek the males in the air.
+Thus, amputation
+<span class="pagebreak" title="243">&nbsp;</span><a name="Page_243" id="Page_243"></a>
+of the wings does not produce sterility in queens.</p>
+
+<hr />
+
+<p>I have frequently cut off one <a name="corr242" id="corr242"></a><a class="correction" href="#cnote242" title="unchanged">antenn&aelig;</a> 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 one still remained, the mutilation was imperfect; and the
+experiment decided nothing. But amputation of both antenn&aelig; produced most
+singular effects. On the fifth of September, I cut both 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 vivacity. Scarcely
+had the workers time to separate and recede before her; she dropped her
+eggs, without attending to deposit them in any cell. The hive not being
+very populous, part was without comb. Hither
+<span class="pagebreak" title="244">&nbsp;</span><a name="Page_244" id="Page_244"></a>
+she seemed particularly
+earnest to repair, and long remained motionless. She appeared to avoid
+the bees; however, several workers followed her into this solitude, and
+treated her with the most evident respect. She seldom required honey
+from them, but, when that occurred, 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 appeared tormented with the desire of leaving her
+habitation. She rushed towards the opening, and entered the glass tube
+adapted there; but the external orific being too small, after fruitless
+exertion, she returned. Notwithstanding these symptoms of delirium, the
+bees did not cease to render her the same attention as they ever pay to
+their queens, but this one received
+<span class="pagebreak" title="245">&nbsp;</span><a name="Page_245" id="Page_245"></a>
+it with indifference. All that I
+describe appeared to me the consequence of amputating the antenn&aelig;.
+However, her organization having already suffered from retarded
+fecundation, and as I had observed her instinct in some degree impaired,
+both causes might possibly concur in producing the same effect. To
+distinguish properly what belonged to the privation of the antenn&aelig;, a
+repetition of the experiment was necessary, in a queen otherwise well
+organised, and capable of laying both kinds of eggs.</p>
+
+<p>This I did on the sixth of September. I amputated both the antenn&aelig; of a
+female which had been several months the subject of observation, and
+being of great fecundity 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 same marks of delirium and agitation, which I think it
+needless to repeat. I shall only add, that
+<span class="pagebreak" title="246">&nbsp;</span><a name="Page_246" id="Page_246"></a>
+to judge better of the
+effect produced by privation of the antenn&aelig;, on the industry and
+instinct of bees, I attentively considered the manner in which these two
+mutilated queens treated each other. You cannot have forgot, Sir, the
+animosity with which queens, possessing all their organs, combat, on
+which account it became extremely interesting to learn whether they
+would experience the same reciprocal aversion after losing their
+antenn&aelig;. We studied these queens a long time; they met several times in
+their courses, and without exhibiting the smallest resentment. This last
+instance is, in my opinion, the most complete evidence of a change
+operated in their instinct.</p>
+
+<p>Another very remarkable circumstance, which this experiment gave me
+occasion to observe, consists in the good reception given by the bees to
+the stranger queen, while they still preserved the first. Having so
+often seen the symptoms of discontent that a plurality of queens
+occasions,
+<span class="pagebreak" title="247">&nbsp;</span><a name="Page_247" id="Page_247"></a>
+after having witnessed the clusters formed around these
+supernumerary queens to confine them, I could not expect they would pay
+the same homage to a second mutilated one they still testified towards
+the first. Is it because after losing the antenn&aelig;, these queens have no
+more any characteristic which distinguishes the one from the other?</p>
+
+<p>I was the more inclined to admit this conjecture from the bad reception
+of a third fertile queen preserving her antenn&aelig;, which was introduced
+into the same hive. The bees seized, bit her, and confined her so
+closely, that she could hardly breath or move. Therefore, if they treat
+two females deprived of antenn&aelig; in the same hive equally well, it is
+probably because they experience the same sensation from these two
+females, and want the means of longer distinguishing them from each
+other.</p>
+
+<p>From all this, I conclude, that the antenn&aelig; are not a frivolous ornament
+to insects, but, according to all appearance, are the
+<span class="pagebreak" title="248">&nbsp;</span><a name="Page_248" id="Page_248"></a>
+organs of touch
+or smell. Yet I cannot affirm which of these senses reside in <a name="corr247" id="corr247"></a><a class="correction" href="#cnote247" title="changed from 'them,'">them.</a> It
+is not impossible that they are organised in such a manner as to fulfil
+both functions at once.</p>
+
+<p>As in the course of this experiment both mutilated females constantly
+endeavoured 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 I removed the first and third queen from the hive,
+leaving the fertile mutilated one, and enlarged the entrance.</p>
+
+<p>The queen left her habitation the same day. At first she tried to fly,
+but, her belly being full of eggs, she fell down and never attempted it
+again. No workers accompanied her. Why, after rendering the queen so
+much 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, carrying
+<span class="pagebreak" title="249">&nbsp;</span><a name="Page_249" id="Page_249"></a>
+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 I can hazard on this question is a conjecture.
+It appears that bees are induced to quit the hives from the increased
+heat which occasions the agitation of their queen, and the tumultuous
+motion which she communicates to them. A mutilated queen,
+notwithstanding her delirium, does not agitate the workers, because she
+seeks the uninhabited parts of the hive, and the glass panes of it: she
+hurries over clusters of bees, but the shock resembles that of any other
+body, and produces only a local and instantaneous motion. The agitation
+arising from it, is not communicated from one place to another, like
+that produced by a queen, which in the natural state wishes to abandon
+her hive and lead out a swarm; there is no increased heat, consequently
+<span class="pagebreak" title="250">&nbsp;</span><a name="Page_250" id="Page_250"></a>
+nothing that renders the hive insupportable to her.</p>
+
+<p>This conjecture, which affords a tolerable explanation why bees persist
+in remaining in the hive, though the mutilated queen has left it, is no
+reason for the motive inducing the queen herself to depart. Her instinct
+is altered; that is the whole that I can perceive. I can discern nothing
+more. It is very fortunate for the hive, that this queen departs, for
+the bees incessantly attend her; nor do they ever think of procuring
+another while she remains; and if she was long of leaving them, it would
+be impossible to replace her; for the workers worms would exceed the
+term at which they are convertible into royal worms, and the hive would
+perish. Observe, that the eggs dropped by the mutilated queen can never
+serve for replacing her, for, not being deposited in cells, they dry and
+produce nothing.</p>
+
+<p>
+<span class="pagebreak" title="251">&nbsp;</span><a name="Page_251" id="Page_251"></a>
+I have yet to say a few words on females laying male eggs only. M.
+Schirach supposes that one branch of their double ovary suffers some
+alteration. He seems to think that one of these branches contains the
+eggs of males, while the other has none but common eggs, and as he
+ascribes the inability of certain queens to lay the latter to some
+disease, his conjecture becomes very plausible. In fact, if the eggs of
+males and workers are indiscriminately mixed in both branches of the
+ovary, it appears at first sight that whatever cause acts on that organ,
+it should equally affect both species of eggs. If on the contrary, one
+branch is occupied by the eggs of drones only, and the other contains
+none but common eggs, we may conceive how disease affects the one, while
+the other remains untouched. Though this conjecture is probable, it is
+confuted by observation. We lately dissected queens, which laid none but
+male eggs, and found both branches of the ovary equally well expanded,
+
+<span class="pagebreak" title="252">&nbsp;</span><a name="Page_252" id="Page_252"></a>
+and equally sound, if I may use the expression. The only difference
+that struck us was that in these two branches, the eggs were apparently
+not so close together as in the ovaries of queens laying both kinds of
+eggs.</p>
+
+<p class="toright italic">
+<span class="smcap">Pregny</span>, 12. September 1791.
+</p>
+
+
+
+<h2>
+<span class="pagebreak" title="253">&nbsp;</span><a name="Page_253" id="Page_253"></a>
+<a name="LETTER_XIII" id="LETTER_XIII"></a>LETTER XIII.<br /><br />
+
+<span class="little italic">ECONOMICAL CONSIDERATIONS ON BEES.</span></h2>
+
+
+<p>In this letter I shall treat of the advantages that may be derived from
+the new invented hives, called <i>book</i> or <i>leaf</i> hives, in promoting the
+<i>economical knowledge</i> of bees.</p>
+
+<p>It is needless to relate the different methods hitherto employed in
+forcing bees to yield up a portion of their honey and wax; all resemble
+each other in being cruel and ill understood.</p>
+
+<p>
+<span class="pagebreak" title="254">&nbsp;</span><a name="Page_254" id="Page_254"></a>
+It is evident, when bees are cultivated for the purpose of sharing the
+produce of their labours, we must endeavour to multiply them as much as
+the nature of the country admits; and consequently to regard their lives
+at the time we plunder them. Therefore it is an absurd custom to
+sacrifice whole hives to get at the riches they contain. The inhabitants
+of this country, who follow no other method, annually lose immense
+numbers of hives; and spring, being generally unfavourable to swarms,
+the loss is irreparable. I well know that at first they will not adopt
+any other method; 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.</p>
+
+<p>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
+<span class="pagebreak" title="255">&nbsp;</span><a name="Page_255" id="Page_255"></a>
+essential
+to success, which I should not omit. Though the bees are indifferent as
+to the position of their combs, and as to their greater or lesser size,
+they are obliged to construct them perpendicular to the horizon, and
+parallel to each other. Therefore, if left entirely to themselves, when
+establishing a colony in one of those new hives, they would frequently
+construct several small combs parallel indeed, but perpendicular to the
+plane of the frames or leaves, and by this disposition prevent the
+advantages which I think to derive from the figure of my hives, since
+they could not be opened without breaking the combs. Thus they must
+previously have a guide to follow; the cultivator himself lays the
+foundation of their edifices, and that by a simple method. A portion of
+comb must be solidly fixed in some of the boxes composing the hive; the
+bees will extend it; and, in prosecution of their work, will accurately
+follow the plan already given them. Therefore on opening the hive, no
+obstacle
+<span class="pagebreak" title="256">&nbsp;</span><a name="Page_256" id="Page_256"></a>
+is to be removed, nor stings to be dreaded, for one of the
+most singular and valuable properties attending this construction, 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, and the tranquillity of the bees has given you great
+surprise. I can desire no other evidence of my assertion. It is in the
+facility of opening these hives at pleasure that all the advantages lie,
+which I expect in perfecting the economical knowledge of bees.</p>
+
+<p>I conceive, when I observe bees may be rendered tractable, that it need
+not be added, I do not arrogate to myself the absurd pretence of
+<i>taming</i> them, for this excites a vague idea of tricks; and I would
+willingly avoid the hazard of exposing myself to any such reproach. I
+ascribe their tranquillity on opening the hives, to the manner that the
+sudden introduction of light affects them; then, they seem rather to
+testify fear than anger. Many retire and enter
+<span class="pagebreak" title="257">&nbsp;</span><a name="Page_257" id="Page_257"></a>
+the cells, and appear to
+conceal themselves. What confirms my conjecture is, 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. The divisions must be separated slowly, and
+care taken not to wound any of the bees. If they cluster too much on the
+combs, they must be brushed off with a feather; and breathing on them
+carefully avoided. The air we expire seems to excite their fury; it
+certainly has some irritating quality, for if bellows are used, they are
+rather disposed to escape than to sting.</p>
+
+<p>Respecting the advantages of leaf hives, I shall observe, they are very
+convenient for forming <i>artificial</i> swarms. In the history of natural
+swarms, I have shewn how many favourable circumstances are necessary for
+their success. From experience I know that they very often fail in our
+climate; and even when a hive is disposed to swarm, it
+<span class="pagebreak" title="258">&nbsp;</span><a name="Page_258" id="Page_258"></a>
+frequently
+happens that the swarm is lost either because the moment of its
+departure has not been foreseen, because it rises out of sight, or
+settles on inaccessible places. Instructing the cultivators of bees how
+to make artificial swarms is a real service, and the form of my hives
+renders this an easy operation. But it requires farther illustration.</p>
+
+<p>Since bees, according to M. Schirach's discovery, can procure another
+queen after having lost their own, provided there is workers brood in
+the combs not above three days old, it results that we can at pleasure
+produce queens, by removing the reigning one. Therefore, if a hive
+sufficiently populous is divided in two, one half will retain the old
+queen, and the other will not be long of obtaining 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
+sufficient to admit of division, if the brood is of the proper
+<span class="pagebreak" title="259">&nbsp;</span><a name="Page_259" id="Page_259"></a>
+age, if
+males exist or are ready to be produced for impregnating the young
+queens.</p>
+
+<p>Supposing the union of all these conditions, the following is the method
+to be pursued. The leaf hive may be divided through the middle without
+any shock. Two empty divisions may be insinuated between the halves,
+which, when exactly applied to each other, are close on the outside. The
+queen must be sought in one of the halves, and marked to avoid mistake.
+If she by chance remains in the division with most brood, she is to be
+transferred to the other with less, that the bees may have every
+possible opportunity 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 occupied. The old entrance, now become useless, will be shut
+up; but as each half requires a new one, it ought to be made at the
+bottom of each
+<span class="pagebreak" title="260">&nbsp;</span><a name="Page_260" id="Page_260"></a>
+division, on purpose that they may be as far asunder as
+possible. Both entrances should not be made on the same day. The bees in
+the half deprived of the queen ought to be confined twenty-four hours,
+and no opening made before then except for admission of air. Without
+this precaution, they would soon search for their queen, and infallibly
+find her in the other division. They will then retire in great numbers
+from their own division, until too few remain to perform the necessary
+labours. But this will not ensue if they are confined twenty-four hours,
+provided that interval is sufficient to make them forget the queen. When
+all these circumstances are favourable, the bees, in the division
+wanting the queen, will the same day begin to labour in procuring
+another, and ten or fifteen days after the operation, their loss will be
+repaired. The young female they have reared, soon issues forth to seek
+impregnation, and in two days commences the laying of workers eggs.
+Nothing more
+<span class="pagebreak" title="261">&nbsp;</span><a name="Page_261" id="Page_261"></a>
+is wanting to the bees of this half hive, and the success
+of the artificial swarm is ensured.</p>
+
+<p>It is to M. Schirach that we are indebted for this ingenious method of
+forming swarms. He supposes, by producing young queens early in spring,
+that early swarms might be procured, which would certainly be
+advantageous in favourable circumstances. But unfortunately this is
+impossible. Schirach believed that queens were impregnated 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 become 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. Thus, if a
+swarm were artificially formed before the usual time of the males
+originating, the bees would be discouraged by the sterility of the young
+female. Or should they remain faithful to
+<span class="pagebreak" title="262">&nbsp;</span><a name="Page_262" id="Page_262"></a>
+her, awaiting the period of
+fecundation, as she could not for three or four weeks receive the
+approaches of the male, she would lay eggs producing males only, and the
+hive in this case would perish. Thus the natural order must not be
+deranged, but we must delay the division of hives until males are about
+to originate or actually exist.</p>
+
+<p>Besides, if M. Schirach did succeed in obtaining artificial swarms,
+notwithstanding the great inconvenience of his hives, it was owing to
+his singular address and unremitting assiduity. He had some pupils in
+the art; these communicated the method of forming artificial swarms to
+others, and there are people now in Saxony who traverse the country
+practising this operation. Those versant in the matter can alone venture
+to undertake it with common hives, whereas, every cultivator can do it
+himself with the leaf hives.</p>
+
+<p>In this construction, another very great advantage will also be found.
+Bees can
+<span class="pagebreak" title="263">&nbsp;</span><a name="Page_263" id="Page_263"></a>
+be forced to work in wax. Here I am led to what I believe is a
+new observation. While naturalists have directed our admiration to the
+parallel position of the combs, they have overlooked another trait in
+the industry of bees, namely, the equal distance uniformly between them.
+On measuring the interval separating the combs, it will generally be
+found four lines. Were they too distant, it is very evident the bees
+would be much dispersed and unable to communicate their heat
+reciprocally; whence the brood would not be exposed to sufficient
+warmth. Were the combs too close, on the contrary, the bees could not
+freely traverse the intervals, and the work of the hive would suffer.
+Therefore, a certain distance always uniform is requisite, which
+corresponds equally well with the service of the hive, and the care
+necessary for the worms. Nature, which has taught bees so much, has
+instructed them regularly to preserve this distance. At the approach of
+winter, they sometimes elongate the
+<span class="pagebreak" title="264">&nbsp;</span><a name="Page_264" id="Page_264"></a>
+cells which are to contain the
+honey, and thus contract the intervals between the combs. But this
+operation is a preparation for a season, when it is important to have
+plentiful magazines, and when their activity being very much relaxed, it
+is unnecessary for their communications to be so spacious and free. On
+the return of spring, the bees hasten to contract these elongated cells,
+that they may become fit for receiving the eggs which the queen will
+lay, and thus re-establish the just distance which nature has ordained.</p>
+
+<p>This being admitted, bees may be forced 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
+build others in the interval. Suppose an artificial swarm is lodged in a
+leaf hive, composed of six divisions, each containing a comb, if the
+young queen is as fertile as she ought to be, the bees will be
+<span class="pagebreak" title="265">&nbsp;</span><a name="Page_265" id="Page_265"></a>
+very
+active in their labours, and disposed to make great collections of wax.
+To induce them towards this an empty box or division must be placed
+between two others, each containing a comb. As all the boxes are of
+equal dimensions, and of the necessary width for receiving a comb, the
+bees having sufficient 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 having more than four lines
+between them. Without any guide, this new comb will be parallel to the
+old ones, to preserve that law which establishes an equal distance
+throughout the whole.</p>
+
+<p>If the hive is strong and the weather good, three empty divisions may at
+first be left between the old combs; one between the first and second,
+another between the third and fourth, and the last between the fifth and
+sixth. The bees will fill them in seven or eight days, and the hive then
+contains nine combs. Should the temperature
+<span class="pagebreak" title="266">&nbsp;</span><a name="Page_266" id="Page_266"></a>
+of the weather continue
+favourable, three new leaves or divisions may be introduced;
+consequently 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 where flowers perpetually blow. But in our
+country, I have reason to think that the labour should not be forced
+more during the first year.</p>
+
+<p>From these details, you are sensible, Sir, how preferable <i>leaf hives</i>
+are to those of any other construction, and even to those ingenious
+stages described by <i>M. Palteau</i>, for the bees cannot by means of them
+be forced to labour more in wax than they would do if left to
+themselves; whereas, they are obliged to do it by inserting empty
+divisions. Next, the combs constructed on those stages cannot be removed
+without destroying considerable portions of brood, deranging the bees,
+and creating real disorder in the hive.</p>
+
+<p>
+<span class="pagebreak" title="267">&nbsp;</span><a name="Page_267" id="Page_267"></a>
+Mine have also this advantage, that what passes within may daily be
+observed, and we may judge of the most favourable moments for depriving
+the bees of part of their stores. With all the combs before us we can
+distinguish those containing brood only, and what it is proper to
+preserve. The scarcity or abundance of provisions is visible, and the
+portion suitable may be taken away.</p>
+
+<p>I should protract this letter too much, if I gave an account of all my
+observations 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 their riches with them. The subject would require a separate
+work; and I may perhaps one day engage in it; but until that arrives I
+shall always feel gratification in communicating to cultivators, who
+wish to follow my method, directions of which long practice has
+demonstrated the utility.</p>
+
+<p>
+<span class="pagebreak" title="268">&nbsp;</span><a name="Page_268" id="Page_268"></a>
+Here I shall only observe, that we hazard absolute ruin of the hives,
+by robbing them of too great a proportion of honey and wax. In my
+opinion, the art of cultivating these animals consists in moderately
+exercising the privilege of sharing their labours; but as a compensation
+for this, every method must be employed which promotes the
+multiplication of bees. Thus, for example, if we desire to procure a
+certain quantity of honey and wax annually, it will be better to seek it
+in a number of hives, managed with discretion, than to plunder a few of
+a great proportion of their treasures.</p>
+
+<p>It is indubitable that the multiplication of these industrious animals
+is much injured by privation of several combs, in a season unfavourable
+to the collection of wax, because the time consumed in replacing them is
+taken from that which should be consecrated to the care of the eggs and
+worms, and by this means the brood suffers. Besides, they must always
+have a
+<span class="pagebreak" title="269">&nbsp;</span><a name="Page_269" id="Page_269"></a>
+sufficient provision of honey left for winter, for although less
+is consumed during this season, they do consume some; because they are
+not torpid, as some authors have conceived.<a name="fnm_N" id="fnm_N"></a><a href="#fn_N" class="fnnum">N</a> Therefore if they have
+not enough, they must be supplied with it,
+<span class="pagebreak" title="270">&nbsp;</span><a name="Page_270" id="Page_270"></a>
+which requires great
+exactness. I admit that in determining to what extent hives may be
+multiplied in a particular country, it is necessary first to know how
+many the country can support, which is a problem yet unsolved. It also
+depends on another, the solution of which is as little known, namely the
+greatest distance that bees fly in collecting their provender. Different
+authors maintain, they can fly several leagues from the hive. But by the
+few observations I have been able to make, this distance seems greatly
+exaggerated. It appears to me that the radius of the circle
+<span class="pagebreak" title="271">&nbsp;</span><a name="Page_271" id="Page_271"></a>
+they
+traverse does not exceed half a league. As they return to the hive with
+the greatest precipitation whenever a cloud passes before the sun, it is
+probable they do not fly far. Nature which has inspired them with such
+terror for a storm, and even for rain, undoubtedly 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 distances bees with the thorax painted, that they might again
+be recognised. But none ever returned that I had carried for twenty-five
+or thirty minutes from their dwelling, while those at a shorter distance
+have found their way and returned. I do not state this experiment as
+decisive. Though bees do not generally fly above half a league, it is
+very possible they go much farther, when flowers are scarce in their own
+vicinity. A conclusive experiment must be made in vast arid or sandy
+plains, separated
+<span class="pagebreak" title="272">&nbsp;</span><a name="Page_272" id="Page_272"></a>
+by a known distance from a fertile region.</p>
+
+<p>Thus, the question yet remains undecided. But without ascertaining the
+number of hives that any district can maintain, I shall remark that
+certain vegetable productions are much more favourable to bees than
+others. More hives, for example, may be kept in a country abounding
+meadows, and where black grain is cultivated, than in a district of
+vineyards or corn.</p>
+
+<hr />
+
+<p>Here 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. Several problems concerning the history of these
+animals still remain unsolved. The experiments I project may perhaps
+throw some light on them; and I shall be animated with much greater
+hopes of success, if you, Sir, will continue your counsels
+<span class="pagebreak" title="273">&nbsp;</span><a name="Page_273" id="Page_273"></a>
+and
+direction. I am, with every sentiment of gratitude and respect,</p>
+
+<p class="toright">
+<span class="smcap">Francis Huber.</span><br /><br />
+<i><span class="smcap">Pregny</span>, 1. October 1791.</i>
+</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_N" id="fn_N"></a><span class="label"><a href="#fnm_N">N</a></span> So far from being torpid in winter, when the thermometer in
+the open air is several degrees below freezing, it stands at (86) and
+(88&deg;), in hives sufficiently populous. The bees then cluster together,
+and move to preserve their heat.</p>
+
+<p>Now that I am on the subject of thermometrical observations, I may
+cursorily remark, that M. Dubois of Bourg en Bresse, in a memoir
+otherwise valuable, is of opinion, that the larv&aelig; cannot be hatched
+below (104). I have repeatedly made the experiment with the most
+accurate thermometers, and obtained a very different result. When the
+thermometer rises to (104&deg;), the heat is so much greater than the eggs
+require, that it is intolerable to the bees. M. Dubois has been
+deceived, I imagine, by too suddenly introducing his thermometer into a
+cluster of bees, and putting them in agitation, the mercury has rose
+higher than it should naturally do. Had he delayed introducing the
+thermometer, he would soon have seen it fall to between 95 and 97, which
+is the usual temperature of hives in summer. In August this year, when
+the thermometer in the open air stood at 94, it did not rise above 99 in
+the most populous hives. The bees had little motion, and a great many
+rested on the board of the hive.
+<span class="pagebreak" title="274">&nbsp;</span><a name="Page_274" id="Page_274"></a>
+</p></div>
+</div>
+
+
+<h2>
+<span class="pagebreak" title="275">&nbsp;</span><a name="Page_275" id="Page_275"></a>
+<a name="APPENDIX" id="APPENDIX"></a>APPENDIX.</h2>
+
+
+<p>[The following passages are chiefly engrossed in the substance of the
+work, but the Translator, as has already been observed, for various
+reasons, judges it expedient to transfer them to an appendix. In his
+opinion these very minute details rather interrupt the connexion of the
+narrative, however interesting they may be considered, and they pertain
+more to researches purely anatomical.</p>
+
+<p>The Translator has likewise in some instances incorporated several long
+and important
+<span class="pagebreak" title="276">&nbsp;</span><a name="Page_276" id="Page_276"></a>
+notes with the text; because it appears to him that they
+actually belong to the substance of the treatise. These are the only
+variations from the original with respect to arrangement.]</p>
+
+<hr />
+
+<p>Swammerdam has given an imperfect description of the ovary of the queen.
+He observes that he has never been able to find the termination of the
+oviducts in the abdomen, nor any other parts excepting those which he
+has described. "Notwithstanding all my exertions, I never could discover
+the site of the vulva, partly because I had not all my apparatus with me
+in the country, when investigating this subject, and partly from my
+apprehension of injuring other parts by pressure, which I had then
+occasion to examine. However, I have clearly observed a muscular
+swelling of the oviduct, where approaching the last ring of the belly;
+that it then contracts and afterwards dilates in becoming membranaceous.
+As I was desirous of preserving the poison bag, which is situated
+exactly here, along with, the muscles aiding the motion of the sting, I
+could follow the oviduct no farther. However, in another female, it
+appeared that the vulva is in the last ring of the abdomen,
+<span class="pagebreak" title="277">&nbsp;</span><a name="Page_277" id="Page_277"></a>
+and under
+the sting. The parts expanding only while the queen lays, renders it
+extremely difficult to penetrate the aperture."</p>
+
+<p>We have attempted to discover what has escaped the indefatigable
+Swammerdam. But his observation that the research can be made to the
+greatest advantage, at the time of laying, has paved the way to us. We
+have remarked that the oviduct did not issue from the body, but that the
+eggs fall into a kind of cavity, where they are retained several seconds
+before being laid.</p>
+
+<p>On the sixth of August, we took a very fertile queen, and holding her
+gently by the wings in a supine position, the whole belly was exposed.
+She seized the extremity with her second pair of legs, and curved it as
+much as possible. This seeming an unfavourable position for laying, we
+forced her to stretch it out. The queen, oppressed with the necessity of
+laying, could no longer retain her eggs. The lower part of the last ring
+then separated so far from the upper part as to leave some of the inside
+discovered. In this cavity the sting lay above in its sheath. As the
+queen now made new efforts, we saw an egg fall into the cavity from the
+end of the oviduct. The lips then closed for several seconds;
+<span class="pagebreak" title="278">&nbsp;</span><a name="Page_278" id="Page_278"></a>
+they
+opened again, and, in a much shorter time, dropped the egg from the
+cavity.</p>
+
+<p>From our own observations we found that the seminal fluid of drones
+coagulated on exposure to the air, and from several experiments had so
+little doubt on the subject, that whenever the female returned with the
+external marks of fecundation, we thought we recognised it in the
+whitish substance filling the sexual organs. It did not then occur to us
+to dissect the females to ascertain the fact more particularly: but this
+year, whether designing to neglect nothing, or to examine the distension
+of the female organs, we determined to dissect several. To our infinite
+surprise, what we had supposed the residue of the prolific fluid,
+actually proved the genital organs of the male, which separate from his
+body during copulation, and remain in the female.</p>
+
+<p>We procured a number of queens according to Schirach's method for the
+purpose of dissection, and set them at liberty that they might seek the
+males. The first which did so, was seized the instant she returned, and
+without dissection spontaneously exhibited what we were so impatient to
+behold. Examining the under part of the belly, we saw the oval end of a
+white substance
+<span class="pagebreak" title="279">&nbsp;</span><a name="Page_279" id="Page_279"></a>
+which distended the sexual organs. The belly was in
+constant motion, by alternate extension and contraction. Already had we
+prepared to sever the rings, and by dissection to ascertain the cause of
+these motions; when the queen curving her belly very much, and
+endeavouring to reach its extremity with her hind legs, seized the
+distending substance with her claws, and evidently made an effort to
+extract it. She at last succeeded, and it fell before us. We expected a
+shapeless mass of coagulated fluid; what therefore was our surprise to
+find it part of the same male that had rendered this queen a mother. At
+first we could not credit our eyes; but after examining it in every
+position, both with the naked eye, and a powerful magnifier, we
+distinctly recognised it to be that part which M. de Reaumur calls the
+<i>lenticular</i> body, or the <i>lentil</i>, in the following description.<a name="fnm_O" id="fnm_O"></a><a href="#fn_O" class="fnnum">O</a></p>
+
+<div class="blockquot">
+<p>'Opening a drone there appears a portion formed by the assemblage of
+several parts, often whiter than milk. This on investigation is found to
+be principally composed of four oblong pieces. The
+<span class="pagebreak" title="280">&nbsp;</span><a name="Page_280" id="Page_280"></a>
+two largest are
+attached to a kind of twisted cord, fig. 4. r, called by Swammerdam the
+root of the penis; and he has denominated seminal vessels, s. s. two
+long bodies that we are about to consider. Other two bodies oblong like
+the preceding, but shorter and not half the diameter, he calls the <i>vasa
+deferentia</i>, d. d. Each communicates with one of the seminal vessels
+near, g. g. where they unite to the twisted cord, r. From the other
+extremity proceeds a very delicate vessel, which, after several
+involutions, terminates in a body, t. a little larger, but difficult to
+disengage from the surrounding trache&aelig;. Swammerdam considers these two
+bodies, t. t. the testicles. Thus there are two parts of considerable
+size, communicating with other two still thicker and longer. These four
+bodies are of a cellular texture, and full of a milky fluid, which may
+be squeezed out. This long twisted cord, r, to which the largest of the
+seminal vessels is connected, this cord, I say, is doubtless the channel
+by which the milky fluid issues. After several plications, it terminates
+in a kind of bladder or fleshy sac, i. i. In different males this part
+is of various length and flatness. By calling it the <i>lenticular</i> body,
+or the lentil, it receives a
+<span class="pagebreak" title="281">&nbsp;</span><a name="Page_281" id="Page_281"></a>
+name descriptive of the figure it presents
+in all males whose internal parts have acquired consistency in spirit of
+wine. The body, l. i. is therefore a lentil, a little thickened, of
+which one half, or nearly so, of the <a name="corr281" id="corr281"></a><a class="correction" href="#cnote281" title="changed from 'cirumference'">circumference</a> is
+edged along the outline by two chesnut coloured scaly plates, e. i. A
+small white cord, the real edge of the lentil, is visible, and separates
+them. This lentil is a little oblong, and, for convenience, we shall
+ascribe two extremities to it, the anterior and posterior. The anterior,
+l, next the head, is where the canal, r, dividing the seminal vessels is
+inserted, and the opposite part; i. next the anus, the posterior. The
+two scaly plates, e. i. e. i, proceed from the vicinity of this last
+part, whence each enlarges to cover part of the lentil. Under the
+broadest part of each plate, there is a division formed by two soft
+points of unequal length; the largest of which is on the circumference
+of the lentil. Besides these two scaly plates, there are two others, n.
+n. of the same colour, narrower, and fully one half shorter, each of
+which is situated very near the preceding, and originates close to the
+origin of that it accompanies, namely, at the posterior part of the
+lentil. The rest of the lentil is
+<span class="pagebreak" title="282">&nbsp;</span><a name="Page_282" id="Page_282"></a>
+white and membranaceous. From behind
+proceeds a tube, k. a canal also white and membranaceous, but it is
+difficult to judge of its diameter, for the membranes, of which it
+consists, are evidently in folds. To one side of this pipe is attached a
+fleshy part, p. somewhat pallet shaped, one side is concave, and the
+edges plaited; the other side is convex. In certain places the plaits
+rise and project from the rest of the outline, and form a kind of rays;
+the pallet appears prettily figured. Though lying with the concave side
+applied to the lentil, it is not fixed to it. Swammerdam seems to
+consider this pallet as the characteristic part of the male.</p>
+<div class="figcenter" style="width: 413px;">
+<a id="fig4" name="fig4"></a>
+<img src="images/fig4.png" width="413" height="700" alt="Parts of a male bee" />
+<span class="caption">Figure 4</span>
+</div>
+
+<p>'Though the parts we have described are the most conspicuous in the
+male, they are neither those which protrude first, nor when protruded
+are the most remarkable. On viewing from the opposite edge of the
+lentil, forming the division of the two great scaly plates, a sac or
+canal, k. proceeding from the posterior part of the lentil, there is
+distinctly visible the body u, which we call the arc; where there are
+five transverse hairy bands of a yellow colour, while the rest is white.
+This arc seems out of the membranaceous canal because it is covered only
+by a very transparent
+<span class="pagebreak" title="283">&nbsp;</span><a name="Page_283" id="Page_283"></a>
+membrane. One end almost reaches the lenticular
+body, and the other terminates where the membranaceous canal joins the
+folded yellow membranes, m. which form a species of sac, that is applied
+to the sides of the aperture, adapted for the genital organs passing
+through. These reddish membranes are those that appear first on
+pressure, and form this elongated portion, at whose end is a kind of
+hairy mask. Finally, with the sac formed by the reddish membranes, there
+are connected two appendages, c. c. of reddish yellow, and red at the
+end, s. These are what appear externally like horns.<a name="fnm_P" id="fnm_P"></a><a href="#fn_P" class="fnnum">P</a>'</p>
+</div>
+
+<p>The lenticular substance, l. i. provided with each scaly lamina, are
+the only parts of those described by M. de Reaumur, that we have found
+engaged in the organs of our queens. The canal, r, by Swammerdam
+denominated the root of the penis, breaks in copulation; and we have
+seen its fragments at the place where it unites to the end of the
+lentil, l. towards the anterior extremity; but we have found no
+traces of the canal, k, formed of involuted membranes, which in the body
+of the male
+<span class="pagebreak" title="284">&nbsp;</span><a name="Page_284" id="Page_284"></a>
+proceeds from the posterior end of the lentil, l. i. nor of
+the plaited pallet, p. adhering to this canal, called by Swammerdam the
+penis from its resemblance to that of other animals, though he is not of
+opinion that this point, which is not perforated, can perform the
+functions of a real penis, and hold the principal part in generation.
+The canal, k, therefore, and all appertaining to it, must break at i,
+quite close to the posterior part of the lentil, since we found no
+remains of the lenticular bodies left by the fecundating males, in the
+body of our females. The canal, r, which Swammerdam calls the root of
+the penis, with greater reason than he was himself aware, is not
+extended in the body of the male as represented by the figure here
+engraved, but this long twilled canal consists of several involutions,
+from the seminal vessels whence it proceeds, into the lenticular body
+where it terminates, and where it conveys the fluid. This canal
+therefore can extend during copulation, and allow the lenticular
+substance to protrude out of the body of the males.</p>
+
+<p>It is evident this may be the case during copulation as is seen on
+opening a drone, for, by endeavouring to displace the lenticular body,
+the involutions of the cord disappear, and it extends much more than
+
+<span class="pagebreak" title="285">&nbsp;</span><a name="Page_285" id="Page_285"></a>
+necessary for the lentil to protrude from the body; and if we attempt
+to separate it farther, the canal breaks at l. close to the lentil,
+and at the same place where it breaks in copulation.</p>
+
+<p>By dissection two nerves are discovered, towards the origin of the
+canal, r. inserted into the seminal vessels and distribute in them, and
+towards the root of the penis many ramifications undoubtedly serving for
+the motion of these parts. Two small parts, perceptible near the nerves,
+are two ligaments for retaining the generative organs in their proper
+place, so that except the root of the penis, they cannot be drawn out
+without some exertion; it and the lenticular body however can protrude,
+and actually do so during copulation. A certain degree of pressure
+forces all these parts from the body of the male, but they spontaneously
+return, and appear reversed.</p>
+
+<p>Swammerdam, and after him M. de Reaumur have admired this mechanism;
+they have thought, indeed, that the return should be occasioned by the
+effect of the air inflating the parts, and they supposed that the male
+organs proceeded from the body, and returned during copulation, the same
+as when forced out by pressure. Following their example, we have pressed
+them
+<span class="pagebreak" title="286">&nbsp;</span><a name="Page_286" id="Page_286"></a>
+from the body of many males; we have a thousand times witnessed
+this wonderful return, which they detail with the greatest precision;
+but our males never survived the operation. We have seen, as M. de
+Reaumur, a few males protrude them spontaneously, even some of the parts
+inverted, but at that moment they died, and were unable to retract the
+parts which a pressure, most likely accidental, had forced out. Thus it
+is improbable that the male organs protrude by turning out of themselves
+in copulation; and the details which follow prove incontestibly, that it
+is otherwise. Had not Swammerdam been prejudiced with this opinion, he
+would have seen that the lenticular body can proceed from the body in
+erection without reversing itself; he could have proportioned the
+<a name="corr286" id="corr286"></a><a class="correction" href="#cnote286" title="changed from 'tortous'">tortuous</a> canal, which he calls the root of the penis; he would have seen
+that, at certain times, it can be sufficiently extended to allows the
+lenticular substance to protrude; he would have discovered the real use
+of the scaly plates; he would have explained that of the canal k, of the
+plaited pallet q, and the movements of all these parts, more admirable
+perhaps than the inversion which he was the first to observe.</p>
+
+<p>
+<span class="pagebreak" title="287">&nbsp;</span><a name="Page_287" id="Page_287"></a>
+Our observations incontestibly prove copulation. The portion of the
+males found engaged in the body of our queens, hitherto called the
+lenticular substance, may be denominated a penis both from its position
+and use. The same surface is presented by it in the queen as in the body
+of the male, which is proved by the position of the lamin&aelig;, e. e.
+attached to the interior of the penis, when found in the queen. It is
+evident, if the supposed inversion took place, the lamin&aelig; would be found
+within the posterior part of the penis; and we should see them through
+its membrane, by their concave side, instead of which the convex surface
+is presented when in the vulva of females, the same as in the body of
+the males. But what is the use of these lamin&aelig;? From their figure,
+hardness, relative position with respect to each other, and their
+situation at the extremity of the penis, we cannot doubt they are real
+pincers. However, to ascertain the fact, we found it necessary to see
+their position, and that of the penis itself in the females. For this
+purpose, we prevented some of the queens from extracting the parts left
+by the impregnating males, and by dissection we discovered that the
+lamin&aelig; were pincers as we had conjectured.</p>
+
+<p>
+<span class="pagebreak" title="288">&nbsp;</span><a name="Page_288" id="Page_288"></a>
+The penis was situated under the sting of the queens, and pressed
+against the upper region of the belly. It was supported by the posterior
+end, against the extremity of the vagina, or excretory canal. There we
+were sensible of the motion and use of the scaly pieces. Their
+extremities were separated a little more than in the male, and pressed
+between them some of the female parts below the excretory canal. The
+extreme minuteness of these parts prevented us from distinguishing them
+clearly, but the effort necessary to separate and remove the penis from
+the female, satisfied us of the use of these lamin&aelig;.</p>
+
+<p>Inspecting a male from above, the convex side of the plates, e. e. is
+presented, and the summit of the angle formed by their origin. When in
+the body of the female, they are in the inverse position; what was above
+in the male is now below, and the extremity of the pincers directed
+upwards. This makes us suspect that in copulation the male mounts on the
+back of the female, but we are far from asserting it positively. It may
+be asked whether that part we call the penis, is the sole part
+introduced into the female during copulation? We have carefully
+investigated this, and can affirm, that it is the only one of all those
+
+<span class="pagebreak" title="289">&nbsp;</span><a name="Page_289" id="Page_289"></a>
+described by M. de Reaumur, which has been found in our females. But we
+have discovered a new part that escaped both him and Swammerdam, which
+appears from the following experiment.</p>
+
+<p>Separating the lenticular substance from the excretory canal, where it
+was attached, we drew along with it a white body, adhering by one
+extremity, and having the other engaged in the vagina. Towards the end
+of the lentil, where the substance adhered, it appeared cylindrical,
+then it swelled, and again contracted, to dilate anew in a greater
+degree than at first; afterwards it contracted and terminated in a
+point. A powerful magnifier was required to see all this. When pulled
+from the lenticular body, the part was commonly broke, and also when
+extracted by the queens from themselves. The figure and situation seemed
+to authorise our considering it the penis itself, and the lenticular
+body only an appendage. But the last queen we examined exhibited a
+peculiarity that induced us to doubt the fact, and led us to suspect
+that this body is nothing else than the seminal fluid itself, moulded
+and coagulated in the vagina,
+<span class="pagebreak" title="290">&nbsp;</span><a name="Page_290" id="Page_290"></a>
+and which from its viscosity adheres to
+the lenticular substance, and accompanies it when separated from the
+vagina. In this queen was found a little extravasated white matter, near
+the opening of the vagina. This, though at first liquid, soon coagulated
+in the air as the seminal fluid of drones does. In separating the
+lenticular body from the vagina, we drew along with it a thread which
+broke near the lentil; and seemed of too little consistence for the
+penis of a male. The lenticular bodies, found in our queens, appeared
+larger than in the males we dissected, and we have remarked with M. de
+Reaumur, that these parts are not of equal size in every male.</p>
+
+
+<p class="gaplet"><i>Experiment 1.</i>&mdash;On the tenth of July, we set successively at liberty
+three virgin queens four or five days old. Two flew away several times;
+their absence was short and fruitless. The third profited better by her
+liberty; she departed thrice; the first and second time her absence was
+short; but the third lasted thirty-five minutes. She returned in a very
+different state; and in such as allowed no doubt of her employment, for
+she exhibited the part of a male that had rendered her a mother. We
+seized her wings with one
+<span class="pagebreak" title="291">&nbsp;</span><a name="Page_291" id="Page_291"></a>
+hand, and in the other received the
+lenticular body, of which she had disengaged herself with her claws. The
+posterior part was armed with two pincers, e. e. shelly and elastic,
+which could be drawn asunder, and then resumed their original position.
+Towards the anterior part of the lentil appeared the fragment of the
+root of the penis; this canal had broke half a line from the lenticular
+body. We allowed the queen to enter her habitation, and adapted the
+entrance so that she could not leave it unknown to us.</p>
+
+<p>On the seventeenth we found no eggs in the hive; the queen was as
+slender as the first day; therefore the male, with which she had
+copulated, had not impregnated her eggs. She was again set at liberty;
+after twice departing, she returned with evidence of a second
+copulation. We then confined her, and the eggs she afterwards laid
+proved that the second copulation had been more successful than the
+first and that there are some males more fit for impregnating queens
+than others. However, it is very rare that the first copulation is
+inefficient; we have only seen two that required it twice; all the rest
+were impregnated by the first.</p>
+
+
+<p class="gaplet">
+<span class="pagebreak" title="292">&nbsp;</span><a name="Page_292" id="Page_292"></a>
+<i>Experiment 2.</i>&mdash;On the eighteenth we put at liberty a virgin queen
+twenty-seven days old, she departed twice. Her second absence was
+twenty-eight minutes, and she returned with the proofs of copulation. We
+prevented her from entering, and put her under a glass to see how she
+would disengage the male organs. This she was unable to accomplish,
+having only the table and sides of the glass for support; therefore we
+introduced a bit of comb; thus providing the same conveniences as are in
+a hive. Fixing herself on <i>it</i> by the first four legs, she stretched out
+the two last, and extending them along her belly seemed to press it
+between them. At length introducing her claws between the two parts of
+the last ring, she seized the lenticular body, and dropped it on the
+table. The posterior part was provided with shelly pincers, under which
+and in the same direction was a grey cylindrical body. The end farthest
+from the lentil was sensibly thicker than that adhering to it, and
+terminated in a point. This point was double, and open like the bill of
+a bird, which induces us to think the body was broken, a conjecture
+supported by the following experiment.</p>
+
+
+<p class="gaplet">
+<span class="pagebreak" title="293">&nbsp;</span><a name="Page_293" id="Page_293"></a>
+<i>Experiment 3.</i>&mdash;On the nineteenth we set at liberty a queen four days
+old; she departed twice; her first absence was short; the second lasted
+thirty minutes, and then she returned with the marks of fecundation. As
+we wished to obtain the male organs entire, it was necessary to prevent
+the queen from breaking them by extracting them with her feet; we
+therefore suddenly killed her, and cut off the last rings in order to
+lay the vulva open. But though deprived of animation, so much life
+remained in these parts that the lenticular body was thrown out
+spontaneously. Under the pincers appeared the remnant of a cylindrical
+body which had broken near the origin and remained in the female. This
+body was very small at the origin; it afterwards sensibly enlarged; next
+contracting by degrees, it terminated in a sharp point. We found the
+point engaged up to the gland in the excretory canal, and the rest in
+the vulva.</p>
+
+
+<p class="gaplet"><i>Experiment 4.</i>&mdash;We set two virgin queens at liberty on the twentieth.
+The first had been abroad on the preceding days, but the scarcity of
+males prevented her from
+<span class="pagebreak" title="294">&nbsp;</span><a name="Page_294" id="Page_294"></a>
+being previously fecundated. She returned with
+the organs of a male. We tried to prevent her from extracting them, but
+she did this so expeditiously with her feet, that we could not
+accomplish it. She was then allowed to enter the hive.</p>
+
+<p>The second queen departed twice. Her first absence was short as usual;
+the second lasted about half an hour, and she returned impregnated.
+Having killed her as suddenly as possible, we laid open the vulva. The
+lenticular body was deposited as in every queen hitherto dissected; the
+pincers were situated under the excretory canal. Some parts not easily
+distinguishable were pressed between the <a name="corr294" id="corr294"></a><a class="correction" href="#cnote294" title="changed from 'laminae'">lamin&aelig;</a>, and their
+office seemed to consist in forcing the extremity of the lentil to
+approach the orifice of the vagina, and apply so forcibly to it that
+some exertion was necessary to separate them. We previously examined
+them, with a very powerful magnifier. Then a peculiarity which had
+escaped us was perceptible. In drawing out the lenticular body, there
+proceeded from the vagina a minute part, v. adhering to the posterior
+end of the lentil, and situated below the plates. It spontaneously
+retracted into the lentil, like
+<span class="pagebreak" title="295">&nbsp;</span><a name="Page_295" id="Page_295"></a>
+the horns of a snail. It appeared
+white, very short, and cylindrical. Under the pincers was a little half
+coagulated seminal fluid at the bottom of the vulva. Though much could
+be expressed, there was none pure; it was almost liquid, but soon
+coagulated, and formed a whitish inorganic mass. This observation
+carefully made removed all our doubts, and demonstrated that what we had
+taken for the penis of males was nothing but the seminal fluid, which
+had coagulated and assumed the interior figure of the vagina. The only
+hard part introduced by the male, was the short cylindrical point which
+retracted into the lentil, when we separated it. Its situation and
+office prove that it is there we must look for the issue of the seminal
+fluid, if we can hope to find an opening, when not engaged in
+copulation.</p>
+
+<p>We found this new part in the first drone we dissected. By pressing the
+seminal vessels, the white liquid then escaped downwards to the root of
+the penis r. and into the lenticular body, l. i. which became
+sensibly swoln. We prevented the fluid from returning, and by new
+pressure of the lentil forced it to advance. However,
+<span class="pagebreak" title="296">&nbsp;</span><a name="Page_296" id="Page_296"></a>
+none escaped, but
+we saw at the posterior end of the lenticular body, and under the scaly
+pincers, a small white cylindrical substance, the same in appearance as
+that we had found engaged in the vagina of the queen. This part
+retracted on pressure, and then returned.</p>
+
+<p>I request you, Sir, while perusing this letter, to inspect the figure of
+the male sexual organs published by M. de Reaumur, and which are copied
+here. The descriptions are most accurate, and present a just idea of the
+situation of these parts when in the male's body. We readily conceive
+how they appear when left in the female by copulation. This detail will
+sufficiently indicate the situation and figure of the new part I have
+discovered.</p>
+
+<p>I suspect that the males perish after losing their sexual organs. But
+why does nature exact so great a sacrifice? This is a mystery which I
+cannot pretend to unveil. I am unacquainted with any analogous fact in
+natural history, but as there are two species of insects whose
+copulation can take place only in the air, namely, ephemer&aelig; and ants, it
+would be extremely interesting to discover whether
+<span class="pagebreak" title="297">&nbsp;</span><a name="Page_297" id="Page_297"></a>
+their males also
+lose their sexual parts, in the same circumstances, and whether, as with
+drones, enjoyment in their flight is the prelude of death.</p>
+
+<p class="center biggap big">FINIS.</p>
+
+
+<div class="footnotes">
+
+<div class="footnote"><p><a name="fn_O" id="fn_O"></a><span class="label"><a href="#fnm_O">O</a></span> Memoires sur les Abeilles, p. 450.</p></div>
+
+<div class="footnote"><p><a name="fn_P" id="fn_P"></a><span class="label"><a href="#fnm_P">P</a></span> Such long and minute descriptions can be very imperfectly
+translated; indeed they are unintelligible without microscopical
+inspections of the parts themselves.&mdash;T.</p></div>
+</div>
+
+
+
+
+
+<h2>ANALYTICAL INDEX.</h2>
+
+
+<div class="center">
+<table summary="topics and page numbers">
+<tr><td class="aindex">Description of a hive invented by the author                   </td><td class="toc2">page&nbsp;<a href="#Page_4">4</a></td></tr>
+<tr><td class="aindex">Swammerdam's opinion on the fecundation of bees                    </td><td class="toc2"><a href="#Page_8">8</a></td></tr>
+<tr><td class="aindex">Sentiments of M. de Reaumur                                       </td><td class="toc2"><a href="#Page_10">10</a></td></tr>
+<tr><td class="aindex">Mr Debraw's opinion                                               </td><td class="toc2"><a href="#Page_11">11</a></td></tr>
+<tr><td class="aindex">Hattorf's opinion                                                 </td><td class="toc2"><a href="#Page_19">19</a></td></tr>
+<tr><td class="aindex">Difficulty of discovering the mode of impregnation                </td><td class="toc2"><a href="#Page_22">22</a></td></tr>
+<tr><td class="aindex">Experiments on the subject                                        </td><td class="toc2"><a href="#Page_23">23</a></td></tr>
+<tr><td class="aindex">Suggestions by M. Bonnet                                          </td><td class="toc2"><a href="#Page_34">34</a></td></tr>
+<tr><td class="aindex">The queen is impregnated by copulation, which never takes place
+    within the hive                                                 </td><td class="toc2"><a href="#Page_41">41</a></td></tr>
+<tr><td class="aindex">Experiments on artificial fecundation have not succeeded          </td><td class="toc2"><a href="#Page_42">42</a></td></tr>
+<tr><td class="aindex">The male loses the sexual organs in copulation                    </td><td class="toc2"><a href="#Page_43">43</a></td></tr>
+<tr><td class="aindex">Regarded impregnation affects the ovaries of the queen            </td><td class="toc2"><a href="#Page_45">45</a></td></tr>
+<tr><td class="aindex">She then lays no eggs but those producing males                   </td><td class="toc2"><a href="#Page_47">47</a></td></tr>
+<tr><td class="aindex">One copulation impregnates all the eggs the queen will lay in
+    two years                                                       </td><td class="toc2"><a href="#Page_54">54</a></td></tr>
+<tr><td class="aindex">Fecundity of a queen                                              </td><td class="toc2"><a href="#Page_63">63</a></td></tr>
+<tr><td class="aindex">Common bees do not transport the queen's eggs                     </td><td class="toc2"><a href="#Page_66">66</a></td></tr>
+<tr><td class="aindex">They sometimes eat them                                           </td><td class="toc2"><a href="#Page_69">69</a></td></tr>
+<tr><td class="aindex">Eggs producing males are sometimes laid in royal cells            </td><td class="toc2"><a href="#Page_71">71</a></td></tr>
+<tr><td class="aindex">Common worms may be converted into queens                         </td><td class="toc2"><a href="#Page_77">77</a></td></tr>
+
+
+<tr><td class="aindex">Operations of the bees when this is done                          </td><td class="toc2"><a href="#Page_78">78</a></td></tr>
+<tr><td class="aindex">Fertile workers sometimes exist                                   </td><td class="toc2"><a href="#Page_89">89</a></td></tr>
+<tr><td class="aindex">They lay none but the eggs of males                               </td><td class="toc2"><a href="#Page_96">96</a></td></tr>
+<tr><td class="aindex">All common bees are originally females                            </td><td class="toc2"><a href="#Page_98">98</a></td></tr>
+<tr><td class="aindex">Receiving the royal food while larv&aelig;, expands their ovaries      </td><td class="toc2"><a href="#Page_105">105</a></td></tr>
+<tr><td class="aindex">Mutual enmity of queens                                          </td><td class="toc2"><a href="#Page_110">110</a></td></tr>
+<tr><td class="aindex">The common bees seem to promote their combats                    </td><td class="toc2"><a href="#Page_117">117</a></td></tr>
+<tr><td class="aindex">A guard is constantly at the entrance of the hive                </td><td class="toc2"><a href="#Page_123">123</a></td></tr>
+<tr><td class="aindex">What ensues when bees lose their queen                           </td><td class="toc2"><a href="#Page_126">126</a></td></tr>
+<tr><td class="aindex">Effects of introducing a stranger queen                          </td><td class="toc2"><a href="#Page_128">128</a></td></tr>
+<tr><td class="aindex">Massacre of the males                                            </td><td class="toc2"><a href="#Page_132">132</a></td></tr>
+<tr><td class="aindex">It never ensues in hives deprived of queens                      </td><td class="toc2"><a href="#Page_135">135</a></td></tr>
+<tr><td class="aindex">A plurality of queens is never tolerated                         </td><td class="toc2"><a href="#Page_142">142</a></td></tr>
+<tr><td class="aindex">The queen bee is oviparous                                       </td><td class="toc2"><a href="#Page_149">149</a></td></tr>
+<tr><td class="aindex">Bees seem occasionally to repose                                 </td><td class="toc2"><a href="#Page_150">150</a></td></tr>
+<tr><td class="aindex">Interval between production of the egg and the perfect state
+    of bees                                                        </td><td class="toc2"><a href="#Page_151">151</a></td></tr>
+<tr><td class="aindex">Mode of spinning the coccoon                                     </td><td class="toc2"><a href="#Page_153">153</a></td></tr>
+<tr><td class="aindex">That of the queen is open at one end                             </td><td class="toc2"><a href="#Page_154">154</a></td></tr>
+<tr><td class="aindex">The size of the bees is not affected by that of the cells        </td><td class="toc2"><a href="#Page_167">167</a></td></tr>
+<tr><td class="aindex">The old queen always conducts the first swarm                    </td><td class="toc2"><a href="#Page_173">173</a></td></tr>
+<tr><td class="aindex">But never before depositing eggs in the royal cells              </td><td class="toc2"><a href="#Page_177">177</a></td></tr>
+<tr><td class="aindex">Singular effect of a sound emitted by perfect queens             </td><td class="toc2"><a href="#Page_189">189</a></td></tr>
+<tr><td class="aindex">The instinct of bees is affected during the period of swarming   </td><td class="toc2"><a href="#Page_208">208</a></td></tr>
+<tr><td class="aindex">Queens are liberated from their cells according to their age     </td><td class="toc2"><a href="#Page_214">214</a></td></tr>
+<tr><td class="aindex">The bees probably judge of this by the sound emitted             </td><td class="toc2"><a href="#Page_217">217</a></td></tr>
+<tr><td class="aindex">Young queens conducting swarms are virgins                       </td><td class="toc2"><a href="#Page_221">221</a></td></tr>
+<tr><td class="aindex">The conduct of bees to old queens is peculiar                    </td><td class="toc2"><a href="#Page_224">224</a></td></tr>
+<tr><td class="aindex">Retarded impregnation affects the instinct of queens             </td><td class="toc2"><a href="#Page_241">241</a></td></tr>
+
+
+<tr><td class="aindex">Amputation of the antenn&aelig; produces singular effects              </td><td class="toc2"><a href="#Page_245">245</a></td></tr>
+<tr><td class="aindex">Advantages of the leaf hive                                      </td><td class="toc2"><a href="#Page_253">253</a></td></tr>
+<tr><td class="aindex">It renders the bees tractable                                    </td><td class="toc2"><a href="#Page_256">256</a></td></tr>
+<tr><td class="aindex">They may there be forced to work in wax                          </td><td class="toc2"><a href="#Page_264">264</a></td></tr>
+<tr><td class="aindex">Uniform distance between the combs                               </td><td class="toc2"><a href="#Page_265">265</a></td></tr>
+<tr><td class="aindex">Natural heat of bees                                             </td><td class="toc2"><a href="#Page_269">269</a></td></tr>
+<tr><td class="aindex">Distance to which they fly                                       </td><td class="toc2"><a href="#Page_271">271</a></td></tr>
+<tr><td class="aindex">Appendix                                                         </td><td class="toc2"><a href="#Page_273">273</a></td></tr>
+<tr><td class="aindex">Anatomical observations on the sexual organs of bees             </td><td class="toc2"><a href="#Page_276">276</a></td></tr>
+<tr><td class="aindex">Experiments proving the copulation of the queen                  </td><td class="toc2"><a href="#Page_290">290</a></td></tr>
+</table>
+</div>
+
+<p class="biggap indent1 little center bt biggapbelow"><span class="smcap">Alex. Smellie</span>, Printer.</p>
+
+<div class="transnote">
+<h4><a name="tnotes" id="tnotes"></a>Transcriber's notes</h4>
+
+<p>The spelling in the original is sometimes idiosyncratic. It has not
+been changed, but a few obvious errors have been corrected. The
+corrections are listed below and marked with a <span class="correction" title="like this">mouse-hover</span> in the text.</p>
+
+<p class="hanging">Inconsistent spellings include:<br />
+  Lusace/Lusaçe, centre/center, choose/chuse,
+  organisation/organization, recognise/recognize</p>
+
+<p class="hanging">Unusual spellings (which have not been changed) include:<br />
+  centinels, coccoon, diaphraghm, encreased, encreasing, groupes,
+  harrassed, inaccessible, incontestible, indispensible, moveable,
+  perceptible, susceptible, uncontrouled, unintelligible</p>
+
+<p class="hanging">Letter I<br />
+  <a name="cnote39" id="cnote39"></a>"secret distinctive characterestics" changed to
+  "secret distinctive <a href="#corr39">characteristics</a>"<br />
+  <a name="cnotefB" id="cnotefB"></a>The "<a href="#corrfB">fig. 4</a>" referred to in the second footnote can not be found.
+</p>
+
+<p class="hanging">Letter II<br />
+
+  <a name="cnote52" id="cnote52"></a>"the copulalation of queens" changed to
+  "the <a href="#corr52">copulation</a> of queens"
+</p>
+
+<p class="hanging">Letter IV<br />
+
+  <a name="cnote82" id="cnote82"></a>"The worms had spun their silk coccons" changed to
+  "The worms had spun their silk <a href="#corr82">coccoons</a>"
+</p>
+
+<p class="hanging">Letter V<br />
+
+  <a name="cnote94" id="cnote94"></a>"characteristics of commo nbees" changed to
+  "characteristics of <a href="#corr94">common bees</a>"
+</p>
+
+<p class="hanging">Letter VI<br />
+
+  <a name="cnote115" id="cnote115"></a>"The result of this rencounter" changed to
+  "The result of this <a href="#corr115">encounter</a>"<br />
+
+  <a name="cnote119" id="cnote119"></a>"genius such as <a href="#corr119">your's</a>" unchanged.<br />
+
+  <a name="cnote123" id="cnote123"></a>"observing that the antennae" changed to
+  "observing that the <a href="#corr123">antennæ</a>"<br />
+
+  <a name="cnote131" id="cnote131"></a>"combats and disastrou scenes" changed to
+  "combats and <a href="#corr131">disastrous</a> scenes"<br />
+
+  <a name="cnote132" id="cnote132"></a>"M. de Reamur speaks of these executions" changed to
+  "M. de <a href="#corr132">Reaumur</a> speaks of these executions"
+</p>
+
+<p class="hanging">Letter IX<br />
+
+  <a name="cnote195" id="cnote195"></a>"Only the few bees that not participated" changed to
+  "Only the few bees that <a href="#corr195">had</a> not participated"
+</p>
+
+<p class="hanging">Letter XI<br />
+
+  <a name="cnote227" id="cnote227"></a>"these tumultous motions" changed to
+  "these <a href="#corr227">tumultuous</a> motions"
+</p>
+
+<p class="hanging">Letter XII<br />
+
+  <a name="cnote242" id="cnote242"></a>"one <a href="#corr242">antennæ</a>" unchanged.<br />
+
+  <a name="cnote247" id="cnote247"></a>"reside in them," changed to
+  "reside in <a href="#corr247">them.</a>"
+
+</p>
+
+<p class="hanging">Appendix<br />
+
+  <a name="cnote281" id="cnote281"></a>"the cirumference is edged" changed to
+  "the <a href="#corr281">circumference</a> is edged"<br />
+
+  <a name="cnote286" id="cnote286"></a>"he could have proportioned the tortous canal" changed to
+  "he could have proportioned the <a href="#corr286">tortuous</a> canal"<br />
+
+  <a name="cnote294" id="cnote294"></a>"pressed between the laminae" changed to
+  "pressed between the <a href="#corr294">laminæ</a>"
+</p>
+
+</div>
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
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@@ -0,0 +1,5329 @@
+The Project Gutenberg EBook of New observations on the natural history of
+bees, by Francis Huber
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: New observations on the natural history of bees
+
+Author: Francis Huber
+
+Translator: Anonymous
+
+Release Date: August 28, 2008 [EBook #26457]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK NEW OBSERVATIONS ON BEES ***
+
+
+
+
+Produced by Louise Pryor, Steven Giacomelli and the Online
+Distributed Proofreading Team at https://www.pgdp.net (This
+file was produced from images produced by Core Historical
+Literature in Agriculture (CHLA), Cornell University)
+
+
+
+
+
+{Transcriber's note
+
+The spelling in the original is sometimes idiosyncratic. It has not
+been changed, but a few obvious errors have been corrected. The
+corrections are listed at the end of this etext.}
+
+
+{Illustration: The figures that are referred to in the text}
+
+
+
+
+  NEW OBSERVATIONS
+  ON THE
+  NATURAL HISTORY
+  OF
+  BEES,
+
+  BY
+  FRANCIS HUBER.
+
+  TRANSLATED FROM THE ORIGINAL.
+
+       *       *       *       *       *
+
+  EDINBURGH:
+
+  PRINTED FOR JOHN ANDERSON,
+  AND SOLD BY
+  LONGMAN, HURST, REES, AND ORME,
+  LONDON.
+
+  ALEX SMELLIE, Printer.
+
+  1806.
+
+
+
+
+  _To
+  SIR JOSEPH BANKS, BART._
+
+  _KNIGHT OF THE MOST HONOURABLE ORDER
+  OF THE BATH, A PRIVY COUNCILLOR,
+  PRESIDENT OF THE ROYAL
+  SOCIETY OF LONDON,
+  &c. &c._
+
+  _THIS TRANSLATION
+  IS INSCRIBED._
+
+
+
+
+CONTENTS.
+
+
+  LETTER 1.--On the impregnation of the queen bee         page 1
+
+  LETTER 2.--Sequel of observations on the impregnation
+             of the queen bee                                 41
+
+  LETTER 3.--The same subject continued; observations
+             on retarding the fecundation of queens           44
+
+  LETTER 4.--On M. Schirach's discovery                       76
+
+  LETTER 5.--Experiments proving that there are
+             sometimes common bees which lay fertile eggs     89
+
+  LETTER 6.--On the combats of queens; the massacre
+             of the males; and what succeeds in a hive
+             where a stranger queen is substituted for
+             the natural one                                 108
+
+  LETTER 7.--Sequel of observations on the reception
+             of a stranger queen; M. de Reaumur's
+             observations on the subject                     137
+
+  LETTER 8.--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 coccoons 145
+
+  LETTER 9.--On the formation of swarms                      171
+
+  LETTER 10.--The same subject continued                     201
+
+  LETTER 11.--The same subject continued                     223
+
+  LETTER 12.--Additional observations on queens
+             that lay only the eggs of drones, and on
+             those deprived of the antennae                   237
+
+  LETTER 13.--Economical considerations on bees              253
+
+  APPENDIX                                                   275
+
+
+
+
+TRANSLATOR'S PREFACE
+
+
+The facts contained in this volume are deeply interesting to the
+Naturalist. They not only elucidate the history of those industrious
+animals, whose nature is the peculiar subject of investigation, but they
+present some singular features in physiology which have hitherto been
+unknown.
+
+The industry of bees has proved a fertile source of admiration in all
+countries and in every age; and mankind have endeavoured to render it
+subservient to their gratifications or emolument. Hence innumerable
+theories, experiments, and observations have ensued, and uncommon
+patience has been displayed in prosecuting the enquiry. But although
+many interesting peculiarities have been discovered, they are so much
+interwoven with errors, that no subject has given birth to more
+absurdities than investigations into the history of bees: and
+unfortunately those treatises which are most easily attained, and the
+most popular, only serve to give such absurdities a wider range, and
+render it infinitely more difficult to eradicate them. A considerable
+portion of the following work is devoted to this purpose. The reader
+will judge of the success which results from the experiments that have
+been employed.
+
+Perhaps this is not the proper place to bestow an encomium on a treatise
+from which so much entertainment and instruction will be derived.
+However, to testify the estimation in which it is held in other nations,
+the remarks upon it by the French philosopher Sue, may be quoted, 'The
+observations are so consistent, and the consequences seem so just, that
+while perusing this work, it appears as if we had assisted the author
+in each experiment, and pursued it with equal zeal and interest. Let us
+invite the admirers of nature to read these observations; few are equal
+to them in excellence, or so faithfully describe the nature, the habits,
+and inclinations of the insects of which they treat.'
+
+It is a remarkable circumstance that the author laboured under a defect
+in the organs of vision, which obliged him to employ an assistant in his
+experiments. Thus these discoveries may be said to acquire double
+authority. But independent of this the experiments are so judiciously
+adapted to the purposes in view, and the conclusions so strictly
+logical, that there is evidently very little room for error. The talents
+of _Francis Burnens_, this philosophic assistant, had long been devoted
+to the service of the author, who, after being many successive years in
+this manner aided in his researches, was at last deprived of him by some
+unfortunate accident.
+
+Whether the author has prosecuted his investigation farther does not
+appear, as no other production of his pen is known in this island.
+
+It is vain to attempt a translation of any work without being to a
+certain degree skilled in the subject of which it treats. Some parts of
+the original of the following treatise, it must be acknowledged, are so
+confused, and some so minute, that it is extremely difficult to give an
+exact interpretation. But the general tenor, though not elegant, is
+plain and perspicuous; and such has it been here retained.
+
+
+
+
+LETTER I.
+
+_ON THE IMPREGNATION OF THE QUEEN BEE._
+
+
+  SIR,
+
+When I had the honour at Genthod of giving you an account of my
+principal experiments on bees, you desired me to transmit a written
+detail, that you might consider them with greater attention. I hasten,
+therefore, to extract the following observations from my journal.--As
+nothing can be more flattering to me than the interest you take in my
+researches, permit me to remind you of your promise to suggest new
+experiments{A}.
+
+After having long studied bees in glass hives constructed on M. de
+Reaumur's principle, you have found the form unfavourable to an
+observer. The hives being too wide, two parallel combs were made by the
+bees, consequently whatever passed between them escaped observation.
+From this inconvenience, which I have experienced, you recommended much
+thinner hives to naturalists, where the panes should be so near each
+other, that only a single row of combs could be erected between them. I
+have followed your admonitions, Sir, and provided hives only eighteen
+lines in width, in which I have found no difficulty to establish swarms.
+However, bees must not be entrusted with the charge of constructing a
+single comb: Nature has taught them to make parallel ones, which is a
+law they never derogate from, unless when constrained by some particular
+arrangement. Therefore, if left to themselves in these thin hives, as
+they cannot form two combs parallel to the plane of the hive, they will
+form several small ones perpendicular to it, and, in that case, all is
+equally lost to the observer. Thus it became essential previously to
+arrange the position of the combs. I forced the bees to build them
+perpendicular to the horizon, and so that the lateral surfaces were
+three or four lines from the panes of the hive. This distance allows the
+bees sufficient liberty, but prevents them from collecting in too large
+clusters on the surface of the comb. By such precautions, bees are
+easily established in very thin hives. There they pursue their labours
+with the same assiduity and regularity; and, every cell being exposed,
+none of their motions can be concealed.
+
+It is true, that by compelling these insects to a habitation where they
+could construct only a single row of combs, I had, in a certain
+measure, changed their natural situation, and this circumstance might
+possibly have affected their instinct. Therefore, to obviate every
+objection, I invented a kind of hives, which, without losing the
+advantages of those very thin, at the same time approached the figure of
+common hives where bees form several rows of combs.
+
+I took several small fir boxes, a foot square and fifteen lines wide,
+and joined them together by hinges, so that they could be opened and
+shut like the leaves of a book{B}. When using a hive of this
+description, we took care to fix a comb in each frame, and then
+introduced all the bees necessary for each particular experiment. By
+opening the different divisions successively, we daily inspected both
+surfaces of every comb. There was not a single cell where we could not
+distinctly see what passed at all times, nor a single bee, I may almost
+say, with which we were not particularly acquainted. Indeed, this
+construction is nothing more than the union of several very flat hives
+which may be separated. Bees, in such habitations, must not be visited
+before their combs are securely fixed in the frames, otherwise, by
+falling out, they may kill or hurt them, as also irritate them to that
+degree that the observer cannot escape stinging, which is always
+painful, and sometimes dangerous: but they soon become accustomed to
+their situation, and in some measure tamed by it; and, 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 exhibiting too formidable
+symptoms of displeasure. You will remember, Sir, that on visiting my
+retreat, I shewed you a hive of this kind that had been a long time in
+experiment, and how much you were surprised that the bees so quietly
+allowed us to open it.
+
+In these hives, I have repeated all my observations, 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
+inconvenience 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 error; and it also appears, that some
+advantages are found in these which may be called _Book_ or
+_Leaf-hives_, as they prove extremely useful in the economical treatment
+of bees, which shall afterwards be detailed.
+
+I now come to the particular object of this letter, the fecundation of
+the queen bee; and I shall, in a few words, examine the different
+opinions of naturalists on this singular problem. Next I shall state the
+most remarkable observations which their conjectures have induced me to
+make, and then describe the new experiments by which I think I have
+solved the problem{C}.
+
+Swammerdam, who studied bees with unremitting attention, and who never
+could see a real copulation between a drone and a queen, was satisfied
+that copulation was unnecessary for fecundation of the eggs: but having
+remarked that, at certain times, the drones exhaled a very strong odour,
+he thought this odour was an emanation of the _aura seminalis_, or the
+_aura seminalis_ itself, which operated fecundation by penetrating the
+body of the female. His conjecture was confirmed on dissecting the male
+organs of generation; for he was so much struck with the disproportion
+between them and those of the female, that he did not believe copulation
+possible. His opinion, concerning the influence of the odour, had this
+farther advantage, that it afforded a good reason for the prodigious
+number of the males. There are frequently fifteen hundred or two
+thousand in a hive; and, according to Swammerdam, it is necessary they
+should be numerous, that the emanation proceeding from them may have an
+intensity or energy sufficient to effect impregnation.
+
+Though M. de Reaumur has refuted this hypothesis by just and conclusive
+reasoning, he has failed to make the sole experiment that could support
+or overturn it. This was to confine all the drones of a hive in a tin
+case, perforated with minute holes, which might allow the emanation of
+the odour to escape, but prevent the organs of generation from passing
+through. Then, this case should have been placed in a hive well
+inhabited, but completely deprived of males, both of large and small
+size, and the consequence attended to. It is evident, 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
+confuted had she produced no eggs, or only sterile ones. However the
+experiment has been made by us, and the queen remained barren;
+therefore, it is undoubted, that the emanation of the odour of the males
+does not impregnate bees.
+
+M. de Reaumur was of a different opinion. He thought that the queen's
+fecundation followed actual copulation. He confined several drones in a
+glass vessel along with a virgin queen: he saw the female make many
+advances to the males; but, unable to observe any union so intimate
+that it could be denominated copulation, he leaves the question
+undecided. We have repeated this experiment: we have frequently confined
+virgin queens with drones of all ages: we have done so at every season,
+and witnessed all their advances and solicitations to the males: we have
+even believed we saw a kind of union between them, but so short and
+imperfect that it was unlikely to effect impregnation. Yet, to neglect
+nothing, we confined the virgin queen, that had suffered the approaches
+of the male, to her hive. During a month that her imprisonment
+continued, she did not lay a single egg; therefore, these momentary
+junctions do not accomplish fecundation.
+
+In the _Contemplation de la Nature_, you have cited the observations of
+the English naturalist Mr Debraw. They appear correct, and at last to
+elucidate the mystery. Favoured by chance, the observer one day
+perceived at the bottom of cells containing eggs, a whitish fluid,
+apparently spermatic, at least, very different from the substance or
+jelly which bees commonly collect around their new hatched worms.
+Solicitous to learn its origin, and conjecturing that it might be the
+male prolific fluid, he began to watch the motions of every drone in the
+hive, on purpose to seize the moment when they would bedew the eggs. He
+assures us, that he saw several insinuate the posterior part of the body
+into the cells, and there deposit the fluid. After frequent repetition
+of the first, he entered on a long series of 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 no brood.
+He saw the queen lay eggs, which were bedewed by the males, and from
+which larvae were hatched, consequently, he could not hesitate advancing
+as a fact demonstrated, that male bees fecundate the queen's eggs in
+the manner of frogs and fishes, 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 number of males in a hive. At the same time,
+the author had neglected to answer one strong objection. Larvae appear
+when there are no drones. From the month of September until April, hives
+are generally destitute of males, yet, notwithstanding their absence,
+the queen then lays fertile eggs. Thus, the prolific fluid cannot be
+required to impregnate them, unless we can suppose that it is necessary
+at a certain time of the year, while at every other season it is
+useless.
+
+To discover the truth amidst these facts apparently so contradictory, I
+wished to repeat Mr Debraw's experiments, and to observe more precaution
+than he himself had done. First, I sought for the fluid, which he
+supposes the seminal, in cells containing eggs. Several were actually
+found with that appearance; and, during the first days of observation,
+neither my assistant nor myself doubted the reality of the discovery.
+But we afterwards found it an illusion arising from the reflection of
+the light, for nothing like a fluid was visible, except when the solar
+rays reached the bottom of the cells. Fragments of the coccoons of
+worms, successively hatched, commonly cover the bottom; and, as they are
+shining, it may easily be conceived that, when much illuminated, an
+illusory effect results from the light. We proved it by the strictest
+examination, for no vestiges of a fluid were perceptible when the cells
+were detached and cut asunder.
+
+Though the first observation inspired us with some distrust of Mr
+Debraw's discovery, we repeated his other experiments with the utmost
+care. On the 6. of August 1787, we immersed a hive, and, with scrupulous
+attention, examined the whole bees while in the bath. We ascertained
+that there was no male, either large or small; and having examined all
+the combs, we found neither male nymph, nor worm. When the bees were
+dry, we replaced them all, 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 necessary that no male should enter the hive during the
+experiment, a glass tube was adapted 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 removed any male that appeared,
+that the result of the experiment might be undisturbed, and I can
+positively affirm that not one was seen. However, from the first day,
+which was the sixth 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 where it was impossible one could be
+introduced, since these eggs, I say, were fertile, it becomes
+indubitable that the fluid of the males is not required for their
+exclusion.
+
+Though it did not appear that any reasonable objection could be started
+against this conclusion, yet, as I had been accustomed in all my
+experiments to seek for the most trifling difficulties that could arise,
+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 fluid to their own habitations for depositing it
+on the eggs.
+
+It was easy to appreciate the force of this objection, for all that was
+necessary was a repetition of the former experiments, and to confine the
+bees so closely to their hives that none could possibly escape. You
+very well know, Sir, that these animals can live three or four months
+confined in a hive well stored with honey and wax, and if apertures are
+left for circulation of the air. This experiment 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.
+
+I extended the precautions so far as to immerse this hive a second time,
+to assure myself that no male had escaped my researches. 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; and that was, his using queens whose
+history he was unacquainted with from their origin. When he observed
+the eggs produced by a queen, confined along with males, were fertile,
+he thence concluded that they had been bedewed by the prolific fluid in
+the cells: but to render his conclusion just, he should first have
+ascertained that the female never had copulated, and this he neglected.
+The truth is, that, without knowing it, he had used, in his experiments,
+a queen after she had commerce with the male. Had he taken a virgin
+queen the moment she came from the royal cell, and confined her along
+with drones in his vessels, the result would have been opposite; for,
+even amidst a seraglio of males, this young queen would never have laid,
+as I shall afterwards prove.
+
+The Lusatian observers, and M. Hattorf in particular, thought the queen
+was fecundated by herself, without concourse with the males. I shall
+here give an abstract of the experiment on which that opinion is
+founded.{D}
+
+M. Hattorf took a queen whose virginity he could not doubt. He excluded
+all the males both of the large and small species, and, in several days,
+he 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
+absent, the queen laid eggs, from which came worms: whence he considers
+she is impregnated by herself.
+
+Reflecting on this experiment, I do not find it sufficiently accurate.
+Males pass with great facility from hive to hive; and M. Hattorf took no
+precaution that none was introduced into his. He says, indeed, there was
+no male, but is silent respecting the means he adopted to prove the
+fact. Though he might be satisfied of no large drone being there, 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, and without greater 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. For several days I
+visited the hive, and found new hatched worms in it. Here then is the
+same result as M. Hattorf obtained? But before deducing the same
+consequence from it, we had to ascertain beyond dispute that no male had
+entered the hive. Thus, it was necessary to immerse the bees, and
+examine each separately. By this operation, we actually found four small
+males. Therefore, to render the experiment decisive, not only was it
+requisite to remove all the drones, but also, by some infallible method,
+to prevent any from being introduced, which the German naturalist had
+neglected.
+
+I prepared to repair this omission, by putting a virgin queen into a
+hive, from which the whole males were carefully removed; and to be
+physically certain that none should enter, a glass tube was adapted at
+the entrance of such dimensions that the working bees could freely pass
+and repass, but too narrow for the smallest male. Matters continued thus
+for thirty days, the workers departing and returning performed their
+usual labours: but the queen remained sterile. At the expiration of this
+time, her belly was equally slender as at the moment of her origin. I
+repeated the experiment several times, and always with the same
+consequence.
+
+Therefore, as a queen, rigorously separated from all commerce with the
+male, remains sterile, it is evident she cannot impregnate herself, and
+M. 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 acquired the
+knowledge of new facts, but these were apparently so contradictory as to
+render the solution of the problem still more difficult. While
+examining Mr. Debraw's hypothesis, I confined a queen in a hive, from
+which all the drones were removed; the queen nevertheless was fertile.
+When considering the opinion of M. Hattorf on the contrary, I put a
+queen, of whose virginity I was perfectly satisfied, in the same
+situation, she remained sterile.
+
+Embarrassed by so many difficulties, I was on the point of abandoning
+the subject of my researches, when at length by more attentive
+reflection, I thought these contradictions might arise from experiments
+made indifferently on virgin queens, and on those with whose history I
+was not acquainted from the origin, and which had perhaps been
+impregnated unknown to me. Impressed with this idea, I undertook a new
+method of observation not on queens fortuitously taken from the hive,
+but on females decidedly in a virgin state, and whose history I knew
+from the instant they left the cell.
+
+From a very great number of hives, I removed all the virgin females,
+and substituted for each a queen taken at the moment of her birth. The
+hives were then divided into two classes. From the first, I took the
+whole males both large and small, and adapted a glass tube at the
+entrance, so narrow, 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 all the drones belonging to them, and even introduced more; and to
+prevent them from escaping, a glass tube, also too narrow for the males,
+was adapted to the entrance of these hives.
+
+For more than a month, I carefully watched this experiment, made on a
+large scale; but much to my surprise, all the queens remained 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 necessary 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 29. June 1788.
+
+Aware, that in summer the males usually leave the hive at the warmest
+time of the day, it was natural for me to conclude that if the queens
+were also obliged to go out for impregnation, instinct would induce them
+to do so at the same time as the males.
+
+At eleven in the forenoon, we placed ourselves opposite a hive
+containing an unimpregnated queen five days old. The sun had shone from
+his rising; the air was very warm; and the males began to leave the
+hives. We then enlarged the entrance of that which we wished to observe,
+and paid great attention to the bees that entered and departed. The
+males appeared, and immediately took flight. Soon afterwards, the young
+queen appeared at the entrance; at first she did not fly, but brushed
+her belly with her hind legs, and traversed the board a little; neither
+workers nor males paid any attention to 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 departure, perhaps judging this precaution
+necessary to recognize it; she then flew away, describing horizontal
+circles twelve or fifteen feet above the earth. We contracted the
+entrance 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 observe all her motions. But she did not remain
+long in a situation favourable for us, and rapidly rose out of sight. We
+resumed our place before the hive; and in seven minutes, the young queen
+returned to the entrance of a habitation which she had left for the
+first time. Having found no external appearance of fecundation, we
+allowed her to enter. 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. Her second
+absence was much longer than the first; twenty-seven minutes elapsed
+before she came back. We then found her in a state very different from
+that in which she was after her first excursion. The sexual organs were
+distended by a white substance, thick and hard, very much resembling the
+fluid in the vessels of the male, completely similar to it indeed in
+colour and consistence{E}.
+
+But more evidence than mere resemblance was requisite to establish that
+the female had returned with the prolific fluid of the males. We allowed
+this queen to enter the hive, and confined her there. In two days, we
+found her belly swoln; and she had already laid near an hundred eggs in
+the worker's cells.
+
+To confirm our discovery, we made several other experiments, and with
+the same success. I shall continue to transcribe my journal.
+
+On the second of July, the weather being very fine, numbers of males
+left the hives. We set at liberty an unimpregnated young queen, eleven
+days old, whose hive had always been deprived of males. Having quickly
+left the hive, she returned to examine it, and then rose out of sight.
+In a few minutes, she returned without any external marks of
+impregnation. In a quarter of an hour, she departed again, but her
+flight was so rapid that we could scarcely follow her a moment. This
+absence continued thirty minutes. On returning, the last ring of the
+body was open, and the sexual organs full of the whitish substance
+already mentioned. She was then replaced in the hive from which all the
+males were excluded. In two days, we found her impregnated.
+
+These observations at length demonstrate why M. Hattorf obtained results
+so different from ours. His queens, though in hives deprived of males,
+had been fecundated, and he thence concludes that sexual intercourse is
+not requisite for their impregnation. But he did not confine the queens
+to their hives, and they had profited by their liberty to unite with the
+males. We, on the contrary, have surrounded our queens with a number of
+males; but they continued sterile; because the precaution of confining
+the males to their hives had also prevented the queens from departing to
+seek that fecundation without, which they could not obtain within.
+
+These experiments were repeated on queens, twenty, twenty-five, and
+thirty days old. All became fertile after a single impregnation;
+however, we have remarked some essential peculiarities in the fecundity
+of those unimpregnated until the twentieth day of their existence; but
+we shall defer speaking of the fact until we can present naturalists
+with observations sufficiently secure and numerous to merit their
+attention: Yet let me add a few words more. Though neither my assistant
+nor myself have witnessed the copulation of a queen and a drone, we
+think that, after the detail which has just been commenced, no doubt of
+it can remain, or of the necessity of copulation to effect impregnation.
+The sequel of experiments, made with every possible precaution, appears
+demonstrative. The uniform sterility of queens in hives wanting males,
+and in those where they were confined along with them; the departure of
+these queens from the hives; and the very conspicuous evidence of
+impregnation with which they return, 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 copulation.
+
+Naturalists have always been very much embarrassed to account for the
+number of males found 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 that extent. As fecundation
+cannot be accomplished within, and as the queen is obliged to traverse
+the expanse of the atmosphere, 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 drones in each hive, there would be little probability
+of their departure at the same instant with the queen, or that they
+would meet in their excursions; and most of the females would thus
+remain sterile.
+
+But why has nature prohibited copulation within the hives? This is a
+secret still unknown to us. It is possible, however, that some
+favourable circumstance may enable us to penetrate it in the course of
+our observations. Various conjectures may be formed; but at this day we
+require facts, and reject gratuitous suppositions. It should be
+remembered, that bees do not form the sole republic among insects
+presenting a similar phenomenon; female ants are also obliged to leave
+the ant-hills previous to fecundation.
+
+I cannot request, Sir, that you will communicate the reflections which
+your genius will excite concerning the facts I have related. This is a
+favour to which I am not yet entitled. But as new experiments will
+unquestionably occur to you, whether on the impregnation 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 encouragement that the
+continuance of my labours can receive.
+
+  _Pregny, 13th August 1789._
+
+       *       *       *       *       *
+
+_Letter from M. Bonnet to M. Huber._
+
+You have most agreeably surprised me, Sir, with your interesting
+discovery of the impregnation of the queen bee. It was a fortunate idea,
+that she left the hive to be fecundated, and your method of ascertaining
+the fact was extremely judicious and well adapted to the object in view.
+
+Let me remind you, that male and female ants copulate in the air; and
+that after impregnation the females return to the ant hills to deposit
+their eggs. _Contemplation de la Nature, Part 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 a copulation in the air. If you have satisfactory evidence
+that the fluid bedewing the last rings of the female is the same with
+that of the male, it is more than mere presumption in favour of
+copulation. Perhaps it may be necessary that the male should seize the
+female under the belly, which cannot easily be done but in the air. The
+large opening at the extremity of the queen, which you have observed in
+so particular a condition, seems to correspond to the singular size of
+the sexual parts of the male.
+
+You wish, my dear Sir, that I should suggest some new experiments on
+these industrious republicans. In doing so, I shall take the greater
+pleasure and interest, as I know to what extent you possess the valuable
+art of combining ideas, and of deducing from this combination 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
+queen, by introducing a little of the male's prolific fluid with a
+pencil, and at the same time observing every precaution to avoid error.
+Artificial fecundation, you are aware, has already succeeded in 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 paint the thorax with some varnish that resists humidity. It will
+also be right to paint the thorax of a considerable number of workers in
+order to discover the duration of their life. This is a more secure
+method than slight mutilations.
+
+For hatching the worm, the egg must be fixed almost vertically by one
+end near the bottom of the cell. Is it true, that it is unproductive
+unless fixed in this manner? I cannot determine the fact; and therefore
+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 queens in the cells, and my
+inclination to suppose them minute worms not yet begun to expand. Their
+elongated figure seems to favour my suspicions. It would therefore be
+proper to watch them with the utmost assiduity, from the instant of
+production until the period of exclusion. If the integument bursts,
+there can be no doubt that these minute substances are real eggs.
+
+I return to the mode of operating copulation. The height that the queen
+and the males rise to in the air prevent us from seeing what passes
+between them. On that account, the hive should be put into an apartment
+with a very lofty ceiling. M. de Reaumur's experiment of confining a
+queen with several males in a glass vessel, merits repetition; and if,
+instead of a vessel, a glass tube, some inches in diameter and several
+feet long, were used, perhaps something satisfactory might be
+discovered.
+
+You have had the fortune to observe the small queens mentioned by the
+Abbe Needham, but which he never saw. It will be of great importance to
+dissect them for the purpose of finding their ovaries. When M. Reims
+informed me that he had confined three hundred workers, along with a
+comb containing no eggs, and afterwards found hundreds in it, I strongly
+recommended that he should dissect the workers. He did so; and informed
+me that eggs were found in three. Probably without being aware of it, he
+has dissected small queens. As small drones exist, it is not surprising
+if small queens are produced also, and undoubtedly by the same external
+causes.
+
+It is of much consequence to be intimately acquainted with this species
+of queens, for they may have great influence on different experiments
+and embarrass the observer: we should ascertain whether they inhabit
+pyramidal cells smaller than the common, or hexagonal ones.
+
+M. Schirach's famous experiment on the supposed conversion of a common
+worm into a royal one, cannot be too often repeated, though the Lusatian
+observers have already done it frequently. I could wish to learn
+whether, as the discoverer maintains, the experiment will succeed only
+with worms, three or four days old, and never with simple eggs.
+
+The Lusatian observers, and those of the Palatinate, affirm, that when
+common bees are confined with combs absolutely void of eggs, they then
+lay none but the eggs of drones. Thus, there must 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 ovaries contain male eggs alone?
+
+According to M. de Reaumur, the life of chrysalids may be prolonged by
+keeping them in a cold situation, such as an ice-house. The same
+experiment should be made on the eggs of a queen; on the nymphs of
+drones and workers.
+
+Another interesting experiment would be to take away all the combs
+composing the common cells, and leave 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.
+Nevertheless, it is likely, by taking away only part of the common
+cells, the workers may be forced to lay common eggs in the cells of
+drones.
+
+I should also wish to have the young larvae 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 influence 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
+adopted to instruct us how bees can regulate their labours, and apply
+them to existing circumstances. This may enable us to discover
+particular facts which we cannot foresee.
+
+The royal eggs and those producing drones, have not yet been carefully
+compared with the eggs from which workers come. But they ought to be so,
+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 endeavour, with the point of
+a pencil, to remove 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 this experiment.
+
+Various facts, which require corroboration, were collected in my Memoirs
+on Bees; of this number are my own observations. You can select what is
+proper, my dear Sir. 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. _Genthod, 18. August 1789._
+
+
+FOOTNOTES:
+
+{A} All these letters are addressed to the celebrated naturalist M.
+Bonnet.--_T._
+
+{B} The leaf or book hive consists of twelve vertical frames or boxes,
+parallel to each other, and joined together. Fig. 1. the sides, f f. f
+g. should be twelve inches long, and the cross spars, f f. g g. nine or
+ten; the thickness of these spars an inch, and their breadth fifteen
+lines. It is necessary that this last measure should be accurate; a a. a
+piece of comb which guides the bees in their work; d. a moveable slider
+supporting the lower part; b b. pegs to keep the comb properly in the
+frame or box; four are in the opposite side; e e. pegs in the sides
+under the moveable slider to support it.
+
+A book hive, consisting of twelve frames, all numbered, is represented
+fig. 2. Between 6 and 7 are two cases with lids, that divide the hive
+into two equal parts, and should only be used to separate the bees for
+forming an artificial swarm; a a. two frames which shut up the two sides
+of the hive, have sliders, b. b.
+
+The entrance appears at the bottom of each frame. All should be close
+but 1 and 12. However it is necessary that they should open at pleasure.
+
+The hive is partly open, fig. 3. and shews how the component parts may
+be united by hinges, and open as the leaves of a book. The two covers
+closing up the sides, a. a.
+
+Fig. 4. is another view of fig. 1. a a. a piece of comb to guide the
+bees; b b. pegs disposed so as to retain the comb properly in the frame;
+c c. parts of two shelves; the one above is fixed, and keeps the comb in
+a vertical position; the under one, which is moveable, supports it
+below.
+
+{C} I cannot insist that my readers, the better to comprehend what is
+here said, shall peruse the Memoirs of M. de Reaumur on Bees, and those
+of the Lusace Society; but I must request them to examine the extracts
+in M. Bonnet's works, tom. 5. 4to edit. and tom. 10. 8vo, where they
+will find a short and distinct abstract of all that naturalists have
+hitherto discovered on the subject.
+
+{D} Vide M. Schirach's History of Bees, in a memoir by M. Hattorf,
+entitled, _Physical Researches whether the Queen Bee requires
+fecundation by Drones?_
+
+{E} It will afterwards appear that what we took for the generative
+fluid, was the male organs of generation, left by copulation in the body
+of the female. This discovery we owe to a circumstance that shall
+immediately be related. Perhaps I should avoid prolixity, by suppressing
+all my first observations on the impregnation of the queen, and by
+passing directly to the experiments 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 think service is
+done to the reader by a candid avowal of my errors. This is an
+additional proof to so many others, of the absolute necessity that an
+observer should repeat all his experiments a thousand times, to obtain
+the certainty of seeing facts as they really exist.
+
+
+
+
+LETTER II.
+
+_SEQUEL OF OBSERVATIONS ON THE IMPREGNATION OF THE QUEEN BEE._
+
+
+  SIR,
+
+All the experiments, related in my preceding letter, were made in 1787
+and 1788. They seem to establish two facts, which had previously been
+the subject of vague conjecture: 1. The queen bee is not impregnated of
+herself, but is fecundated by copulation with the male. 2. Copulation is
+accomplished without the hive, and in the air.
+
+The latter appeared so extraordinary, that notwithstanding all the
+evidence obtained of it, we eagerly desired to take the queen in the
+fact; but, as she always rises to a great height, we never could see
+what passed. On that account you advised us to cut part off the wings of
+virgin queens. We endeavoured to benefit by your advice, in every
+possible manner; but to our great regret, when the wings lost much, the
+bees could no longer fly; and, by cutting off only an inconsiderable
+portion, we did not diminish the rapidity of their flight. Probably
+there is a medium, but we were unable to attain it. On your suggestion,
+we tried to render their vision less acute, by covering the eyes with an
+opaque varnish, which was an experiment equally fruitless.
+
+We likewise attempted artificial fecundation, and took every possible
+precaution to insure success. Yet the result was always unsatisfactory.
+Several queens were the victims of our curiosity; and those surviving
+remained sterile. Though these different experiments were unsuccessful,
+it was proved that queens leave their hives to seek the males, and that
+they return with undoubted evidence of fecundation. Satisfied with this,
+we could only trust to time or accident for decisive proof of an actual
+copulation. We were far from suspecting a most singular discovery, which
+we made in July this year, and which affords complete demonstration of
+the supposed event, namely, that the sexual organs of the male remain
+with the female.{F}
+
+
+FOOTNOTES:
+
+{F} The remainder of this Chapter chiefly consists of anatomical
+details. These may rather be considered an interruption of the
+narrative; and the Translator has judged it expedient to transfer them
+to an Appendix.
+
+
+
+
+LETTER III.
+
+_THE SAME SUBJECT CONTINUED.--OBSERVATIONS ON RETARDING THE FECUNDATION
+OF QUEENS._
+
+
+In my first letter, I remarked, that when queens were prevented from
+receiving the approaches of the male until the twenty-fifth or thirtieth
+day of their existence, the result presented very interesting
+peculiarities. My experiments at that time were not sufficiently
+numerous; but they have since been so often repeated, and the result so
+uniform, that I no longer hesitate to announce, as a certain discovery,
+the singularities which retarded fecundation, produces on the ovaries of
+the queen. If she receives the male during the first fifteen days of her
+life, she remains capable of laying both the eggs of workers and of
+drones; but should fecundation be retarded until the twenty-second day,
+her ovaries are vitiated in such a manner that she becomes unfit for
+laying the eggs of workers, and will produce only those of drones.
+
+In June 1787, being occupied in researches 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. When a hive is ready to swarm, I had before
+observed, that the moment of swarming is always preceded by a very
+lively agitation, which first affects the queen, is then communicated to
+the workers, and excites such a tumult among them, that they abandon
+their labours, and rush in disorder to the outlets of the hive. I then
+knew very well the cause of the queen's agitation, and it is described
+in the history of swarms, but I was ignorant how the delirium
+communicated to the workers; and this difficulty interrupted my
+researches. I therefore thought of investigating, by direct experiments,
+whether at all times, when the queen was greatly agitated, even not in
+the time of the hive swarming, her agitation would in like manner be
+communicated to the workers. The moment a queen 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 it
+would produce a kind of delirium. I had the patience to observe this
+queen thirty-four days. Every morning about eleven o'clock, when the
+weather was fine and the sunshine invited the males to leave their
+hives, I saw her impetuously traverse every corner of her habitation,
+seeking to escape. Her fruitless efforts threw her into an uncommon
+agitation, the symptoms of which I shall elsewhere describe, and all the
+common bees were affected by it. As she never was out all this time, she
+could not be impregnated. At length, on the thirty-sixth day, I 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 this experiment, I was far from
+any hopes that it would lead to the knowledge of another very remarkable
+fact; how great was my astonishment, therefore, on finding that this
+female, which, as usual, began to lay forty-six hours after copulation,
+laid the eggs of drones, but none of workers, and that she continued
+ever afterwards to lay those of drones only.
+
+At first, I exhausted myself with conjectures on this singular fact; the
+more I reflected on it, the more did it seem inexplicable. At length, by
+attentively meditating on the circumstances of the experiment it
+appeared there were two principles, the influence of which I should
+first of all endeavour to appreciate separately. On the one hand, this
+queen had suffered long confinement; on the other, her fecundation had
+been extremely retarded. You know, Sir, that queens generally receive
+the males about the fifth or sixth day, and this queen had not copulated
+until the thirty-sixth. Little weight could be given to the supposition,
+that the peculiarity could be occasioned by confinement. Queens, in the
+natural state, leave their hives only once to seek the males. All the
+rest of their life they remain voluntary prisoners. Thus, it was
+improbable that captivity could produce the effect I wished to explain.
+At the same time, as it was essential to neglect nothing in a subject so
+new, I wished to ascertain whether it was owing to the length of
+confinement, or to retarded fecundation.
+
+Investigating this was no easy matter. To discover whether captivity,
+and not retarded fecundation, vitiated the ovaries, it was necessary to
+allow a female to receive the approaches of a male, and also to keep her
+imprisoned. Now this could not be, for bees never copulate in hives. On
+the same account, it was impossible to retard the copulation of a queen
+without keeping her in confinement. I was long embarrassed by the
+difficulty. At length, I contrived an apparatus, which, though
+imperfect, nearly fulfilled my purpose.
+
+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 allowed free passage 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 the queen could at all times come and fly about, enjoying a
+purer air than was to be 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 between them. By the
+experiments related in my first letter, copulation takes place high in
+the air only: therefore, in this apparatus, I found the advantage of
+retarding fecundation, while the liberty the queen now had, did not
+render her situation too remote from the natural state. I attended to
+the experiment fifteen days. Every fine morning, the young captive left
+her hive; 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 aloft in the air, and soon returned with full evidence of
+impregnation. In two days, she laid, first the eggs of workers, and
+afterwards as many as the most fertile queens.
+
+It thence followed, 1. That captivity did not alter the organs of
+queens. 2. When fecundation took place within the first sixteen days,
+she produced both species of eggs.
+
+This was an important experiment. It rendered my labours much more
+simple, by clearly pointing out the method to be pursued: it absolutely
+precluded the supposed influence of captivity; and left nothing for
+investigation but the consequences of retarded fecundation.
+
+With this view, I repeated the experiment; but, instead of giving the
+virgin queen liberty on the sixteenth day, I retained her until the
+twenty-first. She departed, rose high in the air, was fecundated, and
+returned. Thirty-six hours afterwards, she began to lay: but it was the
+eggs of males only, and, although very fruitful afterwards, she laid no
+other kind.
+
+I occupied myself the remainder of 1787, and the two subsequent years,
+with experiments on retarded fecundation, and had constantly the same
+results. It is undoubted, therefore, that when the copulation of queens
+is retarded beyond the twentieth day, only an imperfect impregnation is
+operated: instead of laying the eggs of workers and males equally, they
+will lay none but those of males.
+
+I do not aspire to the honour of explaining this singular fact. When the
+course of my experiments led me to observe 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
+fecundation. My evidence is demonstrative, for I can always prevent
+queens from laying the eggs of workers, by retarding their fecundation
+until the twenty-second or twenty-third day. But, what is the remote
+cause of this peculiarity; or, in other words, why does the delay of
+impregnation render queens incapable of laying 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 delayed. The
+queen then lays the eggs of workers forty-six hours after copulation,
+and continues for the subsequent eleven months to lay these alone: and
+it is only after this period that a considerable and uninterrupted
+laying of the eggs of drones commences. When, on the contrary,
+impregnation is retarded after the twentieth day, the queen begins, from
+the forty-sixth hour, to lay the eggs of males, and no other kind during
+her whole life. As, in the natural state, she lays the eggs of workers
+only, during the first eleven months, it is clear that these, and the
+male eggs, are not indiscriminately mixed in the oviducts. Undoubtedly
+they occupy a situation corresponding to the principles that regulate
+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 inverted by retarded copulation? How does it happen
+that all the workers eggs which the queen ought to lay, if fecundation
+was in due time, now wither and disappear, yet do not, impede the
+passage of the eggs of drones, which occupy only the second place in the
+ovaries. Nor is this all. I have satisfied myself that a single
+copulation 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 a single copulation will impregnate all the eggs that she will lay
+during her whole life: but I want absolute proof for more than two
+years. This, which is truly a very singular fact in itself, renders the
+influence of retarded fecundation still more difficult to be accounted
+for. Since a single copulation suffices, it is clear that the male fluid
+acts from the first moment on all the eggs that the queen will lay in
+two years. It gives them, according to your principles, that degree of
+_animation_ that afterwards effects their successive expansion. Having
+received the first impressions of life, they grow, they mature, so to
+speak, until the day they are laid: and as the laws of laying are
+constant, because the eggs of the first eleven months are always those
+of workers, it is evident that those which appear first are also the
+eggs that come soonest to maturity. Thus, in the natural state, the
+space of eleven months is necessary for the male eggs to acquire that
+degree of increment they must have attained when laid. This consequence,
+which to me seems immediate, renders the problem insoluble. How can the
+eggs, which should grow slowly for eleven months, suddenly acquire their
+full expansion in forty-eight hours, when fecundation has been retarded
+twenty-one days, and by the effect of this retardation alone? Observe, I
+beseech you, that the hypothesis of successive expansion is not
+gratuitous; it rests on the principles of sound philosophy. Besides, for
+conviction that it is well founded, we have only to look at the figures
+given by Swammerdam of the ovaries of the queen bee. There we see eggs
+in that part of the oviducts contiguous to the vulva, much farther
+advanced, and larger than those contained in the opposite part.
+Therefore the difficulty remains in full force: it is an abyss where I
+am lost.
+
+The only known fact bearing any relation to that now described, is the
+state of certain vegetable seeds, which, although extremely well
+preserved, lose the faculty of germination from age. The eggs of workers
+may also preserve, only for a very short time, the property of being
+fecundated by the seminal fluid; and, after this period, which is about
+fifteen or eighteen days, become disorganised to that degree, that they
+can no longer be animated by it. I am sensible that the comparison is
+very imperfect; besides, it explains nothing, nor does it even put us
+on the way of making any new experiments. I shall add but one reflection
+more.
+
+Hitherto no other effect has been observed from the retarded
+impregnation of animals, but that of rendering them absolutely sterile.
+The first instance of a female still preserving the faculty of
+engendering males, is presented by the queen bee. But as no fact in
+nature is unique, it is most probable that the same peculiarity will
+also be found in other animals. An extremely curious object of research
+would be to consider insects in this new point of view, I say _insects_,
+for I do not conceive that any thing analogous will be found in other
+species of animals. The experiments now suggested would necessarily
+begin with insects the most analogous to bees; as wasps, humble bees,
+mason bees, all species of flies, and the like. Some experiments might
+also be made on butterflies; and, perhaps, an animal might be found
+whose retarded fecundation would be attended with the same effects as
+that of queen bees. Should the animal be larger, dissection will be more
+easily accomplished; and we may discover what happens to the eggs when
+retarded fecundation prevents their expansion. At least, we might hope
+that some fortunate circumstance would lead to solution of the
+problem{G}.
+
+Let us now return to my experiments. In May 1789, I took two queens just
+when they had undergone the last metamorphosis: one was put in a _leaf
+hive_, well provided with honey and wax, and sufficiently inhabited by
+workers and males. The other was put 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, but the common
+bees enjoyed perfect liberty. The queens were imprisoned thirty days;
+and being then set at liberty, they departed, and returned impregnated.
+Visiting the hives in the beginning of July, I found much brood, but
+wholly consisting of the worms and nymphs of males. There actually was
+not a single worker's worm or nymph. Both queens laid uninterruptedly
+until autumn, and constantly the eggs of drones. Their laying ended in
+the first week of November, as that of my other queens.
+
+I was very earnest to learn what would become of them in the subsequent
+spring, whether they would resume laying, or if new fecundation would be
+necessary; and if they did lay, of what species the eggs would be.
+However, the hives being very weak, I dreaded they might perish during
+winter. Fortunately, we were able to preserve them; and from April 1790,
+they recommenced laying. The precautions we had taken prevented them
+from receiving any new approaches of the male. Their eggs were still
+those of males.
+
+It would have been extremely interesting to have followed the history of
+these two females still farther, but, to my great regret, the workers
+abandoned their hives on the fourth of May, and that same day I found
+both queens dead. No weevils were in the hive, which could disturb the
+bees; and the honey was still very plentiful: but as no workers had been
+been produced in the course of the preceding year, and winter had
+destroyed many, they were too few in spring to engage in their wonted
+labours, and, from discouragement, deserted their habitation to occupy
+the neighbouring hives.
+
+In my Journal, I find a detail of many experiments on the retarded
+impregnation of queen bees, so many, that transcribing the whole would
+be tedious. I may repeat, however, that there was not the least
+variation in the principle, and that whenever the copulation of queens
+was postponed beyond the twenty-first day, the eggs of males only were
+produced. Therefore, I shall limit my narrative to those experiments
+that have taught me some remarkable facts.
+
+A queen being hatched on the fourth of October 1789, we put her into a
+leaf-hive. Though the season was well advanced, a considerable number of
+males was still in the hive; and it here became important to learn,
+whether, at this period of the year, they could equally effect
+fecundation; also, in case it succeeded, whether a laying, begun in the
+middle of autumn, would be interrupted or continued during winter. Thus,
+we allowed the queen to leave the hive. She departed, indeed, but made
+four and twenty fruitless attempts before returning with the evidence of
+fecundation. Finally, on the thirty-first of October, she was more
+fortunate: She departed, and returned with the most undoubted proof of
+the success of her amours: She was now twenty-seven days old,
+consequently fecundation had been retarded. She ought to have begun
+laying within forty-six hours, but the weather was cold, and she did not
+lay; which proves, as we may cursorily remark, that refrigeration of the
+atmosphere is the principal agent that suspends the laying of queens
+during winter. I was excessively impatient to learn whether, on the
+return of spring, she would prove fertile, without a new copulation. The
+means of ascertaining the fact was easy; for the entrances of the hives
+only required contraction, so as to prevent her from escaping. She was
+confined from the end of October until May. In the middle of March, we
+visited the combs, and found a considerable number of eggs, but, none
+being yet hatched, we could not know whether they would produce workers
+or males. On the fourth of April, having again examined the state of the
+hive, we found a prodigious quantity of nymphs and worms, all of
+drones; nor had this queen laid a single worker's egg.
+
+Here, as well as in the preceding experiment, 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
+copulation 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
+begin to lay in spring without requiring new copulation.
+
+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, already flies, two thousand four hundred and
+thirty-eight cells, containing either eggs or nymphs of drones. Thus,
+she had laid more than three thousand male eggs during March and April,
+which is above fifty each day. Her death soon afterwards unfortunately
+interrupted my observation, I intended to calculate the total number of
+male eggs that she should lay throughout the year, and compare it with
+those 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 considerable, commences in August, also in the
+interval, that they produce the eggs of workers almost solely. But it is
+otherwise with the females whose copulation has been retarded: they
+produce no workers' eggs. For four or five months following, they lay
+the eggs of males without interruption, and in such numbers, that, in
+this short time, I suppose one queen gives birth to more drones than a
+female, whose fecundation has not been retarded, produces in the course
+of two years. It gives me much regret, that I have not been able to
+verify this conjecture.
+
+I should also describe the very remarkable manner in which queens, that
+lay only the eggs of drones, sometimes deposit them in the cells.
+Instead of being placed in the lozenges forming the bottom, they are
+frequently deposited on the lower side of the cells, two lines from the
+mouth. This arises from the body of such queens being shorter than that
+of those whose fecundation has not been retarded. The extremity remains
+slender, while the first two rings next the thorax are uncommonly swoln.
+Thus, in disposing themselves for laying, the extremity cannot reach the
+bottom of the cells on account of the swoln rings; consequently the eggs
+must remain attached to the part that the extremity reaches. The worms
+proceeding from them pass their vermicular state in the same place where
+the eggs were deposited, which proves that bees are not charged with the
+care of transporting the eggs as has been supposed. But here they
+follow another plan. They extend beyond the surface of the comb those
+cells where they observe the eggs deposited, two lines from the mouth.
+
+Permit me, Sir, to digress a moment from the subject, to give the result
+of an experiment which seems interesting. Bees, I say, are not charged
+with the care of transporting into cells, the eggs misplaced by the
+queen: and, judging by the single instance I have related, you will
+think me well entitled to deny this feature of their industry. However,
+as several authors have maintained the reverse, and even demanded our
+admiration of them in conveying the eggs, I should explain clearly that
+they are deceived.
+
+I had a glass hive constructed of two stages; the higher was filled with
+combs of large cells, and the lower with those of common ones. A kind of
+division, or diaphraghm, separated these two stages from
+each other, having at each side an opening 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 had only those of workers to lay, and
+she was obliged to deposit them in the surrounding large cells from the
+want of others. My object in this arrangement will already be
+anticipated. My reasoning was simple. If the queen laid workers' eggs in
+the large cells, and the bees were charged with transporting them if
+misplaced, they would infallibly take advantage of the liberty allowed
+to pass from either stage: they would seek the eggs deposited in the
+large cells, and carry them down to the lower stage containing the cells
+adapted for that species. If, on the contrary, they left the common eggs
+in the large cells, I should obtain certain proof that they had not the
+charge of transporting them.
+
+The result of this experiment excited my curiosity extremely. We
+observed the queen several days without intermission. During the first
+twenty-four hours, she persisted in not laying a single egg in the
+surrounding cells; she examined them one after another, but passed on
+without insinuating her belly into one. She was restless, and traversed
+the combs in all directions: her eggs appeared an oppressive burden, but
+she persisted 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 mother. I was amused to observe, when she
+approached the edges of the division separating the two stages, that she
+gnawed at them to enlarge the passage: the workers approached her, and
+also laboured with their teeth, and made every exertion to enlarge the
+entrance 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. Then we conceived that the bees would convey them into the
+small cells of the lower stage, and we sought them there with the utmost
+assiduity; but I can safely affirm there was not one. The eggs that the
+queen still laid the third day disappeared as the first. We again sought
+them in the small cells, but none were there. The fact is, they are ate
+by the workers; and this is what has deceived the naturalists, who
+supposed them carried away. They have observed the misplaced eggs
+disappear, and, without farther investigation, have asserted that the
+bees convey them elsewhere: they take them, indeed, not to convey them
+any where, but to devour them. Thus nature has not charged bees with the
+care of placing the eggs in the cells appropriated for them, but she has
+inspired females themselves with sufficient instinct to know the species
+of eggs they are about to lay, and to deposit them in suitable cells.
+This has already been observed by M. de Reaumur, and here my
+observations correspond with his. Thus it is certain that in the natural
+state, when fecundation takes place at the proper time, and the queen
+has suffered from nothing, she is never deceived in the choice of the
+cells where her eggs are to be deposited; she never fails to lay those
+of workers in small cells, and those of males in large ones. The
+distinction is important, for the same certainty of instinct is no
+longer conspicuous in the conduct of those females whose impregnation
+has been deferred. I was oftener than once deceived respecting the eggs
+that such queens laid, for they were deposited indiscriminately in small
+cells and those of drones; and not aware of their instinct having
+suffered, I conceived that the eggs in small cells would produce
+workers; therefore I was very much surprised, when, at the moment they
+should have been hatched, the bees closed up the cells, and
+demonstrated, by anticipation, 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 experiments on queens that lay only the eggs of males, not to
+be deceived by these circumstances, and expect that eggs of males will
+be deposited in the workers cells.
+
+It is a singular fact, that the females, whose fecundation has been
+retarded, sometimes lay the eggs of males in royal cells. I shall prove,
+in the history of swarms, that immediately when queens, in the natural
+state, begin their great laying of male eggs, the workers construct
+numerous royal cells. Undoubtedly, there is some secret relation between
+the appearance of male eggs and the construction of these cells; for it
+is a law of nature from which bees never derogate. It is not surprising,
+therefore, that such cells are constructed in hives governed by queens
+laying the eggs 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 take exactly the same care of the male eggs
+deposited in royal cells, as of those that should become queens. They
+provide them more plentifully with food, they build up the cells as if
+containing a royal worm; in a word, they labour with such regularity
+that we have frequently been deceived. More than once, in the firm
+persuasion of finding royal nymphs, we have opened the cells after they
+were sealed, yet the nymph of a drone always appeared. Here the instinct
+of the workers seemed defective. In the natural state, they can
+accurately distinguish the male worms from those of common bees, as they
+never fail giving 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 to investigate the instinct 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 de la Reine des Abeilles_, translated from the
+German by _Blassiere_, there is printed a letter from M. Schirach to
+you, dated 15 April 1771, where he speaks of some hives, in which the
+whole brood changed into drones. You will remember that he ascribes this
+circumstance to some unknown vice in the ovaries of the queen; but he
+was far from suspecting that retarded fecundation had been the cause of
+vitiation. He justly felicitated himself on discovering a method to
+prevent the destruction of hives in this situation, which was simple,
+for it consisted in removing the queen that laid the eggs of males only,
+and substituting one for her whose ovaries were not impaired. But to
+make the substitution effectual, it was necessary to procure queens at
+pleasure; a secret reserved for M. Schirach, and of which I shall speak
+in the following letter. You observe that the whole experiments of the
+German naturalist tended to the preservation of the hives whose queens
+laid none except male eggs; and that he did not attempt to discover the
+cause of the vice evident in their ovaries.
+
+M. de Reaumur also says a few words, somewhere, of a hive containing
+many more drones than workers, but advances no conjectures on the cause.
+However, he adds, as a remarkable circumstance, that the males were
+tolerated in this hive until the subsequent spring. It is true that bees
+governed 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. I can ascribe no reason for it, but it is a fact I have
+several times witnessed during my long course of observations on
+retarded impregnation. In general it has appeared that while the queen
+lays male eggs, bees do not massacre the males already perfect in the
+hive. PREGNY, _21. August 1791_.
+
+
+FOOTNOTES:
+
+{G} The experiments suggested in this paragraph, recall a singular
+reflection of M. de Reaumur. Where treating of oviparous flies, he says,
+it would not be impossible for a hen to produce a living chicken, if,
+after fecundation, the eggs she should first lay could by any means be
+retained twenty-one days in the oviducts. _Mem. sur. les Insect. tom. 4.
+mem. 10._
+
+
+
+
+LETTER IV.
+
+_ON M. SCHIRACH'S DISCOVERY._
+
+
+When you found it necessary, Sir, in the new edition of your works, to
+give an account of M. Schirach'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
+testimonies. For this reason, I hasten to inform you that all my
+researches establish the reality of the discovery. During ten years
+that I have studied bees, I have repeated M. Schirach's experiment so
+often, and with such uniform success, that I can no longer have the
+least doubt on the subject. Therefore, I consider it an established
+fact, when bees lose their queen, and 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, but a greater quantity of it,
+and 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 a fact, and the philosophical consequences
+you have deduced from it. _Contemplation de la Nature, part. II, chap.
+27._
+
+In this letter I shall content myself with some account of the figure of
+the royal cells constructed by bees around those worms that are destined
+for the royal state, and terminate with discussing some points wherein
+my observations differ from those of M. Schirach.
+
+Bees soon become sensible of having lost their queen, and in a few hours
+commence the labour necessary to repair their loss. First, they select
+the young common worms, which the requisite treatment is to convert into
+queens, and immediately begin with enlarging the cells where they are
+deposited. Their mode of proceeding is curious; and the 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.
+Having chosen a worm, they sacrifice three of the contiguous cells:
+next, they supply it with food, and raise a cylindrical inclosure
+around, by which the cell becomes a perfect tube, with a rhomboidal
+bottom; for the parts forming the bottom are left untouched. If the bees
+damaged it, they would lay open three corresponding cells on the
+opposite surface of the comb, and, consequently, destroy their worms,
+which would be an unnecessary sacrifice, and Nature has opposed it.
+Therefore, leaving the bottom rhomboidal, they are satisfied with
+raising a cylindrical tube around the worm, which, like the other cells
+in the comb, is horizontal. But this habitation remains suitable to the
+worm called to the royal state only during the first three days of its
+existence: another situation is requisite for the other two days it is a
+worm. Then, which is so small a portion of its life, it must inhabit a
+cell nearly of a pyramidal figure, and hanging perpendicularly; we may
+say the workers know it; for, after the worm has completed the third
+day, they prepare the place to be occupied by its new lodging. They gnaw
+away the cells surrounding the cylindrical tube, mercilessly sacrifice
+their worms, and use the wax in constructing a new pyramidal tube, which
+they solder at right angles to the first, and work it downwards. The
+diameter of this pyramid decreases insensibly from the base, which is
+very wide, to the point. During the two days that it is inhabited by
+the worm, a bee constantly keeps its head more or less inserted into the
+cell, and, when this worker quits it, another comes to occupy its place.
+In proportion as the worm grows, the bees labour in extending the cell,
+and bring food, which they place before its mouth, and around its body,
+forming a kind of cord around it. The worm, which can move only in a
+spiral direction, turns incessantly to take the food before its head: it
+insensibly descends, and at length arrives at the orifice of the cell.
+Now is the time of transformation to a nymph. As any farther care is
+unnecessary, the bees close the cell with a peculiar substance
+appropriated for it, and there the worm undergoes both its
+metamorphoses.
+
+Though M. Schirach supposes that none but worms three days old are
+selected for the royal treatment, I am certain of the contrary; and that
+the operation succeeds equally well on those of two days only. I must
+be permitted to relate at length the evidence I have of the fact, which
+will both demonstrate the reality of common worms being converted into
+queens, and the little influence which their age has on the effect of
+the operation.
+
+I put some pieces of comb, with some workers eggs, in the cells, and of
+the same kind as those already hatched, into a hive deprived of the
+queen. The same day several cells were enlarged by the bees, and
+converted into royal cells, and the worms supplied with a thick bed of
+jelly. Five were then removed from those cells, and five common worms,
+which, forty-eight hours before we had seen come from the egg
+substituted for them. The bees did not seem aware of the change; they
+watched over the new worms the same as over those chosen by themselves;
+they continued enlarging the cells, and closed them at the usual time.
+When they had hatched on them seven days{H}, we removed the cells to
+see the queens that were to 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 having elapsed, and no queen appearing, we
+opened them. In one, was a dead queen, but still a nymph; the other two
+were empty. The worms had spun their silk coccoons, but died before
+passing into their nymphine state, and presented only a dry skin. I can
+conceive nothing more conclusive than this experiment. It demonstrates
+that bees have the power of converting the worms of workers into queens;
+since they succeeded in procuring queens, by operating on the worms
+which we ourselves had selected. It is equally demonstrated, that the
+success of the operation does not depend on the worms being three days
+old, as those entrusted to the bees were only two. Nor is this all; bees
+can convert worms still younger into queens. The following experiment
+showed, that when the queen is lost, they destine worms only a few
+hours old to replace her.
+
+I was in possession of a hive, which being long deprived of the female,
+had neither egg nor worm. I provided a queen of the greatest fertility;
+and she immediately began laying in the cells of workers. I removed this
+female before being quite three days in the hive, and before any of her
+eggs were hatched. The following morning, that is, 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 depositing around them a much more abundant provision
+of food than is supplied to common worms. Next day, the worms were near
+forty hours old: the bees had enlarged and converted their hexagonal
+cells into cylindrical ones of the greatest capacity. During the
+subsequent days, they still laboured at them, and closed them on the
+fifth from the origin of the worms. Seven days after sealing of the
+first of these royal cells, a queen of the largest size proceeded from
+it. She immediately rushed towards the other royal cells, and
+endeavoured to destroy their nymphs and worms. In another letter, I
+shall recount the effects of her fury.
+
+From these details, you will observe, Sir, that M. Schirach's
+experiments had not been sufficiently diversified when he affirmed that
+it was essential for the conversion of common 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 existence.
+
+After my researches to corroborate M. Schirach's discovery, I was
+desirous of learning whether, as this observer conceives, the only means
+which the bees have of procuring a queen, is giving the common worms a
+certain kind of aliment, and rearing them in the largest cells. You
+will remember, that M. de Reaumur's sentiments are very different: "The
+mother should lay, and she does lay, eggs from which flies fit for being
+mothers must in their turn proceed. She does 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 peculiar interest in the eggs that one is to
+proceed from, and to consider them of the greatest value. They construct
+particular cells where they are to be deposited.--The figure of a royal
+cell only begun, very much resembles a cup, or, more correctly speaking,
+the cup that has lost its acorn."
+
+M. de Reaumur, though he did not suspect 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 M. Schirach, on the other hand,
+bees always having the power of procuring a queen by bringing up worms
+three days old in a particular manner, it would be needless for nature
+to grant females the faculty of laying royal eggs. Such prodigality is,
+in his eyes, inconsistent with the ordinary laws of nature. Therefore he
+maintains, in direct terms, that she 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 would always be
+too long for the belly of the mother to reach the bottom.
+
+I admit that M. de Reaumur no where says he has seen the queen lay in
+the royal cell. However he did not doubt the fact; and, after all my
+observations, I must esteem his opinion just. It is quite certain that,
+at particular periods of the year, the bees prepare royal cells; that
+the females deposit their eggs in them; and that worms, which shall
+became queens, proceed from these eggs.
+
+M. Schirach's objection, concerning the length of the cells, proves
+nothing; for the queen does not delay depositing her egg till they are
+finished. While only sketched and shaped like the cup of an acorn, she
+lays it. This naturalist, dazzled by the brilliancy of his discovery,
+saw only part of the truth. He was the first to find out the resource
+granted to bees by nature, for repairing the loss of their queen; and
+too soon persuaded himself that she had provided no other resource for
+the production of females. This error arose from not observing bees in
+very flat hives: had he used such as mine, he would have found, on
+opening them in spring, a confirmation of M. de Reaumur's opinion. Then,
+which is the season of swarming, hives in good condition are governed by
+a very fruitful queen: there are royal cells of a figure widely
+different from those constructed around the worms destined by the bees
+for queens. They are large, attached to the comb by a stalk, and
+hanging vertically like stalactites, such, in short, as M. de Reaumur
+has described them. The females lay in them before completion. 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 the egg has been
+laid. In proportion as the worm grows, they are enlarged, and closed by
+the bees when the first transformation approaches. Thus it is true,
+that, in spring, the queen deposits in royal cells, previously prepared,
+eggs from which flies of her own species are to come. Nature has,
+therefore, provided a double means for the multiplication and
+conservation of their race.
+
+  _PREGNY, 24. August 1791._
+
+
+FOOTNOTES:
+
+{H} The author's meaning here is obscure.--T.
+
+
+
+
+LETTER V.
+
+_EXPERIMENTS PROVING THAT THERE ARE SOMETIMES COMMON BEES WHICH LAY
+FERTILE EGGS._
+
+
+The singular discovery of M. Riems, concerning the existence of fertile
+workers, has appeared very doubtful to you, Sir. You have suspected that
+the eggs ascribed to workers by this naturalist had actually been
+produced by small queens, which, on account of their size, were
+confounded with common bees. But you do not positively insist that M.
+Riems is deceived; and, in the letter which you did me the honour to
+address to me, you requested me to investigate, by new experiments,
+whether there are actually working bees capable of laying fertile eggs.
+I have made these experiments with great care: and it is for you to
+judge of the confidence they merit.
+
+On the fifth of August 1788, we found the eggs and worms of large drones
+in two hives, which had both been some time deprived of queens. We also
+observed the rudiments of some royal cells appended like stalactites to
+the edges of the combs. The eggs of males were in them. Being perfectly
+secure that there was no queen of large size among the bees of these two
+hives, the eggs, which daily became more numerous, were evidently laid
+either by queens of small size or by fertile workers. I had reason to
+believe it was actually by common bees, for we had frequently observed
+them inserting the posterior part into the cells; and assuming the same
+attitude as the queen when laying. But, not withstanding every exertion,
+we had never been able to seize one in this situation, 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 observations were continued with equal assiduity, in hopes that, by
+some fortunate chance, or in a moment of address, we could secure one of
+them. More than a month all our endeavours were abortive.
+
+My assistant then offered to perform an operation that required both
+courage and patience, and which I could not resolve to suggest, though
+the same expedient had occurred to myself. He proposed to examine each
+bee in the hive separately, to discover whether some small queen had not
+insinuated herself among them, and escaped our first researches. This
+was an important experiment; for, should no small queen be found, it
+would be demonstrative evidence that the eggs had been laid by simple
+workers.
+
+To perform this operation with all possible exactness, immersing the
+bees was not enough. You know, Sir, that the contact of water stiffens
+their organs, that it produces a certain alteration of their external
+figure: and, from the resemblance of small queens to workers, the
+slightest alteration of shape would prevent us from distinguishing with
+sufficient accuracy to what species those immersed might belong.
+Therefore it was necessary to seize the whole bees of both hives,
+notwithstanding their irritation, and examine their specific character
+with the utmost 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, but what the relief of
+his eyes required. He took every bee in his hand; he attentively
+examined the trunk, the hind limbs, and the sting: there was not one
+without the characteristics of the common bee, that is, the little
+basket on the hind legs, the long trunk, and the straight sting. He had
+previously prepared glass cases containing combs. Into these, he put
+each bee after examination. It is superfluous to observe they were
+confined, which was a precaution indispensible until termination of the
+experiment. Neither was it enough to establish that the whole were
+workers; we had also to continue the experiment, and observe whether any
+would produce eggs. Thus we examined the cells for several days, and
+soon observed new laid eggs, from which the worms of drones came at the
+proper time. My assistant held in his hands the bees that produced them;
+and as he was perfectly certain they were common ones, it is proved that
+there are sometimes fertile workers in hives.
+
+Having ascertained M. Schirach's discovery, by so decisive an
+experiment, we replaced all the bees examined, in very thin glass hives,
+being only eighteen lines thick, and capable of containing but a single
+row of combs, and thus were extremely favourable to the observer. We
+thought, by strictly persisting to watch the bees, we might surprise a
+fertile one in the act of laying, seize and dissect her. This we were
+desirous of doing, for the purpose of comparing her ovaries with those
+of queens, and to ascertain the difference. At length, on the eighth of
+September, we had the good fortune to succeed.
+
+A bee appeared in the position of a female laying. Before she had time
+to leave the cell, we suddenly opened the hive and seized her. She
+presented all the external characteristics of common bees; the only
+difference we could recognise, and that was a very slight one, consisted
+in the belly seeming less and more slender than that of workers. On
+dissection, her ovaries were found more fragile, smaller and composed of
+fewer oviducts than the ovaries of queens. The filaments containing the
+eggs were extremely fine, and exhibited swellings at equal distances. We
+counted eleven eggs of sensible size, some of which appeared ripe for
+laying. This ovary was double like that of queens.
+
+On the ninth of September, we seized another fertile worker the instant
+she laid, and dissected her. The ovary was still less expanded than that
+of the preceding bee, and only four eggs had attained maturity. My
+assistant extracted one from the oviducts, and succeeded in fixing it by
+an end on a glass slider. We may take this opportunity of remarking,
+that it is in the oviducts themselves the eggs are imbued with the
+viscous liquid, with which they are produced, and not in passing through
+the spherical sac as Swammerdam believed. During the remainder of this
+month, we found ten fertile workers in the same hives, and dissected
+them all. In most, the ovaries were easily distinguished, but in some we
+could not discern the faintest traces of them. In these last, the
+oviducts to all appearance were but imperfectly developed, and more
+address than we had acquired in dissection was necessary to distinguish
+them.
+
+Fertile workers never lay the eggs of common bees; they produce none but
+those of males. M. Riems had already observed this singular fact; and
+here all my observations correspond with his. I shall only add to what
+he says, that fertile workers are not absolutely indifferent in the
+choice of cells for depositing their eggs. They always prefer large
+ones; and only use small cells when unable to find those of larger
+diameter. But they so far correspond with queens whose impregnation has
+been retarded, that they sometimes lay in royal cells.
+
+Speaking of females laying male eggs alone, I have already expressed my
+surprise that bees bestow, on those deposited in royal cells, such care
+and attention as to feed the worms proceeding from them, and, at the
+period of transformation, to close them up. But I know not, Sir, why I
+omitted to observe 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 fertile workers
+lay the eggs of drones is very different. They begin indeed with
+bestowing every care on their eggs and worms; they close the cells at a
+suitable time, but never fail to destroy them three days afterwards.
+
+Having finished these first experiments with success, I had still to
+discover the cause of the expansion of the sexual organs of fertile
+workers. M. Riems had not engaged in this interesting problem; and at
+first I dreaded that I should have no other guide towards its solution
+than conjecture. Yet from serious reflection, it appeared, that, by
+connecting the facts contained in this letter, there was some light that
+might elucidate my procedure in this new research.
+
+From M. Schirach's elegant discoveries, it is beyond all doubt that
+common bees are originally of the female sex. They have received from
+nature the germs of an ovary, but she has allowed 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 get 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.
+Therefore, if fertile workers are produced in this situation alone, it
+is evident their origin is only in those hives where bees prepare the
+royal jelly. Towards this circumstance, I bent all my attention. It
+induced me to suspect that when bees give the _royal treatment_ to
+certain worms, they either by accident or a particular instinct, the
+principle of which is unknown to me, drop some particles of royal jelly
+into cells contiguous to those containing the worms destined for queens.
+The larvae of workers that have accidentally received portions of so
+active an aliment, must be more or less affected by it; and their
+ovaries should acquire a degree of expansion. But this expansion will be
+imperfect; why? because the royal food has been administered only in
+small portions, and, besides, the larvae having lived in cells of the
+smallest dimensions, their parts cannot extend beyond the ordinary
+proportions. Thus, the bees produced by them will resemble common
+workers in size and all the external characteristics. Added to that,
+they will have the faculty of laying some eggs, solely from the effect
+of the trifling portion of royal jelly mixed with their aliment.
+
+That we may judge of the justness of this explanation, it is necessary
+to consider fertile workers from their origin; to investigate whether
+the cells, where they are brought up, are constantly in the vicinity of
+the royal cells, and if their food is mixed with particles of the royal
+jelly. Unfortunately, the execution of these experiments is very
+difficult. When pure, the royal jelly is recognised by its sharp and
+pungent taste; but, when mixed with other substances, the peculiar
+savour is very 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{J}.
+
+In June 1790, I observed that one of my thinnest hives had wanted the
+queen several days, and that the bees had no mean 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 worms destined for
+queens, in 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
+destined for replacing the queens, it was very interesting to follow
+their history, and necessary to watch the moment of their last
+transformation. I removed the nineteen cells into a grated box, which
+was introduced 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 experiment. But here another
+precaution was also requisite. It was to be feared, that the bees being
+deprived of the produce of their labour, and the object of their hope
+might be totally discouraged; therefore, I supplied them with another
+piece of comb, containing the brood of workers, reserving power to
+destroy the young brood when necessary. This plan succeeded admirably.
+The bees, in bestowing all their attention on these last worms, forgot
+those that had been removed.
+
+When the moment of transformation of the nymphs in the nineteen cells
+arrived, I examined the grated box frequently every day, and at length
+found six bees exactly similar to _common bees_. The worms of the
+remaining thirteen had perished without changing.
+
+The portion of brood comb that had been put into the hive to prevent the
+discouragement of the bees was then removed. I put 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 six
+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 produced by some of the six bees?
+But, to attain absolute certainty, it was necessary to take them in the
+act of laying. Some ineffaceable mark was also required for
+distinguishing them in particular.
+
+This proceeding was attended with the most ample success. We 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
+perceived one introducing the posterior part into 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 privation of the right antenna, instantly
+recognised that it was one of the six that had passed to the vermicular
+state in the vicinity of the royal cells.
+
+I could no longer doubt the truth of my conjecture; at the same time, I
+know not whether the truth will appear as rigorous to you, Sir, as it
+does to myself. But I reason in the following manner: If it is certain
+that fertile workers are always produced in the vicinity of royal cells,
+it is no less true, that in itself, the vicinity is indifferent; for the
+size and figure of these cells can produce no effect on the worms in
+those surrounding them; there must be something more; we 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 the worms in the
+adjacent cells have had a portion of the same food. This is what they
+gain, therefore, by vicinity to the royal cells. The bees, in their
+course thither, will pass in numbers over them, stop and drop some
+portion of the jelly destined for the royal larvae. This reasoning, I
+presume, is consistent with the principles of sound logic.
+
+I have repeated the experiment now described so often, and weighed all
+the concomitant circumstances with so much care, that whenever I please,
+I can produce fertile workers in my hives. The method is simple. 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
+supplying the included worms with the royal jelly. Portions of this
+aliment also fall on the young larvae deposited in the adjacent cells,
+and it unfolds the ovaries to a certain degree. 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. Therefore, to
+save them, all their enemies must be removed, and the larvae of the royal
+cells taken away before undergoing their last metamorphoses. Then the
+fertile workers, being without rivals at the time of their origin, will
+be well received, and, by taking the precaution to mark them, it will be
+seen, in a few days, that they produce the eggs of males. Thus, the
+whole secret of this proceeding consists in removing the royal cells at
+the proper time; that is, after being sealed, and previous to the young
+queens leaving them{K}.
+
+I shall add but a few words to this long letter. There is nothing so
+very surprising in the production of fertile workers, when we consider
+the consequences of M. Shirach's beautiful discovery. But why do they
+lay male eggs only? I can conceive, indeed, that the reason of their
+laying few is from their ovaries being but imperfectly expanded, but I
+can form no idea why all the eggs should be those of males, neither can
+I any better account for their use in hives; and hitherto, I have made
+no experiments on their mode of fecundation.
+
+  _PREGNY, 25. August 1791._
+
+
+FOOTNOTES:
+
+{I} It is difficult to discover whether the author thinks, as some
+naturalists, that bees are instrumental in hatching the eggs.--T.
+
+{J} The original is extremely confused in the preceding passages.--T.
+
+{K} I have frequently seen queens, at the moment of production, begin
+first by attacking the royal cells and then the common ones beside them.
+As I had not seen fertile workers when I first observed this fact, I
+could not conceive from what motive the fury of the queen was thus
+directed towards the common cells. But now I know they can distinguish
+the species included, and have the same instinctive jealousy or aversion
+towards them as against the nymphs of queens properly so denominated.
+
+
+
+
+LETTER VI.
+
+_ON THE COMBATS OF QUEENS: THE MASSACRE OF THE MALES: AND WHAT SUCCEEDS
+IN A HIVE WHERE A STRANGER QUEEN IS SUBSTITUTED FOR THE NATURAL ONE._
+
+
+M. de Reaumur had not witnessed every thing relative to bees when he
+composed his history of these industrious animals. Several observers,
+and those of Lusace in particular, have discovered many important facts
+that escaped him; and I, in my turn, have made various observations of
+which he had no suspicion: at the same time, and this is a very
+remarkable circumstance, not only has all that he expressly declares he
+saw been verified by succeeding naturalists, but all his conjectures are
+found just. The German naturalists, Schirach, Hattorf, and Riems
+sometimes contradict him, indeed, in their memoirs; but I can maintain
+that, while combating the opinion of M. de Reaumur, it is they who are
+almost always wrong; of which several instances might be adduced.
+
+What I shall now proceed to say will give me an opportunity of detailing
+some interesting facts.
+
+It was observed by M. de Reaumur, that when any supernumerary queen is
+either produced in a hive, or comes into it, one of the two soon
+perishes. He has not actually witnessed 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. M. Schirach, on the other
+hand, and, after him, M. Riems, thinks that the working bees assail the
+stranger, and sting her to death. I cannot comprehend by what means they
+have been able to make this observation: as they used very thick hives
+only, with several rows of combs, they could at most but observe the
+commencement of hostilities. While the combat lasts, the bees move with
+great rapidity; they fly on all sides; and, gliding between 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 queens and
+workers, but I have very often beheld one between the queens themselves.
+
+In one of my hives in particular, there were five or six royal cells,
+each including a nymph. The eldest first underwent its transformation.
+Scarcely did ten minutes elapse from the time of this young queen
+leaving her cradle, when she visited the other royal cells still close.
+She furiously attacked the nearest; and, by dint of labour, succeeded
+in opening the top: we saw her tearing the silk of the coccoon with her
+teeth; but her efforts were probably inadequate to the object, for she
+abandoned this end of the cell, and began at the other, where she
+accomplished a larger aperture. When it was sufficiently enlarged, she
+endeavoured to introduce her belly, and made many exertions until she
+succeeded in giving her rival a deadly wound with her sting. Then having
+left the cell, all the bees that had hitherto been spectators of her
+labour, began to increase the opening, and drew out the dead body of a
+queen scarcely come from its envelope of a nymph.
+
+Meanwhile, the victorious young queen attacked another royal cell, but
+did not endeavour to introduce her extremity into it. There was only a
+royal nymph, and no queen, come to maturity, as in the first cell. In
+all probability, nymphs of queens inspire their rivals with less
+animosity; still they do not escape destruction: because, whenever a
+royal cell has been opened before the proper time, the bees extract the
+contents in whatever form they may be, whether worm, nymph, or queen.
+Lastly, the young queen attacked the third cell, but could not succeed
+in penetrating it. She laboured languidly, and appeared as if exhausted
+by her first exertions. As we now required queens for some particular
+experiments, we resolved to remove the other royal cells, yet in safety,
+to secure them from her fury.
+
+After this observation, we wished to see what ensued on two queens
+leaving their cells at the same time, and in what manner one perished. I
+find an observation on this head in my Journal, 15. May 1790.
+
+In one of our thinnest hives, two queens left their cells almost at the
+same moment. Whenever they observed each other, they rushed together,
+apparently with great fury, and were in such a position that the antennae
+of each was seized by the teeth of the other: the head, breast, and
+belly of the one were exposed to the head, breast, and belly of the
+other: the extremity of their bodies were curved; they were reciprocally
+pierced with the stings; and both fell dead at the same instant. But it
+seems as if nature has not ordained that both combatants should perish
+in the duel; but rather that, when finding themselves in the situation
+described, namely, opposite, and belly to belly, they fly at that moment
+with the utmost precipitation. Thus, when these two rivals felt the
+extremities about to meet, they disengaged 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 think that we may here
+penetrate the intention of nature.
+
+There ought to be none but one queen in a hive: therefore it is
+necessary, if by chance a second is either produced or comes into the
+hive, that one of the two must be destroyed. This cannot be committed to
+the working bees, because, in a republic composed of so many
+individuals, an equal consent cannot be supposed always to exist; it
+might frequently happen that one group of bees destroyed one of the
+queens, while a second would massacre the other; and the hive thus be
+deprived of queens. Therefore it was necessary that the queens
+themselves should be entrusted with the destruction of their rivals: but
+as, in these combats, nature demands but 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 trifling facts. But here the object and the means
+seem so plain, that I have ventured to advance my conjectures. You will
+judge better than I can, whether they are well founded.--Let me now
+return from this digression.
+
+A few minutes after the two queens separated, their terror ceased, and
+they again began to seek each other. Immediately on coming in sight,
+they rushed together, seized one another, and resumed exactly their
+former position. The result of this encounter was the same. When their
+bellies approached, they hastily disengaged themselves, and fled with
+precipitation. During all this time, the workers seemed in great
+agitation; and the tumult appeared to increase when the adversaries
+separated. Two different times, we observed them stop the flight of the
+queens, seize their limbs, and retain them prisoners above a minute. At
+last, the queen, which was either the strongest 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, brought the
+extremity of her own body under the belly of the other; and, by this
+means, easily pierced her with the sting. Then she withdrew her sting
+after losing hold of the wing. The vanquished 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 fecundated, and mothers, had the
+same animosity.
+
+On the 22. of July, we selected a flat hive, containing a very fertile
+queen: and being curious to learn whether, as virgin queens, she would
+destroy the royal cells, three were introduced into the middle of the
+comb. Whenever she observed this, _she_ sprung forward on the whole, and
+pierced them towards the bottom; nor did she desist until the included
+nymphs were exposed. The workers which had hitherto been spectators of
+this destruction, now came to carry the nymphs away. They greedily
+devoured the food remaining at the bottom of the cells, and also sucked
+the fluid from the abdomen of the nymphs: and then terminated with
+destroying the cells from which they had been drawn.
+
+In the next place, we introduced a very fertile queen into this hive;
+after painting the thorax to distinguish her from the reigning queen. A
+circle of bees quickly formed around the stranger, but their intention
+was not to caress and receive her well; for they insensibly accumulated
+so much, and surrounded her so closely, that in scarcely a minute she
+lost her liberty and became a prisoner. It is a remarkable circumstance,
+that other workers at the same time collected round the reigning queen
+and restrained all her motions; we instantly saw her confined like the
+stranger. Perhaps it may be said, 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 way to allow her full liberty for the attack; then if the queens
+testified a disposition to fly, they returned to enclose them.
+
+We have repeatedly witnessed this fact, but it presents so new and
+singular a characteristic in the policy of bees, that it must be seen
+again a thousand times before any positive assertion can be made on the
+subject. I would therefore recommend that naturalists should attentively
+examine the combat of queens, and particularly ascertain what part is
+taken by the workers. Is their object to accelerate the combat? Do they
+by any secret means excite the fury of the combatants? Whence does it
+happen that accustomed to bestow every care on their queen, in certain
+circumstances, they oppose her preparations to avoid impending danger?
+
+A long series of observations are necessary to solve these problems. It
+is an immense field for experiment, which will afford infinitely curious
+results. I intreat you to pardon my frequent digressions. The subject is
+deeply philosophical, genius such as your's is required to treat it
+properly; and I shall now be satisfied with proceeding in the
+description of the combat.
+
+The cluster of bees that surrounded the reigning queen having allowed
+her some freedom, she seemed to advance towards that part of the comb
+where her rival stood; then, all the bees receded before her, the
+multitude of workers, separating the two adversaries, gradually
+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 combination, we had still to examine
+whether a combat would ensue between two queens, one impregnated, and
+the other a virgin; and what circumstances attended it.
+
+On the 18. of September, we introduced a very fruitful queen into a
+glass hive, already containing a virgin queen, and put her on the
+opposite side of the comb, that we might have time to see how the
+workers would receive her. She was immediately surrounded, but they
+confined her only a moment. Being oppressed with the necessity of
+laying, she dropped some eggs; however, 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 advanced towards the edge of the
+comb, and soon approached very near the virgin queen. When in sight,
+they rushed together; the virgin queen got on the back of the other, and
+gave her several stings in the belly, but, having aimed at the scaly
+part, they did not injure her, and the combatants separated. In a few
+minutes, they returned to the charge; but this time the impregnated
+queen mounted on her rival; however, she sought in vain to pierce her,
+for the sting did not enter; the virgin queen then disengaged herself
+and fled; she also succeeded in escaping another attack, where her
+adversary 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 successful exertion, the virgin queen
+mortally wounded the stranger, and she expired in a moment. The sting
+had penetrated so far that the victorious queen was unable to extract
+it, and she was overthrown by the fall of her enemy. She made great
+exertions to disengage the sting: but could succeed by no other means
+than turning on the extremity of the belly, as on a pivot. Probably the
+barbs of the sting fell by this motion, and, closing like a spiral
+around the stem, came more easily from the wound.
+
+These observations, Sir, I think will satisfy you, respecting the
+conjecture of our celebrated 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 will
+at no time attempt to employ their stings against the stranger queen. I
+can conceive what has misled M. Riems and Schirach; but it is necessary
+for 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 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 supernumerary queens; and if an observer wishes 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 a natural state,
+for the following 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
+antennae every individual endeavouring to penetrate the hive, and also
+the various substances put within their reach; which affords us an
+opportunity of observing that the antennae are certainly the organs of
+feeling. If a stranger queen appears, she is instantly seized by the
+bees on guard, which prevent her entry by laying hold of her legs or
+wings with their teeth, and crowd so closely around her, that she cannot
+move. Other bees, from the interior of the hive, gradually come to their
+assistance, and confine her still more narrowly, all having their heads
+towards the centre where the queen is inclosed; and they remain with
+such evident anxiety, eagerness, and attention, that the cluster they
+form may be carried about for some time, without their being sensible
+of it. A stranger queen, so closely confined and hemmed in, cannot
+possibly penetrate the hive. If the bees retain her too long imprisoned,
+she perishes. Her death probably ensues from hunger, or the privation of
+air; it is undoubted, at least, that she is never stung. We never saw
+the bees direct their stings against her, except a single time, and then
+it was owing to ourselves. We endeavoured, from compassion for a queen's
+situation, to remove her from the center 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 several of the workers were themselves killed; and
+surely they could not intend destroying one another. Had we not
+interfered, they would have been content with confining the queen, and
+would not have massacred her.
+
+It was in similar circumstances that M. Riems saw the workers anxiously
+pursue a queen. He thought they designed to sting her, and thence
+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 II, chap. 27, note 7_. But you are sensible, Sir, from
+these details, that he has been mistaken. He did not know the attention
+that bees bestow on what passes at the entrance of their hive, and he
+was entirely ignorant of the means they take to prevent supernumerary
+queens from penetrating it.
+
+       *       *       *       *       *
+
+After ascertaining that the workers in no situation sting the
+supernumerary queens, we were curious to learn how a stranger queen
+would be received in a hive wanting a 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.
+
+Bees do not immediately observe the removal of their queen; their
+labours are uninterrupted; they watch over the young, and perform all
+their ordinary occupations. 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 a
+delirious impetuosity. Then they discover their queen is no longer among
+them. But how do they become sensible of it? How do the bees on the
+surface of the comb discover that the queen is not on the next comb? In
+treating of another characteristic of these animals, you have yourself,
+Sir, proposed the same question; I am incapable of answering it indeed,
+but I have collected some facts, that may perhaps facilitate the
+elucidation of this 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
+interpret this expression strictly. Substitution of another queen is not
+attended with the same effect, if she is introduced into the hive within
+the first twelve hours after removal of the reigning one. Here the
+agitation continues; and the bees treat the stranger the same as when
+the presence of their own leaves them nothing to desire. They surround,
+seize, and keep her captive, a very long time, in an impenetrable
+cluster; and she commonly dies either from hunger or privation of air.
+
+If eighteen hours elapse before substitution of a stranger queen for the
+native one removed, she is at first treated 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 moves
+languidly; and sometimes 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 where they had originally been ill
+received.
+
+If, before substituting the stranger queen, twenty-four hours elapse,
+she 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 word reception is very
+indefinite, it is proper to enter into some detail for explaining the
+exact sense in which I use it. On the 15. of August, I introduced a
+fertile queen, eleven months old, into a glass hive. The bees were
+twenty-four hours deprived of their queen, and had already begun the
+construction of twelve royal-cells, such as described in the preceding
+chapter. 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 others that treated her exactly in the same manner. All
+vibrated their wings at once, and ranged themselves in a circle around
+their sovereign. Hence resulted a kind of agitation which gradually
+communicated to the workers situated on the same surface of the comb,
+and induced them to come and reconnoitre, in their turn, what was going
+on. They soon arrived; and, having broke through the circle formed by
+the first, approached the queen, touched her with the antennae, and gave
+her honey. After this little ceremony they retired; and, placing
+themselves behind the others, enlarged the circle. There they vibrated
+their wings, and buzzed without tumult or disorder, and as if
+experiencing some very agreeable sensation. The queen had not yet moved
+from the place where I had put her, but in a quarter of an hour she
+began to move. The bees, far from opposing her, opened the circle at
+that part to which she turned, followed her, and formed a guard around.
+She was oppressed with the necessity of laying, and dropped eggs.
+Finally, after four hours abode, she began to deposit male eggs in the
+cells she met.
+
+While these events passed on the surface of the comb where the queen
+stood, all was quiet on the other side. Here the workers were apparently
+ignorant of a queen's arrival in the hive. They laboured with great
+activity at the royal cells, as if ignorant that they no longer stood in
+need of them: they watched over the royal worms, supplied them with
+jelly and the like. But the queen having at length come to this side,
+she was received with the same respect that she had experienced from
+their companions on the other side of the comb. They encompassed her;
+gave her honey; and touched her with their antennae: and what proved
+better that they treated her as a mother, was their immediately
+desisting from 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 that bees receive
+a stranger queen; when they have time to forget their own, she is
+treated exactly as if she was their natural one, except that there is
+perhaps at first greater interest testified in her, or more conspicuous
+demonstrations of it. I am sensible of the impropriety of these
+expressions, but M. de Reaumur in some respect authorises them. He does
+not scruple to say, that bees pay _attention_, _homage_, and _respect_,
+to their queen, and from his example the like expressions have escaped
+most authors that 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 full of combats and disastrous
+scenes, I should, perhaps, give you an account of some more pleasing
+and interesting facts relative to their industry. However, to avoid
+returning to duels and massacres, I shall here subjoin my observations
+on the massacre of the males.
+
+You will remember, Sir, it is agreed by all observers, that at a certain
+period of the year, the workers kill and expel the drones. M. de Reaumur
+speaks of these executions as a horrible massacre. He does not expressly
+affirm, indeed, that he has himself witnessed it, but what we have seen
+corresponds so well with his account, that there can be no doubt he has
+beheld the peculiarities of the massacre.
+
+It is usually in the months of July and August, that the bees free
+themselves of the males. Then they are drove away and pursued to the
+inmost parts of the hive, where they collect in numbers; and as at the
+same time we find many dead drones on the ground before the hives, it is
+indubitable that after being expelled, the bees sting them to death.
+Yet on the surface of the comb, we do not see the sting used against
+them; there the bees are content to pursue and drive them away. You
+observe this, Sir, yourself, in the new notes added to _la Contemplation
+de la Nature_; and you seem disposed to think, that the drones forced to
+retire to the extremity of the hive, perish from hunger. Your conjecture
+was extremely probable. Still it was possible the carnage might take
+place in the bottom of the hive, and had been unobserved, because that
+part is dark, and escapes the observer's eye.
+
+To appreciate the justice of this suspicion, we thought of making the
+support of the hive of glass, and of placing ourselves below to see what
+passed in the scene of action. Therefore, a glass table was constructed,
+on which were put six hives with swarms of the same year; and, lying
+under it, we endeavoured to discover how the drones were destroyed. The
+invention succeeded to admiration. On the 4 of July, we saw the workers
+actually massacre the males, in the whole six swarms, at the same hour,
+and with the same peculiarities.
+
+The glass table was covered with bees full of animation, which flew upon
+the drones, as they came from the bottom of the hive; seized them by the
+antennae, the limbs, and the wings, and after having dragged them about,
+or, so to speak, after quartering them, they killed them by repeated
+stings directed between the rings of the belly. The moment that this
+formidable weapon reached them, was the last of their existence; they
+stretched 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 stuck
+the sting so deep, that it could hardly be withdrawn, and these bees
+were obliged to turn upon themselves before the stings could be
+disengaged.
+
+Next day, having resumed our former position, we witnessed new scenes of
+carnage. During three hours, the bees furiously destroyed the males.
+They had massacred all their own on the preceding evening, but now
+attacked those which, driven from the neighbouring hives, had taken
+refuge amongst them. We saw them also tear some remaining nymphs from
+the combs; they greedily sucked all the fluid from the abdomen, and then
+carried them away. The following days no drones remained in the hives.
+
+These two observations seem to me decisive. It is incontestible that
+nature has charged the workers with the destruction of the males at
+certain seasons of the year. But what means does she use to excite their
+fury against them? This is a question that I cannot pretend to answer.
+However, an observation I have made may one day lead to solution of the
+problem. The males are never destroyed in hives deprived of queens, on
+the contrary, while a savage massacre prevails in other places, they
+there find an asylum. They are tolerated and fed, and many are seen even
+in the middle of January. They are also preserved in 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 but in hives where the queens are completely
+fertile, and it never begins until the season of swarming is past.
+
+  _PREGNY, 28 August 1791._
+
+
+
+
+LETTER VII.
+
+_SEQUEL OF EXPERIMENTS ON THE RECEPTION 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 a sensible pleasure in acknowledging that
+if I have made any progress in the art of observation, I am indebted for
+it to profound study of the works of this naturalist. In general his
+authority has such weight, that I can scarcely trust my own experiments
+when the results are different from his. Likewise, on finding myself in
+opposition to the _historian of bees_, I repeat my experiments. I vary
+the mode of conducting them; I examine with the utmost caution all the
+circumstances that might mislead me, and never are my labours
+interrupted before acquiring the moral certainty of avoiding error. With
+the aid of these precautions, I have discovered the justice of M. de
+Reaumur's suggestions, and I have a thousand times seen, if certain
+experiments seemed to combat them, it was from incorrectness of
+execution. Yet I must except some facts where my results have constantly
+been different from his. Those respecting the reception of a stranger
+queen substituted for the natural one, are of the number.
+
+If, after removing the natural queen, a stranger is immediately
+substituted, the usurper is ill received. I never could succeed in
+making them adopt her, but by allowing an interval of twenty or
+twenty-four hours to elapse. Then they seemed to have forgot their own
+queen; and respectfully received any female put in her place. M. de
+Reaumur, on the contrary, asserts, that should the original queen be
+removed, and another presented, this new one will be perfectly well
+received from the beginning. As evidence of this assertion, he gives the
+detail of an experiment which must be read in his work, for I shall here
+give only an extract of it{L}. 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 this moment, the
+tumult ceased, and the stranger queen was received with all respect.
+
+I do not dispute the truth of this experiment; but, in my opinion, it
+does not warrant the conclusion that M. de Reaumur deduces from it. His
+apparatus removed the bees too much from their natural condition, to
+allow him to judge of their instinct and dispositions. In other
+situations, he has himself observed, that these animals, reduced to
+small numbers, lost their industry and activity, and feebly continued
+their ordinary labours. Thus their instinct is affected by every
+operation that too much diminishes their number. To render such an
+experiment truly conclusive, it must be made in a populous hive; and on
+removing the native queen, a stranger must immediately be substituted in
+her place. Had this been done, I am fully persuaded, that M. de Reaumur
+would have seen the bees imprison the usurper, confine her at least
+twelve or fifteen hours among them, and frequently suffocate her: nor
+would he have witnessed any favourable reception before an interval of
+twenty-four hours after removal of the original queen. No variation has
+occurred in my experiments regarding this fact. Their number, and the
+attention bestowed on them, make me presume they merit your confidence.
+
+M. de Reaumur, in another passage of the same Memoir, affirms, that
+_bees, which have a queen they are satisfied with, are nevertheless
+disposed to give the best possible reception to any female that seeks
+refuge among them_. In the preceding letter, I have related my
+experiments on this head: their success has been very different from
+that of M. de Reaumur's. I have proved that the workers never employ
+their stings against the queen; but this cannot be called the welcome
+reception of a stranger. They retain her within their ranks, and seem to
+allow her liberty only when she prepares to combat the reigning queen.
+This observation cannot be made except in the thinnest hives. Those used
+by M. de Reaumur had always two parallel combs at least, which must
+have prevented him from observing some very important circumstances that
+influence the conduct of workers when supplied with several females. The
+first circles formed around a stranger queen he has taken for caresses;
+and, from the little that this queen could advance between the combs, it
+must have been impossible for him to observe that the circles, which
+always continued contracting, ended in restraint of the females there
+inclosed. Had he used thinner hives, he would have discovered that what
+he supposed indication of a favourable reception was the prelude of
+actual imprisonment.
+
+I feel reluctant to assert that M. de Reaumur was deceived. Yet I cannot
+admit that, on certain occasions, bees tolerate a plurality of females
+in their hives. The experiment on which this affirmation rests will not
+be considered decisive. In the month of December, he introduced a
+stranger queen into a glass hive, in his cabinet, and confined her
+there. The bees had no opportunity of going out. This stranger was well
+received; her presence awakened the workers from their lethargic state,
+into which they did not relapse; she excited no carnage; 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 queen was living when the
+new one was introduced into the hive: however the author neglected this;
+and it is very probable 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 observations coincide with his, and still
+more, when my experiments justify his conjectures. But I think it
+proper to point out those cases where the imperfections of his hives
+have led him into error, and to explain from what causes I have not seen
+certain facts in the same manner he did. I feel particular anxiety to
+merit your confidence, and I am aware that the greatest exertions are
+necessary, when I have to combat the historian of bees. I confide in
+your judgment; and pray you to be assured of my respect.
+
+  _PREGNY, 30. August 1791._
+
+
+FOOTNOTES:
+
+{L} Edit. 4to, Tom. V. p. 258.
+
+
+
+
+LETTER 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 COCCOONS._
+
+
+In this letter I shall collect some isolated observations relative to
+various points in the history of bees, concerning which you wished me to
+engage.
+
+You desired me to investigate whether the queen is really _oviparous_.
+M. de Reaumur leaves this question undecided. He observes, that he has
+never seen the worm hatched; and he only asserts that worms are found in
+those cells where eggs have been deposited three days preceding. If we
+attempt to catch the moment when the worm leaves the egg, we must extend
+our observations beyond the interior of the hive; for there the
+continual motion of the bees obscures what passes at the bottom of
+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 to hatch the worms, should the eggs
+be too soon deprived of it they wither and perish. The sole method of
+succeeding in seeing the worm come out, consists in watching the queen
+while she lays, in marking the egg so as to be recognised, and removing
+it from the hive to the microscope only an hour or two before the three
+days elapse. The worm will certainly be hatched, provided it has been
+exposed as long as possible to the full degree of heat. Such is the
+course I have pursued; and the following are 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 the cell, and put
+the pyramidal bottom, where the egg was fixed, on a glass slider. Slight
+motions were soon perceptible in the eggs. At first, we could observe no
+external organization: the worm was entirely concealed from us by its
+pellicle. We then prepared to examine the egg with a powerful magnifier;
+however, during the interval, the worm burst its surrounding membrane,
+and cast off part of the envelope, which was torn and ragged on
+different parts of the body, and more evidently so towards the last
+rings. The worm alternately curved and stretched itself, with very
+lively action. Twenty minutes were occupied in casting off the spoil;
+when this exertion ceased: the worm lay down, curved, and seemed to take
+that rest which it required. An egg laid in a worker's cell produced
+this animal, which would have become a worker itself.
+
+We next directed our attention to the moment when a male worm would be
+hatched. An egg was exposed to the sun on a glass slider; and, with a
+good magnifier, nine rings of the worm were perceptible within the
+transparent pellicle. This membrane was still entire, and the worm
+perfectly motionless. The two longitudinal lines of tracheae were visible
+on the surface, and many ramifications. We never lost sight of the egg a
+single instant, and now succeeded in observing the first motions of the
+worm. The thick end alternately straightened and curved, and almost
+reached the part where the sharp extremity was fixed. These exertions
+burst the membrane, first on the upper part, towards the head, then on
+the back, and afterwards on all the rest successively. The ragged
+pellicle remained in folds on different parts of the body, and then fell
+off. Thus it is beyond dispute, that the queen is oviparous.
+
+Some observers affirm, that the workers attend to the eggs before the
+worms are hatched; and it is certain that, at whatever time a hive is
+examined, we always see some workers with the head and thorax inserted
+into cells containing eggs, and remaining motionless several minutes in
+this position. It is impossible to discover what they do, for the
+interior of the cell is concealed by their 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 in a strange hive, where there is the necessary degree of
+heat, the worms come out at the usual time, just as if they had been
+left in the cells. Thus no extraordinary aid or attention is required
+for their exclusion.
+
+When the workers penetrate the cells, and remain fifteen or twenty
+minutes motionless, I have reason to believe, it is only to repose from
+their labours. My observations on the subject seem correct. You know,
+Sir, that a kind of irregular shaped cells, are frequently constructed
+on the panes of the hive. These, being glass on one side, are
+exceedingly convenient to the observer, since all that passes within is
+exposed. I have often seen bees enter these cells when nothing could
+attract them. The cells contained neither eggs nor honey, nor did they
+need further completion. Therefore the workers repaired thither only to
+enjoy some moments of repose. Indeed, they were fifteen or twenty
+minutes so perfectly motionless, that had not the dilatation of the
+rings shewed their respiration, we might have concluded them dead. The
+queen also sometimes penetrates the large cells of the males, and
+continues very long motionless in them. Her position prevents the bees
+from paying their full homage to her, yet even then the workers do not
+fail to form a circle around her, and brush the part of her belly that
+remains exposed.
+
+The drones do not enter the cells while reposing, but cluster together
+on the combs; and sometimes retain this position eighteen or twenty
+hours without the slightest motion.
+
+As it is important, in many experiments, to know the exact time that the
+three species of bees exist before assuming their ultimate form, I shall
+here subjoin my own observations 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. The worm
+now begins spinning its coccoon, in which operation thirty-six hours are
+consumed. In three days, it changes to a nymph, and passes six days in
+this form. It is only on the twentieth day of its existence, counting
+from the moment the egg is laid, that it attains the fly state.
+
+The royal worm also passes three days in the egg, and is five a worm;
+the bees then close its cell; and it immediately begins spinning the
+coccoon, which occupies twenty-four hours. The tenth and eleventh day it
+remains in complete repose, and even sixteen hours of the twelfth. Then
+the transformation to a nymph takes place, in which state four days and
+a third are passed. Thus it is not before the sixteenth day that the
+perfect state of queen is attained.
+
+The male worm passes three days in the egg, six and a half as a worm,
+and metamorphoses into a fly on the twenty-fourth day after the egg is
+laid.
+
+Though the larvae of bees are apodal, they are not condemned to absolute
+immobility in their cells; for they can move by a spiral motion. During
+the first three days, this motion is so slow as scarcely to be
+perceptible, but it afterwards becomes more evident. I have then
+observed them perform two complete revolutions in an hour and three
+quarters. When the period of transformation arrives, they are only two
+lines from the orifice of the cells. As their position is constantly the
+same, bent in an arc, those in the workers' and drones' cells are
+perpendicular to the horizon, while those in the royal cells lie
+horizontally. It might be thought, that the difference of position has
+much influence on the increment of the different larvae; yet it has none.
+By reversing combs containing common cells full of brood, I have put the
+worms in a horizontal position; but they were not injured. I have also
+turned the royal cells, so that the worms came into a horizontal
+direction; however their increment was neither slower nor less perfect.
+
+       *       *       *       *       *
+
+I have been much surprised at the mode of bees spinning their coccoons,
+and 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. The royal larvae,
+on the other hand, spin imperfect coccoons, open behind, and enveloping
+only the head, thorax, and first ring of the abdomen. The discovery of
+this difference, which at first may seem trifling, has given me extreme
+pleasure, for it evidently demonstrates the admirable art with which
+nature connects the various characteristics in the industry of bees.
+
+You will remember, Sir, the evidence I gave you of the mutual aversion
+of queens, of the combats in which they engage, 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 a complete coccoon, she 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 retract it, would
+become the victim of her own fury. Thus, that the queen might destroy
+her rivals, it was necessary the last rings of the body should remain
+uncovered, therefore the royal nymphs must only form imperfect 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 in her care
+of preserving and multiplying the species. But from the facts I relate,
+we must admire her precautions in exposing certain individuals to a
+mortal danger.
+
+The detail on which I have just entered clearly indicates the final
+cause of the opening left by the royal worms in their coccoons; but it
+does not shew whether it is in consequence of a particular instinct that
+they leave this opening, or whether the wideness of their cells prevents
+them from stretching the thread up to the top. This question interested
+me very much; the only method of deciding it was to observe the worms
+while spinning, which cannot be done in their opaque cells. 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 operation
+consisted in extracting worms and introducing them here; but my
+assistant accomplished it with much address. He opened several sealed
+royal cells, where we knew the larvae were about to begin their coccoons,
+and, taking them gently out, introduced one into each of my glass cells
+without the smallest injury.
+
+They soon prepared to work; and commenced by stretching the anterior
+part of the body in a straight line, while the other was bent in a
+curve. This formed a curve of which the longitudinal sides of the cells
+were tangents, and afforded two points of support. The head was next
+conducted to the different parts of the cell which it could reach, and
+it carpeted the surface with a thick bed of silk. We remarked that the
+threads were not carried from one side to another, and that this would
+have been impracticable, for the worms being obliged to support
+themselves, and to keep the posterior rings curved, the free and
+moveable part of the body was not long enough for the mouth to reach the
+sides diametrically opposite, and fix the threads to them. You will
+remember, Sir, that the royal cells are of a pyramidal form, with a wide
+base, and a long contracted top. These cells hang perpendicularly in the
+hive, the point downwards, from which position the royal worm can be
+supported in the cell, only when the curvature of the posterior part
+forms two points of support; and that it cannot obtain this support
+without resting on the lower part, or towards the extremity. Therefore
+if it attempted to stretch out and spin towards the wide end of the
+cell, it could not reach both sides from being too distant. One part
+would be touched by its extremity, the other by its back, and it would
+consequently 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
+support themselves.
+
+These first experiments obviated the suspicion of any particular
+instinct in the royal worms. They proved, if the worms spun incomplete
+coccoons, it was because they were forced to do so by the figure of
+their cells. However, I wished to have evidence still more direct. I put
+them into cylindrical glass cells, or portions of glass tubes resembling
+common cells, and I had the satisfaction of seeing them spin complete
+coccoons, as the worms of workers do. Lastly, I put common worms in very
+wide cells, and they left the coccoon open. Thus it is demonstrated,
+that the royal worms, and those of workers, have the same instinct and
+the same industry, or in other words, when situated in the same
+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 coccoons, undergo all their metamorphoses equally well. Thus
+the necessity imposed on them by nature, of having the coccoons open, is
+not necessary for their increment; nor has it any other object than that
+of exposing them to the certainty of perishing by the wounds of their
+natural enemy; an observation new and truly singular.
+
+       *       *       *       *       *
+
+I ought to 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 find 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, influence their size. With this view, you have
+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 large cells produced workers of
+larger size, we were bound to conclude that the size of the cells had a
+sensible influence on the size of the bees. The first time I made this
+experiment, it did not succeed, because weevils lodged in the hive
+discouraged the bees. But I repeated it afterwards, and the result was
+very remarkable.
+
+I removed the whole comb, consisting of common cells, from one of my
+best glass hives, and left that composed of males' cells alone: and to
+avoid vacuities, I supplied others of the same kind. This was in June,
+the season most favourable to bees. I expected that the bees would
+quickly have repaired the ravages produced by this operation 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 they would resume their activity, I continued
+observing them several days; however, my hopes were disappointed. Their
+homage to the queen was not interrupted indeed; but except in this,
+their conduct to the queen was quite different from what it usually is;
+they clustered on the combs without exciting any sensible heat. A
+thermometer among them rose only to 81 deg., though standing at 77 deg.
+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 eggs, hesitated long before depositing them in the
+large cells; she chose rather to drop them at random than lay in cells
+unsuitable. However, on the second day, we found six that had been
+deposited there with all regularity. The worms were hatched three days
+afterwards, and then we began to study their history. Though the bees
+provided them with food, they did not carefully attend to it; yet I was
+in hopes they might be reared. I was again disappointed; for next
+morning all the worms had disappeared, and their cells were left empty.
+Profound silence reigned in the hive; few bees left it, and these
+returned without pellets of wax on the limbs; all was cold and
+inanimate. To promote a little motion, I thought of supplying the hive
+with a comb, composed of large cells, full of male brood of all ages.
+The bees, which had twelve days obstinately refused working in wax, did
+not unite this comb to their own. However, their industry was awakened
+in a way that I had not anticipated. They removed all the brood from
+this comb, cleaned out the whole cells, and prepared them for receiving
+new eggs. I cannot determine whether they expected the queen 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 removed 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, and
+this order cannot be inverted. We have seen that another fact led me to
+the same consequence; and as that was extremely important, I am
+delighted to have it confirmed by a new observation. 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 certain 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 large cells; for, by the
+experiment just described, 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 has instructed her that their size is
+neither proportioned to the size nor necessities of common worms, would
+not she also have been instructed not to lay 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 bees is not very conspicuous. Here the most prominent
+feature of industry appears in the common bees. When I supplied them
+with a comb of small cells, full of male brood, their activity was
+awakened; but instead of bestowing the necessary care on this brood, as
+they would have done in every other situation, they destroyed the whole
+nymphs and larvae, and cleaned out their cells, that the queen, now
+oppressed with the necessity of laying, might suffer no delay in
+depositing her eggs. Could we allow them either reason or reflection,
+this would be an interesting proof of their affection for her.
+
+The experiment, now detailed at length, not having fulfilled my object
+in determining the influence of the size of the cells on that of the
+worms, I invented another which proved more successful.
+
+Having selected a comb of large cells, containing the eggs and worms of
+males, I removed all the worms from their farina, and my assistant
+substituted those of workers a day old in their place. Then he
+introduced this comb into a hive that had the queen. The bees did not
+abandon these substituted worms; they covered their cells with a top
+almost flat, a kind quite different from what is put on the cells of
+males; which proves, that they were well aware that these, though
+inhabiting large cells, were not males. This comb remained eight days
+in the hive, counting from the time the cells were sealed. I then
+removed it to examine the included nymphs, which proved those of workers
+in different stages of advancement; but, as to size and figure, they
+perfectly resembled what had grown in the smallest cells. I thence
+concluded, that the larvae of workers do not acquire greater size in
+large than in small cells. Although this 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; undoubtedly
+she has ordained that their organs should be fully expanded, and there
+is sufficient space for that purpose; therefore more would be useless.
+Their expansion ought to be no greater in the most spacious cells than
+in those appropriated 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 will probably be less than that of common workers,
+because they have been cramped in the cells; but it does not thence
+ensue, that a larger cell will admit of them growing 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 circumstances. The large cells of males are
+sufficiently capacious for the perfect expansion of their organs. Thus,
+although reared in cells of still greater capacity, they will grow no
+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 them, and reared 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
+will be checked, they will not attain the usual size, of which there is
+proof in the following experiment. I had a comb consisting of the cell
+of large drones, and one with those of workers, which also served for
+the male worms. Of these, my assistant took a certain number from the
+smallest cells, and deposited them on a quantity of food purposely
+prepared in the large ones; and in return he introduced into the small
+cells the worms that had been hatched in the other, and then committed
+both to the care of the workers 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 metamorphosis arrived,
+gave both kinds that convex covering usually put on those of the 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.
+
+You suggested another experiment which I carefully made, but it met with
+an unforeseen obstacle. To appreciate the influence of the royal food on
+the expansion 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
+without success. Nor do I think it can ever succeed. 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 deprived of a queen,
+they change the cells containing worms into cells of the largest kind.
+Then the worms will infallibly be converted to queens.
+
+But there is another situation where we can judge of the influence of
+the royal food administered to worms in common cells. I have spoken at
+great length in my letter on the existence of fertile workers. You
+cannot forget, Sir, that the expansion of their sexual organs is owing
+to the reception of some particles of royal jelly, while in the
+vermicular form. For want of new observations, I must refer you to what
+is previously said on the subject.
+
+  _PREGNY, 4 September 1791._
+
+
+
+
+LETTER IX.
+
+_ON THE FORMATION OF SWARMS._
+
+
+I can add but a few facts to the information M. de Reaumur has
+communicated relative to swarms.
+
+A young queen, according to this celebrated naturalist, is always or
+almost always at the head of a swarm; but he does not assert the fact
+positively, and had some doubts on the subject. "Is it certain," says
+he "as we have hitherto supposed, in coincidence with all who have
+treated of bees, that the new colony is always conducted by a young
+mother? May not the old mother be disgusted with her habitation? or may
+she not be influenced by some particular 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 conducted the young swarms. By the following
+details, you will observe, that his suspicions 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 related.
+
+But previous to entering on this subject, I should repeat what has
+already been frequently observed, that the _leaf_ or flat hives are
+indispensible in studying the industry and instinct of bees. When they
+are left at liberty to conduct several rows of parallel combs, we can no
+longer observe what is continually passing between them, or they must be
+dislodged by water or smoke, for examining what has been constructed; a
+violent proceeding, which has a material influence on their instinct,
+and consequently exposes an observer to the risk of supposing simple
+accidents permanent laws.
+
+I now proceed to experiments proving that an old queen always conducts
+the first swarm.
+
+One of my glass hives consisted of three parallel combs, placed in
+squares that opened like the leaves of a book. It was well peopled and
+abundantly provided with honey, wax, and brood, of every age. On the
+fifth of May 1788, I removed the queen, and on the sixth, transferred
+all the bees into another hive, with a fertile queen at least a year
+old. They entered easily and without fighting, and were in general 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 formed regular
+circles around her. However, there was a little agitation in the
+evening, but confined to the surface of the comb where we had put the
+queen, and which she had not quitted. All was perfectly quiet on the
+other side of this comb.
+
+In the morning of the seventh, the bees had destroyed the twelve royal
+cells, but, independent of that, order continued prevalent in the hive;
+the queen laid the eggs of males in the large cells, and those of
+workers 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
+perpendicularly 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, and
+the longest were only about two lines and a half in depth from the
+bottom to the orifice.
+
+On the thirteenth, the queen seemed already more slender than when
+introduced into the hive; however she still laid some eggs, both in
+common cells and those of males. We also surprised her this day laying
+in a royal cell: she first dislodged the worker there employed, by
+pushing it away with her head, and then supported herself by the
+adjoining cells while depositing the egg.
+
+On the fifteenth, the queen was still more slender: the bees continued
+their attention to the royal cells, which were all unequally advanced;
+some to three or four lines in height, while others were already an inch
+long; which proved 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 hastened to collect
+and put the bees into a hive purposely prepared. Though we had
+overlooked the facts attending the departure of the swarm, the object of
+this experiment was fulfilled; for, on examination of all the bees, we
+were convinced they had been conducted by the old queen; by that we
+introduced on the sixth of the month, and which had been deprived of one
+of the antennae. Observe, there was no other queen in this colony. In the
+hive she had left, we found seven 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.
+
+The new swarm next became the object of our attention: we observed it
+during the rest of the year, during winter and the subsequent spring;
+and, in April, we had the satisfaction of seeing a new swarm depart with
+the same queen at its head that had conducted the former swarm in May
+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 royal cells; she laid in
+one of them before us; and in the last place led forth the swarm.
+
+We have several times repeated the same experiment with equal success.
+Thus it appears incontestible, that the old queen always 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.
+The bees prepare these cells only while the queen lays male eggs; and a
+remarkable fact attends it, that after this laying terminates, her belly
+being considerably diminished, she can easily fly, whereas, her belly is
+previously so heavy she can hardly drag it along. Therefore it is
+necessary she should lay in order to be in a condition for undertaking
+her journey, which may sometimes be very long.
+
+But this single condition is not enough. It is also requisite that the
+bees be very numerous: they should even be superabundant, and a person
+might say they are aware of it: for, if the hive is thin, no royal cells
+are constructed when the male eggs are laid, which is solely at the
+period that the queen is able to conduct a colony. This fact was proved
+by the following experiment on a large scale.
+
+On the third of May 1788, we divided eighteen hives into two portions;
+all the queens were about a year old. Thus each portion of the hives
+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 of the hives threw a swarm.--Therefore, if the
+hive containing the old queen is not very populous, she remains in it
+until the subsequent spring; and if the population is then sufficient,
+royal cells will be constructed: the queen 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 very brief abstract of my observations on swarms conducted by
+old queens. You must excuse the long detail on which I am about to
+enter, concerning 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
+hitherto been very obscure. A long course of observations, protracted
+even during several years, was necessary to remove, in some degree, the
+veil that concealed these mysteries. I have been indemnified for the
+trouble, indeed, by the pleasure of seeing my experiments reciprocally
+confirmed; but, considering the assiduity 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 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
+queens; and I shall now extract the chief experiments from my journal.
+
+On the fourteenth of May, we introduced two portions of bees, from the
+straw hives, into a large glass hive very flat; and allowed them only
+one queen of the preceding year, and which had already commenced laying
+in its native hive. We introduced her on the fifteenth. She was very
+fertile. The bees received her well, and she soon began to lay in large
+and small cells alternately.
+
+On the twentieth, we saw the formation of twelve royal cells, all on the
+edges of the communications, or passages through the combs, and shaped
+liked stalactites.
+
+On the twenty-seventh, ten were much but unequally enlarged; 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 became more lively, yet she still continued examining
+the cells as when about to lay; sometimes introducing half her belly,
+but suddenly withdrawing it, without having laid. At other times she
+deposited an egg, which lay in an irregular position, on one side of the
+hexagon, and not fixed by an end to the bottom of the cell. The queen
+produced no distinct sound in her course, and we heard nothing different
+from the ordinary humming of bees. She passed over those in her way;
+sometimes when she stopped, the bees meeting her also stopped; and
+seemed to consider her. They advanced briskly, struck her with their
+antennae, and mounted on her back. She then went on carrying some of the
+workers on her back. None gave her honey, but she voluntarily took it
+from the cells in her way. The bees no longer inclosed and formed
+regular circles around her. The first, aroused by her motions, followed
+her running in the same manner, and in their passage excited those still
+tranquil on the combs. The way the queen had traversed was evident after
+she left it, by the agitation created, which was never afterwards
+quelled: she had soon visited every part of the hive, and occasioned a
+general agitation; if some places still remained tranquil, the bees in
+agitation arrived, and communicated their motion. The queen no longer
+deposited her eggs in cells; she let them fall fortuitously: nor did the
+bees any longer watch over the young; they ran about in every different
+direction; even those returning from the fields, before the agitation
+came to its height, no sooner entered the hive than they participated in
+these tumultuous motions. They neglected to free themselves of the waxen
+pellets on their limbs, and ran blindly about. At last the whole rushed
+precipitately towards the outlets 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, and, for that reason, as I wished it to continue very populous, I
+removed the queen, at the moment she came out, that the bees might not
+fly too far, and 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 removed its queen also.
+
+All these facts were certain, and appeared susceptible of no error.
+Notwithstanding this, I was particularly earnest to learn whether old
+queens always followed the same course; which induced me, on the
+twenty-ninth, to replace, in the glass hive, the queen a year old, which
+had hitherto been the subject of my experiments, and had just began to
+lay the eggs of males. On the same day, we found one of the royal cells
+left by the preceding queen larger than the rest; and, from its length,
+supposed the included worm two days old: the egg had, therefore, been
+laid on the twenty-fourth by that queen, and the worm was hatched on the
+twenty-seventh. On the thirtieth, the queen laid a great deal in the
+large and small cells alternately. Now, and the two following 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,
+and on the second another. The bees also commenced some new ones. All
+was perfectly quiet at eleven in the morning; but, at mid-day, the
+queen, from the utmost tranquillity, became evidently agitated; and her
+agitation insensibly communicated to the workers in every part of their
+dwelling. In a few minutes they precipitately crowded to the entrances,
+and, along with the queen, left the hive. After they had settled on the
+branch of a neighbouring tree, I sought for the queen; thinking that, by
+removing her, the bees might return to the hive, which actually ensued.
+Their first care seemed to consist in seeking the female; they were
+still in great agitation, but gradually calmed; and in three hours
+complete tranquillity was restored.
+
+They had resumed their usual occupations on the third: 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, not by applying wax
+cordons, but by removing wax from the surface. Towards the top this
+waved work is almost imperceptible; it becomes deeper above, and the
+workers excavate it still more from thence to the base of the pyramid.
+The cell, when once shut, also becomes thinner; and is so much so,
+immediately preceding the queen's metamorphosis from a nymph, that all
+its motions are perceptible through the thin covering of wax on which
+the waved work is founded. It is a very remarkable circumstance, that in
+making the cells thinner, from the moment they are closed, the bees know
+to regulate their labour so that it terminates only when the nymph is
+ready to undergo its last metamorphosis.
+
+On the seventh day the coccoon is almost completely _unwaxed_, if I may
+use the expression, at the part next to the head and thorax of the
+queen. This operation facilitates her exit; for she has nothing to do
+but cut the silk that forms the coccoon. Most probably the object is, to
+promote evaporation of the superabundant fluids of the nymph. I have
+made some direct experiments to ascertain the fact, but 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 closing the second. The like
+was done to other royal cells successively, during the subsequent days.
+
+Every moment of the seventh, we expected the queen to leave the royal
+cell shut on the thirtieth of May. The seven days had elapsed. 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 being
+unwilling that she should yet quit her cell, they had soldered the
+covering to it with some particles of wax. What seemed most singular
+was, that this female emitted a very distinct sound, or clacking from
+her prison. It was still more audible in the evening, and even consisted
+of several monotonous notes in rapid succession.
+
+The same sound proceeded from the royal cell on the eighth. Several
+bees kept guard round each royal cell.
+
+The first cell opened on the ninth. The young queen was lively, slender,
+and of a brown colour. Now, we understood why bees retain the female
+captive in their cells, after the period for transformation has elapsed;
+it is, that they may be able to fly the instant they are hatched. The
+new queen occupied all our attention. When she approached the other
+royal cells, the bees on guard pulled, bit her, and chased her away;
+they seemed to be greatly irritated against her, and she enjoyed
+tranquillity only when at a good distance from these cells. This
+procedure was frequently repeated through the day. She twice emitted the
+sound; in doing so she stood, her thorax against a comb, and the wings
+crossed on her back; they were in motion but without being unfolded or
+further opened. Whatever might be the cause of her assuming this
+attitude, the bees were affected 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 approached, all the guards became agitated,
+surrounded her on all sides, bit, and commonly drove her away; sometimes
+when in these circumstances, she emitted her sound, assuming the
+position just described, from that moment the bees became motionless.
+
+The queen confined in the second cell had not yet left it, and was heard
+to hum several times. We accidentally discovered how the bees fed her.
+On attentive examination, a small aperture was perceptible in the end of
+the coccoon which she had cut to escape, and which her guards had again
+covered with wax, to confine her still longer. She thrust her trunk
+through the cleft; at first the bees did not observe it alternately
+thrust out and drawn in, but one at length perceiving it, came to apply
+its trunk to that of the captive queen, and then gave way to others that
+also approached her with honey. When satisfied she retracted her trunk,
+and the bees again closed up the opening with wax.
+
+The queen this day between twelve and one became extremely agitated. The
+royal cells had multiplied very much; she could go no where without
+meeting them, and on approaching she was very roughly treated. Then she
+fled, but to obtain no better reception. At last, these things agitated
+the bees; they precipitately rushed through the outlets of the hive, and
+settled on a tree in the garden. It singularly happened that the queen
+was herself unable to follow or conduct the swarm. She had attempted to
+pass between two royal cells before they were abandoned by the bees
+guarding them, and she was so confined and maltreated as to be incapable
+of moving. We then removed her into a separate hive prepared for a
+particular experiment; the bees, which had clustered on a branch, soon
+discovered their queen was not present, and returned of their own accord
+to the hive. Such is an account of the second colony of this hive.
+
+We were extremely solicitous to ascertain what would become of the other
+royal cells. Four of the close ones had attained complete maturity, and
+the queens would have left them had not the bees prevented it. They were
+not open either previous to the agitation of the swarms, or at the
+moment 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 formed the swarm. We could not
+discover what occasioned the difference in their captivity.
+
+On the twelfth, the queen was at last liberated, 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. Returning at mid-day, we were greatly surprised to find it
+almost totally deserted. During our absence, it had thrown a prodigious
+swarm, which still clustered on the branch of a neighbouring tree. We
+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, escaped. Thus, there was no doubt
+of both queens being in the swarm. We found it so; and removed them,
+that the bees might return to the hive, which they did very soon.
+
+While we were occupied in this operation, the fourth captive queen left
+her prison, 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 guard around the royal cells, and took
+great care to remove the queen whenever she attempted to approach.
+Eighteen royal cells now remained to be guarded.
+
+The fifth queen left her cell at ten at night; therefore two queens were
+now in the hive. They immediately began fighting, but came to disengage
+themselves from each other. However they fought several times during the
+night without any thing decisive. Next day, the thirteenth, we witnessed
+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 strikes the bees motionless. For some
+minutes, we conceived, that taking advantage of the dread exhibited by
+the workers on guard, she would open it, and destroy the young female;
+also she prepared to mount the cell; but in doing so she ceased the
+sound, and quitted that attitude which paralyses 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 described. The agitation
+was so considerable, that a sufficient number of bees did not remain to
+guard the royal cells, and several of the imprisoned queens were thus
+enabled to make their escape. Three were in the cluster formed by the
+swarm, and other three remained in the hive. We removed those that had
+led the colony, to force the bees to return. They entered the hive,
+resumed their post around the royal cells, and maltreated the queens
+when approaching.
+
+A duel took place in the night of the fifteenth, in which one queen
+fell. We found her dead next morning before the hive; but three still
+remained, as one had been hatched during night. Next morning we saw a
+duel. Both combatants were extremely agitated, either with the desire of
+fighting, or the treatment of the bees, when they came near the royal
+cells. Their agitation quickly communicated to the rest of the bees, and
+at mid-day they departed impetuously 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, which I shall
+give you no account of, as it shewed nothing new.
+
+Unfortunately we lost this, which was a very strong swarm; the bees flew
+out of sight, and could never be found. The hive was now very thinly
+inhabited. Only the few bees that had not participated in the general
+agitation remained, and those that returned from the fields after the
+swarm had departed. The cells were, therefore, slenderly guarded; the
+queens escaped from them, and engaged in several combats, until the
+throne remained with the most successful.
+
+Notwithstanding the victories of this queen, she was treated with great
+indifference from the sixteenth to the nineteenth, that is, the three
+days that she preserved her virginity. At length, having gone to seek
+the males, she returned with all the external signs of fecundation, and
+was henceforth received with every mark of respect; she laid her first
+eggs 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
+experiments which I made at the same time for 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, 6. September 1791._
+
+_P. S._--In revising this letter, I find I have neglected taking notice
+of an objection that may embarrass my readers, and which ought to be
+answered.
+
+After the first five swarms had thrown, I had always returned the bees
+to the hive: it is not surprising, therefore, that it was continually so
+sufficiently stocked that each colony was numerous. But things are
+otherwise in the natural state: the bees composing a swarm do not return
+to the hive; and it will undoubtedly be asked, What 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 the 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 workers' 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 affording a second colony without
+being too much weakened. The third and fourth swarm weaken it more
+sensibly; but the inhabitants always remain in sufficient numbers to
+preserve the course of their labours uninterrupted; and the losses are
+soon repaired by the great fecundity of the queen, as she lays above an
+hundred eggs a day.
+
+If, in some cases, the agitation of swarming 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, cannot share in the agitation; when returned to the
+hive, they quietly resume their labours; and their number is not small,
+for, when the weather is fine, at least a third of the bees 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 endeavouring to depart
+become members of the new colony. When this agitation or delirium 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 the rest, seem
+drenched in perspiration; their wings grow moist; they are incapable of
+flight; and even when able to escape, they advance no farther than the
+board of the hive, and soon return.
+
+Those that have lately left their cells remain behind the swarm, still
+feeble, they could not support themselves in flight. Here then are also
+many recruits to people what we should have thought a deserted
+habitation.
+
+
+
+
+LETTER X.
+
+_THE SAME SUBJECT CONTINUED._
+
+
+To preserve greater regularity in continuing the history of swarms, I
+think it proper to recapitulate in a few words the principal points of
+the preceding letter, and to expatiate on each, concerning the result of
+new experiments, respecting which I have still been silent.
+
+In the first place. _If at the return of spring, we examine a hive well
+peopled, and governed by a fertile queen, we shall see her lay a
+prodigious number of male eggs in the course of May, and the workers
+will chuse that moment for constructing several royal cells of the kind
+described by M. de Reaumur._ Such is the result of several long
+continued observations, among which there has not been the slightest
+variation, and I cannot hesitate in announcing it as demonstrated.
+However, I should here add the necessary explanation. It is necessary
+that the queen, before commencing her _great_ laying of the eggs of
+males, be eleven months old; when young she lays only those of workers.
+A queen, hatched in spring, will perhaps lay fifty or sixty eggs of
+drones in whole, but before beginning her great laying of them, which
+should be two thousand in a month, she must have completed her eleventh
+month in age. In the course of our experiments, which more or less
+disturbed the natural state of things, it often happened that the queen
+did not attain this age until October, and immediately began laying male
+eggs. The workers, as if induced by some emanation from the eggs, also
+adopted this time for building the royal cells. No swarm resulted
+thence, it is true, because in autumn all the necessary circumstances
+are absolutely wanting, but it is not less evident, that there is a
+secret relation between the production of the eggs of males, and the
+construction of royal cells.
+
+This laying commonly continues thirty days. The bees on the twentieth or
+twenty-first lay the foundation of several royal cells. Sometimes they
+build sixteen or twenty; we have even had twenty-seven. When the cells
+are three or four lines high, the queen lays those eggs from which her
+own species will come, but not the whole in one day. That the hive may
+throw several swarms, it is essential that the young females conducting
+them be not all produced at the same time. One may affirm, that the
+queen anticipates the fact, for she takes care to allow at least the
+interval of a day between every egg deposited in the cells. It is proved
+by the bees knowing to close the cells the moment the worms are ready to
+metamorphose to nymphs. Now, as they close all the royal cells at
+different periods, it is evident the included worms are not all of an
+equal age.
+
+The queen's belly is very turgid before she begins laying the eggs of
+drones; but it sensibly decreases as she advances, and when terminated
+is very small. Thus she finds herself in a condition to undertake a
+journey which circumstances may prolong; thus this condition was
+necessary; 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 larvae hatched from the eggs laid by the queen, in
+the royal cells, are ready to transform to nymphs, this queen leaves the
+swarm conducting a swarm along with her; and the first swarm that
+proceeds from the hive is uniformly conducted by the old queen._{M} I
+think I can divine the reason of it.
+
+That there may never be a plurality of females in a hive, nature has
+inspired queens with a natural horror against each other; they never
+meet without endeavouring to fight, and to accomplish their mutual
+destruction. Thus, the chance of combat is equal between them, and
+fortune will decide to which the empire shall pertain. But if one
+combatant is older than the rest, she is stronger, and the advantage
+will be with her. She will destroy her rivals successively as produced.
+Thus, if the old queen did not leave the hive, when the young ones
+undergo their last metamorphosis, it could produce no more swarms, and
+the species would perish. Therefore, to preserve the species, it is
+necessary that the old queen 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 old queen; and that may
+happen thus. Nature has not willed that the queen shall quit the first
+hive before her production of male eggs is finished. It is necessary 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 workers. The bees in that case would have no means of replacing
+her, and the colony would go to ruin.
+
+All these things have been with infinite wisdom foreseen. The first
+operation of the bees of a swarm is to construct the cells of workers.
+They labour at them with great ardour, and as the ovaries of the queen
+have been disposed 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. It may be affirmed, that the bees
+know their queen will also lay the eggs of drones; she actually does
+begin to deposit some, though in much smaller number than at first;
+enough however to encourage the bees to construct royal cells. Now, if
+in these circumstances the weather is favourable, it is not impossible
+that a second colony may be formed, and that the queen may depart at the
+head of it within three weeks of conducting the first swarm. But I
+repeat, the fact is rare in our climate. Let me now return to the hives
+from which the queen has led the first colony.
+
+Thirdly. _After the old queen has conducted the first swarm from the
+hive, the remaining bees take particular care of the royal cells, and
+prevent the young queens successively hatched from leaving them, unless
+at an interval of several days between each._
+
+In the preceding letter, I have given you the detail and proof of this
+fact, and I shall here add some reflexions. During the period of
+swarming, the conduct or instinct of bees seems to receive a particular
+modification. At all other times, when they have lost their queen, they
+appropriate workers worms to replace her; they prolong and enlarge the
+cells of these worms; they supply them with aliment more abundantly, and
+of a more pungent taste; and by this alteration, the worms that would
+have changed to common bees are transformed to queens. We have seen
+twenty-seven 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 may perhaps leave her cell, and attack all
+the other royal cells successively, which she will tear open to destroy
+her rivals, without the workers taking any part in their defence. Should
+several queens be hatched at once, they will pursue each other, and
+fight until the throne remain with her that is victorious. Far from
+opposing such duels, the other bees rather seem to excite the
+combatants.
+
+Things are quite reversed during the period of swarming. The royal cells
+then constructed are of a different figure from the former. They
+resemble stalactites, and in the beginning are like the cup of an acorn.
+The bees assiduously guard the cells when the young queens are ready for
+their last metamorphosis. At length the female hatched from the first
+egg laid by the old queen leaves her cell; the workers at first treat
+her with indifference. But she, immediately 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
+drive her away, so that she is forced to remove; but the royal cells
+being numerous, scarce can she find a place of rest. Incessantly
+harassed with the desire of attacking the other queens, and incessantly
+repelled, she becomes agitated, and hastily traverses the different
+groupes of workers, to which she communicates her agitation. At this
+moment numbers of bees rush towards the aperture of the hive, and, with
+the young queen at their head, depart to seek another habitation.
+
+After the departure of the colony, the remaining workers set another
+queen at liberty, and treat her with equal indifference as the first.
+They drive her from the royal cells; being perpetually harrassed, she
+becomes agitated; departs, and carries a new swarm along with her. In a
+populous hive this scene is repeated three or four times during spring.
+As the number of bees is so much reduced, that they are no longer
+capable of preserving a strict watch over the royal cells, several
+females then leave their confinement at once; they seek each other,
+fight, and the queen at last victorious 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 after the first colony is led out by the old queen,
+until the next swarm is conducted by the first young queen set at
+liberty. The interval between the second and third is still shorter; and
+the fourth sometimes departs 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 the day when the sun shines, and the air is calm. Sometimes
+we have observed all the precursors of swarming, disorder and agitation,
+but a cloud passed before the sun, and tranquillity was restored; the
+bees thought no more of swarming. An hour afterwards, the sun having
+again appeared, the tumult was renewed; it rapidly augmented; and the
+swarm departed.
+
+Bees generally seem much alarmed at the prospect of bad weather. While
+ranging in the fields the passage of a cloud before the hive induces
+them precipitately to return. I am induced to think they are disquieted
+by the sudden diminution of light. For if the sky is uniformly obscured,
+and there is no alteration in clearness or in the clouds dispelling,
+they proceed to the fields for their ordinary collections, and the first
+drops of a soft rain does not 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 sometimes seem to use this right in an arbitrary manner. However
+the confinement of the queens is always longer when bad weather lasts
+several days together. Here the final object cannot be mistaken. If the
+young females were at liberty to leave their cradles during these bad
+days, there would be a plurality of queens in the hive, consequently
+combats; and victims would fall. Bad weather might continue so long,
+that all the queens might at once have undergone their last
+metamorphosis, or attained their liberty. One victorious over the whole
+would enjoy the throne, and the hive, which should naturally produce
+several swarms, could give only one. Thus the multiplication of the
+species would have been left to the chance of rain, or fine weather,
+instead of which it is rendered independent of either, by the wise
+dispositions of nature. By allowing only a single female to escape at
+once, the formation of swarms is secured. This explanation appears so
+simple, that it is superfluous to insist farther on it.
+
+But I should mention another important circumstance resulting from the
+captivity of queens; which is, that they are in a condition to fly, when
+the bees have given them liberty, and by this means are capable of
+profiting by the first moment of sunshine to depart at the head of a
+colony.
+
+You well know, Sir, that all drones and workers are not in a condition
+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 hours must elapse before the acquisition of perfect strength, and
+the development of all their faculties. It would be the same with the
+females was not their confinement protracted after the period of
+transformation; 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 used to retain the queens in
+captivity. The bees solder the covering to the sides of the cell by a
+cordon 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 according to their age. Immediately when the royal
+cells were sealed, we marked them all with numbers, and we chose this
+period because it indicated the age of the queens exactly. The oldest
+was first liberated, then the one immediately younger, and so on with
+the rest. None of the younger queens were set at liberty before the
+older ones.
+
+I have a thousand times asked myself how the bees could so accurately
+distinguish the age of their captives. Undoubtedly I should do better to
+answer this question by a simple avowal of my ignorance. At the same
+time, I must be permitted to state a conjecture. You will admit, that I
+have not, as some authors, abused the right of giving myself up to
+hypothesis; may not the humming 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 queens? It is certain that the female, whose
+cell is first sealed, is also the first to emit this sound. That in the
+next emits it sooner than the rest, and so on with those immediately
+subsequent. As their captivity may continue six days, it is possible
+that the bees in this space of time may forget which has emitted it
+first; but it is also possible, that the queens diversify the sounds,
+encreasing the loudness as they become older, and that the bees can
+distinguish these variations. We have even ourselves been able to
+distinguish differences in the sound, either with relation to the
+succession of notes, or their intensity; and probably there are
+gradations still more imperceptible that escape our organs, but may be
+sensible to those of the workers.
+
+What gives weight to this conjecture is, that the queens brought up by
+M. Schirach's method, are perfectly mute; neither do the workers form
+any guard around their cells, nor do they retain them in captivity a
+moment beyond the period of transformation, and, when they have
+undergone it, they are allowed to combat until one has become
+victorious over all the rest. Why? Because the object is only to replace
+the last queen. Now, provided that among the worms reared as queens,
+only one succeeds, the fate of the others is uninteresting to the bees,
+whereas, during the period of swarming, it is necessary to preserve a
+succession of queens, for conducting the different colonies; and to
+ensure the safety of the queens, it is necessary to avert the
+consequences of the mutual horror by which they are animated against
+each other. Behold the evident cause of all the precautions that bees,
+instructed by nature, take during the period of swarming; behold an
+explanation of the captivity of females; and that the duration of their
+captivity 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 have never been able to discover the
+situation of the organ which produces the sound. But I have instituted a
+new course of experiments on the subject, which are still unfinished.
+
+Another problem still remains for solution. Why are the queens reared,
+according to M. Schirach's method, mute, whilst those bred in 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 M. Schirach's method,
+receive it only the second or third day of their existence. It appears
+to me that this circumstance may have an influence on the different
+parts of organisation, and particularly on the organ of voice.
+Experiment has not confirmed this conjecture. I constructed glass cells
+in perfect imitation of royal cells, that the metamorphosis of the worms
+into nymphs, and of the nymphs to queens, might be visible. These
+experiments are related in a preceding letter. Into one of these
+artificial cells we introduced the nymph of a worm, reared according to
+M. Schirach's method, twenty-four hours before it could naturally
+undergo its last metamorphosis; and we replaced the glass cell in the
+hive, that the nymph might have the necessary degree of heat. Next 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 had contrived an aperture for her thrusting out her
+trunk, and that the bees might feed her. I expected that she would have
+been completely mute; but it was otherwise; for she emitted sounds
+similar to those already described, therefore my conjecture was
+erroneous.
+
+I next conceived that the queen being restrained in her motions, and in
+her desire for liberty, was induced to emit certain sounds. All queens,
+in this new point of view, are equally capable of emitting the sound,
+but to induce them to it, they must be in a confined situation. In the
+natural state, the queens that come from workers are not a single
+instant in restraint; 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 are 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 something more probable.
+
+I do not ascribe to myself the credit of having discovered the humming
+of the queen bee. Old authors speak of it. M. de Reaumur cites a Latin
+work published 1671, _Monarchia Femina_, by Charles Butler. He gives a
+very brief abstract of this naturalist's observations, who we easily see
+has exaggerated or rather disguised the truth, by mixing it with the
+most absurd fancies; but it is not the less evident that Butler has
+heard this peculiar humming of queens, and that he did not confound it
+with the confused humming sometimes heard in hives.
+
+Fourthly. _The young queens conducting swarms from their native hive are
+still in a virgin state._ The day after, being settled in their new
+abode, they generally depart in quest of the males; and this is usually
+the fifth day of their existence as queens; for two or three pass in
+captivity, one in their native hive, and a fifth in their new dwelling.
+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 they begin to lay. The old queen,
+which leads the first swarm in spring, requires no farther commerce with
+the males, for preservation of her fecundity. A single copulation is
+sufficient to impregnate all the eggs she will lay for at least two
+years.
+
+  _PREGNY, 8. September 1791._
+
+
+FOOTNOTES:
+
+{M} Schirach seems to have been aware of this fact.--T.
+
+
+
+
+LETTER XI.
+
+_THE SAME SUBJECT CONTINUED._
+
+
+I have collected my chief observations on swarms in the two preceding
+letters; those most frequently repeated, and of which the uniformity of
+result leads me to apprehend no error. I have deduced what seem the most
+direct consequences; and in all the theoretical part, I have sedulously
+avoided going beyond facts. What is yet to be mentioned is more
+hypothetical, but it engrosses several curious experiments.
+
+It has been demonstrated, that the principal motive of the young females
+departing when hives swarm, is their insuperable antipathy to each
+other. I have repeatedly observed that they cannot gratify their
+aversion, because the workers with the utmost care prevent them from
+attacking the royal cells. This perpetual opposition at length creates a
+visible inquietude, and excites a degree of agitation that induces 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 different. Always
+accustomed to respect fertile queens, they do not forget what they owe
+to her; they allow her the most uncontrouled liberty. She is permitted
+to approach the royal cells; and if she even attempts to destroy them,
+no opposition is presented 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. Therefore, I candidly confess
+myself ignorant of the motives of her departure.
+
+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 young ones, have the greatest aversion to 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 departure of
+old queens, seven royal cells opened at one side were destroyed by the
+queen. If rain continues several days, the whole are destroyed; in this
+case, there is no swarm, which too often happens in our climate, where
+spring is generally rainy. Queens never attack cells containing an egg
+or a very young worm; but only when the worm is ready for transforming
+to a nymph, or when it has undergone its last metamorphosis.
+
+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 is 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
+immediate success, her inquietude during this work becomes a terrible
+agitation. If the weather continues favourable, while she remains in
+this condition, she is naturally disposed to depart.
+
+It may easily be understood, that the workers 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
+respect. But you will undoubtedly ask, Sir, What motive can induce the
+workers to follow their queen from the hive, while they treat the young
+queens very ill, and, even in their most amicable moments, testify
+perfect indifference towards them. Probably it is to escape the heat to
+which the hive is then exposed. The extreme agitation of the females
+leads them to traverse the combs in all directions. They pass through
+groupes of bees, injure and derange them; they communicate a kind of
+delirium, and these tumultuous motions raise the temperature to an
+insupportable degree. We have frequently proved it by the thermometer.
+In a populous hive it commonly stands between 92 deg. and 97 deg., in a
+fine day of spring; but during the tumult which precedes swarming, it
+rises above 104 deg. And this is heat intolerable to bees. When exposed
+to it, they rush impetuously towards the outlets of the hive and depart.
+In general they cannot endure the sudden augmentation of heat, and in
+that case quit their dwelling; neither do those returning from the
+fields enter when the temperature is extraordinary.
+
+I am certain, from direct experiments, that the impetuous courses of the
+queen over the combs, actually throws the workers into agitation; and I
+was able to ascertain it in the following manner. I wished to avoid a
+complication of causes. It was particularly important to learn, whether
+the queen would impart her agitation but not at the time of swarming.
+Therefore I took two females still virgins, but capable of fecundation
+for above five days, and put one in a glass hive sufficiently populous;
+the other I put into a different hive of the same kind. Then I shut the
+hives in such a way that there was no possibility of their escape. The
+air had free circulation. I then prepared to observe the hives every
+moment that the fineness of the weather would invite both males and
+females to go abroad, for the purpose of fecundation. Next morning, the
+weather being gloomy, no male left the hive, and the bees were tranquil;
+but towards eleven of the following day, the sun shining bright, both
+queens began to run about seeking an exit from every part of their
+dwelling; 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 that part of the
+hive where the openings were placed; unable to escape they ascended with
+equal rapidity, and ran heedlessly over the cells until four in the
+afternoon. It is nearly about this time that the sun declining in the
+horizon recalls the males; queens requiring fecundation never remain
+later abroad. The two females became calmer, and tranquillity was in a
+short time restored. This was repeated 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
+always in a tumultuous state when the queen herself is in agitation.
+
+I have but one fact more to mention. It has already been observed, that
+on losing the female, bees give the larvae of simple workers the royal
+treatment, and, according to M. Schirach, in five or six days they
+repair the loss of their queen. In this case there are no swarms. All
+the females leave their cells almost at the same moment, and after a
+bloody combat the throne remains with the most fortunate.
+
+I can very well comprehend that the object 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 metamorphosis almost at
+the same time? Since they are enabled to retain the young females in
+their cells, why do they allow all the queens, reared according to
+Schirach's method, to escape at once. By prolonging their captivity
+more or less, they would fulfil two most important objects at once, in
+repairing the loss of their females and preserving a succession of
+queens to conduct several swarms.
+
+At first it was my opinion, that this difference of conduct proceeded
+from the difference of circumstances in which they found themselves
+situated. They are induced to make all their dispositions relative to
+swarming only when in great numbers, and when they have a queen occupied
+with her principal laying of male eggs; whereas, having lost their
+female, the eggs of drones are no longer in the combs to influence their
+instinct. They are in a certain degree restless and discouraged.
+
+Therefore, after removing the queen from a hive, I thought of rendering
+all the other circumstances as similar as possible to the situation of
+bees preparing to swarm. By introducing a great many workers, I
+encreased the population to excess, and supplied them with combs of
+male brood in every stage. Their first occupation was to construct royal
+cells after Schirach's method, and to rear 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 this they discontinued, as there was
+no queen to deposit her eggs. Finally, I gave them several close royal
+cells, taken indifferently from hives preparing to swarm. However, all
+these precautions were fruitless; the bees were occupied only with
+replacing their lost queen; they neglected the royal cells entrusted to
+their care; the included queens came out at the ordinary time, without
+being detained prisoners a moment; they engaged in several combats, and
+there were no swarms.
+
+Recurring to subtleties, we may perhaps suggest a cause for this
+apparent contradiction. 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 beautiful system, and the more must we distrust that
+facility of imagination from which we think by embellishment to
+elucidate facts.
+
+In general, Naturalists who have long observed animals, and those in
+particular who have chose insects for their favourite study, have too
+readily ascribed to them our sentiments, our passions, and even our
+intentions and designs. Incited 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 painted different traits of the industry of these
+minute animals, with the colours inspired by an exalted imagination. Nor
+is even the celebrated Reaumur to be acquitted of such a charge. He
+frequently ascribes combined intentions to bees; love, anticipation, and
+other faculties of too elevated a kind. I think I can perceive that
+although he formed very just ideas of their operations, he would be
+well pleased that his reader should admit they were sensible of their
+own interests. He is a painter who by a happy interest 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 association of ideas _simply sensible_. It is
+not my object here to penetrate those depths, or to insist on the
+details.
+
+But, on the whole, facts relative to the formation of swarms perhaps
+present more subjects of admiration than any other part of the history
+of bees. I think it proper to state, in a few words, the simplicity of
+the methods by which the wisdom of nature guides their instinct. It
+cannot allow them the slightest portion of understanding; it leaves them
+no precautions to be taken, no combination to be followed, no foresight
+to exercise, no knowledge to acquire. But having adapted their
+sensorium to the different operations with which they are charged, it is
+the impulse 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 larvae, and collect
+provisions, we must not seek for method, affection, or foresight. The
+only inducement must be sought for in some pleasing sensation attached
+to each of these operations. I address a philosopher; 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 works which treats of the industry of animals. Let me
+add but another sentence. The inducement of pleasure is not the sole
+agent; there is another principle, the prodigious influence of which, at
+least with regard to bees, has hitherto been unknown, that is the
+sentiment of aversion which all females continually feel against each
+other, a sentiment whose existence is so fully demonstrated by my
+experiments, and which explains many important facts in the theory of
+swarms.
+
+  _PREGNY, 10. September 1791._
+
+
+
+
+LETTER XII.
+
+_ADDITIONAL OBSERVATIONS ON QUEENS THAT LAY ONLY THE EGGS OF DRONES, AND
+ON THOSE DEPRIVED OF THE ANTENNAE._
+
+
+In relating my first observations on queens that lay male eggs alone, I
+have proved that they lay 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 workers appeared defective. It is
+indeed most singular, that bees which know the worms of males so well
+when the eggs are laid in small cells, and never fail to give them a
+convex 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
+irregularity depends on something I cannot comprehend.
+
+In revising what is said on this subject, I observe still wanting an
+interesting experiment to complete the history of queens that lay only
+the eggs of drones. I had to investigate whether these females could
+themselves distinguish that the eggs they deposit in the royal cells
+would not produce queens. I have already observed that they do not
+endeavour to destroy these cells when close, and I thence concluded,
+that in general the presence of royal cells in their hive does not
+inspire them with the same aversion to females whose fecundation has
+been retarded; but to ascertain the fact more correctly, it was
+essential to examine how the presence of a cell containing a royal nymph
+would affect a queen that had never laid any other than the eggs of
+drones.
+
+This experiment was easy; and I put it in practice on the fourth of
+September, in a hive some time deprived of its queen. The bees had not
+failed to construct several royal cells for replacing their females. I
+chose this opportunity for supplying them with a queen, 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. One remaining was
+enough to shew the impression it would make on the stranger queen
+introduced; had she endeavoured to destroy it; this, in my opinion,
+would have proved that she anticipated the origin of a dangerous rival.
+You must admit the use I make of the word anticipate; it saves a long
+paraphrase; I feel the impropriety of it. If, on the contrary, she did
+not attack the cell I would thence conclude that the delay of
+fecundation, which deprived her of the power of laying workers eggs, had
+also impaired her instinct. This was the fact; the queen passed several
+times over the royal cell, both 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 bestowed 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 honey 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 is no longer preserved against their own
+sex in the nymphine state, nor do they longer 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, should be so well
+treated and become as dear to them as females laying both kinds of eggs.
+But I remember to have observed a fact more astonishing still. I have
+seen workers bestow every attention on a queen though sterile; and after
+her death treat her dead body as they had treated herself when alive,
+and long prefer this inanimate body to the most fertile queens I had
+offered them. This sentiment, which assumes the appearance of so lively
+an affection, is probably the effect of some agreeable sensation
+communicated to bees by their queen, independent of fertility. Those
+laying only the eggs of males probably excite the same sensation in the
+workers.
+
+I now recollect that the celebrated Swammerdam somewhere observes, that
+when a queen is blind, sterile, or mutilated, she ceases to lay, and the
+workers of her hive no longer labour or make any collections, as if
+aware that it was now useless to work. He cites no experiment that led
+him to the discovery. Those made by myself have afforded some very
+singular results.
+
+I frequently amputated the four wings of queens; and not only did they
+continue laying, but the same confederation of them was testified by the
+workers as before. Therefore, Swammerdam has no foundation for
+asserting, that mutilated queens cease to lay. Indeed, from his
+ignorance of fecundation taking place without the hives, it is possible
+he cut the wings off virgin queens, and they, becoming incapable of
+flight, remained sterile from inability to seek the males in the air.
+Thus, amputation of the wings does not produce sterility in queens.
+
+       *       *       *       *       *
+
+I have frequently cut off one antennae 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 one still remained, the mutilation was imperfect; and the
+experiment decided nothing. But amputation of both antennae produced most
+singular effects. On the fifth of September, I cut both 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 vivacity. Scarcely
+had the workers time to separate and recede before her; she dropped her
+eggs, without attending to deposit them in any cell. The hive not being
+very populous, part was without comb. Hither she seemed particularly
+earnest to repair, and long remained motionless. She appeared to avoid
+the bees; however, several workers followed her into this solitude, and
+treated her with the most evident respect. She seldom required honey
+from them, but, when that occurred, 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 appeared tormented with the desire of leaving her
+habitation. She rushed towards the opening, and entered the glass tube
+adapted there; but the external orific being too small, after fruitless
+exertion, she returned. Notwithstanding these symptoms of delirium, the
+bees did not cease to render her the same attention as they ever pay to
+their queens, but this one received it with indifference. All that I
+describe appeared to me the consequence of amputating the antennae.
+However, her organization having already suffered from retarded
+fecundation, and as I had observed her instinct in some degree impaired,
+both causes might possibly concur in producing the same effect. To
+distinguish properly what belonged to the privation of the antennae, a
+repetition of the experiment was necessary, in a queen otherwise well
+organised, and capable of laying both kinds of eggs.
+
+This I did on the sixth of September. I amputated both the antennae of a
+female which had been several months the subject of observation, and
+being of great fecundity 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 same marks 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 considered the manner in which these two
+mutilated queens treated each other. You cannot have forgot, Sir, the
+animosity with which queens, possessing all their organs, combat, on
+which account it became extremely interesting to learn whether they
+would experience the same reciprocal aversion after losing their
+antennae. We studied these queens a long time; they met several times in
+their courses, and without exhibiting the smallest resentment. This last
+instance is, in my opinion, the most complete evidence of a change
+operated in their instinct.
+
+Another very remarkable circumstance, which this experiment gave me
+occasion to observe, consists in the good reception given by the bees to
+the stranger queen, while they still preserved the first. Having so
+often seen the symptoms of discontent that a plurality of queens
+occasions, after having witnessed the clusters formed around these
+supernumerary queens to confine them, I could not expect they would pay
+the same homage to a second mutilated one they still testified towards
+the first. Is it because after losing the antennae, these queens have no
+more any characteristic which distinguishes the one from the other?
+
+I was the more inclined to admit this conjecture from the bad reception
+of a third fertile queen preserving her antennae, which was introduced
+into the same hive. The bees seized, bit her, and confined her so
+closely, that she could hardly breath or move. Therefore, if they treat
+two females deprived of antennae in the same hive equally well, it is
+probably because they experience the same sensation from these two
+females, and want the means of longer distinguishing them from each
+other.
+
+From all this, I conclude, that the antennae are not a frivolous ornament
+to insects, but, according to all 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 organised in such a manner as to fulfil
+both functions at once.
+
+As in the course of this experiment both mutilated females constantly
+endeavoured 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 I removed the first and third queen from the hive,
+leaving the fertile mutilated one, and enlarged the entrance.
+
+The queen left her habitation the same day. At first she tried to fly,
+but, her belly being full of eggs, she fell down and never attempted it
+again. No workers accompanied her. Why, after rendering the queen so
+much 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, carrying 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 I can hazard on this question is a conjecture.
+It appears that bees are induced to quit the hives from the increased
+heat which occasions the agitation of their queen, and the tumultuous
+motion which she communicates to them. A mutilated queen,
+notwithstanding her delirium, does not agitate the workers, because she
+seeks the uninhabited parts of the hive, and the glass panes of it: she
+hurries over clusters of bees, but the shock resembles that of any other
+body, and produces only a local and instantaneous motion. The agitation
+arising from it, is not communicated from one place to another, like
+that produced by a queen, which in the natural state wishes to abandon
+her hive and lead out a swarm; there is no increased heat, consequently
+nothing that renders the hive insupportable to her.
+
+This conjecture, which affords a tolerable explanation why bees persist
+in remaining in the hive, though the mutilated queen has left it, is no
+reason for the motive inducing the queen herself to depart. Her instinct
+is altered; that is the whole that I can perceive. I can discern nothing
+more. It is very fortunate for the hive, that this queen departs, for
+the bees incessantly attend her; nor do they ever think of procuring
+another while she remains; and if she was long of leaving them, it would
+be impossible to replace her; for the workers worms would exceed the
+term at which they are convertible into royal worms, and the hive would
+perish. Observe, that the eggs dropped by the mutilated queen can never
+serve for replacing her, for, not being deposited in cells, they dry and
+produce nothing.
+
+I have yet to say a few words on females laying male eggs only. M.
+Schirach supposes that one branch of their double ovary suffers some
+alteration. He seems to think that one of these branches contains the
+eggs of males, while the other has none but common eggs, and as he
+ascribes the inability of certain queens to lay the latter to some
+disease, his conjecture becomes very plausible. In fact, if the eggs of
+males and workers are indiscriminately mixed in both branches of the
+ovary, it appears at first sight that whatever cause acts on that organ,
+it should equally affect both species of eggs. If on the contrary, one
+branch is occupied by the eggs of drones only, and the other contains
+none but common eggs, we may conceive how disease affects the one, while
+the other remains untouched. Though this conjecture is probable, it is
+confuted by observation. We lately dissected queens, which laid none but
+male eggs, and found both branches of the ovary equally well expanded,
+and equally sound, if I may use the expression. The only difference
+that struck us was that in these two branches, the eggs were apparently
+not so close together as in the ovaries of queens laying both kinds of
+eggs.
+
+  _PREGNY, 12. September 1791._
+
+
+
+
+LETTER XIII.
+
+_ECONOMICAL CONSIDERATIONS ON BEES._
+
+
+In this letter I shall treat of the advantages that may be derived from
+the new invented hives, called _book_ or _leaf_ hives, in promoting the
+_economical knowledge_ of bees.
+
+It is needless to relate the different methods hitherto employed in
+forcing bees to yield up a portion of their honey and wax; all resemble
+each other in being cruel and ill understood.
+
+It is evident, when bees are cultivated for the purpose of sharing the
+produce of their labours, we must endeavour to multiply them as much as
+the nature of the country admits; and consequently to regard their lives
+at the time we plunder them. Therefore it is an absurd custom to
+sacrifice whole hives to get at the riches they contain. The inhabitants
+of this country, who follow no other method, annually lose immense
+numbers of hives; and spring, being generally unfavourable to swarms,
+the loss is irreparable. I well know that at first they will not adopt
+any other method; 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. Though the bees are indifferent as
+to the position of their combs, and as to their greater or lesser size,
+they are obliged to construct them perpendicular to the horizon, and
+parallel to each other. Therefore, if left entirely to themselves, when
+establishing a colony in one of those new hives, they would frequently
+construct several small combs parallel indeed, but perpendicular to the
+plane of the frames or leaves, and by this disposition prevent the
+advantages which I think to derive from the figure of my hives, since
+they could not be opened without breaking the combs. Thus they must
+previously have a guide to follow; the cultivator himself lays the
+foundation of their edifices, and that by a simple method. A portion of
+comb must be solidly fixed in some of the boxes composing the hive; the
+bees will extend it; and, in prosecution of their work, will accurately
+follow the plan already given them. Therefore on opening the hive, no
+obstacle is to be removed, nor stings to be dreaded, for one of the
+most singular and valuable properties attending this construction, 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, and the tranquillity of the bees has given you great
+surprise. I can desire no other evidence of my assertion. It is in the
+facility of opening these hives at pleasure that all the advantages lie,
+which I expect in perfecting the economical knowledge of bees.
+
+I conceive, when I observe bees may be rendered tractable, that it need
+not be added, I do not arrogate to myself the absurd pretence of
+_taming_ them, for this excites a vague idea of tricks; and I would
+willingly avoid the hazard of exposing myself to any such reproach. I
+ascribe their tranquillity on opening the hives, to the manner that the
+sudden introduction of light affects them; then, they seem rather to
+testify fear than anger. Many retire and enter the cells, and appear to
+conceal themselves. What confirms my conjecture is, 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. The divisions must be separated slowly, and
+care taken not to wound any of the bees. If they cluster too much on the
+combs, they must be brushed off with a feather; and breathing on them
+carefully avoided. The air we expire seems to excite their fury; it
+certainly has some irritating quality, for if bellows are used, they are
+rather disposed to escape than to sting.
+
+Respecting the advantages of leaf hives, I shall observe, they are very
+convenient for forming _artificial_ swarms. In the history of natural
+swarms, I have shewn how many favourable circumstances are necessary for
+their success. From experience I know that they very often fail in our
+climate; and even when a hive is disposed to swarm, it frequently
+happens that the swarm is lost either because the moment of its
+departure has not been foreseen, because it rises out of sight, or
+settles on inaccessible places. Instructing the cultivators of bees how
+to make artificial swarms is a real service, and the form of my hives
+renders this an easy operation. But it requires farther illustration.
+
+Since bees, according to M. Schirach's discovery, can procure another
+queen after having lost their own, provided there is workers brood in
+the combs not above three days old, it results that we can at pleasure
+produce queens, by removing the reigning one. Therefore, if a hive
+sufficiently populous is divided in two, one half will retain the old
+queen, and the other will not be long of obtaining 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
+sufficient to admit of division, if the brood is of the proper age, if
+males exist or are ready to be produced for impregnating the young
+queens.
+
+Supposing the union of all these conditions, the following is the method
+to be pursued. The leaf hive may be divided through the middle without
+any shock. Two empty divisions may be insinuated between the halves,
+which, when exactly applied to each other, are close on the outside. The
+queen must be sought in one of the halves, and marked to avoid mistake.
+If she by chance remains in the division with most brood, she is to be
+transferred to the other with less, that the bees may have every
+possible opportunity 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 occupied. The old entrance, now become useless, will be shut
+up; but as each half requires a new one, it ought to be made at the
+bottom of each division, on purpose that they may be as far asunder as
+possible. Both entrances should not be made on the same day. The bees in
+the half deprived of the queen ought to be confined twenty-four hours,
+and no opening made before then except for admission of air. Without
+this precaution, they would soon search for their queen, and infallibly
+find her in the other division. They will then retire in great numbers
+from their own division, until too few remain to perform the necessary
+labours. But this will not ensue if they are confined twenty-four hours,
+provided that interval is sufficient to make them forget the queen. When
+all these circumstances are favourable, the bees, in the division
+wanting the queen, will the same day begin to labour in procuring
+another, and ten or fifteen days after the operation, their loss will be
+repaired. The young female they have reared, soon issues forth to seek
+impregnation, and in two days commences the laying of workers eggs.
+Nothing more is wanting to the bees of this half hive, and the success
+of the artificial swarm is ensured.
+
+It is to M. Schirach that we are indebted for this ingenious method of
+forming swarms. He supposes, by producing young queens early in spring,
+that early swarms might be procured, which would certainly be
+advantageous in favourable circumstances. But unfortunately this is
+impossible. Schirach believed that queens were impregnated 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 become 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. Thus, if a
+swarm were artificially formed before the usual time of the males
+originating, the bees would be discouraged by the sterility of the young
+female. Or should they remain faithful to her, awaiting the period of
+fecundation, as she could not for three or four weeks receive the
+approaches of the male, she would lay eggs producing males only, and the
+hive in this case would perish. Thus the natural order must not be
+deranged, but we must delay the division of hives until males are about
+to originate or actually exist.
+
+Besides, if M. Schirach did succeed in obtaining artificial swarms,
+notwithstanding the great inconvenience of his hives, it was owing to
+his singular address and unremitting assiduity. He had some pupils in
+the art; these communicated the method of forming artificial swarms to
+others, and there are people now in Saxony who traverse the country
+practising this operation. Those versant in the matter can alone venture
+to undertake it with common hives, whereas, every cultivator can do it
+himself with the leaf hives.
+
+In this construction, another very great advantage will also be found.
+Bees can be forced to work in wax. Here I am led to what I believe is a
+new observation. While naturalists have directed our admiration to the
+parallel position of the combs, they have overlooked another trait in
+the industry of bees, namely, the equal distance uniformly between them.
+On measuring the interval separating the combs, it will generally be
+found four lines. Were they too distant, it is very evident the bees
+would be much dispersed and unable to communicate their heat
+reciprocally; whence the brood would not be exposed to sufficient
+warmth. Were the combs too close, on the contrary, the bees could not
+freely traverse the intervals, and the work of the hive would suffer.
+Therefore, a certain distance always uniform is requisite, which
+corresponds equally well with the service of the hive, and the care
+necessary for the worms. Nature, which has taught bees so much, has
+instructed them regularly to preserve this distance. At the approach of
+winter, they sometimes elongate the cells which are to contain the
+honey, and thus contract the intervals between the combs. But this
+operation is a preparation for a season, when it is important to have
+plentiful magazines, and when their activity being very much relaxed, it
+is unnecessary for their communications to be so spacious and free. On
+the return of spring, the bees hasten to contract these elongated cells,
+that they may become fit for receiving the eggs which the queen will
+lay, and thus re-establish the just distance which nature has ordained.
+
+This being admitted, bees may be forced 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
+build others in the interval. Suppose an artificial swarm is lodged in a
+leaf hive, composed of six divisions, each containing a comb, if the
+young queen is as fertile as she ought to be, the bees will be very
+active in their labours, and disposed to make great collections of wax.
+To induce them towards this an empty box or division must be placed
+between two others, each containing a comb. As all the boxes are of
+equal dimensions, and of the necessary width for receiving a comb, the
+bees having sufficient 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 having more than four lines
+between them. Without any guide, this new comb will be parallel to the
+old ones, to preserve that law which establishes an equal distance
+throughout the whole.
+
+If the hive is strong and the weather good, three empty divisions may at
+first be left between the old combs; one between the first and second,
+another between the third and fourth, and the last between the fifth and
+sixth. The bees will fill them in seven or eight days, and the hive then
+contains nine combs. Should the temperature of the weather continue
+favourable, three new leaves or divisions may be introduced;
+consequently 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 where flowers perpetually blow. But in our
+country, I have reason to think that the labour should not be forced
+more during the first year.
+
+From these details, you are sensible, Sir, how preferable _leaf hives_
+are to those of any other construction, and even to those ingenious
+stages described by _M. Palteau_, for the bees cannot by means of them
+be forced to labour more in wax than they would do if left to
+themselves; whereas, they are obliged to do it by inserting empty
+divisions. Next, the combs constructed on those stages cannot be removed
+without destroying considerable portions of brood, deranging the bees,
+and creating real disorder in the hive.
+
+Mine have also this advantage, that what passes within may daily be
+observed, and we may judge of the most favourable moments for depriving
+the bees of part of their stores. With all the combs before us we can
+distinguish those containing brood only, and what it is proper to
+preserve. The scarcity or abundance of provisions is visible, and the
+portion suitable may be taken away.
+
+I should protract this letter too much, if I gave an account of all my
+observations 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 their riches with them. The subject would require a separate
+work; and I may perhaps one day engage in it; but until that arrives I
+shall always feel gratification in communicating to cultivators, who
+wish to follow my method, directions of which long practice has
+demonstrated the utility.
+
+Here I shall only observe, that we hazard absolute ruin of the hives,
+by robbing them of too great a proportion of honey and wax. In my
+opinion, the art of cultivating these animals consists in moderately
+exercising the privilege of sharing their labours; but as a compensation
+for this, every method must be employed which promotes the
+multiplication of bees. Thus, for example, if we desire to procure a
+certain quantity of honey and wax annually, it will be better to seek it
+in a number of hives, managed with discretion, than to plunder a few of
+a great proportion of their treasures.
+
+It is indubitable that the multiplication of these industrious animals
+is much injured by privation of several combs, in a season unfavourable
+to the collection of wax, because the time consumed in replacing them is
+taken from that which should be consecrated to the care of the eggs and
+worms, and by this means the brood suffers. Besides, they must always
+have a sufficient provision of honey left for winter, for although less
+is consumed during this season, they do consume some; because they are
+not torpid, as some authors have conceived.{N} Therefore if they have
+not enough, they must be supplied with it, which requires great
+exactness. I admit that in determining to what extent hives may be
+multiplied in a particular country, it is necessary first to know how
+many the country can support, which is a problem yet unsolved. It also
+depends on another, the solution of which is as little known, namely the
+greatest distance that bees fly in collecting their provender. Different
+authors maintain, they can fly several leagues from the hive. But by the
+few observations I have been able to make, this distance seems greatly
+exaggerated. It appears to me that the radius of the circle they
+traverse does not exceed half a league. As they return to the hive with
+the greatest precipitation whenever a cloud passes before the sun, it is
+probable they do not fly far. Nature which has inspired them with such
+terror for a storm, and even for rain, undoubtedly 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 distances bees with the thorax painted, that they might again
+be recognised. But none ever returned that I had carried for twenty-five
+or thirty minutes from their dwelling, while those at a shorter distance
+have found their way and returned. I do not state this experiment as
+decisive. Though bees do not generally fly above half a league, it is
+very possible they go much farther, when flowers are scarce in their own
+vicinity. A conclusive experiment must be made in vast arid or sandy
+plains, separated by a known distance from a fertile region.
+
+Thus, the question yet remains undecided. But without ascertaining the
+number of hives that any district can maintain, I shall remark that
+certain vegetable productions are much more favourable to bees than
+others. More hives, for example, may be kept in a country abounding
+meadows, and where black grain is cultivated, than in a district of
+vineyards or corn.
+
+       *       *       *       *       *
+
+Here 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. Several problems concerning the history of these
+animals still remain unsolved. The experiments I project may perhaps
+throw some light on them; and I shall be animated 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 HUBER.
+  _PREGNY, 1. October 1791._
+
+
+FOOTNOTES:
+
+{N} So far from being torpid in winter, when the thermometer in the open
+air is several degrees below freezing, it stands at (86) and (88 deg.),
+in hives sufficiently populous. The bees then cluster together, and move
+to preserve their heat.
+
+Now that I am on the subject of thermometrical observations, I may
+cursorily remark, that M. Dubois of Bourg en Bresse, in a memoir
+otherwise valuable, is of opinion, that the larvae cannot be hatched
+below (104). I have repeatedly made the experiment with the most
+accurate thermometers, and obtained a very different result. When the
+thermometer rises to (104 deg.), the heat is so much greater than the
+eggs require, that it is intolerable to the bees. M. Dubois has been
+deceived, I imagine, by too suddenly introducing his thermometer into a
+cluster of bees, and putting them in agitation, the mercury has rose
+higher than it should naturally do. Had he delayed introducing the
+thermometer, he would soon have seen it fall to between 95 and 97, which
+is the usual temperature of hives in summer. In August this year, when
+the thermometer in the open air stood at 94, it did not rise above 99 in
+the most populous hives. The bees had little motion, and a great many
+rested on the board of the hive.
+
+
+
+
+APPENDIX.
+
+
+[The following passages are chiefly engrossed in the substance of the
+work, but the Translator, as has already been observed, for various
+reasons, judges it expedient to transfer them to an appendix. In his
+opinion these very minute details rather interrupt the connexion of the
+narrative, however interesting they may be considered, and they pertain
+more to researches purely anatomical.
+
+The Translator has likewise in some instances incorporated several long
+and important notes with the text; because it appears to him that they
+actually belong to the substance of the treatise. These are the only
+variations from the original with respect to arrangement.]
+
+       *       *       *       *       *
+
+Swammerdam has given an imperfect description of the ovary of the queen.
+He observes that he has never been able to find the termination of the
+oviducts in the abdomen, nor any other parts excepting those which he
+has described. "Notwithstanding all my exertions, I never could discover
+the site of the vulva, partly because I had not all my apparatus with me
+in the country, when investigating this subject, and partly from my
+apprehension of injuring other parts by pressure, which I had then
+occasion to examine. However, I have clearly observed a muscular
+swelling of the oviduct, where approaching the last ring of the belly;
+that it then contracts and afterwards dilates in becoming membranaceous.
+As I was desirous of preserving the poison bag, which is situated
+exactly here, along with, the muscles aiding the motion of the sting, I
+could follow the oviduct no farther. However, in another female, it
+appeared that the vulva is in the last ring of the abdomen, and under
+the sting. The parts expanding only while the queen lays, renders it
+extremely difficult to penetrate the aperture."
+
+We have attempted to discover what has escaped the indefatigable
+Swammerdam. But his observation that the research can be made to the
+greatest advantage, at the time of laying, has paved the way to us. We
+have remarked that the oviduct did not issue from the body, but that the
+eggs fall into a kind of cavity, where they are retained several seconds
+before being laid.
+
+On the sixth of August, we took a very fertile queen, and holding her
+gently by the wings in a supine position, the whole belly was exposed.
+She seized the extremity with her second pair of legs, and curved it as
+much as possible. This seeming an unfavourable position for laying, we
+forced her to stretch it out. The queen, oppressed with the necessity of
+laying, could no longer retain her eggs. The lower part of the last ring
+then separated so far from the upper part as to leave some of the inside
+discovered. In this cavity the sting lay above in its sheath. As the
+queen now made new efforts, we saw an egg fall into the cavity from the
+end of the oviduct. The lips then closed for several seconds; they
+opened again, and, in a much shorter time, dropped the egg from the
+cavity.
+
+From our own observations we found that the seminal fluid of drones
+coagulated on exposure to the air, and from several experiments had so
+little doubt on the subject, that whenever the female returned with the
+external marks of fecundation, we thought we recognised it in the
+whitish substance filling the sexual organs. It did not then occur to us
+to dissect the females to ascertain the fact more particularly: but this
+year, whether designing to neglect nothing, or to examine the distension
+of the female organs, we determined to dissect several. To our infinite
+surprise, what we had supposed the residue of the prolific fluid,
+actually proved the genital organs of the male, which separate from his
+body during copulation, and remain in the female.
+
+We procured a number of queens according to Schirach's method for the
+purpose of dissection, and set them at liberty that they might seek the
+males. The first which did so, was seized the instant she returned, and
+without dissection spontaneously exhibited what we were so impatient to
+behold. Examining the under part of the belly, we saw the oval end of a
+white substance which distended the sexual organs. The belly was in
+constant motion, by alternate extension and contraction. Already had we
+prepared to sever the rings, and by dissection to ascertain the cause of
+these motions; when the queen curving her belly very much, and
+endeavouring to reach its extremity with her hind legs, seized the
+distending substance with her claws, and evidently made an effort to
+extract it. She at last succeeded, and it fell before us. We expected a
+shapeless mass of coagulated fluid; what therefore was our surprise to
+find it part of the same male that had rendered this queen a mother. At
+first we could not credit our eyes; but after examining it in every
+position, both with the naked eye, and a powerful magnifier, we
+distinctly recognised it to be that part which M. de Reaumur calls the
+_lenticular_ body, or the _lentil_, in the following description.{O}
+
+    'Opening a drone there appears a portion formed by the assemblage of
+    several parts, often whiter than milk. This on investigation is
+    found to be principally composed of four oblong pieces. The two
+    largest are attached to a kind of twisted cord, fig. 4. r, called by
+    Swammerdam the root of the penis; and he has denominated seminal
+    vessels, s. s. two long bodies that we are about to consider. Other
+    two bodies oblong like the preceding, but shorter and not half the
+    diameter, he calls the _vasa deferentia_, d. d. Each communicates
+    with one of the seminal vessels near, g. g. where they unite to the
+    twisted cord, r. From the other extremity proceeds a very delicate
+    vessel, which, after several involutions, terminates in a body, t. a
+    little larger, but difficult to disengage from the surrounding
+    tracheae. Swammerdam considers these two bodies, t. t. the testicles.
+    Thus there are two parts of considerable size, communicating with
+    other two still thicker and longer. These four bodies are of a
+    cellular texture, and full of a milky fluid, which may be squeezed
+    out. This long twisted cord, r, to which the largest of the seminal
+    vessels is connected, this cord, I say, is doubtless the channel by
+    which the milky fluid issues. After several plications, it
+    terminates in a kind of bladder or fleshy sac, i. i. In different
+    males this part is of various length and flatness. By calling it the
+    _lenticular_ body, or the lentil, it receives a name descriptive of
+    the figure it presents in all males whose internal parts have
+    acquired consistency in spirit of wine. The body, l. i. is therefore
+    a lentil, a little thickened, of which one half, or nearly so, of
+    the circumference is edged along the outline by two chesnut coloured
+    scaly plates, e. i. A small white cord, the real edge of the lentil,
+    is visible, and separates them. This lentil is a little oblong, and,
+    for convenience, we shall ascribe two extremities to it, the
+    anterior and posterior. The anterior, l, next the head, is where the
+    canal, r, dividing the seminal vessels is inserted, and the opposite
+    part; i. next the anus, the posterior. The two scaly plates, e. i.
+    e. i, proceed from the vicinity of this last part, whence each
+    enlarges to cover part of the lentil. Under the broadest part of
+    each plate, there is a division formed by two soft points of unequal
+    length; the largest of which is on the circumference of the lentil.
+    Besides these two scaly plates, there are two others, n. n. of the
+    same colour, narrower, and fully one half shorter, each of which is
+    situated very near the preceding, and originates close to the origin
+    of that it accompanies, namely, at the posterior part of the lentil.
+    The rest of the lentil is white and membranaceous. From behind
+    proceeds a tube, k. a canal also white and membranaceous, but it is
+    difficult to judge of its diameter, for the membranes, of which it
+    consists, are evidently in folds. To one side of this pipe is
+    attached a fleshy part, p. somewhat pallet shaped, one side is
+    concave, and the edges plaited; the other side is convex. In certain
+    places the plaits rise and project from the rest of the outline, and
+    form a kind of rays; the pallet appears prettily figured. Though
+    lying with the concave side applied to the lentil, it is not fixed
+    to it. Swammerdam seems to consider this pallet as the
+    characteristic part of the male.
+
+    'Though the parts we have described are the most conspicuous in the
+    male, they are neither those which protrude first, nor when
+    protruded are the most remarkable. On viewing from the opposite edge
+    of the lentil, forming the division of the two great scaly plates, a
+    sac or canal, k. proceeding from the posterior part of the lentil,
+    there is distinctly visible the body u, which we call the arc; where
+    there are five transverse hairy bands of a yellow colour, while the
+    rest is white. This arc seems out of the membranaceous canal because
+    it is covered only by a very transparent membrane. One end almost
+    reaches the lenticular body, and the other terminates where the
+    membranaceous canal joins the folded yellow membranes, m. which form
+    a species of sac, that is applied to the sides of the aperture,
+    adapted for the genital organs passing through. These reddish
+    membranes are those that appear first on pressure, and form this
+    elongated portion, at whose end is a kind of hairy mask. Finally,
+    with the sac formed by the reddish membranes, there are connected
+    two appendages, c. c. of reddish yellow, and red at the end, s.
+    These are what appear externally like horns.{P}'
+
+The lenticular substance, l. i. provided with each scaly lamina, are the
+only parts of those described by M. de Reaumur, that we have found
+engaged in the organs of our queens. The canal, r, by Swammerdam
+denominated the root of the penis, breaks in copulation; and we have
+seen its fragments at the place where it unites to the end of the
+lentil, l. towards the anterior extremity; but we have found no traces
+of the canal, k, formed of involuted membranes, which in the body of the
+male proceeds from the posterior end of the lentil, l. i. nor of the
+plaited pallet, p. adhering to this canal, called by Swammerdam the
+penis from its resemblance to that of other animals, though he is not of
+opinion that this point, which is not perforated, can perform the
+functions of a real penis, and hold the principal part in generation.
+The canal, k, therefore, and all appertaining to it, must break at i,
+quite close to the posterior part of the lentil, since we found no
+remains of the lenticular bodies left by the fecundating males, in the
+body of our females. The canal, r, which Swammerdam calls the root of
+the penis, with greater reason than he was himself aware, is not
+extended in the body of the male as represented by the figure here
+engraved, but this long twilled canal consists of several involutions,
+from the seminal vessels whence it proceeds, into the lenticular body
+where it terminates, and where it conveys the fluid. This canal
+therefore can extend during copulation, and allow the lenticular
+substance to protrude out of the body of the males.
+
+It is evident this may be the case during copulation as is seen on
+opening a drone, for, by endeavouring to displace the lenticular body,
+the involutions of the cord disappear, and it extends much more than
+necessary for the lentil to protrude from the body; and if we attempt
+to separate it farther, the canal breaks at l. close to the lentil, and
+at the same place where it breaks in copulation.
+
+By dissection two nerves are discovered, towards the origin of the
+canal, r. inserted into the seminal vessels and distribute in them, and
+towards the root of the penis many ramifications undoubtedly serving for
+the motion of these parts. Two small parts, perceptible near the nerves,
+are two ligaments for retaining the generative organs in their proper
+place, so that except the root of the penis, they cannot be drawn out
+without some exertion; it and the lenticular body however can protrude,
+and actually do so during copulation. A certain degree of pressure
+forces all these parts from the body of the male, but they spontaneously
+return, and appear reversed.
+
+Swammerdam, and after him M. de Reaumur have admired this mechanism;
+they have thought, indeed, that the return should be occasioned by the
+effect of the air inflating the parts, and they supposed that the male
+organs proceeded from the body, and returned during copulation, the same
+as when forced out by pressure. Following their example, we have pressed
+them from the body of many males; we have a thousand times witnessed
+this wonderful return, which they detail with the greatest precision;
+but our males never survived the operation. We have seen, as M. de
+Reaumur, a few males protrude them spontaneously, even some of the parts
+inverted, but at that moment they died, and were unable to retract the
+parts which a pressure, most likely accidental, had forced out. Thus it
+is improbable that the male organs protrude by turning out of themselves
+in copulation; and the details which follow prove incontestibly, that it
+is otherwise. Had not Swammerdam been prejudiced with this opinion, he
+would have seen that the lenticular body can proceed from the body in
+erection without reversing itself; he could have proportioned the
+tortuous canal, which he calls the root of the penis; he would have seen
+that, at certain times, it can be sufficiently extended to allows the
+lenticular substance to protrude; he would have discovered the real use
+of the scaly plates; he would have explained that of the canal k, of the
+plaited pallet q, and the movements of all these parts, more admirable
+perhaps than the inversion which he was the first to observe.
+
+Our observations incontestibly prove copulation. The portion of the
+males found engaged in the body of our queens, hitherto called the
+lenticular substance, may be denominated a penis both from its position
+and use. The same surface is presented by it in the queen as in the body
+of the male, which is proved by the position of the laminae, e. e.
+attached to the interior of the penis, when found in the queen. It is
+evident, if the supposed inversion took place, the laminae would be found
+within the posterior part of the penis; and we should see them through
+its membrane, by their concave side, instead of which the convex surface
+is presented when in the vulva of females, the same as in the body of
+the males. But what is the use of these laminae? From their figure,
+hardness, relative position with respect to each other, and their
+situation at the extremity of the penis, we cannot doubt they are real
+pincers. However, to ascertain the fact, we found it necessary to see
+their position, and that of the penis itself in the females. For this
+purpose, we prevented some of the queens from extracting the parts left
+by the impregnating males, and by dissection we discovered that the
+laminae were pincers as we had conjectured.
+
+The penis was situated under the sting of the queens, and pressed
+against the upper region of the belly. It was supported by the posterior
+end, against the extremity of the vagina, or excretory canal. There we
+were sensible of the motion and use of the scaly pieces. Their
+extremities were separated a little more than in the male, and pressed
+between them some of the female parts below the excretory canal. The
+extreme minuteness of these parts prevented us from distinguishing them
+clearly, but the effort necessary to separate and remove the penis from
+the female, satisfied us of the use of these laminae.
+
+Inspecting a male from above, the convex side of the plates, e. e. is
+presented, and the summit of the angle formed by their origin. When in
+the body of the female, they are in the inverse position; what was above
+in the male is now below, and the extremity of the pincers directed
+upwards. This makes us suspect that in copulation the male mounts on the
+back of the female, but we are far from asserting it positively. It may
+be asked whether that part we call the penis, is the sole part
+introduced into the female during copulation? We have carefully
+investigated this, and can affirm, that it is the only one of all those
+described by M. de Reaumur, which has been found in our females. But we
+have discovered a new part that escaped both him and Swammerdam, which
+appears from the following experiment.
+
+Separating the lenticular substance from the excretory canal, where it
+was attached, we drew along with it a white body, adhering by one
+extremity, and having the other engaged in the vagina. Towards the end
+of the lentil, where the substance adhered, it appeared cylindrical,
+then it swelled, and again contracted, to dilate anew in a greater
+degree than at first; afterwards it contracted and terminated in a
+point. A powerful magnifier was required to see all this. When pulled
+from the lenticular body, the part was commonly broke, and also when
+extracted by the queens from themselves. The figure and situation seemed
+to authorise our considering it the penis itself, and the lenticular
+body only an appendage. But the last queen we examined exhibited a
+peculiarity that induced us to doubt the fact, and led us to suspect
+that this body is nothing else than the seminal fluid itself, moulded
+and coagulated in the vagina, and which from its viscosity adheres to
+the lenticular substance, and accompanies it when separated from the
+vagina. In this queen was found a little extravasated white matter, near
+the opening of the vagina. This, though at first liquid, soon coagulated
+in the air as the seminal fluid of drones does. In separating the
+lenticular body from the vagina, we drew along with it a thread which
+broke near the lentil; and seemed of too little consistence for the
+penis of a male. The lenticular bodies, found in our queens, appeared
+larger than in the males we dissected, and we have remarked with M. de
+Reaumur, that these parts are not of equal size in every male.
+
+       *       *       *       *       *
+
+_Experiment 1._--On the tenth of July, we set successively at liberty
+three virgin queens four or five days old. Two flew away several times;
+their absence was short and fruitless. The third profited better by her
+liberty; she departed thrice; the first and second time her absence was
+short; but the third lasted thirty-five minutes. She returned in a very
+different state; and in such as allowed no doubt of her employment, for
+she exhibited the part of a male that had rendered her a mother. We
+seized her wings with one hand, and in the other received the
+lenticular body, of which she had disengaged herself with her claws. The
+posterior part was armed with two pincers, e. e. shelly and elastic,
+which could be drawn asunder, and then resumed their original position.
+Towards the anterior part of the lentil appeared the fragment of the
+root of the penis; this canal had broke half a line from the lenticular
+body. We allowed the queen to enter her habitation, and adapted the
+entrance so that she could not leave it unknown to us.
+
+On the seventeenth we found no eggs in the hive; the queen was as
+slender as the first day; therefore the male, with which she had
+copulated, had not impregnated her eggs. She was again set at liberty;
+after twice departing, she returned with evidence of a second
+copulation. We then confined her, and the eggs she afterwards laid
+proved that the second copulation had been more successful than the
+first and that there are some males more fit for impregnating queens
+than others. However, it is very rare that the first copulation is
+inefficient; we have only seen two that required it twice; all the rest
+were impregnated by the first.
+
+       *       *       *       *       *
+
+_Experiment 2._--On the eighteenth we put at liberty a virgin queen
+twenty-seven days old, she departed twice. Her second absence was
+twenty-eight minutes, and she returned with the proofs of copulation. We
+prevented her from entering, and put her under a glass to see how she
+would disengage the male organs. This she was unable to accomplish,
+having only the table and sides of the glass for support; therefore we
+introduced a bit of comb; thus providing the same conveniences as are in
+a hive. Fixing herself on _it_ by the first four legs, she stretched out
+the two last, and extending them along her belly seemed to press it
+between them. At length introducing her claws between the two parts of
+the last ring, she seized the lenticular body, and dropped it on the
+table. The posterior part was provided with shelly pincers, under which
+and in the same direction was a grey cylindrical body. The end farthest
+from the lentil was sensibly thicker than that adhering to it, and
+terminated in a point. This point was double, and open like the bill of
+a bird, which induces us to think the body was broken, a conjecture
+supported by the following experiment.
+
+       *       *       *       *       *
+
+_Experiment 3._--On the nineteenth we set at liberty a queen four days
+old; she departed twice; her first absence was short; the second lasted
+thirty minutes, and then she returned with the marks of fecundation. As
+we wished to obtain the male organs entire, it was necessary to prevent
+the queen from breaking them by extracting them with her feet; we
+therefore suddenly killed her, and cut off the last rings in order to
+lay the vulva open. But though deprived of animation, so much life
+remained in these parts that the lenticular body was thrown out
+spontaneously. Under the pincers appeared the remnant of a cylindrical
+body which had broken near the origin and remained in the female. This
+body was very small at the origin; it afterwards sensibly enlarged; next
+contracting by degrees, it terminated in a sharp point. We found the
+point engaged up to the gland in the excretory canal, and the rest in
+the vulva.
+
+       *       *       *       *       *
+
+_Experiment 4._--We set two virgin queens at liberty on the twentieth.
+The first had been abroad on the preceding days, but the scarcity of
+males prevented her from being previously fecundated. She returned with
+the organs of a male. We tried to prevent her from extracting them, but
+she did this so expeditiously with her feet, that we could not
+accomplish it. She was then allowed to enter the hive.
+
+The second queen departed twice. Her first absence was short as usual;
+the second lasted about half an hour, and she returned impregnated.
+Having killed her as suddenly as possible, we laid open the vulva. The
+lenticular body was deposited as in every queen hitherto dissected; the
+pincers were situated under the excretory canal. Some parts not easily
+distinguishable were pressed between the laminae, and their office seemed
+to consist in forcing the extremity of the lentil to approach the
+orifice of the vagina, and apply so forcibly to it that some exertion
+was necessary to separate them. We previously examined them, with a very
+powerful magnifier. Then a peculiarity which had escaped us was
+perceptible. In drawing out the lenticular body, there proceeded from
+the vagina a minute part, v. adhering to the posterior end of the
+lentil, and situated below the plates. It spontaneously retracted into
+the lentil, like the horns of a snail. It appeared white, very short,
+and cylindrical. Under the pincers was a little half coagulated seminal
+fluid at the bottom of the vulva. Though much could be expressed, there
+was none pure; it was almost liquid, but soon coagulated, and formed a
+whitish inorganic mass. This observation carefully made removed all our
+doubts, and demonstrated that what we had taken for the penis of males
+was nothing but the seminal fluid, which had coagulated and assumed the
+interior figure of the vagina. The only hard part introduced by the
+male, was the short cylindrical point which retracted into the lentil,
+when we separated it. Its situation and office prove that it is there we
+must look for the issue of the seminal fluid, if we can hope to find an
+opening, when not engaged in copulation.
+
+We found this new part in the first drone we dissected. By pressing the
+seminal vessels, the white liquid then escaped downwards to the root of
+the penis r. and into the lenticular body, l. i. which became sensibly
+swoln. We prevented the fluid from returning, and by new pressure of the
+lentil forced it to advance. However, none escaped, but we saw at the
+posterior end of the lenticular body, and under the scaly pincers, a
+small white cylindrical substance, the same in appearance as that we had
+found engaged in the vagina of the queen. This part retracted on
+pressure, and then returned.
+
+I request you, Sir, while perusing this letter, to inspect the figure of
+the male sexual organs published by M. de Reaumur, and which are copied
+here. The descriptions are most accurate, and present a just idea of the
+situation of these parts when in the male's body. We readily conceive
+how they appear when left in the female by copulation. This detail will
+sufficiently indicate the situation and figure of the new part I have
+discovered.
+
+I suspect that the males perish after losing their sexual organs. But
+why does nature exact so great a sacrifice? This is a mystery which I
+cannot pretend to unveil. I am unacquainted with any analogous fact in
+natural history, but as there are two species of insects whose
+copulation can take place only in the air, namely, ephemerae and ants, it
+would be extremely interesting to discover whether their males also
+lose their sexual parts, in the same circumstances, and whether, as with
+drones, enjoyment in their flight is the prelude of death.
+
+FINIS.
+
+
+FOOTNOTES:
+
+{O} Memoires sur les Abeilles, p. 450.
+
+{P} Such long and minute descriptions can be very imperfectly
+translated; indeed they are unintelligible without microscopical
+inspections of the parts themselves.--T.
+
+
+
+
+ANALYTICAL INDEX.
+
+
+  Description of a hive invented by the author                   page 4
+  Swammerdam's opinion on the fecundation of bees                     8
+  Sentiments of M. de Reaumur                                        10
+  Mr Debraw's opinion                                                11
+  Hattorf's opinion                                                  19
+  Difficulty of discovering the mode of impregnation                 22
+  Experiments on the subject                                         23
+  Suggestions by M. Bonnet                                           34
+  The queen is impregnated by copulation, which never takes place
+    within the hive                                                  41
+  Experiments on artificial fecundation have not succeeded           42
+  The male loses the sexual organs in copulation                     43
+  Regarded impregnation affects the ovaries of the queen             45
+  She then lays no eggs but those producing males                    47
+  One copulation impregnates all the eggs the queen will lay in
+    two years                                                        54
+  Fecundity of a queen                                               63
+  Common bees do not transport the queen's eggs                      66
+  They sometimes eat them                                            69
+  Eggs producing males are sometimes laid in royal cells             71
+  Common worms may be converted into queens                          77
+  Operations of the bees when this is done                           78
+  Fertile workers sometimes exist                                    89
+  They lay none but the eggs of males                                96
+  All common bees are originally females                             98
+  Receiving the royal food while larvae, expands their ovaries       105
+  Mutual enmity of queens                                           110
+  The common bees seem to promote their combats                     117
+  A guard is constantly at the entrance of the hive                 123
+  What ensues when bees lose their queen                            126
+  Effects of introducing a stranger queen                           128
+  Massacre of the males                                             132
+  It never ensues in hives deprived of queens                       135
+  A plurality of queens is never tolerated                          142
+  The queen bee is oviparous                                        149
+  Bees seem occasionally to repose                                  150
+  Interval between production of the egg and the perfect state
+    of bees                                                         151
+  Mode of spinning the coccoon                                      153
+  That of the queen is open at one end                              154
+  The size of the bees is not affected by that of the cells         167
+  The old queen always conducts the first swarm                     173
+  But never before depositing eggs in the royal cells               177
+  Singular effect of a sound emitted by perfect queens              189
+  The instinct of bees is affected during the period of swarming    208
+  Queens are liberated from their cells according to their age      214
+  The bees probably judge of this by the sound emitted              217
+  Young queens conducting swarms are virgins                        221
+  The conduct of bees to old queens is peculiar                     224
+  Retarded impregnation affects the instinct of queens              241
+  Amputation of the antennae produces singular effects               245
+  Advantages of the leaf hive                                       253
+  It renders the bees tractable                                     256
+  They may there be forced to work in wax                           264
+  Uniform distance between the combs                                265
+  Natural heat of bees                                              269
+  Distance to which they fly                                        271
+  Appendix                                                          273
+  Anatomical observations on the sexual organs of bees              276
+  Experiments proving the copulation of the queen                   290
+
+ALEX. SMELLIE, Printer.
+
+
+
+
+{Transcriber's notes
+
+The spelling in the original is sometimes idiosyncratic. It has not
+been changed, but a few obvious errors have been corrected. The
+corrections are listed below.
+
+Inconsistent spellings include:
+  Lusace/Lusace, centre/center, choose/chuse,
+  organisation/organization, recognise/recognize
+
+Unusual spellings (which have not been changed) include:
+  centinels, coccoon, diaphraghm, encreased, encreasing, groupes,
+  harrassed, inaccessible, incontestible, indispensible, moveable,
+  perceptible, susceptible, uncontrouled, unintelligible
+
+Letter I
+
+  "secret distinctive characterestics" changed to
+  "secret distinctive characteristics"
+
+Letter II
+
+  "the copulalation of queens" changed to
+  "the copulation of queens"
+
+Letter IV
+
+  "The worms had spun their silk coccons" changed to
+  "The worms had spun their silk coccoons"
+
+Letter V
+
+  "characteristics of commo nbees" changed to
+  "characteristics of common bees"
+
+Letter VI
+
+  "The result of this rencounter" changed to
+  "The result of this encounter"
+
+  "genius such as your's" unchanged.
+
+  "observing that the antennae" changed to
+  "observing that the antennae"
+
+  "combats and disastrou scenes" changed to
+  "combats and disastrous scenes"
+
+  "M. de Reamur speaks of these executions" changed to
+  "M. de Reaumur speaks of these executions"
+
+Letter IX
+
+  "Only the few bees that not participated" changed to
+  "Only the few bees that had not participated"
+
+Letter XI
+
+  "these tumultous motions" changed to
+  "these tumultuous motions"
+
+Letter XII
+
+  "one antennae" unchanged.
+
+  "reside in them," changed to
+  "reside in them."
+
+Appendix
+
+  "the cirumference is edged" changed to
+  "the circumference is edged"
+
+  "he could have proportioned the tortous canal" changed to
+  "he could have proportioned the tortuous canal"
+
+  "pressed between the laminae" changed to
+  "pressed between the laminae"
+}
+
+
+
+
+
+End of the Project Gutenberg EBook of New observations on the natural
+history of bees, by Francis Huber
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