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New observations on the natural history of bees
by Francis Huber
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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" }

THE END

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