Underneath the wasp's brown paper manor house, the ground is channeled into a sort of drain for the refuse of the nest. Here are shot the dead or weakly larvae which a continual inspection roots out from the cells to make room for fresh occupants; here, at the time of the autumn massacre, are flung the backward grubs; here, lastly, lies a good part of the crowd killed by the first touch of winter. During the rack and ruin of November and December, this sewer becomes crammed with animal matter.
Such riches will not remain unemployed. The world's great law which says that nothing edible shall be wasted provides for the consumption of a mere ball of hair disgorged by the owl. How shall it be with the vast stores of a ruined wasps' nest! If they have not come yet, the consumers whose task it is to salve this abundant wreckage for nature's markets, they will not tarry in coming and waiting for the manna that will soon descend from above. That public granary, lavishly stocked by death, will become a busy factory of fresh life. Who are the guests summoned to the banquet?
If the wasps flew away, carrying the dead or sickly grubs with them, and dropped them on the ground round about their home, those banqueters would be, first and foremost, the insect-eating birds, the warblers, all of whom are lovers of small game. In this connection, we will allow ourselves a brief digression. We all know with what jealous intolerance the nightingales occupy each his own cantonment. Neighborly intercourse among them is tabooed. The males frequently exchange defiant couplets at a distance; but, should the challenged party draw near, the challenger makes him clear off. Now, not far from my house, in a scanty clump of holly oaks which would barely give the woodcutter the wherewithal for a dozen faggots, I used, all through the spring, to hear such full-throated warbling of nightingales that the songs of those virtuosi, all giving voice at once and with no attempt at order, degenerated into a deafening hubbub.
Why did those passionate devotees of solitude come and settle in such large numbers at a spot where custom decrees that there is just room enough for one household only? What reasons have made the recluse become a congregation? I asked the owner of the spinney about the matter.
'It's like that every year,' he said. 'The clump is overrun by Nightingales.'
'And the reason? '
'The reason is that there is a hive close by, behind that wall.'
I looked at the man in amazement, unable to understand what connection there could be between a hive and the thronging nightingales.
'Why, yes,' he added, 'there are a lot of nightingales because there are a lot of bees.
Another questioning look from my side. I did not yet understand. The explanation came: 'The bees,' he said, 'throw out their dead grubs. The front of the hive is strewn with them in the mornings; and the nightingales come and collect them for themselves and their families. They are very fond of them.'
This time I had solved the puzzle. Delicious food, abundant and fresh each day, had brought the songsters together. Contrary to their habit, numbers of nightingales are living on friendly terms in a cluster of bushes, in order to be near the hive and to have a larger share in the morning distribution of plump dainties.
In the same way, the nightingale and his gastronomical rivals would haunt the neighborhood of the wasps' nests, if the dead grubs were cast out on the surface of the soil; but these delicacies fall inside the burrow and no little bird would dare to enter the murky cave, even if the entrance were not too small to admit it. Other consumers are needed here, small in size and great in daring; the fly is called for and her maggot, the king of the departed. What the greenbottles, the bluebottles and the flesh flies do in the open air, at the expense of every kind of corpse, other flies, narrowing their province, do underground at the Wasps' expense.
Let us turn our attention, in September, to the wrapper of a wasps' nest. On the outer surface and there alone, this wrapper is strewn with a multitude of big, white, elliptical dots, firmly fixed to the brown paper and measuring about two millimeters and a half long by one and a half wide. Flat below, convex above and of a lustrous white, these dots resemble very neat drops fallen from a tallow candle. Lastly, their backs are streaked with faint transversal lines, an elegant detail perceptible only with the lens. These curious objects are scattered all over the surface of the wrapper, sometimes at a distance from one another, sometimes gathered into more or less dense groups. They are the eggs of the Volucella, or bumblebee fly (Volucella zonaria, LIN.)
Also stuck to the brown paper of the outer wrapper and mixed up with the Volucella's are a large number of other eggs, chalk white, spear-shaped and ridged lengthwise with seven or eight thin ribs, after the manner of the seeds of certain Umbelliferae. The finishing touch to their delicate beauty is the fine stippling all over the surface. They are smaller by half than the others. I have seen grubs come out of them which might easily be the earliest stage of some pointed maggots which I have already noticed in the burrows. My attempts to rear them failed; and I am not able to say which fly these eggs belong to. Enough for us to note the nameless one in passing. There are plenty of others, which we must make up our minds to leave unlabelled, in view of the jumbled crowd of feasters in the ruined wasps' nest. We will concern ourselves only with the most remarkable, in the front rank of which stands the bumblebee Fly.
She is a gorgeous and powerful fly; and her costume, with its brown and yellow bands, shows a vague resemblance to that of the wasps. Our fashionable theorists have availed themselves of this brown and yellow to cite the Volucella as a striking instance of protective mimicry. Obliged, if not on her own behalf, at least on that of her family, to introduce herself as a parasite into the wasp's home, she resorts, they tell us, to trickery and craftily dons her victim's livery. Once inside the wasps' nest, she is taken for one of the inhabitants and attends quietly to her business.
The simplicity of the wasp, duped by a very clumsy imitation of her garb, and the depravity of the fly, concealing her identity under a counterfeit presentment, exceed the limits of my credulity. The wasp is not so silly nor the Volucella so clever as we are assured. If the latter really meant to deceive the Wasp by her appearance, we must admit that her disguise is none too successful. Yellow sashes round the abdomen do not make a wasp. It would need more than that and, above all, a slender figure and a nimble carriage; and the Volucella is thickset and corpulent and sedate in her movements. Never will the wasp take that unwieldy insect for one of her own kind. The difference is too great.
Poor Volucella, mimesis has not taught you enough. You ought this is the essential point to have adopted a wasp's shape; and that you forgot to do: you remained a fat fly, easily recognizable. Nevertheless, you penetrate into the terrible cavern; you are able to stay there for a long time, without danger, as the eggs profusely strewn on the wrapper of the wasps' nest show. How do you set about it?
Let us, first of all, remember that the bumblebee fly does not enter the enclosure in which the combs are heaped: she keeps to the outer surface of the paper rampart and there lays her eggs. Let us, on the other hand, recall the Polistes (1) placed in the company of the wasps in my vivarium. Here of a surety is one who need not have recourse to mimicry to find acceptance. She belongs to the guild, she is a wasp herself. Any of us that had not the trained eye of the entomologist would confuse the two species. Well, this stranger, as long as she does not become too importunate, is quite readily tolerated by the caged wasps. None seeks to pick a quarrel with her. She is even admitted to the table, the strip of paper smeared with honey. But she is doomed if she inadvertently sets foot upon the combs. Her costume, her shape, her size, which tally almost exactly with the costume, shape and size of the wasp, do not save her from her fate. She is at once recognized as a stranger and attacked and slaughtered with the same vigor as the larvae of the Hylotoma sawfly and the Saperda beetle, neither of which bears any outward resemblance to the larva of the wasps.
Seeing that identity of shape and costume does not save the Polistes, how will the Volucella fare, with her clumsy imitation? The wasp's eye, which is able to discern the dissimilar in the like, will refuse to be caught. The moment she is recognized, the stranger is killed on the spot. As to that there is not the shadow of a doubt.
In the absence of bumblebee flies at the moment of experimenting, I employ another fly, Milesia fulminans, who, thanks to her slim figure and her handsome yellow bands, presents a much more striking likeness to the wasp than does the fat Volucella zonaria. Despite this resemblance, if she rashly venture on the combs, she is stabbed and slain. Her yellow sashes, her slender abdomen deceive nobody. The stranger is recognized behind the features of a double.
My experiments under glass, which varied according to the captures which I happened to make, all lead me to this conclusion: as long as there is more propinquity, even around the honey, the other occupants are tolerated fairly well; but, if they touch the cells, they are assaulted and often killed, without distinction of shape or costume. The grubs' dormitory is the sanctum sanctorum which no outsider must enter under pain of death.
With these caged captives I experiment by daylight, whereas the free wasps work in the absolute darkness of their underground retreat. Where light is absent, color goes for nothing. Once, therefore, that she has entered the cavern, the bumblebee fly derives no benefit from her yellow bands, which are supposed to be her safeguard. Whether garbed as she is or otherwise, it is easy for her to effect her purpose in the dark, on condition that she avoids the tumultuous interior of the wasps' nest. So long as she has the prudence not to hustle the passers by, she can dab her eggs, without danger, on the paper wall. No one will know of her presence. The dangerous thing is to cross the threshold of the burrow in broad daylight, before the eyes of those who go in and out. At that moment alone, protective mimicry would be convenient. Now does the entrance of the Volucella into the presence of a few wasps entail such very great risks? The wasps' nest in my enclosure, the one which was afterwards to perish in the sun under a bell glass, gave me the opportunity for prolonged observations, but without any result upon the subject of my immediate concern. The bumblebee fly did not appear. The period for her visits had doubtless passed; for I found plenty of her grubs when the nest was dug up.
Other flies rewarded me for my assiduity. I saw some at a respectful distance, I need hardly say entering the burrow. They were insignificant in size and of a dark gray color, not unlike that of the housefly. They had not a patch of yellow about them and certainly had no claim to protective mimicry. Nevertheless, they went in and out as they pleased, calmly, as though they were at home. As long as there was not too great a number at the door, the wasps left them alone. When there was anything of a crowd, the gray visitors waited near the threshold for a less busy moment. No harm came to them.
Inside the establishment, the same peaceful relations prevail. In this respect I have the evidence of my excavations. In the underground charnel house, so rich in Fly grubs, I find no corpses of adult flies. If the strangers had been slaughtered in passing through the entrance hall, or lower down, they would fall to the bottom of the burrow anyhow, with the other rubbish. Now in this charnel house, as I said, there are never any dead bumblebee flies, never a fly of any sort. The incomers are respected. Having done their business, they go out unscathed.
This tolerance on the part of the wasps is surprising. And a suspicion comes to one's mind: can it be that the Volucella and the rest are not what the accepted theories of natural history call them, namely, enemies, grub killers sacking the wasps' nest? We will look into this by examining them when they are hatched. Nothing is easier, in September and October, than to collect the Volucella's eggs in such numbers as we please. They abound on the outer surface of the wasps' nest. Moreover, as with the larvae of the wasp, it is some time before they are suffocated by the petroleum fumes; and so most of them are sure to hatch. I take my scissors, cut the most densely populated bits from the paper wall of the nest and fill a jar with them. This is the warehouse from which I shall daily, for the best part of the next two months, draw my supply of nascent grubs.
The Volucella's egg remains where it is, with its white color always strongly marked against the brown of the background. The shell wrinkles and collapses; and the fore end tears open. From it there issues a pretty little white grub, thin in front, swelling slightly in the rear and bristling all over with fleshy protuberances. The creature's papillae are set on its sides like the teeth of a comb; at the rear, they lengthen and spread into a fan; on the back, they are shorter and arranged in four longitudinal rows. The last section but one carries two short, bright red breathing tubes, standing aslant and joined to each other. The fore part, near the pointed mouth, is of a darker, brownish color. This is the biting and motor apparatus, seen through the skin and consisting of two fangs. Taken all round, the grub is a pretty little thing, with its bristling whiteness, which gives it the appearance of a tiny snowflake. But this elegance does not last long: grown big and strong, the bumblebee fly's grub becomes soiled with sanies, turns a russety brown and crawls about in the guise of a hulking porcupine.
What becomes of it when it leaves the egg? This my warehousing jar tells me, partly. Unable to keep its balance on sloping surfaces, it drops to the bottom of the receptacle, where I find it, daily, as hatched, wandering restlessly. Things must happen likewise at the wasps'. Incapable of standing on the slant of the paper wall, the newborn grubs slide to the bottom of the underground cavity, which contains, especially at the end of the summer, a heaped up provender of deceased wasps and dead larvae removed from the cells and flung outside the house, all nice and gamy, as proper maggot's food should be.
The Volucella's offspring, themselves maggots, notwithstanding their snowy apparel, find in this charnel house victuals to their liking, incessantly renewed. Their fall from the high walls might well be not accidental, but rather a means of reaching, quickly and without searching, the good things down at the bottom of the cavern. Perhaps, also, some of the white grubs, thanks to the holes that make the wrapper resemble a spongy cover, manage to slip inside the Wasps' nest. Still, most of the Volucella's grubs, at whatever stage of their development, are in the basement of the burrow, among the carrion remains. The others, those settled in the wasps' home itself, are comparatively few.
These returns are enough to show us that the grubs of the bumblebee fly do not deserve the bad reputation that has been given them. Satisfied with the spoils of the dead, they do not touch the living; they do not ravage the wasps' nest: they disinfect it.
Experiment confirms what we have learnt in the actual nests. Over and over again, I bring wasp grubs and Volucella grubs together in small test tubes, which are easy to observe. The first are well and strong; I have just taken them from their cells. The others are in various stages, from that of the snowflake born the same day to that of the sturdy porcupine. There is nothing tragic about the encounter. The grubs of the bumblebee fly roam about the test-tube without touching the live tidbit. The most that they do is to put their mouths for a moment to the morsel; then they take it away again, not caring for the dish.
They want something different: a wounded, a dying grub; a corpse dissolving into sanies. Indeed, if I prick the wasp grub with a needle, the scornful ones at once come and sup at the bleeding wound. If I give them a dead grub, brown with putrefaction, the worms rip it open and feast on its humors. Better still: I can feed them quite satisfactorily with wasps that have turned putrid under their horny rings; I see them greedily suck the juices of decomposing Rosechafer grubs; I can keep them thriving with chopped up butcher's meat, which they know how to liquefy by the method of the common maggot. And these unprejudiced ones, who accept anything that comes their way, provided it be dead, refuse it when it is alive. Like the true flies that they are, frank body snatchers, they wait, before touching a morsel, for death to do its work.
Inside the wasps' nest, robust grubs are the rule and weaklings the rare exception, because of the assiduous supervision which eliminates anything that is diseased and like to die. Here, nevertheless, Volucella grubs are found, on the combs, among the busy wasps. They are not, it is true, so numerous as in the charnel house below, but still pretty frequent. Now what do they do in this abode where there are no corpses? Do they attack the healthy? Their continual visits from cell to cell would at first make one think so; but we shall soon be undeceived if we observe their movements closely; and this is possible with my glass roofed colonies.
I see them fussily crawling on the surface of the combs, curving their necks from side to side and taking stock of the cells. This one does not suit, nor that one either; the bristly creature passes on, still in search, thrusting its pointed fore part now here, now there. This time, the cell appears to fulfil the requisite conditions. A larva, glowing with health, opens wide its mouth, believing its nurse to be approaching. It fills the hexagonal chamber with its bulging sides.
The gluttonous visitor bends and slides its slender fore part, a blade of exquisite suppleness, between the wall and the inhabitant, whose slack rotundity yields to the pressure of this animated wedge. It plunges into the cell, leaving no part of itself outside but its wide hind quarters, with the red dots of the two breathing tubes.
It remains in this posture for some time, occupied with its work at the bottom of the cell. Meanwhile, the wasps present do not interfere, remain impassive, showing that the grub visited is in no peril. The stranger, in fact, withdraws with a soft, gliding motion. The chubby babe, a sort of India rubber bag, resumes its original volume without having suffered any harm, as its appetite proves. A nurse offers it a mouthful, which it accepts with every sign of unimpaired vigor. As for the Volucella grub, it licks its lips after its own fashion, pushing its two fangs in and out; then, without further loss of time, goes and repeats its probing elsewhere.
What it wants down there, at the bottom of the cells, behind the grubs, cannot be decided by direct observation; it must be guessed at. Since the visited larva remains intact, it is not prey that the Volucella grub is after. Besides, if murder formed part of its plans, why descend to the bottom of the cell, instead of attacking the defenseless recluse straight way? It would be much easier to suck the patient's juices through the actual orifice of the cell. Instead of that, we see a dip, always a dip and never any other tactics.
Then what is there behind the wasp grub? Let us try to put it as decently as possible. In spite of its exceeding cleanliness, this grub is not exempt from the physiological ills inseparable from the stomach. Like all that eats, it has intestinal waste matter with regard to which its confinement compels it to behave with extreme discretion. Like so many other close-cabined larvae of Wasps and Bees, it waits until the moment of the transformation to rid itself of its digestive refuse. Then, once and for all, it casts out the unclean accumulation whereof the pupa, that delicate, reborn organism, must not retain the least trace. This is found later, in any empty cell, in the form of a dark purple plug. But, without waiting for this final purge, this lump, there are, from time to time, slight excretions of fluid, clear as water. We have only to keep a Wasp grub in a little glass tube to recognize these occasional discharges. Well, I see nothing else to explain the action of the Volucella's grubs when they dip into the cells without wounding the larvae. They are looking for this liquid, they provoke its emission. It represents to them a dainty which they enjoy over and above the more substantial fare provided by the corpses.
The bumblebee fly, that sanitary inspector of the Vespine city, fulfils a double office: she wipes the wasp's children and she rids the nest of its dead. For this reason, she is peacefully received, as an auxiliary, when she enters the burrow to lay her eggs; for this reason, her grub is tolerated, nay more, respected, in the very heart of the dwelling, where none might stray with impunity. I remember the brutal reception given to the Saperda and Hylotoma grubs when I place them on a comb. Forthwith grabbed, bruised and riddled with stings, the poor wretches perish. It is quite a different matter with the offspring of the Volucella. They come and go as they please, poke about in the cells, elbow the inhabitants and remain unmolested. Let us give some instances of this clemency, which is very strange in the irascible Wasp.
For a couple of hours, I fix my attention on a Volucella grub established in a cell, side by side with the Wasp grub, the mistress of the house. The hind quarters emerge, displaying their papillae. Sometimes also the fore part, the head, shows, bending from side to side with sudden, snake-like motions. The wasps have just filled their crops at the honey pot; they are dispensing the rations, are very busily at work; and things are taking place in broad daylight, on the table by the window.
As they pass from cell to cell, the nurses repeatedly brush against and stride across the Volucella grub. There is no doubt that they see it. The intruder does not budge, or, if trodden on, curls up, only to reappear the next moment. Some of the wasps stop, bend their heads over the opening, seem to be making inquiries and then go off, without troubling further about the state of things. One of them does something even more remarkable: she tries to give a mouthful to the lawful occupant of the cell; but the larva, which is being squeezed by its visitor, has no appetite and refuses. Without the least sign of anxiety on behalf of the nursling which she sees in awkward company, the wasp retires and goes to distribute its ration elsewhere. In vain I prolong my examination: there is no fluster of any kind. The Volucella grub is treated as a friend, or at least as a visitor that does not matter. There is no attempt to dislodge it, to worry it, to put it to flight. Nor does the grub seem to trouble greatly about those who come and go. Its tranquillity, tells us that it feels at home.
Here is some further evidence: the grub has plunged, head downwards, into an empty cell, which is too small to contain the whole of it. Its hindquarters stick out, very visibly. For long hours, it remains motionless in this position. At every moment, wasps pass and repass close by. Three of them, at one time together, at another separately, come and nibble at the edges of the cell; they break off particles which they reduce to paste for a new piece of work. The passers by, intent upon their business, may not perceive the intruder; but these three certainly do. During their work of demolition, they touch the grub with their legs, their antennae, their palpi; and yet none of them minds it. The fat grub, so easily recognized by its queer figure, is left alone; and this in broad daylight, where everybody can see it. What must it be when the profound darkness of the burrows protects the visitor with its mysteries!
I have been experimenting all along with big Volucella grubs, colored with the dirty red which comes with age. What effect will pure white produce? I sprinkle on the surface of the combs some larvae that have lately left the egg. The tiny, snow-white grubs make for the nearest cells, go down into them, come out again and hunt elsewhere. The wasps peaceably let them go their way, as heedless of the little white invaders as of the big red ones. Sometimes, when it enters an occupied cell, the little creature is seized by the owner, the wasp grub, which nabs it and turns and returns it between its mandibles. Is this a defensive bite? No, the wasp grub has merely blundered, taking its visitor for a proffered mouthful. There is no great harm done. Thanks to its suppleness, the little grub emerges from the grip intact and continues its investigations.
It might occur to us to attribute this tolerance to some lack of penetration in the wasps' vision. What follows will undeceive us: I place separately, in empty cells, a grub of Saperda scalaria and a Volucella grub, both of them white and selected so as not to fill the cell entirely. Their presence is revealed only by the paleness of the hind part which serves as a plug to the opening. A superficial examination would leave the nature of the recluse undecided. The wasps make no mistake: they extirpate the Saperda grub, kill it, fling it on the dust heap; they leave the Volucella grub in peace.
The two strangers are quite well recognized in the secrecy of the cells: one is the intruder that must be turned out; the other is the regular visitor that must be respected. Sight helps, for things take place in the daylight, under glass; but the wasps have other means of information in the dimness of the burrow. When I produce darkness by covering the apparatus with a screen, the murder of the trespassers is accomplished just the same. For so say the police regulations of the wasps' nest: any stranger discovered must be slain and thrown on the midden.
To thwart this vigilance, the real enemies need to be masters of the art of deceptive immobility and cunning disguise. But there is no deception about the Volucella grub. It comes and goes, openly, wheresoever it will; it looks round amongst the wasps for cells to suit it. What has it to make itself thus respected? Strength? Certainly not. It is a harmless creature, which the wasp could rip open with a blow of her shears, while a touch of the sting would mean lightning death. It is a familiar guest, to whom no denizen of a wasps' nest bears any ill will. Why? Because it renders good service: so far from working mischief, it does the scavenging for its hosts. Were it an enemy or merely an intruder, it would be exterminated; as a deserving assistant, it is respected.
Then what need is there for the Volucella to disguise herself as a wasp? Any fly, whether clad in drab or motley, is admitted to the burrow directly she makes herself useful to the community. The mimicry of the bumblebee fly, which was said to be one of the most conclusive cases, is, after all, a mere childish notion. Patient observation, continually face to face with facts, will have none of it and leaves it to the armchair naturalists, who are too prone to look at the animal world through the illusive mists of theory.