New observations on the natural history of bees
Chapter 18
_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.
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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.]
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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.
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_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