CHAPTER II.
KINDS OF BEES COMPOSING A COLONY—BEE PRODUCTS AND DESCRIPTION OF COMBS—DEVELOPMENT OF BROOD.

KINDS OF BEES IN A COLONY.

Fig. 5.—Ovaries of queen and workers: A, abdomen of queen—under side (magnified eight times); P, petiole; O, O, ovaries; hs, position filled by honey sac; ds, position through which digestive system passes; od, oviduct; co.d, common oviduct; E, egg-passing oviduct; s, spermatheca; i, intestine; po, poison bag; p.g, poison gland; st, sting; p, palpi. B, rudimentary ovaries of ordinary worker; sp, rudimentary spermatheca. C, partially developed ovaries of fertile worker; sp, rudimentary spermatheca. (From Cheshire.)

Each colony of bees in good condition at the opening of the season contains a laying queen and some 30,000 to 40,000 worker bees, or six to eight quarts by measurement. Besides this there should be four, five, or even more combs fairly stocked with developing brood, with a good supply of honey about it. Drones may also be present, even several hundred in number, although it is better to limit their production to selected hives, which in the main it is not difficult to accomplish.

Under normal conditions the queen lays all of the eggs which are deposited in the hive, being capable of depositing under favorable conditions as many as 4,000 in twenty-four hours. Ordinarily she mates but once, flying from the hive to meet the drone—the male bee—high in the air, when five to nine days old generally, although this time varies under different climatic conditions as well as with different races. Seminal fluid sufficient to impregnate the greater number of eggs she will deposit during the next two or three years (sometimes even four or five years) is stored at the time of mating in a sac—the spermatheca, opening into the oviduct or egg-passage (fig. 5, s). The queen seems to be able to control this opening so as to fertilize eggs or not as she wills at the time of depositing them. If fertilized they develop into workers or queens according to the character of the food given, the size and shape of the cell, etc.; if unfertilized, into drones. The queen's life may extend over a period of four or five years, but three years is quite as long as any queen ought to be kept, unless a particularly valuable one for breeding purposes and not easy to replace. Indeed, if full advantage be taken of her laying powers it will rarely be found profitable to retain a queen longer than two years.

Upon the workers, which are undeveloped females, devolves all the labor of gathering honey, pollen, propolis, and bringing water, secreting wax, building combs, stopping up crevices in the hive, nursing the brood, and defending the hives. To enable them to do all this they are furnished with highly specialized organs. These will be more fully referred to in connection with the description of the products gathered and prepared by the workers.

Fig. 6.—A, Head of queen, magnified ten times, showing smaller compound eyes at sides, and three ocelli on vertex of head; n, jaw notch. B, head of drone, magnified ten times, showing larger compound eyes at sides, with three ocelli between; n, jaw notch. (From Cheshire.)

The drones, aside from contributing somewhat to the general warmth of the hive necessary to the development of the brood, seem to have no other office but that connected with reproduction. In the wild state colonies of bees are widely separated, being located wherever the swarms chance to have found hollow trees or rock cavities, hence the production of many drones has been provided for, so young queens flying out to mate will not run too many risks from bird and insect enemies, storms, etc. Mating in the hive would result in too continuous in-and-in breeding, producing loss of vigor. As we find it arranged, the most vigorous are the most likely to reproduce their species.

At the time of the queen's mating there are in the hive neither eggs nor young larvæ from which to rear another queen; thus, should she be lost, no more fertilized eggs would be deposited, and the old workers gradually dying off without being replaced by young ones, the colony would become extinct in the course of a few months at most, or meet a speedier fate through intruders, such as wax-moth larvæ, robber bees, wasps, etc., which its weakness would prevent its repelling longer; or cold is very likely to finish such a decimated colony, especially as the bees, because queenless, are uneasy and do not cluster compactly.

The loss of queens while flying out to mate is evidently one of the provisions in nature to prevent bees from too great multiplication, for were there no such checks they would soon become a pest in the land. On the other hand, the risk to the queen is not uselessly increased, for she mates but once during her life.

Fig. 7.—Modifications of the legs of different bees: A, Apis: a, wax pincer and outer view of hind leg; b, inner aspect of wax pincer and leg; c, compound hairs holding grains of pollen; d, anterior leg, showing antenna cleaner; e, spur on tibia of middle leg. B, Melipona: f, peculiar group of spines at apex of tibia of hind leg; g, inner aspect of wax pincer and first joint of tarsus. C, Bombus: h, wax pincer; i, inner view of same and first joint of tarsus—all enlarged. (From Insect Life.)

BEE PRODUCTS AND ORGANS USED IN THEIR PREPARATION.

Fig. 8.—Head and tongue of Apis mellifera worker (magnified twelve times), a, Antenna, or feeler; m, mandibula, or outer jaw; g, gum flap, or epipharynx; mxp, maxillary palpus; pg, paraglossa; mx, maxilla, or inner jaw; lp, labial palpus, l, ligula, or tongue; b, bouton, or spoon of the same. (Reduced from Cheshire.)

Pollen and honey form the food of honey bees and their developing brood. Both of these are plant products which are only modified somewhat by the manipulation to which they are subjected by the bees and are then stored in waxen cells if not wanted for immediate use. Pollen, the fertilizing dust of flowers, is carried home by the bees in small pellets held in basket-like depressions on each of the hind legs. The hairs covering the whole surface of the bee's body are more or less serviceable in enabling the bee to collect pollen, but those on the under side of the abdomen are most likely to get well dusted, and the rows of hairs, nine in number, known as pollen brushes, located on the inner surface of the first tarsal joint (fig. 7, b), are then brought into use to brush out this pollen. When these brushes are filled with pollen the hind legs are crossed during flight and the pollen combed out by the spine-like hairs that fringe the posterior margin of the tibial joint—that above a in fig. 7. The outer surface of this joint is depressed, and this, with the rows of curved hairs on the anterior margin and the straighter ones just referred to forms a basket like cavity known as the corbiculum or pollen basket, represented by the longest joints of the legs, A, B, and C, fig. 7. Into this the pollen falls, and with the middle pair of legs is tamped down for transportation to the hive. Having arrived there, the bee thrusts its hind legs into a cell located as near to the brood nest as may be, and loosening the pellets lets them fall into the bottom of the cell. The tibial spur (fig. 7, e) on each middle leg is, as Professor Cheshire has pointed out, probably of use in prying the pellets out. The latter are simply dropped into cells and left for some other bee to pack down by kneading or pressing with its mandibles. Various colors—yellow, brown, red, slate, etc., according to the kinds of flowers from which gathered—frequently show in layers in the same cell. Often when partly filled with pollen the cell is then filled up with honey and sealed more or less hermetically with wax. The bees store the pollen, for convenience in feeding, above and at the sides of the brood and as near to it as possible, the comb on each side of the brood nest being generally well stored with it.

NECTAR AND HONEY.

The liquid secreted in the nectaries of flowers is usually quite thin, containing, when just gathered, a large per centage of water. Bees suck or lap it up from such flowers as they can reach with their flexible, sucking tongue, 0.25 to 0.28 inch long. (Fig. 8, l.) This nectar is taken into the honey sac (Plate II, h.s.) located in the abdomen, for transportation to the hive. It is possible that part of the water is eliminated by the gatherers before they reach the hive. A Russian bee keeper, M. Nassanoff, while dissecting a worker, discovered between the fifth and sixth abdominal segments a small canal, to which he attributed an excretory function, and Zoubareff, having noticed bees ejecting a watery substance while returning from the fields, suggested that this gland probably served to separate a portion of the water from the nectar, the liquid deposited in the cells appearing to contain less of it than that just secreted by the flowers.


Bul. 1, new series, Div. of Entomology, U. S. Dept. of Agriculture. Plate II.

Digestive System of Bee (magnified ten times).

A, Horizontal section of body; lp, labial palpus; mx,, maxilla: e, eye; dv, dv, dorsal
vessel: v, ventricles of the same; No. 1. No. 2, No. 3, salivary gland systems, 1, 2, 3;
œ, œsophagus; pro.t, prothorax; mesa.t, mesathorax; meta.t, metathorax; g, g,
ganglia of chief nerve chain; n, nerves; hs, honey sac; p, petaloid stopper of honey
sac or stomach mouth; c.s, chyle stomach: bt, biliary or Malpighian vessels: si,
small intestine; l, lamellæ or gland plates of colon; li, large intestine.


However this maybe, evaporation takes place rapidly in the heat of the hive after the nectar or thin honey has been stored, as it is temporarily, in open cells. Besides being thin, the nectar has at first a raw, rank taste, generally the flavor and odor peculiar to the plant from which gathered, and these are frequently far from agreeable. To make from this raw product the healthful and delicious table luxury which honey constitutes—"fit food for the gods"—is another of the functions peculiar to the worker bee. The first step is the stationing of workers in lines near the hive entrances. These, by incessant buzzing of their wings, drive currents of air into and out of the hive and over the comb surfaces. If the hand be held before the entrance at such a time a strong current of warm air may be felt coming out. The loud buzzing heard at night during the summer time is due to the wings of workers engaged chiefly in ripening nectar. Instead of being at rest, as many suppose, the busy workers are caring for the last lot of gathered nectar and making room for further accessions. This may go on far into the night, or even all night, to a greater or less extent, the loudness and activity being proportionate to the amount and thinness of the liquid. Frequently the ripening honey is removed from one set of cells and placed in others. This may be to gain the use of certain combs for the queen, or possibly it is merely incidental to the manipulation the bees wish to give it. When, finally, the process has been completed, it is found that the water content has usually been reduced to 10 or 12 per cent, and that the disagreeable odors and flavors, probably due to volatile oils, have also been driven off in a great measure, if not wholly, by the heat of the hive, largely generated by the bees. During the manipulation an antiseptic—formic acid—secreted by glands in the head of the bee, and it is also possible other glandular secretions, have been added. The finished product is stored in waxen cells above and around the brood nest and the main cluster of bees, as far from the entrance as it can be and still be near to the brood and bees. The work of sealing with waxen caps then goes forward rapidly, the covering being more or less porous.

Each kind of honey has its distinctive flavor and aroma, derived, as already indicated, mainly from the particular blossoms by which it was secreted, but modified and softened by the manipulation given it in the hives. When the secretion is abundant in a flower having a short or open corolla, hence one from which the bees find it easy to obtain the honey, they will confine their visits to that kind if the latter is present in sufficient numbers. Thus it is that linden, white clover, buckwheat, white sage, mesquite, sourwood, aster, tulip tree, mangrove, orange, and other kinds of honey may be harvested separately, and each be readily recognizable by its color, flavor, consistency, and aroma. When, however, no great honey yielder is present in large quantity and the source is miscellaneous, all manner of combinations of qualities may exist, introducing great and often agreeable variety. Thus the medicinal qualities and the food value of different kinds of honey differ as greatly as do their prices on the market.

PROPOLIS.

This substance, commonly known as "bee glue," is obtained by the bees from the buds and crevices of trees, and is carried to the hives in the corbicula or basket-like cavities on the outside of the tibial joints of the workers' hind legs, the same as they carry pollen. The workers with their mandibles scrape together and bite off the particles of propolis, and with the front and middle legs pass them back to the baskets, where the middle legs and feet are used to tamp them down. The pellets can be readily distinguished from those of pollen, the latter being dull and granular in appearance, while the freshly gathered propolis is compact and shiny. This resinous material, which becomes hard soon after it is gathered, is at first quite sticky, and the bee bringing it requires aid in unloading. Another worker takes hold of the mass with its jaws, and by united exertion they get it out of the pocket, though often by piecemeal and in long threads. It is not stored in cells, but is used at once to stop up crevices in the hives and to varnish the whole interior surface, as well as to glue movable portions fast, also in strengthening the combs at their attachments, and if the latter are designed exclusively for honey, and especially if not filled at once, the edges of their completed cells receive a thin coating of propolis, which adds considerably to their strength. The bees often make the flight hole smaller by filling a part of it with masses of propolis, sometimes mixed with old wax. Carniolans gather the least and Tunisians the most propolis of any of the different races. On this account the former are better suited than the latter to the production of fancy white comb honey.

BEE POISON AND THE STING.

The worker and the queen are supplied with another organ which is of great importance to them, namely, the sting; for without this the hard-earned stores of the hive would soon be a prey to all manner of marauders, and the queen would be deprived of an organ of occasional use to her in dispatching rivals, and of daily use to her during the working season in the deposition of eggs. The darts work independently and alternately, and are connected at the base with the poison sac, without whose powerful contents such a tiny weapon would be wholly ineffective. Poison glands pour an acid secretion—largely formic acid—into this sac, whence it is conveyed to the tip of the sting along the groove or canal formed by the junction of the sheath and the darts. The sting being but an ovipositor modified also another purpose in addition to oviposition, in the perfect female (the queen) its main use is in placing the eggs in their proper position in the bottoms of the cells.

Formic acid is known to have considerable antiseptic properties. Chemical tests show its presence in well-ripened honey, but not in freshly gathered nectar. The natural conclusion is that it has been added by the bees to assist in the preservation of the honey. In what manner it is supplied has frequently been questioned. Tests applied to the blood of the bee show its presence there, and the secretions of the head glands show still larger quantities. It is therefore reasonable to suppose that these glands, as well as the poison glands themselves secrete formic acid, and that the honey receives its portion from the former, the head glands, upon being disgorged from the honey-sac or during the manipulation to which it is subjected in the hive.

WATER.

During cold or cool weather much condensation of moisture takes place in wooden hives as these are usually arranged. The water, collecting in drops on the interior walls of the hive and on the cold, sealed honey, often trickles down over the cluster of bees, to their great injury. It has been claimed that when brood rearing begins this condensed moisture will be utilized in the preparation of brood food. Very possibly it may, yet its use is probably detrimental, since it is charged with waste products of the hive—those of respiration, etc. In its absence the water contained in the honey, if the latter has not granulated, seems to be sufficient. Later, however, when no condensation takes place in the hive and the greater number of developing larvæ require considerable supplies of water in their food, special trips are made to brooks and pools for it, and dew is often gathered from leaves.

SILK.

The larval bee produces a small amount of silk from glands in its head. The pupal cell is partially lined with this. Later, as the bee develops, there being no further use for the glands, they become atrophied.

WAX.

The light colored pellets which are carried into the hive on the hind legs of the workers, and which have been described as pollen, are often mistaken for wax. The fact is, wax is not gathered in the form in which we see it, except in rare instances, when, bits of comb having been left about, small quantities will be loaded up and taken in as pellets on the legs. Ordinarily it comes into the hive in the shape of honey and is transformed by the workers within their own bodies into wax. This production is wholly confined to the workers, for although the queen has wax plates 011 the underside of the abdomen and wax glands beneath them, yet both are less developed than in the workers and are never used. The wax plates of the worker overlying the secreting glands are well shown in fig. 9, those of the queen and of the related genera, Bombus and Melipona, being shown for comparison. During wax secretion, that is, when combs are being built or honey cells sealed over, a high temperature is maintained in the hive, and many workers may be seen to have small scales of wax protruding from between the segments of the abdomen on the underside. The molds or plates, eight in number, in which the scales appear are concealed by the overlapping of the abdominal segments, but when exposed to view (fig. 9, a) are seen to be five-sided depressions lined with a transparent membrane. The wax glands themselves are beneath this membrane, and through it the wax comes in a liquid form. As the scales harden they are pushed out by the addition of wax beneath. The bees pluck them out with neat pincers (fig. 7, a and b) formed by the articulation of the hind tibiæ with the adjacent tarsal joints, pass them forward to the mandibles, and mold them into the shape of hexagonal cells, meanwhile warming and moistening them with the secretions of the head glands to render the wax more pliable.

Fig. 9.—Wax disks of social bees: a, Apis mellifera worker; b. A. mellifera queen; c, Melipona worker; d, Bombus worker—all enlarged. (From Insect Life.)

COMBS.

Wax is fashioned by the workers into cells of various sizes and shapes, according to the use to be made of them. The most regular in shape and size are the cells designed for brood (fig. 4). These combs in which workers are bred show nearly 29 cells on a square inch of surface, the combs being seven-eighths inch thick and the cells generally quite regular hexagons in outline. Drone cells are larger, there being but 18 of them to the square inch of surface, and the comb is 1¼ inches thick. The cells of combs designed only for honey are frequently more irregular in shape, generally curve upward somewhat, and are often deepened as the honey is stored in them, so that these combs sometimes reach a thickness of 2 or 3 inches.

The cells in which queens are bred bear in size and shape some resemblance to a ground or pea nut. They are often irregular in form, being sometimes curved, or short and thick, according to the space below their point of attachment, which is most frequently the lower edge or the side edge of a comb, or sometimes a mere projection or angularity in the general surface of a comb. Queen cells open downward instead of being built horizontally like drone and worker cells (figs. 62 and 63).

Fig. 10.—Comb building—side of hive removed. (Original.)

Into the material used in constructing brood combs bees often incorporate bits of wax and fiber-like gnawings of cocoons from old combs in which brood has been reared, and if given cappings or trimmings of combs they will work them all over and utilize most of the material. Also when the bees have abundant supplies of pollen much of this is incorporated into the material of brood combs, thus saving the costlier substance—wax. Such combs show at once by their brownish or straw color, even when first constructed, that they are not made of wax alone. It will readily be seen from the above that the quantity of honey consumed by the bees in producing a pound of comb must vary greatly, for if the comb is designed for surplus honey it will be made of newly secreted wax, while if for brood other material will, as just stated, replace a portion of the wax. The amount of honey coming in varies from day to day, and it is difficult to estimate how much is consumed in feeding and keeping warm the brood: moreover, a high temperature is required in the hive to facilitate the secretion and working of wax, necessitating, of course, extra food consumption when the outside temperature is low. Accordingly estimates as to the amount of honey required to produce 1 pound of comb range from 5 pounds to 25 pounds. More accurate experiments are needed in this direction before anything positive can be stated. Until then 18 to 20 pounds might be looked upon as nearest the correct figure for white surplus combs, and half as much for dull straw-colored or brownish combs built for brood rearing.

DEVELOPMENT OF BROOD.

Ordinarily the winter cluster in a hive of bees occupies the more central combs, four or five in number. Near the middle of this cluster the queen deposits the first eggs of the season (which are fertilized eggs) in the small-sized or worker cells. Under favorable circumstances, that is, in a strong colony amply protected against inclement weather, this deposition usually occurs in January, though in a very mild climate some brood is generally present during every month of the year, and the cessation of egg-laying is very short. The eggs hatch on the third day after deposition into minute white larvæ, to which the workers supply food in abundance. The composition of this food has been the subject of much attention and more theorizing. It may be considered as pretty certain that during the first three days of the life of the larva its food is a secretion from glands located in the heads of the adult workers—a sort of bee milk, to which, after the third day, honey is added in the case of the worker larvæ, and honey and pollen in the case of drone larva?. As this weaning proceeds both worker and drone larvæ receive pollen, and in constantly increasing proportions, in place of the secretion. But this rich albuminous substance is continued to the queen larvæ throughout their whole period of feeding; moreover, the quantity of this food supplied to each queen larva is apparently super-abundant, for after it ceases to feed quite a mass of the food somewhat dried out will be found in the bottom of the cell from which a well-developed queen has issued. After assuming the pupa form the young queen is attached to this food by means of the tip of the abdomen, and it very likely continues for some time to receive nourishment from the mass.

The following table shows approximately the time occupied in the development of worker, drone, and queen:

Egg. Larva. Pupa. From deposition
of egg to imago.
Days. Days. Days. Days.
Queen 3 7 15½
Worker 3 5 13 21
Drone 3 6 15 24

The original circles of brood are gradually increased by the deposition of eggs in the cells next outside those already occupied, and circles are soon begun in the adjoining combs. In this way the space occupied by the developing bees is gradually increased, while preserving the general spherical shape of the brood nest thus formed, which, however, the shape of the hive often modifies somewhat. As already mentioned, each circle of brood has rows of pollen cells about it, chiefly above and at the sides, and the combs on either side contiguous to the brood are usually well packed with pollen. Outside of the pollen most of the honey on hand is stored. Thus (fig. 11) a cross section made in any direction through the middle of a hive in normal condition at the opening of the active season should show this relative arrangement of brood, pollen, and honey, which economizes most the heat of the hive and the labors of the nurse bees, favoring in this way the rapid increase of the population.

Fig. 11.—Cross section of brood apartment: s, s, sides of hive; t, t, top-bars of frames; h, p, l, sb, combs containing (h) honey, (p) pollen, (l) larvæ and eggs, and (sb) sealed brood. (Original.)

THE WORKER.

The worker larvæ are fed five days, and then the cell is given by the adult bees a covering which is quite porous by reason of numerous pollen grains incorporated into its mass, this openness of texture being necessary to give the developing bee air to breathe. The larva strength ens this capping by a loose webwork of silk within, extending down the side but slightly and attached at its edges to the last skin cast by the molting larva. This skin, extremely delicate and pressed closely against the inside of the cell, forms the lining of its sides and bottom. In about twelve days after sealing, that is, twenty-one days from the time the egg was deposited, the imago, or perfect bee, bites its way through the brown covering.

In the course of a couple of days it takes up the work of a nurse, and in a week to ten days may appear at the entrance on pleasant days, taking, however, but short flights for exercise, as ordinary field work is not undertaken until it has passed about two weeks in the care of brood. The worker then takes up also wax secretion, if honey is to be capped over or combs built, although old bees can and do to a certain extent engage in wax production.

THE DRONE.

Eggs left unfertilized produce drones and require twenty-four days from the time they are deposited until the perfect insect appears. They are normally deposited in the larger-sized horizontal cells, and when the latter are sealed, the capping is more convex as well as lighter-colored than that of worker brood, which is brown and nearly flat.

The fact that drones develop from unfertilized eggs is to be noted as having an important practical bearing in connection with the introduction of new strains of a given race or of new races of bees into an apiary. From a single choice home-bred or imported mother, young queens of undoubted purity of blood may be reared for all of the colonies of the apiary, and since the mating of these young queens does not affect their drone progeny, thereafter only drones of the desired strain or race and pure in blood will be produced, rendering, therefore, the pure mating of future rearings fairly certain if other bees are not numerous within a mile or two. Eventually also all of the colonies will be changed to the new race and without admixture of impure blood, provided always that the young queens be reared from mothers of pure blood mated to drones of equal purity.