# Creation by Evolution/The Evolution of the Bee and the Beehive

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THE EVOLUTION OF THE BEE AND THE BEEHIVE
By Sir Arthur Everett Shipley
Master of Christ’s College, Cambridge
For so work the honey-bees, Creatures that, by a rule in nature, teach The act of order to a peopled kingdom. They have a king and officers of sorts, Where some, like magistrates, correct at home. Others, like merchants, venture trade abroad; Others, like soldiers armed in their stings, Make boot upon the summer’s velvet buds, Which pillage they with merry march bring home.
Shakespeare.
In a primitive and savage state of society each individual of a tribe is a host in himself. He is at once a
Tinker, tailor, soldier, sailor, ’Pothecary, ploughboy, thief,
and except that he cannot very well be his own undertaker he performs all the functions of the various traders and professional experts that in a more civilized state of society are carried on by numerous men, each suitably trained for one pursuit, and generally for only one. The North American Indians built their own wigwams, tilled the soil, fished, hunted, fought in tribal wars, and engaged in other activities. The women took a large part in the drudgery of life—cooking, tending the young, helping in shifting the camp. But as affairs became more complicated a higher social order was ​established. The medicine man, who acted both as priest and doctor, was evolved. A chieftain was set up. The old men became counsellors. Still each family lived in its own wigwam and not with others in an apartment house or a hotel.
Certain social communities other than human have in a similar way evolved from simple beginnings, and one of the highest of these is undoubtedly that of the honey-bee; and the society of the honey-bee is even more complex than anything in our own civilization. “The bee in its own line,” writes J. A. Thomson, “is hardly inferior to man, and represents an achievement that angels might desire to look into.”
In a beehive there are three ranks of individuals. First, there is the queen bee (Fig. 1), who is indeed the mother of her people, for she alone lays eggs; and as a rule she is a solitary monarch, and tolerates no rivals. Then there are the workers (Fig. 2), which in structure are females, though they have ceased laying eggs. Like Martha, they are cumbered with much serving. Third, there are the males, or drones (Fig. 3), quite useless in the conduct of the affairs of the hive except that one of them will ultimately fertilize the queen bee.
The cells of the drones, having to accommodate a rather larger larva, are made slightly bigger, and in some wild honey-bees they are all placed together in a special drone comb. The cell in which the queen is reared is, however, altogether different. It is about the size of an acorn, and its walls are much thicker than those of the other cells and are usually rounded. As soon as the queen bee is hatched out these walls are destroyed and their wax is used to add more worker cells to the comb. An average hive (Fig. 7) will contain some 30,000 working bees, some 2,000 drones, and but one queen, who alone is ​a functional female and produces a continuous flow of eggs.
Fig. 7.—Comb of hive bee (Natural Size).
A, empty queen cell; B, the same, torn open; C, the same, cut down; D, drone larva; E, F, sealed drone cells; G, sealed worker cells; H, old queen cell; I, sealed honey; K, masses of pollen; L, pollen cells; M, abortive queen cell; N, emerging bee; O, eggs and larva. (After Cheshire.)
The queen moves along on her egg-laying journey, exploring every empty waxen cell with her feelers and inserting a single egg into each cell. She never seems to tire, and she never misses a cell. During her progress she is surrounded by a small court of worker bees, who act as courtiers, walking backward. Some of them fan her with their wings; others stroke her with their tongues; still others feed her with half-digested pap, or “royal jelly,” and all are humming most agreeably and soothingly. During May and June ​the queen will lay three thousand to four thousand eggs every four and twenty hours, and in the course of her life of four or five years she produces hundreds of thousands of eggs. But should the number of bees in the hive decrease she will cease laying eggs, as there will not then be sufficient workers to attend the resultant larvae.
As soon as an egg is placed in a cell the worker bees get busy. They push their heads into the cell and seem to do something to the egg, though what it is is not clearly known. Within three or four days a very small white maggot-like grub (Fig. 8) emerges from the eggshell. It has no legs and is devoid of everything we associate with insects—it has no wings, no stings, no feelers, no eyes, and its intestine ends blindly.
For the first day or two the young larvae are fed from the secretion of the salivary glands of the workers. This is known as pap, or “royal jelly.” The larvae not only lap this up, but float in it. On the fourth day this food is mixed with honey, and henceforward the drones are completely weaned and feed entirely on honey and pollen. The queen bee, on the other hand, lives on nothing but royal pap. After about six days the larvae cease to feed. They are then sealed up in their cells (see Fig. 8) by the worker bees and each larva makes a cocoon case, in which it forms a chrysalis or pupa.
“Three facts emerge from a study of this community:
“1. The lesson of solidarity, of the social spirit, to which the interests of the individual are subservient.
“2. The distribution of labour in accordance with the law of mutual help, each doing his work like an instrument in a vast orchestra and all producing a beautiful harmony.
“3. The law of sacrifice for the sake of the future race.”
Each individual is so wrapped up in the community that if isolated from its fellows it dies. The constant sense of mutual help, of self-sacrifice for the future race, is the dominating characteristic of all bees, and there is something that Maeterlinck calls the “spirit of the hive,” which in some way guides, directs, and controls the work of this strange, self-sacrificing community. Here there is no private property. As Dryden says in his translation of Vergil’s book about the bee: “All is the State’s; the State provides for all.” In a passionate devotion to duty and in an energy expended solely for others, in a single-minded purpose, the queen and the worker honey-bee are unique among animals. Now, how has ​this wonderful socialistic life come about? How has it been evolved? How can we discover the steps in its evolution? We can trace the social bees and wasps back to solitary bees and wasps, and we can trace a steady growth of complexity in the habits of life of these solitary insects and in the complexity of their homes until we reach the stage that is briefly described above.
One of the characteristics of the bee, as anyone can observe, is a hairy body. The body is so completely covered with hair that it has a furry appearance. Now the simplest form of bee, which has no common English name but is known scientificially as Prosopis, has hardly any hair. Its tongue is rudimentary, its hind legs are not adapted for collecting pollen, as are those of the honey-bee, and it does not lead a social life. It makes separate cells, each lined with a silken membrane, in the stems of such plants as brambles (Fig. 10) ; or it burrows in the earth, or even in the mortar of walls. It collects little if any pollen and it stores in its separate cells a very weak honey, in which the egg is ​laid. This bee is common in America as well as in Great Britain.
Here we have a bee that has not developed the typical hairs of a honey-bee, that collects little or no pollen, that stores the cells in which eggs are laid with thin honey, which it brings straight from flowers and does not first deposit in honey cells—a bee that produces separate and distinct cells, which may or may not be in contact.
A little higher up in the scale of progress we find another group of bees, which burrow tunnels in sandy soil, some of them nearly a foot in length. The tunnels and the cells are lined with a paper-like material, and the cells are divided by partitions, which may or may not be in contact. These cells are furnished with a fluid mixture of pollen and honey, both of which have been swallowed by the mother bee. All this shows an advance over the work of the bee first described, inasmuch as pollen forms a conspicuous part of the food of the larvae and there is a common entrance through a tunnel to the cells (Fig. 11). The nourishing fluid is more liquid than that supplied by the higher bees, and the papery lining ​is formed from a slime that gradually hardens. In this group, as in many others, the male is considerably smaller than the female.
Another group of bees falls under the common name of leaf-cutting bees (Fig. 13) . This bee is more robust than the ordinary hive bee and has a broader head. It makes nests in hollows in stems, in wood, or in the soil. The cell is made of leaves or of parts of leaves or petals of roses and other plants, which are moulded into a thimble-like form that has a lid composed of a smaller round piece of leaf. The cells are placed end to end and not side by side, and the pieces of leaves are gummed together. The string of cells thus made rarely exceeds seven. When completed each cell is half ​filled with pollen, on which an egg is laid. Other species of this group enter houses in India, and both sexes there take part in making cells of clay, which may be set in any hollow tube, such as the barrel of a gun or the hollow in the back of a book which is lying open, or in the interior of a piece of bamboo.
Then we have the mason bees, which construct nests of sand or soil or clay moulded together with some sticky substance. Externally each cell is rough and untidy, but inside it is smooth and polished. Generally ten to twenty cells form a nest, and each cell is stored with a mixture of honey and pollen. Some of these mason bees are very hairy, and the two sexes differ from each other in colour. In its general appearance this bee is something between a humble-bee and a honey-bee, but it is solitary in its habits. Each cell may be an inch deep, and here we see pollen being carried on ​the hairs of the under surface of the body. To place this pollen in the cell the bee enters backward and, with the aid of its hind-legs, brushes and scrapes and combs the pollen off from the under surface of its body so that it falls into the cell. This is a distinct advance on what we had at the beginning of our series, where the pollen is swallowed and brought up again. The pollen and honey are, however, not kept separate, but are worked up by the jaws of the bee into a paste, on which the egg is laid, and the cell is then closed with cement. The work of building this cell takes about two days, and after it is finished the bee will begin to make a second cell close to the first, and will continue its work until it has made eight or nine cells, when it places a thick, dome-like layer of mortar over the whole series. The result is a nest about the size of half an orange. The larvae live in these nests for months; they do not pass through their life-history so rapidly as the honey-bee.
An equally ingenious insect is the carder bee, which has developed the habit of making nests of wool or cotton, obtained from plants that grow in the neighbourhood. This bee is referred to by Gilbert White in his “Natural History of Selborne.” The male, like that of the honey-bee, is conspicuously larger than the female. These carder bees build their nests in any hollow, such as a cavity in wood or a deserted nest of other bees, or in an empty snail shell. In order to retain in the cell the fluid mixture of pollen and honey they line the cell with a thin cement. A few allied species form their cells of resin instead of wool or cotton.
The last of the solitary bees we shall consider are the carpenter-bees. These are big, burly black or bluish-black bees. They have powerful jaws, with which they carve their way into dried wood. They avoid living timber, but they will bore a hole into a beam or a rafter, and this hole will lead ​into three or four parallel galleries, in which they place their broad cells. Between the cells they make partitions formed of fragments of wood cemented together by their saliva. These bees pass the winter in the adult stage, hibernating in the imago condition. Both sexes reappear in the spring, and some species may take two years to complete the cycle of their life history. They are very hairy, and some of the females closely resemble the bumble-bee. The cells are provisioned with pollen, and the bees apparently produce little or no honey.
If we now turn to the social bees we find three groups. One is known as the mosquito bee, from its very small size. These bees are also sometimes spoken of as stingless bees, though they have a rudimentary sting, which they do not use. Little is known about them, but they form communities consisting of a large number of individuals. We do not certainly know whether these bees are all the product of a single queen or whether there may be more than one egg-producer in each colony, but the evidence seems to show that every colony has its own queen. The nests are rich in honey, and to prevent them from being robbed the workers, who are usually occupied in collecting pollen, also collect clay, with which they build a wall to protect the nest, which is generally placed on a bank or in the trunk of a tree. Every nest is thus completely surrounded with clay. The honey is stored in separate cells or in clusters of cells, each cluster about the size of a pigeon’s egg, and these are placed at the bottom of the hive, away from the cells where the larvae are growing. The comb made by some species resembles a spiral staircase, and there are special cells for the pollen as well as for the honey; and here, for the first time, we find wax used to form the comb. Here also we find the three separate castes, the queen or queens, producing eggs; the working bees, or bar​ren females; and the drones, or males. In some nests the wax is mixed with resin or gum, which makes it darker. The inside of the nest, like the inside of the hive of the honeybee, is dark. Sufficient food, consisting of pollen and honey, is placed in each cell, and on this the egg is deposited by the queen bee or queen bees. Occasionally a bee leaves the nest, and apparently in many nests all three castes are reared in identical cells on a similar diet. There is no such specialization as that shown in the hive of the honey-bee. Another point of difference is that among the mosquito-bees the cells in which the larvae mature are sealed up. After the egg is laid there is no contact between the larva and the mother or the workers. The drone has not degenerated into the “waster” that he becomes in the hive of the honey-bee but takes part in cementing the wax for the cell walls. The entrance to the hive is guarded during the day by certain sentinels and is closed at night by a mixture of wax and gum.
When we come to the bumble or humble bee we find still further progress toward the state of things we find in the hive of the honey-bee. The bumble-bee has a sting but seldom uses it, and as the poison is weak the pain it inflicts is much less than that produced by the sting of the honey-bee. On the other hand, when once the bee has stung it can withdraw the sting and use it again. This the honey-bee cannot do; its stinging results in its death. The life of the bumble-bee is less orderly than that of the honey-bee. There is less of that irritating efficiency, and there is much more litter; after all what would life be without litter! Bumble-bees are found nearly everywhere in the world except in Africa and Australia, but they prefer a temperate climate. There are hundreds of species of this genus, and seventeen of these are found in Great Britain.
To describe the life of the bumble-bee we may begin with ​the queen late in the summer. The nest is now dying down; in fact, the activity of the hive lasts only a few months, not all the year round, as does that of the stingless bee. Late in July or August the community begins to rear up queens. Once grown up the queen leaves the nest and hides in some cranny or among some debris. Here she is sought by the male. Once fertilized, the queen abandons the nest, which falls into a state of “death, damnation, and decay.” She now seeks winter quarters and, having filled her crop with honey, she goes into retreat for eight or nine months, hiding high up in banks or in burrows under trees. At first she sleeps lightly and can be easily aroused. Later she sinks into a deep lethargy and appears to be dead. But as the spring advances she gradually resumes her activities. She emerges and begins to collect pollen. As the days lengthen her desire to start a colony becomes overwhelming and she seeks a home. She may find some burrow abandoned by a fieldmouse, which is commonly approached by a tunnel.
Having found her home, she flies backward and forward from it, gradually increasing the length of her trial flights. This she does so that she may find her way home after raiding the flowers for pollen and honey. She mixes the two, and in the centre of the nest constructs a small pillar of the resultant paste (Fig. 14), and on this she moulds a circular wall of wax. In this rough, irregularly-shaped cell she lays a batch of eggs, usually about a dozen, and seals them in with wax. She then broods like a hen over the cell and does not leave her offspring night or day except to gather food. But she has to provision the nest, and for this purpose she prepares a waxen spherical honey-pot, which may be as big as a thimble. This is a frail affair of thin, soft wax, but it is water-tight and is capable of lasting some weeks. Arriving at the entrance to the nest, the queen refreshes herself as she ​is passing in and out, and by night the honey-pot may be quite empty of its thin and watery contents. In about four days the larvae hatch out as whitish grubs and begin to feed upon the pollen bed upon which they have been lying. At first they feed upon any mixed pollen and honey provided by the queen. As they grow older they are individually and compulsorily fed.
Should the first three or four batches of larvae hatch healthy and vigorous workers, the queen, who is now evidently tired, ceases to leave the hive and confines herself to laying eggs and helping with the necessary indoor work. When fully grown the workers cease to use the queen’s ​honey-pot, which now falls into decay, and they store in the papery cocoons they have vacated the honey they have themselves collected, strengthening the edges with wax.
Some species of bumble-bees construct special honey-pots of their own, as many as twenty or more in a hive. These contain a very watery syrup, which is eaten up daily, but the honey in the cocoons is thicker and seems to be used to feed the younger queens. A few species make special receptacles for the pollen, which is mixed with honey. The comb made by these species is irregular and rough compared with that of the honey-bee, which shows mathematical rigidity. It is placed on the basal irregular waxen layer of vacated cocoons, on which also are placed cells containing larvae and pupae. Sometimes the whole comb may be covered by a waxen dome, but there is always room left for the bumble-bees to circulate. These bees have evidently an acute sense of smell, and human breath is particularly distasteful to them. They are almost as clever as honey-bees in their power of scenting out nectar and, owing to the length of the tongue, a bumble-bee can probe flowers to reach nectar that lies beyond the reach of the honey-bee. The bumble-bee fertilizes the honeysuckle, the horehound, and the red clover, whose introduction into New Zealand proved a failure until bumble-bees were brought in to fertilize it.
The hive of the bumble-bee is kept up for only three or four months. But the inmates are very busy; in fact, they work themselves to death. They begin foraging earlier in the morning than the honey-bee and they continue foraging till dusk. They spend the night in attending the young and brooding over the cocoons, for they never sleep. After laying from 200 to 400 eggs and slaving to bring up her progeny, the queen, as the season closes, begins to lay special eggs that are destined to turn into males and fertile females. ​The cocoons for future queens are larger and may be readily recognized. But at present there is no evidence that the queen larva is fed on a special diet. Royalty seems to be inherent in the egg and not induced by special feeding. Unlike the useless and swaggering drone of the honey-bee the male bumble-bee leaves the hive and finds flowers for itself. It is no charge on the resources of the community. Several scores of males and queens are produced, and when hatched out they also leave the hive, are fertilized, and go into winter quarters. The queen ages rapidly; her hair drops off and she gradually ceases to lay eggs. As the new queens grow up on the rich and ample store of food provided in the hive the workers become listless. Flowers are becoming scarce, and one by one the bees grow torpid and drop asleep, and from this sleep there is no awakening.
The bumble-bee is certainly more human and less exasperating than the honey-bee. It has none of its monotonous perfection of organization. The queen has something of a mother in her. She is not reduced to a mere egg-laying apparatus, which lays eggs with the regularity and inevitableness of a recurring decimal. The bumble-bee queen broods over her young and nurses them with “a mother's tender care.” The workers work as hard as do the honey-bees, but they are less self-conscious and less self-satisfied, and the drones at any rate have the grace to provide for themselves during their brief life. One has a feeling that one might appeal to the better instincts of a bumble-bee, but that it would be perfectly useless to make such an appeal to a honey-bee.
Now let us summarize the results of the research we have made to discover the steps in the evolution of the honey-bee, with its wonderful social system. The most primitive bee makes a small cell or nest in the ground (Fig. 16), packs ​it with pollen, usually mixed with honey, deposits an egg, covers the cell in, and leaves the young larva to eat up the food provided. The grub or larva then turns into a pupa, from which emerges the active adult insect, which makes its way out into the world. The next stage higher is shown in Fig. 17.
The cells are placed side by side or end to end, as are those of the leaf-cutting bee or the carpenter-bee; but each individual cell of both these bees is furnished with food and an egg and then left alone. The young bee does not ​receive any care or help from its mother. In the bees named the male may be smaller than the female and may not be differentiated into a lazy, idle drone.
In the next stage of progress we have a small colony, which inhabits a nest that has a common entrance, marked by an arrow in Fig. 17. In this stage the cells may be side by side, as in Fig. 11, or end to end, as in Fig. 18. In Fig. 17 the colony is surrounded by a specially protected case, such as we find in more complicated hives of bees and wasps. Finally, in Fig. 19, we find a number of cells side by side, which by pressure may become six-sided. Most primitive bees collect more pollen than honey and secrete no wax. Some bees make cells of leaves or of a substance that they secrete, which becomes papery; some carve cells out of wood; some cover their cells with a dome-like layer of mortar; and in many of these primitive nests the larvae spend months and months before hatching out.
When we reach the social bees—that is, the bees that live together in societies—we find that the most primitive are the mosquito-bees; but whether their communities are the product of a single queen or whether there is more than one egg-producer in their midst is not clear. Here we find, for the first time, special cells or collections of cells set apart for storing honey and other special cells set apart for storing pollen; and here, for the first time, we find wax, of which a comb is built up. This wax is a special secretion of the bee's body. Here again we find that the colonies have separated into queen, or queens; working bees, or females that do not lay eggs; and drones, or males. The fertile and unfertile females are reared on a similar diet. The larva is always sealed up, and once the egg is laid the young are deprived of a mother's care. And here again the drone takes part in the common activities of the hive.
​The next stage toward the final product—that is, the honey-bee with its hive—is that of the bumble-bee. The hive of the bumble-bee, unlike that of the honey-bee, dies down during the winter, and the life instincts of the community are carried along in the body of the queen or queens, which retire into winter quarters early in the autumn. In the bumble-bee's hive as first formed there are no regular waxen cells, but the larvae grow up in an irregularly-shaped cell, which is sealed into a waxen covering. Over this the queen bee broods like a hen. Here also we have a specially prepared honey-pot situated near the entrance to the hive. This is not a modification of the ordinary cell, as in the comb of the honey-bee. In the bumble-bees the males take part in the work of the hive. The honeycomb is irregular and rough and may or may not be covered by a waxen dome. The queen bumble-bee lays only 200 to 400 eggs, which is a small number compared with the tens of thousands laid by the queen honey-bee. In the hive of the honey-bee we have true and exact hexagonal cells, each wall of which takes part in forming one side of the surrounding cells. Some of the cells are rather bigger, and these contain the heavy, overgrown drones. Others (but only a few) are still bigger and form irregular lumps of thick wax. Each of these big cells houses an egg, which is destined to become a queen. Other cells are set apart for the storage of pollen and still others for the storage of honey; but of course most of the cells that form the comb contain a single egg, which produces a grub or larva that receives hourly attention from the sterile workers, who act as foster mothers.
It has now been shown that there is a gradual development or evolution from a single pair of bees that make a single cell, isolated and self-contained, through a series of grades. The cells become more and more packed together till they ​reach the stage of the comb of a honey-bee, and there is developed a community of insects that rivals in complexity and in division of labour anything that we meet with in human communities. A clearer example of evolution could hardly be imagined—the gradual development from a simple primitive state of life to one of the highest complexity.
REFERENCES
* Edwardes, Tickner. The Lore of the Honey-Bee. London, Methuen & Co.
* Maeterlinck, Maurice. Le Vie des Abeilles. Paris, Bibliothèque Charpentier.
* Maeterlinck, Maurice. The Life of the Bee (translated by Alfred Sutro) . London, George Allen & Sons.
* Shipley, A. E. Life. Cambridge University Press.
* Shipley, A. E. Studies in Insect Life. London, T. Fisher Unwin.
* Sladen, F. W. The Humble Bee. London, Macmillan & Co., Ltd.
* Stabler, Hans von. Die Biologie der Biene. Wurzburg, H. Sturtz.
“The bees have existed many thousands of years; we have watched them for ten or twelve lustres. And if it could even be proved that no change has occurred in the hive since we first opened it, should we have the right to conclude that nothing had changed before our first questioning glance? Do we not know that in the evolution of species a century is but as a drop of rain that is caught in the whirl of the river, and that millenaries glide as swiftly over the life of universal matter as single years over the history of a people?”— Maeterlinck’s The Life of the Bee.