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This latter observation is important, as proving that bees can remember for at least a whole day the locality where they have found honey only once before, and that they so far think about their past experiences as to return to that locality when foraging.
As the a.s.sociation of ideas by contiguity is the principle which forms the basis of all psychology, it is desirable to consider still more attentively this the earliest manifestation that we have of it in the memory of the Hymenoptera. That it is not exercised with exclusive reference to _locality_ is proved by the following observation of Sir John Lubbock:--
I kept a specimen of _Polistes Gallica_ for no less than nine months.[49] ... I had no difficulty in inducing her to feed on my hand; but at first she was shy and nervous. She kept her sting in constant readiness... . Gradually she became quite used to me, and when I took her on my hand apparently expected to be fed. She even allowed me to stroke her without any appearance of fear, and for some months I never saw her sting.
One other observation which goes to prove that other things besides locality are noted and remembered by bees may here be quoted. Sir John placed a bee in a bell jar, the closed end of which he held towards a window. The bee buzzed about at that end trying to make for the open air. He then showed her the way out of the open end of the jar, and after having thus learnt it, she was able to find the way out herself.
This seems to show that the bee, like the wasp on the closed window-pane, was able to appreciate and to remember the difference between the quality of gla.s.s as resisting and air as permeable, although to her sense of vision the difference must have been very slight. In other words, the bee must have remembered that by first flying _away_ from the window, round the edge of the jar, and then _towards_ the window, she could surmount the transparent obstacle; and this implies a somewhat different act of memory from that of a.s.sociating a particular object--such as honey--with a particular locality. It is noteworthy that a fly under similar circ.u.mstances did not require to be taught to find its way out of the jar, but spontaneously found its own way out. This, however, may be explained by the fact that flies do not always direct their flight towards windows, and therefore the escape of this one was probably not due to any act of intelligence.
While upon the subject of memory in the Hymenoptera, it is indispensable that we should again refer to the observation of Messrs. Belt and Bates already alluded to on pages 150-51. For it is from that observation rendered evident that these sand-wasps took definite pains, as it were, to _teach themselves_ the localities to which they desired to return.
Mr. Bates further observed that after thus taking a careful mental note of the place, they would return to it without a moment's hesitation after an absence of an hour. The observation of Mr. Belt, already quoted _in extenso_, proves that these mental notes may be taken with the utmost minuteness, so that even in the most intricate places the insect, on its return, is perfectly confident that it has not made a mistake.
With regard to the duration of memory, Stickney relates a case in which some bees took possession of a hollow place beneath a roof, and having been then removed into a hive, continued for several years to return and occupy the same hole with their successive swarms.[50]
Similarly Huber relates an observation of his own showing the duration of memory in bees. One autumn he put some honey in a window, which the bees visited in large numbers. During the winter the honey was taken away and the shutters shut. When they were again opened in the spring the bees returned, although there was no honey in the window.
These two cases amply prove that the memory of bees is comparable with that of ants, which, as we have seen from a.n.a.logous facts, also extends at least over a period of many months.
_Emotions._
Sir John Lubbock's experiments on this head go to show that the social sympathies of bees are even less developed than he found them to be in certain species of ants. Thus he says:--
I have already mentioned with reference to the attachment which bees have been said to show for one another, that though I have repeatedly seen them lick a bee which had smeared herself in honey, I never observed them show the slightest attention to any of their comrades who had been drowned in water. Far, indeed, from having been able to discover any evidence of affection among them, they appear to be thoroughly callous and utterly indifferent to one another. As already mentioned, it was necessary for me occasionally to kill a bee; but I never found that the others took the slightest notice. Thus on the 11th of October I crushed a bee close to one which was feeding--in fact, so close that their wings touched; yet the survivor took no notice whatever of the death of her sister, but went on feeding with every appearance of composure and enjoyment, just as if nothing had happened. When the pressure was removed, she remained by the side of the corpse without the slightest appearance of apprehension, sorrow, or recognition. It was, of course, impossible for her to understand my reason for killing her companion; yet neither did she feel the slightest emotion at her sister's death, nor did she show any alarm lest the same fate should befall her also. In a second case exactly the same occurred. Again, I have several times, while a bee has been feeding, held a second bee by the leg close to her; the prisoner, of course, struggled to escape, and buzzed as loudly as she could; yet the selfish eater took no notice whatever.
So far, therefore, from being at all affectionate, I doubt whether bees are in the least fond of one another.
Reaumur, however ('Insects,' vol. v., p. 265), narrates a case in which a hive-bee was partly drowned and so rendered insensible; the others in the hive carefully licked and otherwise tended her till she recovered.
This seems to show that bees, like ants, are more apt to have their sympathies aroused by the sight of ailing or injured companions than by that of healthy companions in distress; but Sir John Lubbock's observations above quoted go to prove that even in this case display of sympathy is certainly not the rule.
_Powers of Communication._
Huber says that when one wasp finds a store of honey 'it returns to its nest, and brings off in a short time a hundred other wasps;' and this statement is confirmed by Dujardin, who witnessed a somewhat similar performance in the case of bees--the individual which first found a concealed store informing other individuals of the fact, and so on till numberless individuals had found it.
Although the systematic experiments of Sir John Lubbock have not tended to confirm these observations with regard to bees and wasps, we must not too readily allow his negative results to discredit these positive observations--more especially as we have seen that his _later_ experiments have fully confirmed the opinion of these previous authors with respect to ants. His experiments on bees and wasps consisted in exposing honey in a hidden situation, marking a bee or wasp that came to it, and observing whether it afterwards brought any companions to share the booty. He found that although the same insect would return over and over again, strangers came so rarely that their visits could only be attributed to accidental and independent discovery. Only if the honey were in an exposed situation, where the insects could _see_ one another feeding, would one follow the other to the food.
But we have the more reason not to accept unreservedly the conclusion to which these experiments in themselves might lead, because the very able observer F. Muller states an observation of his own which must be considered as alone sufficient to prove that bees are able to communicate information to one another:--
Once (he says[51]) I a.s.sisted at a curious contest, which took place between the queen and the other bees in one of my hives, which throws some light on the intellectual faculties of these animals. A set of forty-seven cells have been filled, eight on a newly completed comb, thirty-five on the following, and four around the first cell of a new comb. When the queen had laid eggs in all the cells of the two older combs she went several times round their circ.u.mference (as she always does, in order to ascertain whether she has not forgotten any cell), and then prepared to retreat into the lower part of the breeding-room. But as she had overlooked the four cells of the new comb, the workers ran impatiently from this part to the queen, pus.h.i.+ng her, in an odd manner, with their heads, as they did also other workers they met with. In consequence the queen began again to go around on the two older combs; but as she did not find any cell wanting an egg she tried to descend, but everywhere she was pushed back by the workers. This contest lasted for a rather long while, till the queen escaped without having completed her work. Thus the workers knew how to advise the queen that something was as yet to be done, but they knew not how to show her _where_ it had to be done.
Again, Mr. Josiah Emery, writing to 'Nature,'[52] with reference to Sir John Lubbock's experiments, says that the faculty of communication which bees possess is so well and generally known to the 'bee-hunters' of America, that the recognised method of finding a bees' nest is to act upon the faculty in question:--
Going to a field or wood at a distance from tame bees, with their box of honey they gather up from the flowers and imprison one or more bees, and after they have become sufficiently gorged, let them out to return to their home with their easily gotten load.
Waiting patiently a longer or shorter time, according to the distance of the bee-tree, the hunter scarcely ever fails to see the bee or bees return accompanied with other bees, which are in like manner imprisoned till they in turn are filled, when one or more are let out at places distant from each other, and the direction in each case in which the bee flies noted, and thus, by a kind of triangulation, the position of the bee-tree proximately ascertained.
Those who have stored honey in their houses understand very well how important it is to prevent a single bee from discovering its location. Such discovery is sure to be followed by a general onslaught from the hive unless all means of access is prevented. It is possible that our American are more intelligent than European bees, but hardly probable; and I certainly shall not ask an Englishman to admit it. Those in America who are in the habit of playing first, second, and third fiddle to instinct will probably attribute this seeming intelligence to that principle.
According to De Fraviere, bees have a number of different notes or tones which they emit from the stigmata of the thorax and abdomen, and by which they communicate information. He says:--
As soon as a bee arrives with important news, it is at once surrounded, emits two or three shrill notes, and taps a comrade with its long, flexible, and very slender feelers, or antennae. The friend pa.s.ses on the news in similar fas.h.i.+on, and the intelligence soon traverses the whole hive. If it is of an agreeable kind--if, for instance, it concerns the discovery of a store of sugar or of honey, or of a flowering meadow--all remains orderly. But, on the other hand, great excitement arises if the news presages some threatened danger, or if strange animals are threatening invasion of the hive. It seems that such intelligence is conveyed first to the queen, as the most important person in the state.
This account, which is quoted from Buchner, no doubt bears indications of imaginative colouring; but if the observation as to the emission of sounds is correct--and, as we shall see, this point is well confirmed by other observers--it is most likely concerned in communicating by tone a general idea of good or harm: probably in the former case it acts as a sign, 'follow me;' and in the latter as a signal of danger. Buchner further says that, according to Landois, if a saucer of honey is placed before a hive, a few bees come out, which emit a cry of tut, tut, tut.
This note is rather shrill, and resembles the cry of an attacked bee.
Hereupon a large number of bees come out of the hive to collect the offered honey.
Again,--
The best way to observe the power of communication possessed by bees by means of their interchange of touches, is to take away the queen from a hive. In a little time, about an hour afterwards, the sad event will be noticed by a small part of the community, and these will stop working and run hastily about over the comb. But this only concerns part of the hive, and the side of a single comb. The excited bees, however, soon leave the little circle in which they at first revolved, and when they meet their comrades they cross their antennae and lightly touch the others with them.
The bees which have received some impression from this touch now become uneasy in their turn, and convey their uneasiness and distress in the same way to the other parts of the dwelling. The disorder increases rapidly, spreads to the other side of the comb, and at last to all the people. Then arises the general confusion before described.
Huber tested this communication by the antennae by a striking experiment. He divided a hive into two quite separate parts by a part.i.tion wall, whereupon great excitement arose in the division in which there was no queen, and this was only quieted when some workers began to build royal cells.
He then divided a hive in similar fas.h.i.+on by a trellis, through which the bees could pa.s.s their feelers. In this case all remained quiet, and no attempt was made to build royal cells: the queen could also be clearly seen crossing her antennae with the workers on the other side of the trellis.
Apparently the feelers are also connected with the exceedingly fine scent of the bees, which enables them, wonderful as it may seem, to distinguish friend and foe, and to recognise the members of their own hive among the thousands and thousands of bees swarming around, and to drive back from the entrance stranger or robber bees. The bee-masters, therefore, when they want two separate colonies or the members of them to unite in one hive, sprinkle water over the bees, or stupefy them with some fumigating substance, so as to make them to a certain extent insensible to smell, in order to attain their object. It is always possible to unite colonies by making the bees smell of some strong-smelling stuff, such as musk.[53]
Lastly, under the present heading I shall quote one other observation, for which I am also indebted to Buchner's very admirable collection of facts relating to the psychology of Hymenoptera:--
Herr L. Brofft relates, in 'der Zoologische Garten'
(XVIII. Year, No. 1, p. 67), that a poor and a rich hive stood next each other on his father's bee stand, and the latter suddenly lost its queen. Before the owner had come to a decision thereupon the bees of the two hives came to a mutual understanding as to the condition of their two states. The dwellers in the queenless hive, with their stores of provisions, went over into the less populous or poorer hive, after they had a.s.sured themselves, by many influential deputations, as to the state of the interior of the poor hive, and, as appeared, especially as to the presence of an egg-laying queen!
_General Habits._
The active life of bees is divided between collecting food and rearing young. We shall therefore consider these two functions separately.
The food collected consists of two kinds, honey (which, although stored in the 'crop' for the purpose of carriage from the flowers to the cells, appears to be but the condensed nectar of flowers) and so-called 'bee-bread.' This consists of the pollen of flowers, which is worked into a kind of paste by the bees and stored in their cells till it is required to serve as food for their larvae. It is then partly digested by the nurses with honey, so that a sort of chyle is formed. It is observable that in each flight the 'carrier bees' collect only one kind of pollen, so that it is possible for the 'house bees' (which, by the way, are the younger bees left at home to discharge domestic duties with only a small proportion of older ones, left probably to direct the more inexperienced young) to sort it for storage in different cells. In the result there are several different kinds of bee-bread, some being more stimulating or nutritious than others. The most nutritious has the effect, when given to any female larva, of developing that larva into a queen or fertile female. This fact is well known to the bees, who only feed a small number of larvae in this manner, and the larvae which they select so to feed they place in larger or 'royal' cells, with an obvious foreknowledge of the increased dimensions to which the animal will grow under the influence of this food. Only one queen is required for a single hive; but the bees always raise several, so that if any mishap should occur to one, other larvae may be ready to fall back upon.
Besides honey and bee-bread two other substances are found in beehives.
These are propolis and beeswax. The former is a kind of sticky resin collected for the most part from coniferous trees. This is used as mortar in building, &c. It adheres so strongly to the legs of the bee which has gathered it, that it can only be detached by the help of comrades. For this purpose the loaded bee presents her legs to her fellow-workers, who clean it off with their jaws, and while it is still ductile, apply it round the inside of the hive. According to Huber, who made this observation, the propolis is applied also to the insides of the cells. The workers first planed the surfaces with their mandibles, and one of them then pulled out a thread of propolis from the heap deposited by the carrier bees, severed it by a sudden throwing back of the head, and returned with it to the cell which it had previously been planing. It then laid the thread between the two walls which it had planed; but, proving too long, a portion of the thread was bitten off.
The properly measured portion was then forced into the angle of the cell by the fore-feet and mandibles. The thread, now converted into a narrow ribbon, was next found to be too broad. It was therefore gnawed down to the proper width. Other bees then completed the work which this one had begun, till all the walls of the cells were framed with bands of propolis. The object of the propolis here seems to be that of giving strength to the cells.
The wax is a secretion which proceeds from between the segments of the abdomen. Having ingested a large meal of honey, the bees hang in a thick cl.u.s.ter from the top of their hive in order to secrete the wax. When it begins to exude, the bees, a.s.sisted by their companions, rub it off into heaps, and when a sufficient quant.i.ty of the material has been thus collected, the work begins of building the cells. As the cells are used both for storing food and rearing young, I shall consider them later on. Now we have to pa.s.s to the labours incidental to propagation.
All the eggs are laid by one queen, who requires during this season a large amount of nourishment, so much, indeed, that ten or twelve working bees (_i.e._ sterile females) are set apart as her feeders. Leaving the 'royal cell,' she walks over the nursery-combs attended by a retinue of workers, and drops a single egg into each open cell. It is a highly remarkable fact that the queen is able to control the s.e.x of the eggs which she lays, and only deposits drone or male eggs in the drone cells, and worker or female eggs in the worker cells--the cells prepared for the reception of drone larvae being larger than those required for the worker larvae. Young queens lay more worker eggs than old queens, and when a queen, from increasing age or any other cause, lays too large a proportion of drone eggs, she is expelled from the community or put to death. It is remarkable, also, under these circ.u.mstances, that the queen herself seems to know that she has become useless, for she loses her propensity to attack other queens, and so does not run the risk of making the hive virtually queenless. There is now no doubt at all that the determining cause of an egg turning out male or female is that which Dzierzon has shown, namely, the absence or presence of fertilisation--unfertilised eggs always developing into males, and fertilised ones into females. The manner, therefore, in which a queen controls the s.e.x of her eggs must depend on some power that she has of controlling their fertilisation.
The eggs hatch out into larvae, which require constant attention from the workers, who feed them with the chyle or bee-bread already mentioned. In three weeks from the time that the egg is deposited, the white worm-like larva has pa.s.sed through its last metamorphosis. When it has emanc.i.p.ated itself its nurses a.s.semble round it to wash and caress it, as well as to supply it with food. They then clean out the cell which it has left.
When so large a number of the larvae hatch out as to overcrowd the hive, it is the function of the queen to lead forth a swarm. Meanwhile several larval queens have been in course of development, and matters are so arranged by the foresight of the bees, that one or more young queens are ready to emerge at a time when otherwise the hive would be left queenless. But the young queen or queens, although perfectly formed, must not escape from their royal prison-houses until the swarm has fairly taken place; the worker bees will even strengthen the coverings of these prison-houses if, owing to bad weather or other causes, swarming is delayed. The prisoner queens, which are fed through a small hole in the roof of their cells, now continually give vent to a plaintive cry, called by the bee-keepers 'piping,' and this is answered by the mother queen. The tones of the piping vary. The reason why the young queens are kept such close prisoners till after the departure of the mother queen with her swarm, is simply that the mother queen would destroy all the younger ones, could she get the chance, by stinging them. The workers, therefore, never allow the old queen to approach the prisons of the younger ones. They establish a guard all round these prisons or royal cells, and beat off the old queen whenever she endeavours to approach. But if the swarming season is over, or anything should prevent a further swarm from being sent out, the worker bees offer no further resistance to the jealousy of the mother queen, but allow her in cold blood to sting to death all the young queens in their nursery prisons. As soon as the old queen leaves with a swarm, the young queens are liberated in succession, but at intervals of a few days; for if they were all liberated at once they would fall upon and destroy one another. Each young queen as it is liberated goes off with another swarm, and those which remain unliberated are as carefully guarded from the liberated sister queen as they were previously guarded from the mother queen. When the season is too late for swarming the remaining young queens are liberated simultaneously, and are then allowed to fight to the death, the survivor being received as sovereign.
The bees, far from seeking to prevent these battles, appear to excite the combatants against each other, surrounding and bringing them back to the charge when they are disposed to recede from each other; and when either of the queens shows a disposition to approach her antagonist, all the bees forming the cl.u.s.ter instantly give way to allow her full liberty of attack. The first use which the conquering queen makes of her victory is to secure herself against fresh dangers by destroying all her future rivals in the royal cells; while the other bees, which are spectators of the carnage, share in the spoil, greedily devouring any food which may be found at the bottom of the cells, and even sucking the fluid from the abdomen of the pupae before they toss out the carca.s.ses.[54]
Similarly, when a strange queen is put into a hive already provided with a queen--
A circle of bees instinctively crowd around the invader, not, however, to attack her--for a worker never a.s.saults a queen--but to respectfully prevent her escape, in order that a combat may take place between her and their reigning monarch. The lawful possessor then advances towards the part of the comb where the invader has established herself, the attendant workers clear a s.p.a.ce for the encounter, and, without interfering, wait the result. A fearful encounter then ensues, in which one is stung to death, the survivor mounting the throne. Although the workers of a _de facto_ monarch will not fight for her defence, yet, if they perceive a strange queen _attempting_ to enter the hive, they will surround her, and hold her until she is starved to death; but such is their respect for royalty that they never attempt to sting her.[55]
All these facts display a wonderful amount of apparently sagacious purpose on the part of the workers, although they may not seem to reflect much credit on the intelligence of the queens. But in this connection we must remember the observation of F. Huber, who saw two queens, which were the only ones left in the hive, engaged in mortal combat; and when an opportunity arose for each to sting the other simultaneously, they simultaneously released each other's grasp, as if in horror of a situation that might have ended in leaving the hive queenless. This, then, is the calamity to avert which all the instincts both of workers and queens are directed. And that these instincts are controlled by intelligence is suggested, if not proved, by the adaptations which they show to special circ.u.mstances. Thus, for instance, F. Huber smoked a hive so that the queen and older bees effected their escape, and took up their quarters a short distance away.
The bees which remained behind set about constructing three royal cells for the purpose of rearing a new queen. Huber now carried back the old queen and ensconced her in the hive. Immediately the bees set about carrying away all the food from the royal cells, in order to prevent the larvae contained therein from developing into queens. Again, if a strange queen is presented to a hive already provided with one, the workers do not wait for their own queen to destroy the pretender, but themselves sting or smother her to death. When, on the other hand, a queen is presented to a hive which is without one, the bees adopt her, although it is often necessary for the bee-master to protect her for a day or two in a trellis cage, until her subjects have become acquainted with her.
When a hive is queenless, the bees stop all work, become restless, and make a dull complaining noise. This, however, is only the case if there is likewise a total absence of royal pupae, and of ordinary pupae under three days of age--_i.e._ the age during which it is possible to rear an ordinary larva into a queen.