Manual of the apiary

by Albert John Cook

ANATOMY AND PHYSIOLOGY OF THE HONEY-BEE.

With a knowledge of the anatomy and some glimpses of the physiology of insects in general, we shall now find it easy to learn the special anatomy and physiology of the highest insects of the order.

THREE KINDS OF BEES IN EACH FAMILY.

As we have already seen, a very remarkable feature in the economy of the honey-bee, described even by Aristotle, which is true of many other bees, and also of ants and many wasps, is the presence in each family of three distinct kinds, which differ in form, color, structure, size, habits and function. Thus we have the queen, a number of drones, and a far greater number of workers. Huber, Bevan, Munn and Kirby also speak of a fourth kind blacker than the usual workers. These are accidental, and are, as conclusively shown by Von Berlepsch, ordinary workers, more deeply colored by loss of hair, dampness, or some other atmospheric condition. American apiarists are too familiar with these black bees, for after our severe winters they prevail in the colony, and, as remarked by the noted Baron, "They quickly disappear." Munn also tells of a fifth kind, with a top-knot, which appears at swarming seasons. I am at a great loss to know what he refers to, unless it be the pollen masses of the asclepias or milk-weed, which sometimes fasten to our bees and become a severe burden.

THE QUEEN BEE.

The queen ([Fig, 14]), although referred to as the mother bee, was called the king by Virgil, Pliny, and by writers as late as the last century, though in the ancient "Bee Master's Farewell," by John Hall, published in London in 1796, I find an admirable description of the queen bee, with her function correctly stated. Réaumur as quoted by "Wildman on Bees," published in London in 1770, says "this third sort has a grave and sedate walk, is armed with a sting, and is mother of all the others."

Huber, to whom every apiarist owes so much, and who, though blind, through the aid of his devoted wife and intelligent servant, Frances Burnens, developed so many interesting facts, demonstrated the fact of the queen's maternity. This author's work, second edition, published in Edinburgh, in 1808, gives a full history of his wonderful observations and experiments, and must ever rank with Langstroth as a classic, worthy of study by all.

Fig. 14.

The queen, then, is the mother bee, in other words, a fully developed female. Her ovaries ([Fig, 11, a, a]) are very large, nearly filling her long abdomen. The tubes already described as composing them are very numerous, while the spermatheca ([Fig. 11, e]) is plainly visible. This is muscular, receives abundant nerves, and thus, without doubt, may or may not be compressed to force the sperm cells in contact with the eggs as they pass by the duet. Leuckart estimates that the spermatheca will hold more than 25,000,000 spermatozoa.

The possession of the ovaries and attendant organs, is the chief structural peculiarity which marks the queen, as these are the characteristic marks of females among all animals. But she has other peculiarities worthy of mention She is longer than either drones or workers, being more than seven-eighths of an inch in length, and, with her long tapering abdomen, is not without real grace and beauty. The queen's mouth organs, too, are developed to a less degree than are those of the worker-bees. Her jaws ([Fig, 21, b]) or mandibles are weaker, with a rudimentary tooth, and her tongue or ligula ([Fig, 15, a]), as also the labial palpi ([Fig, 15, b]) and maxillæ are considerably shorter. Her eyes, like the same in the worker-bee ([Fig, 5]), are smaller than those of the drones, and do not meet above. So the three ocelli are situated above and between. The queen's wings, too, ([Fig, 14]) are relatively shorter than those either of the workers or drones, for instead of attaining to the end of the body, they reach but little beyond the third joint of the abdomen. The queen, though she has the characteristic posterior tibia and basal tarsus ([Fig, 16, p]), in respect to breadth, has not the cavity and surrounding hairs, which form the pollen baskets of the workers.

Fig. 15.

a—Ligula. d, d—Paraglossæ.

b—Labial palpi.

The queen possesses a sting ([Fig, 11, d]) which is longer than that of the workers, and resembles that of the bumble-bees in being curved, and that of bumble-bees and wasps in having few and short barbs—the little projections which point back like the barb of a fish-hook, and which, in case of the workers, prevent the withdrawing of the instrument, when once fairly inserted. While there are seven quite prominent barbs on each shaft of the worker's sting, there are only three on those of the queen, and these are very short, and, as in a worker's sting, they are successively shorter as we recede from the point of the weapon. Aristotle says that the queen will seldom use her sting, which I have found true. I have often tried to provoke a queen's anger, but never with any evidence of success. Neighbour ([page 14, note]) gives three cases where queens used their stings, in one of which cases she was disabled from farther egg-laying. She stings with slight effect.

Fig. 16.

t—Tibia. p—Broadened tibia and basal tarsus. t s—Tarsal joints.

The queen, like the neuters, is developed from an impregnated egg, which, of course, could only come from a queen that had previously mated. These eggs are not placed in a horizontal cell, but in one specially prepared for their reception ([Fig, 26, i]). These queen cells are usually built on the edge of the comb, or around an opening in it, which is necessitated from their size and form, as usually the combs are too close together to permit their location elsewhere. These cells extend either vertically or diagonally downward, are composed of was mixed with pollen, and in size and form much resemble a pea-nut. The eggs must be placed in these cells, either by the queen or workers. Huber, who though blind had wondrous eyes, also witnessed the act. I have frequently seen eggs in these cells, and without exception in the exact position in which the queen always places her eggs in the other cells. John Hall, in the old work already referred to, whose descriptions, though penned so long ago, are wonderfully accurate, and indicate great care, candor, and conscientious truthfulness, asserts that the queen is five times as long laying a royal egg as she is the others. From the character of his work, and its early publication, I can but think that he had witnessed this rare sight. Some candid apiarists of our own time and country—E. Gallup among the rest—claim to have witnessed the act. The eggs are so well glued, and are so delicate, that, with Neighbour, I doubt the possibility of a removal. The opponents to this view base their belief on a supposed discord between the queen and neuters. This antagonism is inferred, and I have but little faith in the inference, or the argument from it. I know that when royal cells are to be torn down, and inchoate queens destroyed, the workers aid the queen in this destruction. I have also seen queens pass by unguarded queen-cells, and yet respect them. I have also seen several young queens dwelling amicably together in the same hive. Is it not probable that the bees are united in whatever is to be accomplished, and that when queens are to be destroyed all spring to the work, and when they are to live all regard them as sacred? It is true that the actions of bees are controlled and influenced by the surrounding conditions or circumstances, but I have yet to see satisfactory proof of the old theory that these conditions impress differently the queen and the workers. The conditions which lead to the building of queen-cells and the peopling of the same are—loss of queen, when a worker larva from one to four days old will be surrounded by a cell inability of a queen to lay impregnated eggs, her spermatheca having become emptied; great number of worker-bees in the hive; restricted quarters; the queen not having place to deposit eggs, or the workers little or no room to store honey and lack of ventilation, so that the hive becomes too close. These last three conditions are most likely to occur at times of great honey secretion.

A queen may be developed from an egg, or from a worker larva less than three days old. Mr. Doolittle has known queens to be reared from worker larvæ taken at four-and-a-half days from hatching. In this latter case, the cells adjacent to the one containing the selected larva are removed, and the larva surrounded by a royal cell. The development of the queen larva is much like that of the worker, soon to be detailed, except that it is more rapid, and is fed richer and more plenteous food, called royal jelly. This peculiar food, as also its use and abundance in the cell, was first described by Schirach, a Saxon clergyman, who wrote a work on bees in 1771. According to Hunter, this royal pabulum is richer in nitrogen than that of the common larvæ. It is thick, like rich cream; slightly yellow, and so abundant that the queen larva not only floats in it during all its period of growth, but quite a large amount remains after her queenship vacates the cell. We often find this royal jelly in incomplete queen-cells, without larvæ. Mr. Quinby suggests that this is stored for future use.

What a mysterious circumstance is this: These royal scions simply receive a more abundant and sumptuous diet, and occupy a more ample habitation—for I have more than once confirmed the statement of Mr. Quinby, that the direction of the cell is immaterial—and yet what a marvelous transformation. Not only are the ovaries developed and filled with eggs, but the mouth-organs, the wing's, the legs, and the sting, aye, and the size, form and habits are all wondrously changed. That the development of parts should be accelerated, and the size increased is not so surprising—as in breeding other insects I have frequently found that kind and amount of food, would hasten or retard growth, and might even cause a dwarfed imago—but that it should so essentially modify the structure, is certainly a rare and unique circumstance, hardly to be found except here and in related animals. Bevan has suggested that fertile workers', while larvæ, have received some of this royal jelly, from their position near a developing queen. Langstroth supposes that they receive some royal jelly, purposely given by the workers, and I had previously thought this reasonable, and probably true. But these pests of the apiarist, and especially of the breeder, almost always, so far as I have observed, make their appearance in colonies long queenless, and I have noticed a case similar to that given by Quinby, where these occurred in a nucleus where no queen had been developed. May it not be true, that a desire for eggs stimulates growth of the ovaries, growth of eggs in the ovarian tubes, and consequent ability to deposit. The common high-holder, Colaptes auratus—a bird belonging to the woodpecker family, usually lays five eggs, and only five; but let cruel hands rob her of these promises of future loved ones—and wondrous to relate, she continues to lay more than a score. One thus treated, here on the College campus, actually laid more than thirty eggs. So we see that animal desires may influence and move organs that are generally independent of the will.

The larval queen is longer and more rapid of development than the other larvæ. When developed from the egg—as in case of normal swarming—the larva feeds for five days, when the cell is capped by the workers. The infant queen then spins her cocoon, which occupies about one day. The end of the cocoon is left open. Some one has suggested that this is an act of thoughtful generosity on the part of the queen larva, thus to render her own destruction more easy, should the welfare of the colony demand it, as now a sister queen may safely give the fatal sting. The queen now spends nearly three days in absolute repose. Such rest is common to all cocoon-spinning larvæ. The spinning, which is done by a rapid motion to and fro of the head, always carrying the delicate thread, much like the moving shuttle of the weaver, seems to bring exhaustion and need of repose. She now assumes the nymph or pupa state ([Fig, 26, i]). At the end of the sixteenth day she comes forth a queen. Huber states that when a queen emerges, the bees are thrown into a joyous excitement, so that he noted a rise in the temperature of the hive from 92° F. to 104° F. I have never tested this matter accurately, but I have failed to notice any marked demonstration on the natal day of her lady-ship the queen, or extra respect paid her as a virgin. When queens are started from worker larvæ, they will issue as images in ten or twelve days from the date of their new prospects. Mr. Doolittle writes me that he has known them to issue in eight and one-half days.

As the queen's development is probably due to superior quality and increased quantity of food, it would stand to reason that queens started from eggs are preferable; the more so, as under normal circumstances, I believe, they are almost always thus started. The best experience sustains this position. As the proper food and temperature could best be secured in a full colony—and here again the natural economy of the hive adds to our argument—we should infer that the best queens would be reared in strong colonies, or at least kept in such colonies till the cells were capped. Experience also confirms this view. As the quantity and quality of food, and the general activity of the bees is directly connected with the full nourishment of the queen-larva, and as these are only at the maximum in times of active gathering—the time when queen-rearing is naturally started by the bees—we should also conclude that queens reared at such seasons are superior. My experience—and I have carefully observed in this connection—most emphatically sustains this view.

Five or six days after issuing from the cell—Neighbour says the third day—if the day is pleasant, the queen goes forth on her "marriage flight" otherwise she will improve the first pleasant day thereafter for this purpose. Huber was the first to prove that impregnation always takes place on the wing. Bonnet also proved that the same is true of ants, though in this case millions of queens and drones often swarm out at once. I have myself witnessed several of these wholesale matrimonial excursions among ants. I have also frequently taken bumble-bees in copulo while on the wing. I have also noticed both ants and bumble-bees to fall while united probably borne down by the expiring males. That butterflies! moths, dragon-flies, etc., mate on the wing is a matter of common observation. That it is possible to impregnate queens when confined, I think very doubtful. The queens will caress the drones, but the latter seem not to heed their advances. That this ever has been done I also question, though many think they have positive proof that it has occurred. Yet, as there are so many chances to be mistaken, and as experience and observation are so excessive against the possibility, I think that these may be cases of hasty or inaccurate judgment. Many, very many, with myself, have followed Huber in clipping the queen's wing, only to produce a sterile or drone-laying queen. Prof Leuckart believes that successful mating demands that the large air-sacks ([Fig, 2, f]) of the drones shall be filled, which he thinks is only possible during flight. The demeanor of the drones leads me to think, that the excitement of flight, like the warmth of the hand, is necessary to induce the sexual impulse.

I presume, that in all the future, Huber's statement that the queen must take wing to be impregnated, will remain unrefuted. Yet it will do no harm to keep trying. Success may come. Mating, too, in green-houses or rooms is also impracticable. I have given this thorough trial. The drones are incorrigible cowards, and their inordinate fear seems even to overcome the sexual desires.

If the queen fails to find an admirer the first day, she will go forth again and again till she succeeds. Huber stated that after twenty-one days the case is hopeless. Bevan states that if impregnated from the fifteenth to the twenty-first she will be largely a drone-laying queen. That such absolute dates can be fixed in either of the above cases is very questionable. Yet, all experienced breeders know that queens kept through the winter as virgins are sure to remain so. It is quite likely that the long inactivity of the spermatheca wholly or in part paralyzes it, so that queens that are late in mating cannot impregnate the eggs as she desires. This would accord with what we know of muscular organs. Berlepsch believed that a queen that commenced laying as a virgin could never lay impregnated eggs, even though she afterwards mated. Langstroth thought that he had observed to the contrary.

If the queen be observed after a successful "wedding tour," she will be seen to bear the marks of success in the pendant drone appendages, consisting of the penis, the yellow cul-de-sacks, and the hanging thread-like ducts.

It is not at all likely that a queen, after she has met a drone, ever leaves the hive again except that she leaves with a swarm. Some of the observing apiarists think that an old queen may be again impregnated. The fact that queens, with clipped wings, are as long fertile as others, makes me think that cases which have led to such conclusions are capable of other explanation.

If the queen lays eggs before meeting the drones, or if for any reason she fails to mate, her eggs will only produce male bees. This strange anomaly—development of the eggs without impregnation—was discovered and proved by Dzierzon, in 1845. Dr. Dzierzon, who, as a student of practical and scientific apiculture, must rank with the great Huber, is a Roman Catholic priest of Carlesmarkt, Germany. This doctrine—called parthenogenesis, which means produced from a virgin—is still doubted by some quite able bee-keepers, though the proofs are irrefragable: 1st. Unmated queens will lay eggs that will develop, but drones always result. 2d. Old queens often become drone-layers, but examination shows that the spermatheca is void of seminal fluid. Such an examination was first made by Prof. Siebold, the great German anatomist, in 1813, and later by Leuckart and Leidy. I have myself made several such examinations. The spermatheca can easily be seen by the unaided vision, and by crushing it on a glass slide, by compressing with a thin glass cover, the difference between the contained fluid in the virgin and impregnated queen is very patent, even with a low power. In the latter it is more viscid and yellow, and the vesicle more distended. By use of a high power, the active spermatozoa or germ-cells become visible. 3d. Eggs in drone-cells are found by the microscopist to be void of the sperm-cells, which are always found in all other fresh-laid eggs. This most convincing, and interesting observation, was first made by Von Siebold, at the suggestion of Berlepsch. It is quite difficult to show this. Leuckart tried before Von Siebold, at Berlepsch's apiary, but failed. I have also tried to discover these germ-cells in worker-eggs, but as yet have been unsuccessful. Siebold has noted the same facts in eggs of wasps. 4th. Dr. Dönhoff, of Germany, in 1855, took an egg from a drone-cell, and by artificial impregnation produced a worker-bee. Such an operation, to be successful, must be performed as soon as the egg is laid.

Parthenogenesis, in the production of males, has also been found by Siebold to be true of other bees and wasps, and of some of the lower moths, in the production of both males and females. While the great Bonnet first discovered what may be noticed on any summer day, all about us, even on the house-plants at our very windows, that parthenogenesis is best illustrated by the aphides or plant lice. In the fall males and females appear, which mate, when the female lays eggs, which in the spring produce only females; these again produce only females, and thus on, for several generations, till with the cold of autumn come again the males and females. Bonnet observed seven successive generations of productive virgins. Duval noted nine generations in seven months, while Kyber observed production exclusively by parthenogenesis in a heated room for four years. So, we see, that this strange and almost incredible method of increase, is not rare in the great insect world.

About two days after she is impregnated, the queen, under normal circumstances, commences to lay, usually worker-eggs, and as the condition of the hive seldom impels to swarming the same summer, so that no drones are required, she usually lays no others the first season.

It is frequently noticed that the young queen at first lays quite a number of drone-eggs. Queen-breeders often observe this in their nuclei. This continues for only a few days. This does not seem strange. The act of forcing the sperm-cells from the spermatheca is muscular and voluntary, and that these muscles should not always act promptly at first, is not strange, nor is it unprecedented. Mr. Wagner suggested that the size of the cell determined the sex, as in the small cells the pressure on the abdomen forced the fluid from the spermatheca. Mr. Quinby also favored this view. I greatly question this theory. All observing apiarists have known eggs to be laid in worker-cells, ere the cell was hardly commenced, when there could be no pressure. In case of queen-cells, too, if the queen does lay the eggs—as I believe—these would be unimpregnated, as the cell is very large. I know the queen sometimes passes from drone to worker-cells very abruptly while laying, as I have witnessed such a procedure—the same that so greatly rejoiced the late Baron of Berlepsch, after weary hours of watching—but that she can thus control at the instant this process of adding or withholding the sperm-cells, certainly seems not so strange as that the spermatheca, hardly bigger than a pin-head, could supply these cells for months, yes, and for years. Who that has seen the bot-fly dart against the horse's legs, and as surely leave the tiny yellow egg, can doubt but that insects possess very sensitive oviducts, and can extrude the minute eggs just at pleasure. That a queen may force single eggs, at will, past the mouth of the spermatheca, and at the same time add or withhold the sperm-cells, is, I think, without question, true. What gives added force to this view, is the fact that other bees, wasps and ants exercise the same volition, and can have no aid from cell-pressure, as all the eggs are laid in receptacles of the same size. But the Baron of Berlepsch, worthy to be a friend of Dzierzon, has fully decided the matter. He has shown that old drone cells are as small as new worker-cells, and yet each harbors its own brood. Very small queens, too, make no mistakes. With no drone-cells, the queen will sometimes lay drone-eggs in worker-cells, in which drones will then be reared. And will, if she must, though with great reluctance, lay worker-eggs in drone-cells.

Before laying an egg, the queen takes a look into the cell, probably to see if all is right. If the cell contains any honey, pollen, or an egg, she usually passes it by, though when crowded, a queen will sometimes, especially if young, insert two or three eggs in a cell, and sometimes, in such cases, she drops them, when the bees show their dislike of waste, and appreciation of good living, by making a breakfast of them. If the queen finds the cell to her liking, she turns about, inserts her abdomen, and in an instant the tiny egg is glued, in position ([Fig, 26, b]) to the bottom of the cell.

The queen, when considered in relation to the other bees of the colony, possesses a surprising longevity. It is not surprising for her to attain the age of three years in the full possession of her powers, while they have been known to do good work for five years. Queens, often at the expiration of one, two, three or four years, depending on their vigor and excellence, either cease to be fertile, or else become impotent to lay impregnated eggs—the spermatheca having become emptied of its sperm-cells. In such cases the workers usually supersede the queen; that is they destroy the old queen, ere all the worker-eggs are gone, and take of the few remaining ones to start queen-cells, and thus rear young, fertile and vigorous queens.

It sometimes happens, though rarely, that a fine-looking queen, with full-formed ovaries, and large spermatheca, well-filled with male fluid, will deposit freely, but none of the eggs will hatch. Readers of the bee-publications know that I have frequently received such for dissection. The first I ever got was a remarkably fine-looking Italian, received from the late Dr. Hamlin, of Tennessee. All such queens that I have examined seem perfect, even though scrutinized with a high-power objective. We can only say that the egg is at fault, as frequently transpires with higher animals, even to the highest. These females are barren; through some fault with the ovaries, the eggs grown therein are sterile. To detect just what is the trouble with the egg is a very difficult problem, if it is capable of solution at all. I have tried to determine the ultimate cause, but without success.

The function of the queen is simply to lay eggs, and thus keep the colony populous; and this she does with an energy that is fairly startling. A good queen in her best estate will lay two or three thousand eggs a day. I have seen a queen in my observing hive, lay for some time at the rate of four eggs per minute, and have proved by actual computation of brood cells, that a queen may lay over three thousand eggs in a day. Langstroth and Berlepsch both saw queens lay at the rate of six eggs a minute.

The latter had a queen that laid three thousand and twenty-one eggs in 24 hours, by actual count, and in 20 days she laid fifty-seven thousand. This queen continued prolific for five years, and must have laid, says the Baron, at a low estimate, more than 1,300,000 eggs. Dzierzon says queens may lay 1,000,000 eggs, and I think these authors have not exaggerated. Yet, with even these figures as an advertisement, the queen bee cannot boast of superlative fecundity, as the queen white-ant—an insect closely related to the bees in habits, though not in structure, as the white-ants are lace-wings and belong to the sub-order Neuroptera, which includes our day-flies, dragon-flies, etc.—is known to lay over 80,000 eggs daily. Yet this poor helpless thing, whose abdomen is the size of a man's thumb, and composed almost wholly of eggs, while the rest of her body is not larger than the same in our common ants, has no other amusement; she cannot walk; she cannot even feed herself or care for her eggs. What wonder then that she should attempt big things in the way of egg-laying? She has nothing else to do, or to feel proud of.

Different queens vary as much in fecundity as do different breeds of fowls. Some queens are so prolific that they fairly demand hives of India rubber to accommodate them, keeping their hives gushing with bees and profitable activity while others are so inferior, that the colonies make a poor, sickly effort to survive at all, and usually succumb early, before those adverse circumstances which are ever waiting to confront all life on the globe. The activity of the queen, too, is governed largely by the activity of the workers. The queen will either lay sparingly, or stop altogether, in the interims of storing honey, while, on the other hand, she is stimulated to lay to her utmost capacity, when all is life and activity in the hive.

It would seem that the queen either reasons from conditions, is taught by instinct, or else that without her volition the general activity of the worker-bees stimulates the ovaries, how, we know not, to grow more eggs. We know that such a stimulus is born of desire, in case of the high-holder, already referred to. That the queen may have control of the activity of her ovaries, either directly or indirectly, through reflex nervous action induced by the general excitement of the bees, which always follows active storing, is not only possible, but quite likely.

The old poetical notion that the queen is the revered and admired sovereign of the colony, whose pathway is ever lined by obsequious courtiers, whose person is ever the recipient of loving caresses, and whose will is law in this bee-hive kingdom, controlling all the activities inside the hive, and leading the colony whithersoever they may go, is unquestionably mere fiction. In the hive, as in the world, individuals are valued for what they are worth. The queen, as the most important individual, is regarded with solicitude, and her removal or loss noted with consternation, as the welfare of the colony is threatened; yet, let the queen become useless, and she is despatched with the same absence of emotion that characterizes the destruction of the drones when they have become supernumeraries. It is very doubtful if emotion or sentimentality are ever moving forces among the lower animals. There are probably certain natural principles that govern in the economy of the hive, and aught that conspires against, or tends to intercept the action of these principles, becomes an enemy to the bees. All are interested, and doubtless more united than is generally believed, in a desire to promote the free action of these principles. No doubt the principle of antagonism among the various bees has been overrated. Even, the drones, when they are being killed off in the autumn, make a sickly show of defense, as much as to say, the welfare of the colony demands that such worthless vagrants should be exterminated; "so mote it be;" go ahead. The statement, too, that there is often serious antagonism between the queen and workers, as to the destruction or preservation of inchoate queens, yet in the cell, is a matter which may well be investigated. It is most probable that what tends most for the prosperity of the colony is well understood by all, and without doubt there is harmonious action among all the denizens of the hive, to foster that which will advance the general welfare, or to make war on whatever may tend to interfere with it. If the course of any of the bees seems wavering and inconsistent, we may rest assured that circumstances have changed, and that could we perceive the bearing of all the surrounding conditions, all would appear consistent and harmonious.

Fig. 17.

THE DRONES.

These are the male bees, and are generally found in the hive only from May to November: though they may remain all winter, and are not infrequently absent during the summer. Their presence or absence depends on the present and prospective condition of the colony. If they are needed, or likely to be needed, then they are present. There are in nature several hundred in each colony. The number may and should be greatly reduced by the apiarist. These ([Fig, 17]) are shorter than the queen, being less than three-fourths of an inch in length, are more robust and bulky than either the queen or workers, and are easily recognized when flying by their loud, startling hum. As in other societies, the least useful make the most noise. This loud hum is caused by the less rapid vibration of their large, heavy wings. Their flight is more heavy and lumbering than that of the workers. Their ligula, labial palpi, and maxillæ—like the same in the queen bee—are short, while their jaws ([Fig, 21, a]) possess the rudimentary tooth, and are much the same in form as those of the queen, but are heavier, though not so strong as those of the workers. Their eyes ([Fig, 4]) are very prominent, meet above, and thus the simple eyes are thrown forward. Their posterior legs are convex on the outside ([Fig, 18]), so, like the queens, they have no pollen baskets. The drones are without the defensive organ, having no sting, while their special sex-organs ([Fig, 10]) are not unlike those of other insects, and have already been sufficiently described.

Fig. 18.

t—Tibia. p—Broadened tibia and basal tarsus. t s—Joints of Tarsus, c—Claws.

It was discovered by Dzierzon, in 1845, that the drones hatch from unimpregnated eggs. This strange phenomenon, seemingly so incredible, is as has been shown in speaking of the queen, easily proved and beyond question. These eggs may come from an unimpregnated queen, a fertile worker—which will soon be further described—or from an impregnated queen, which may voluntarily prevent impregnation. Such eggs may be placed in the larger horizontal cells ([Fig, 28, a]), in manner already described. As stated by Bevan, the drone feeds six and a half days as a larva, before the cell is capped. The capping of the drone-cells is very convex, and projects beyond the plane of the same in worker-cells, so that the drone brood is easily distinguished from worker, and from the darker color—the wax being thicker and less pure—the capping of both drone and worker brood-cells enable us easily to distinguish them from honey-cells. In twenty-four days from the laying of the egg, the drones come forth from the cells. Of course variation of temperature, and other conditions, as variable amount of diet, may slightly retard or advance the development of any brood, in the different stages. The drones—in fact all bees—when they first emerge from the cells, are gray, soft, and appear generally unsophisticated.

Just what the longevity of the male bee is, I am unable to state. It is probable, judging from analogy, that they live till accident, the worker bees, or the performance of their natural function causes their death. The worker-bees are liable to kill off the drones, which they do by constantly biting and worrying them. They may also destroy the drone-brood. It is not very rare to see workers carrying out immature drones even in mid-summer. At the same time, too, they may destroy inchoate queens. Such action is prompted by a sudden check in the yield of honey, and with the drones is most common at the close of the season. The bees seem very cautious and far-sighted. If the signs of the times presage a famine, they stay all proceedings looking to the increase of colonies. On the other hand, unlimited honey, rapid increase of brood, crowded quarters—whatever the age of the queen—is sure to bring many of the male bees. While any circumstances that indicate a future need of drones will prevent their destruction even in late autumn.

The function of the drones is solely to impregnate the queen, though when present they may add animal heat. That their nutrition is active, is suggested by the fact, that upon dissection, we always find their capacious stomachs filled with honey.

Impregnation of the queen always takes place, as before stated, while on the wing, outside the hive, usually during the heat of warm sunshiny days. After mating, the drone organs adhere to the queen, and may be seen hanging to her for some hours. The copulatory act is fatal to the drones. By holding a drone in the hand, the ejection of the sex-organs is often produced, and always followed by immediate death. As the queen only meets a single drone, and that only once, it might be asked why nature was so improvident as to decree hundreds of drones to an apiary or colony, whereas a score would suffice as well. Nature takes cognizance of the importance of the queen, and as she goes forth amidst the myriad dangers of the outer world, it is safest and best that her stay abroad be not protracted; that the experience be not repeated, and especially, that her meeting a drone be not delayed. Hence the superabundance of drones—especially under natural conditions, isolated in forest homes, where ravenous birds are ever on the alert for insect game—is most wise and provident. Nature is never "penny wise and pound foolish." In our apiaries the need is wanting, and the condition, as it exists in nature, is not enforced.

The fact that parthenogenesis prevails in the production of the drones, has led to the theory that from a pure queen, however mated, must ever come a pure drone. My own experience and observation, which I believe are those of all apiarists, has confirmed this theory. Yet, if the impure mating of our cows, horses, and fowls, renders the females of mixed blood ever afterward, as is believed and taught by many who would seem most competent to judge—though I must say I am somewhat skeptical in the matter—then we must look closely as to our bees, for certainly, if a mammal, and especially a fowl, is tainted by impure mating, then we may expect the same of insects. In fowls such influence, if it exists, must come simply from the presence in the female generative organs of the germ-cells, or spermatozoa, and in mammals, too, there is little more than this, for though they are viviparous, so that the union and contact of the offspring and mother seems very intimate, during fœtal development, yet there is no intermingling of the blood, for a membrane ever separates that of the mother from that of the fœtus, and only the nutritious and waste elements pass from one to the other. To claim that the mother is tainted through the circulation, is like claiming that the same result would follow her inhaling the breath of her progeny after birth. I can only say, that I believe this whole matter is still involved in doubt, and still needs more careful, scientific and prolonged observation.

THE NEUTERS, OR WORKER-BEES.

These, called "the bees," by Aristotle, and even by Wildman and Bevan, are by far the most numerous individuals of the hive—there being from 15,000 to 40,000 in every good colony. It is possible for a colony to be even much more populous than this. These are also the smallest bees of the colony, as they measure but little more than one-half of an inch in length ([Fig, 19]).

Fig. 19.

The workers—as taught by Schirach, and proved by Mlle. Jurine, of Geneva, Switzerland, who, at the request of Huber, sought for and found, by aid of her microscope, the abortive ovaries—are undeveloped females. Rarely, and probably very rarely, except that a colony is long or often queenless, as is frequently true of our nuclei, these bees are so far developed as to produce eggs, which, of course, would always be drone eggs. Such workers—known as fertile—were first noticed by Riem, while Huber actually saw one in the act of egg-laying. Except in the power to produce eggs, they seem not unlike the other workers. Huber supposed that these were reared in cells contiguous to royal cells, and thus received royal food by accident. The fact, as stated by Mr. Quinby, that these occur in colonies where queen-larvæ were never reared, is fatal to the above theory. Langstroth and Berlepsch thought that these bees, while larvæ, were fed, though too sparingly, with the royal aliment, by bees in need of a queen, and hence the accelerated development. Such may be the true explanation. Yet if, as some apiarists aver, these appear where no brood has been fed, and so must be common workers, changed after leaving the cell, as the result of a felt need, then we must conclude that development and growth—as with the high-holder—spring from desire. The generative organs are very sensitive, and exceedingly susceptible to impressions, and we may yet have much to learn as to the delicate forces which will move them to growth and activity. Though these fertile workers are a poor substitute for a queen, as they are incapable of producing any but drones, and are surely the harbingers of death and extinction to the colony, yet they seem to satisfy the workers, for they will not brook the presence of a queen when a fertile worker is in the hive, nor will they suffer the existence in the hive of a queen-cell, even though capped. They seem to be satisfied, though they have very slight reason to be so. These fertile workers lay indifferently in large or small cells—often place several eggs in a single cell, and show their incapacity in various ways.

Fig. 20.

a—Ligula. b, b—Labial palpi.

c, c—Maxillæ. d—Paraglossæ.

The workers, as might be surmised by the importance and variety of their functions, are structurally very peculiar Their tongues ([Fig, 20, a]), labial palpi ([Fig. 20, b, b]), and maxillæ ([Fig. 20, c, c]), are very much elongated, while the former is very hairy, and doubles under the throat when not in use. The length of the ligula enables them to reach into flowers with long tubes, and by aid of the hairs they lap up the nectar. When the tongue is big with its adhering load of sweet, it is doubled back, enclosed by the labial palpi and maxillæ, and then extended, thus losing its load of nectar, which at the same time is sucked into the large honey-stomach. The bees, at will, can force the honey back from the honey-stomach, when it is stored in the honey-cells or given to the other bees.

Fig. 21.

a—Jaw of drone.

b—Jaw of queen.

c—Jaw of worker.

The jaws ([Fig, 21, c]) are very strong, without the rudimentary tooth, while the cutting edge is semi-conical, so that when the jaws are closed they form an imperfect cone. Thus these are well formed to cut comb, knead wax, and perform their various functions. Their eyes ([Fig, 5]) are like those of the queen, while their wings, like those of the drones, attain the end of the body. These organs ([Fig, 3]), as in all insects with rapid flight, are slim and strong, and, by their more or less rapid vibrations, give the variety of tone which characterizes their hum. Thus we have the rapid movements and high pitch of anger, and the slow motion and mellow note of content and joy.

Fig. 22.

t—Tibia. b—Rim of hairs. p—Pollen basket.

t s—Joint of tarsi, c—Claws.

On the outside of the posterior tibia and basal tarsus is a cavity, made more deep by its rim of hairs, known as the pollen basket ([Fig, 22, p]). In these pollen baskets is compacted the pollen, which is gathered by the mouth organs, and carried back by the four anterior legs. Opposite the pollen baskets are regular rows of golden hairs ([Fig. 23, e]), which probably aid in storing and compacting the pollen balls.

On the anterior legs of the workers, between the femur and tibia, is a curious notch ([Fig. 24, C]), covered by a spur ([Fig. 24, B]). For several years this has caused speculation among my students, and has attracted the attention of observing apiarists. Some have supposed that it aided bees in reaching deeper down into tubular flowers, others that it was used in scraping off pollen, and still others that it enabled bees to hold on when clustering. The first two functions may belong to this, though other honey and pollen-gathering bees do not possess it. The latter function is performed by the claws at the end of the tarsi.

Fig. 23.

e—Rows of hairs. t—Tibia. c—Claws.

Fig. 24.

Fig. 25.

a—Tube. b b—Barbed spears drawn out of tube and turned back, c—Poison sack. d—Muscles.

The workers, too, possess an organ of defense ([Fig, 25]), which they are quick to use if occasion requires. This is not curved as in the queen, but straight. The gland which secretes the poison is double, and the sack ([Fig, 25, c]), in which it is stored, is as large as a flax-seed. The sting proper, is a triple organ, consisting of three sharp spears, very smooth and of exquisite polish. The most highly-wrought steel instruments, under a high magnifier, look rough and unfinished, while the parts of the sting show no such inequalities. One of these spears ([Fig, 25, a]) is canaliculate—that is, it forms an imperfect tube—and in this canal work the other two ([Fig. 25, b, b]), which fill the vacant space, and thus the three make a complete tube, and through this tube, which connects with the poison sack, passes the poison. The slender spears which work in the tube are marvelously sharp, and project beyond it when used, and are worked alternately by small but powerful muscles ([Fig. 25, d]), so they may pass through buckskin, or even the thick scarf-skin of the hand. These are also barbed at the end with teeth, seven of which are prominent, which extend out and back like the barb of a fish-hook. Hence the sting cannot be withdrawn, if it penetrates any firm substance, and so when used, it is drawn from the bee, and carries with it a portion of the alimentary canal, thus costing the poor bee its life. Darwin suggests that bees and wasps were developed from the saw-flies, and that the barbs on the sting are the old-time saws, transformed into the spear-like barbs. He does not explain why these are so much shorter and more obscure in the queen, and in other bees and wasps. The honey-stomach or crop in the workers ([Fig, 9, o]) is well developed, though no larger than those in the drones. Whether it is more complex in structure, I do not know.

The workers hatch from an impregnated egg, which can only come from a queen that has met a drone, and is always laid in the small, horizontal cell. These eggs are in no wise different, so far as we can see, from those which are laid in the drone or queen-cells. All are cylindrical and slightly curved ([Fig, 26, b, c]) and are fastened by one end to the bottom of the cell, and a little to one side of the centre. As already shown, these are voluntarily fertilized by the queen as she extrudes them, preparatory to fastening them in the cells. These eggs, though so small—one-sixteenth of an inch long—may be easily seen by holding the comb so that the light will shine into the cells. With experience, they are detected almost at once, but I have often found it quite difficult to make the novice see them, though very plainly visible to my experienced eye.

Fig. 26.

b and c—Eggs. d, e, f and g—Various sizes of larvæ. h—Pupa. i—Pupa of queen, in queen-cell. k, k—Caps.

The egg hatches in three days. The larva ([Fig, 26, d, e, f, g]), incorrectly called grub, maggot—and even caterpillar, by Hunter—is white, footless, and lies coiled up in the cell till near maturity. It is fed a whitish fluid, though this seems to be given grudgingly, as it never seems to have more than it wishes to eat, so it is fed quite frequently by the mature workers. It would seem that the workers fear an excessive development, which, as we have seen, is most mischievous and ruinous, and work to prevent the same, by a mean and meager diet. The food is composed of pollen and honey. Certainly of pollen, for, as I have repeatedly proved, without pollen, no brood will be reared. Probably some honey is incorporated, as sugar is an essential in the nutrition of all animals, and we could hardly account for the excessive amount of honey consumed, while breeding, by the extra amount consumed by the bees, consequent upon the added exercise required in caring for the brood. M. Quinby, Doolittle, and others, say water is also an element of this food. But bees often breed very rapidly when they do not leave the hive at all, and so water, other than that contained in the honey, etc., cannot be added. This makes it a question if water is ever added. The time when bees seem to need water, and so repair to the rill and the pond, is during the heat of summer, when they are most busy. May this not be quaffed to slake their own thirst?

In six days the cell is capped over by the worker-bees. This cap is composed of pollen and wax, so it is darker, more porous, and more easily broken than the caps of the honey-cells; it is also more convex ([Fig, 26, k]). The larva, now full grown, having lapped up all the food placed before it, surrounds itself with a silken cocoon, so excessively thin that it requires a great number to appreciably reduce the size of the cells. These always remain in the cell, after the bees, escape, and give to old comb its dark color and great strength. Yet they are so thin, that cells used even for a dozen years, seem to serve as well for brood as when first used. In three days the insect assumes the pupa state ([Fig, 26, h]). In all insects the spinning of the cocoon seems an exhaustive process, for so far as I have observed, and that is quite at length, this act is succeeded by a variable period of repose. The pupa is also called a nymph. By cutting open cells it is easy to determine just the date of forming the cocoon, and of changing to the pupa state. The pupa looks like the mature bee with all its appendages bound close about it, though the color is still whitish:

In twenty-one days the bee emerges from the cell. The old writers were quite mistaken in thinking that the advent of these was an occasion of joy and excitement among the bees. All apiarists have noticed how utterly unmoved the bees are, as they push over and crowd by these new-comers in the most heedless and discourteous manner imaginable. Wildman tells of seeing the workers gathering pollen and honey the same day that they came forth from the cells. This idea is quickly disproved if we Italianize black-bees. We know that for some days these young bees do not leave the hive at all, except in case of swarming, when bees even too young to fly will essay to go with the crowd. These young bees, like the young drones and queens, are much lighter for the first few days.

The worker-bees never attain a great age. Those reared in autumn may live for eight or nine months, and if in queenless stocks, where little labor is performed, even longer; while those reared in spring will wear out in three, and when most busy, will often die in from thirty to forty-five days. None of these bees survive the year through, so there is a limit to the number which may exist in a colony. As a good queen will lay, when in her best estate, three thousand eggs daily, and as the workers live from one to three months, it might seem that forty thousand was too small a figure for the number of workers. Without doubt a greater number is possible. That it is rare is not surprising, when we remember the numerous accidents and vicissitudes that must ever attend the individuals of these populous communities.

The function of the worker-bees is to do all the manual labor of the hives. They secrete the wax, which forms in small pellets ([Fig, 27, a, a]) under the over-lapping rings under the abdomen. I have found these wax-scales on both old and young. According to Fritz Müller, the admirable German observer, so long a traveler in South America, the bees of the genus melipona secrete the wax on the back.

The young bees build the comb, ventilate the hive, feed the larvæ and cap the cells. The older bees—for, as readily seen in Italianizing, the young bees do not go forth for the first one or two weeks—gather the honey, collect the pollen, or bee-bread, as it is generally called, bring in the propolis or bee glue, which is used to close openings, and as a cement, supply the hive with water(?), defend the hive from all improper intrusion, destroy drones when their day of grace is past, kill and arrange for replacing worthless queens, destroy inchoate queens, drones, or even workers, if circumstances demand it, and lead forth a portion of the bees when the conditions impel them to swarm.

When there are no young bees, the old bees will act as house-keepers and nurses, which they otherwise refuse to do. The young bees, on the other hand, will not go forth to glean, even though there be no old bees to do this necessary part of bee-duties. An indirect function of all the bees is to supply animal heat, as the very life of the bees require that the temperature inside the hive be maintained at a rate considerably above freezing. In the chemical processes attendant upon nutrition, much heat is generated, which, as first shown by Newport, may be considerably augmented at the pleasure of the bees, by forced respiration. The bees, too, by a rapid vibration of their wings, have the power to ventilate their hives, and thus reduce the temperature, when the weather is hot. Thus they moderate the heat of summer, and temper the cold of winter.

Under Surface of Bee, showing Wax between Segments.

CHAPTER III.
SWARMING OR NATURAL METHOD OF INCREASE.

The natural method by which an increase of colonies among bees is secured, is of great interest, and though it has been closely observed, and assiduously studied for a long period, and has given rise to theories as often absurd as sound, yet, even now, it is a fertile field for investigation, and will repay any who may come with the true spirit of inquiry, for there is much concerning it which is involved in mystery. Why do bees swarm at unseemly times? Why is the swarming spirit so excessive at times and so restrained at other seasons? These and other questions we are too apt to refer to erratic tendencies of the bees, when there is no question but that they follow naturally upon certain conditions, perhaps intricate and obscure, which it is the province of the investigator to discover. Who shall be first to unfold the principles which govern these, as all other actions of the bees?

In the spring or early summer, when the hive has become populous, and storing very active, the queen, as if conscious that a home could be overcrowded, and foreseeing such danger, commences to deposit drone-eggs in drone-cells, which the worker-bees, perhaps moved by like considerations, begin to construct, if they are not already in existence. In fact, drone comb is almost sure of construction at such times. No sooner is the drone brood well under way, than the large, awkward, queen-cells are commenced, often to the number of ten or fifteen, though there may be not more than three or four. In these, eggs are placed, and the rich royal jelly added, and soon, often before the cells are even capped—and very rarely before a cell is built, if the bees are crowded, the hives unshaded, the ventilation insufficient, or the honey-yield very bountiful—some bright day, usually about ten o'clock, after an unusual disquiet both inside and outside the hive, a large part of the worker-bees—being off duty for the day, and having previously loaded their honey-sacks—rush forth from the hive as if alarmed by the cry of fire, the queen among the number, though she is by no means among the first, and frequently is quite late in her exit. The bees, thus started on their quest for a new home, after many uproarious gyrations about the old one, dart forth to alight upon some bush, limb, or fence, though in one case I have known the first swarm of bees to leave at once, for parts unknown, without even waiting to cluster. After thus meditating for the space of from one to three hours, upon a future course, they again take wing and leave for their new home, which they have probably already sought out.

Some suppose the bees look up a home before leaving the hive, while others claim that scouts are in search of one while the bees are clustered. The fact that bees take a right-line to their new home, and fly too rapidly to look as they go, would argue that a home is preêmpted, at least, before the cluster is dissolved. The fact that the cluster remains sometimes for hours—even over night—and at other times for a brief period, would lead us to infer that the bees cluster, in waiting for a new home to be found. Yet, why do bees sometimes alight after flying a long distance, as did a first swarm the past season, upon our College grounds? Was their journey long, so that they must needs stop to rest, or were they flying at random, not knowing whither they were going?

If for any reason the queen should fail to join the bees, and perhaps rarely, when she is among them, they will, after having clustered, return to their old home. The youngest bees will remain in the old hive, to which those bees, if there are any such, which are abroad in quest of stores will return. The presence of young bees on the ground—those with flight too feeble to join the rovers—will always mark the previous home of the emigrants. Soon, in seven or eight days, perhaps rarely a little later, the first queen will come forth from her cell, and in two or three days she will or may lead a new colony forth, but before she does this, the peculiar note, known as the piping of the queen, may be heard. This piping sounds like peep, peep, is shrill and clear, and can be plainly heard by placing the ear to the hive, nor would it be mistaken. It is followed by a lower, hoarser note, made by a queen still within the cell.

Some have supposed that the cry of the liberated queen was that of hate, while that by the queen still imprisoned was either of enmity or fear. Never will an after-swarm leave, unless preceded by this peculiar note.

At successive periods of one or two days, one, two, or even three more colonies may issue from the old home. These last swarms will all be heralded by the piping of the queen. They will be less particular as to the time of day when they issue, as they have been known to leave before sun-rise, and even after sun-set. The well-known apiarist, Mr. A. F. Moon, once knew a swarm to issue by moon-light. They will, too, as a rule, cluster farther from the hive. The after swarms are preceded by the queen, and in case swarming is delayed, may be attended by a plurality of queens. Berlepsch and Langstroth both saw eight queens issue with a swarm, while, others report even more. These virgin queens fly very rapidly, so the swarm will seem more active and definite in their course than will first swarms.

The cutting short of swarming preparations before the second, third, or even the first swarm issues, is by no means a rare occurrence. This is effected by the bees' destroying the queen-cells, and sometimes by a general extermination of the drones, and is generally to be explained by a cessation of the honey yield. Cells thus destroyed are easily recognized, as they are torn open from the side, and not cut back from the end.

Swarming out at other times, especially in late winter and spring, is sometimes noticed by apiarists. This is due to famine, mice, or some other disturbing circumstance, which makes the hive intolerable to the bees.