CHAPTER 3. A DANGEROUS DIET.
The Scolia's egg is in no way exceptional in shape. It is white, cylindrical, straight and about four millimetres long by one millimetre thick. (About.156 x.039 inch.—Translator's Note.) It is fixed, by its fore-end, upon the median line of the victim's abdomen, well to the rear of the legs, near the beginning of the brown patch formed by the mass of food under the skin.
I watch the hatching. The grub, still wearing upon its hinder parts the delicate pellicle which it has just shed, is fixed to the spot to which the egg itself adhered by its cephalic extremity. A striking spectacle, that of the feeble creature, only this moment hatched, boring, for its first mouthful, into the paunch of its enormous prey, which lies stretched upon its back. The nascent tooth takes a day over the difficult task. Next morning the skin has yielded; and I find the new-born larva with its head plunged into a small, round, bleeding wound.
In size the grub is the same as the egg, whose dimensions I have just given. Now the Cetonia-larva, to meet the Scolia's requirements, averages thirty millimetres in length by nine in thickness (1.17 x.35 inch.—Translator's Note.), whence follows that its bulk is six or seven hundred times as great as that of the newly-hatched grub of the Scolia. Here certainly is a quarry which, were it active and capable of wriggling and biting, would expose the nurseling to terrible attacks. The danger has been averted by the mother's stiletto; and the fragile grub attacks the monster's paunch with as little hesitation as though it were sucking the breast.
Day by day the young Scolia's head penetrates farther into the Cetonia's belly. To pass through the narrow orifice made in the skin, the fore-part of the body contracts and lengthens out, as though drawn through a die-plate. The larva thus assumes a rather strange form. Its hinder half, which is constantly outside the victim's belly, has the shape and fulness usual in the larvae of the Digger-wasps, whereas the front half, which, once it has dived under the skin of the exploited victim, does not come out again until the time arrives for spinning the cocoon, tapers off suddenly into a snake-like neck. This front part is moulded, so to speak, by the narrow entrance-hole made in the skin and henceforth retains its slender formation. As a matter of fact, a similar configuration recurs, in varying degrees, in the larvae of the Digger-wasps whose ration consists of a bulky quarry which takes a long time to consume. These include the Languedocian Sphex, with her Ephippiger, and the Hairy Ammophila, with her Grey Worm. There is none of this sudden constriction, dividing the creature into two disparate halves, when the victuals consist of numerous and comparatively small items. The larva then retains its usual shape, being obliged to pass, at brief intervals, from one joint in its larder to the next.
From the first bite of the mandibles, until the whole head of game is consumed, the Scolia-larva is never seen to withdraw its head and its long neck from inside the creature which it is devouring. I suspect the reason of this persistence in attacking a single point; I even seem to perceive the need for a special art in the manner of eating. The Cetonia-larva is a square meal in itself, one large dish, which has to retain a suitable freshness until the end. The young Scolia, therefore, must attack with discretion, at the unvarying point chosen by the mother on the ventral surface, for the entrance-hole is at the exact point where the egg was fixed. As the nurseling's neck lengthens and dives deeper, the victim's entrails are nibbled gradually and methodically: first, the least essential; next, those whose removal leaves yet a remnant of life; lastly, those whose loss inevitably entails death, followed very soon by putrefaction.
At the first bites we see the victim's blood oozing through the wound. It is a highly-elaborated fluid, easy of digestion, and forms a sort of milk-diet for the new-born grub. The little ogre's teat is the bleeding paunch of the Cetonia-larva. The latter will not die of the wound, at least not for some time. The next thing to be tackled is the fatty substance which wraps the internal organs in its delicate folds. This again is a loss which the Cetonia can suffer without dying then and there. Now comes the turn of the muscular layer which lines the skin; now, that of the essential organs; now, that of the nerve-centres and the trachean network, whereupon the last gleam of light is extinguished and the Cetonia reduced to a mere bag, empty but intact, save for the entrance-hole made in the middle of the belly. From now onwards, these remains may rot if they will: the Scolia, by its methodical fashion of consuming its victuals, has succeeded in keeping them fresh to the very last; and now you may see it, replete, shining with health, withdraw its long neck from the bag of skin and prepare to weave the cocoon in which its development will be completed.
It is possible that I may not be quite accurate as to the precise order in which the organs are consumed, for it is not easy to perceive what happens inside the exploited larva's body. The ruling feature in this scientific method of eating, which proceeds from the parts less to the parts more necessary to preserve a remnant of life, is none the less obvious. If direct observation did not already to some degree confirm it, a mere examination of the half-eaten larva would do so in the most positive fashion.
The Cetonia-larva is at first a plump grub. Drained by the Scolia's tooth, it gradually becomes limp and wrinkled. In a few days' time it resembles a shrivelled bit of bacon-fat and then a bag whose two sides have fallen in. Yet this bit of bacon and this bag have the same characteristic look of fresh meat as had the grub before it was bitten into. Despite the persistent nibbling of the Scolia, life continues, holding at bay the inroads of putrefaction until the mandibles have given their last bites. Does not this remnant of tenacious vitality in itself show that the organs of primary importance are the last to be attacked? Does it not prove that there is a progressive dismemberment passing from the less essential to the indispensable?
Would you like to see what becomes of a Cetonia-larva when the organism is wounded in its vital centres at the very beginning? The experiment is an easy one; and I made a point of trying it. A sewing-needle, first softened and flattened into a blade, then retempered and sharpened, gives me a most delicate scalpel. With this instrument I make a fine incision, through which I remove the mass of nerves whose remarkable structure we shall soon have occasion to study. The thing is done: the wound, which does not look serious, has left the creature a corpse, a real corpse. I lay my victim on a bed of moist earth, in a jar with a glass lid; in fact, I establish it in the same conditions as those of the larvae on which the Scoliae feed. By the next day, without changing shape, it has turned a repulsive brown; presently it dissolves into noisome putrescence. On the same bed of earth, under the same glass cover, in the same moist, warm atmosphere, the larvae three-quarters eaten by the Scoliae retain, on the contrary, the appearance of healthy flesh.
If a single stroke of my dagger, fashioned from the point of a needle, results in immediate death and early putrefaction; if the repeated bites of the Scolia gut the creature's body and reduce it almost to a skin without completely killing it, the striking contrast between these two results must be due to the relative importance of the organs injured. I destroy the nerve-centres and inevitably kill my larva, which is putrid by the following day; the Scolia attacks the reserves of fat, the blood, the muscles and does not kill its victim, which will provide it with wholesome food until the end. But it is clear that, if the Scolia were to set to work as I did, there would be nothing left, after the first few bites, but an actual corpse, discharging fluids which would be fatal to it within twenty-four hours. The mother, it is true, in order to assure the immobility of her prey, has injected the poison of her sting into the nerve-centres. Her operation cannot be compared with mine in any respect. She practises the method of the skilful physiologist who induces anaesthesia; I go to work like the butcher who chops, cuts and disembowels. The sting leaves the nerve-centres intact. Deprived of sensibility by the poison, they have lost the power of provoking muscular contractions; but who can say that, numbed as they are, they no longer serve to maintain a faint vitality? The flame is extinguished, but there is still a glowing speck upon the wick. I, a rough blunderer, do more than blow out the lamp: I throw away the wick and all is over. The grub would do the same if it bit straight into the mass of nerves.
Everything confirms the fact: the Scolia and the other Hunting Wasps whose provisions consist of bulky heads of game are gifted with a special art of eating, an exquisitely delicate art which saves a remnant of life in the prey devoured, until it is all consumed. When the prey is a small one, this precaution is superfluous. Consider, for instance, the Bembex-grubs in the midst of their heap of Flies. The prey seized upon is broached on the back, the belly, the head, the thorax, indifferently. The larva munches a given spot, which it leaves to munch a second, passing to a third and a fourth, at the bidding of its changing whims. It seems to taste and select, by repeated trials, the mouthfuls most to its liking. Thus bitton at several points, covered with wounds, the Fly is soon a shapeless mass which would putrefy very quickly if the meagre dish were not devoured at a single meal. Allow the Scolia-grub the same unlicensed gluttony: it would perish beside its corpulent victim, which should have kept fresh for a fortnight, but which almost from the beginning would be no more than a filthy putrescence.
This art of careful eating does not seem easy to practise: at least, the larva, if ever so little diverted from its usual courses, is no longer able to apply its talent as a capable trencherman. This will be proved by experiment. I must begin by observing that, when I spoke of my larva which turned putrid within twenty-four hours, I adopted an extreme case for the sake of greater clearness. The Scolia, taking its first bite, does not and cannot go to such lengths. Nevertheless it behooves us to enquire whether, in the consumption of the victuals, the initial point of attack is a matter of indifference and whether the rummaging through the entrails of the victim entails a determined order, without which success is uncertain or even impossible. To these delicate questions no one, I think, can reply. Where science is silent, perhaps the grub will speak. We will try.
I move from its position a Scolia-grub which has attained a quarter or a third of its full growth. The long neck plunged into the victim's belly is rather difficult to extract, because of the need of molesting the creature as little as possible. I succeed, by means of a little patience and repeated strokes with the tip of a paint-brush. I now turn the Cetonia-larva over, back uppermost, at the bottom of the little hollow made by pressing my finger in the layer of mould. Lastly, I place the Scolia on its victim's back. Here is my grub under the same conditions as just now, with this difference, that the back and not the belly of its victim is presented to its mandibles.
I watch it for a whole afternoon. It writhes about; it moves its little head now in this direction, now in that, frequently laying it on the Cetonia, but without fixing it anywhere. The day draws to a close; and still it has accomplished nothing. There are restless movements, nothing more. Hunger, I tell myself, will eventually induce it to bite. I am wrong. Next morning I find it more anxious than the day before and still groping about, without resolving to fix its mandibles anywhere. I leave it alone for half a day longer without obtaining any result. Yet twenty-four hours of abstinence must have awakened a good appetite, above all in a creature which, if left undisturbed, would not have ceased eating.
Excessive hunger cannot induce it to nibble at an unlawful spot. Is this due to feebleness of the teeth? By no means: the Cetonia's skin is no tougher on the back than on the belly; moreover, the grub is capable of perforating the skin when it leaves the egg; a fortiori, it must be more capable of doing so now that it has attained a sturdy growth. Thus we see no lack of ability, but an obstinate refusal to nibble at a point which ought to be respected. Who knows? On this side perhaps the grub's dorsal vessel would be wounded, its heart, an organ indispensable to life. The fact remains that my attempts to make the grub tackle its victim from the back have failed. Does this mean that it entertains the least suspicion of the danger which it might incur were it to produce putrefaction by awkwardly carving its victuals from the back? It would be absurd to give such an idea a moment's consideration. Its refusal is dictated by a preordained decree which it is bound to obey.
My Scolia-grubs would die of starvation if I left them on their victim's back. I therefore restore matters as they were, with the Cetonia-larva belly uppermost and the young Scolia on top. I might utilise the subjects of my previous experiments; but, as I have to take precautions against the disturbance which may have been caused by the test already undergone, I prefer to operate on new patients, a luxury in which the richness of my menagerie allows me to indulge. I move the Scolia from its position, extract its head from the entrails of the Cetonia-larva and leave it to its own resources on its victim's belly. Betraying every symptom of uneasiness, the grub gropes, hesitates, casts about and does not insert its mandibles anywhere, though it is now the ventral surface which it is exploring. It would not display greater hesitation if placed on the back of the larva. I repeat, who knows? On this side it might perhaps injure the nervous plexus, which is even more essential than the dorsal vessel. The inexperienced grub must not drive in its mandibles at random; its future is jeopardized if it gives a single ill-judged bite. If it gnaws at the spot where I myself operated with my needle wrought into a scalpel, its victuals will very soon turn putrid. Once more, then, we witness an absolute refusal to perforate the skin of the victim elsewhere than at the very point where the egg was fixed.
The mother selects this point, which is undoubtedly that most favourable to the future prosperity of the larva, though I am not able clearly to discern the reasons for her choice; she fixes the egg to it; and the place where the opening is to be made is henceforth determined. It is here that the grub must bite: only here, never elsewhere. Its invincible refusal to tackle the Cetonia in any other part, even though it should die of starvation, shews us how rigorous is the rule of conduct with which its instinct is inspired.
As it gropes about, the grub laid on the victim's ventral surface sooner or later rediscovers the gaping wound from which I have removed it. If this takes too long for my patience, I can myself guide its head to the place with the point of a paint-brush. The grub then recognizes the hole of its own making, slips its neck into it and little by little dives into the Cetonia's belly, so that the original state of affairs appears to be exactly restored. And yet its successful rearing is henceforth highly problematical. It is possible that the larva will prosper, complete its development and spin its cocoon; it is also possible—and the case is not unusual—that the Cetonia-larva will soon turn brown and putrid. We then see the Scolia itself turn brown, distended as it is with putrescent foodstuffs, and then cease all movement, without attempting to withdraw from the sanies. It dies on the spot, poisoned by its excessively high game.
What can be the meaning of this sudden corruption of the victuals, followed by the death of the Scolia, when everything appeared to have returned to its normal condition? I see only one explanation. Disturbed in its activities and diverted from its usual courses by my interference, the grub, when replaced on the wound from which I extracted it, was unable to rediscover the lode at which it was working a few minutes earlier; it thrust its way at random into the victim's entrails; and a few untimely bites extinguished the last sparks of vitality. Its confusion rendered it clumsy; and the mistake cost it its life. It dies poisoned by the rich food which, if consumed according to the rules, should have made it grow plump and lusty.
I was anxious to observe the deadly effects of a disturbed meal in another fashion. This time the victim itself shall disorder the grub's activities. The Cetonia-larva, as served up to the young Scolia by its mother, is profoundly paralysed. Its inertia is complete and so striking that it constitutes one of the leading features of this narrative. But we will not anticipate. For the moment, the thing is to substitute for this inert larva a similar larva, but one not paralysed, one very much alive. To ensure that it shall not double up and crush the grub, I confine myself to reducing it to helplessness, leaving it otherwise just as I extracted it from its burrow. I must also be careful of its legs and mandibles, the least touch of which would rip open the nurseling. With a few turns of the finest wire I fix it to a little slab of cork, with its belly in the air. Next, to provide the grub with a ready-made hole, knowing that it will refuse to make one for itself, I contrive a slight incision in the skin, at the point where the Scolia lays her egg. I now place the grub upon the larva, with its head touching the bleeding wound, and lay the whole on a bed of mould in a transparent beaker protected by a pane of glass.
Unable to move, to wriggle, to scratch with its legs or snap with its mandibles, the Cetonia-larva, a new Prometheus bound, offers its defenceless flanks to the little Vulture destined to devour its entrails. Without too much hesitation, the young Scolia settles down to the wound made by my scalpel, which to the grub represents the wound whence I have just removed it. It thrusts its neck into the belly of its prey; and for a couple of days all seems to go well. Then, lo and behold, the Cetonia turns putrid and the Scolia dies, poisoned by the ptomaines of the decomposing game! As before, I see it turn brown and die on the spot, still half inside the toxic corpse.
The fatal issue of my experiment is easily explained. The Cetonia-larva is alive in every sense. True, I have, by means of bonds, suppressed its outward movements, in order to provide the nurseling with a quiet meal, devoid of danger; but it was not in my power to subdue its internal movements, the quivering of the viscera and muscles irritated by its forced immobility and by the Scolia's bites. The victim is in possession of its full power of sensation; and it expresses the pain experienced as best it may, by contractions. Embarrassed by these tremors, these twitches of suffering flesh, incommoded at every mouthful, the grub chews away at random and kills the larva almost as soon as it has started on it. In a victim paralysed by the regulation sting, the conditions would be very different. There are no external movements, nor any internal movements either, when the mandibles bite, because the victim is insensible. The grub, undisturbed in any way, is then able, with an unfaltering tooth, to pursue its scientific method of eating.
These marvellous results interested me too much not to inspire me with fresh devices when I pursued my investigations. Earlier enquiries had taught me that the larvae of the Digger-wasps are fairly indifferent to the nature of the game, though the mother always supplies them with the same diet. I had succeeded in rearing them on a great variety of prey, without paying regard to their normal fare. I shall return to this subject later, when I hope to demonstrate its great philosophical significance. Let us profit by these data and try to discover what happens when we give the Scolia food which is not properly its own.
I select from my heap of garden-mould, that inexhaustible mine, two larvae of the Rhinoceros Beetle, Oryctes nasicornis, about one-third full-grown, so that their size may not be out of proportion to the Scolia's. It is in fact almost identical with the size of the Cetonia. I paralyse one of them by giving an injection of ammonia in the nerve-centres. I make a fine incision in its belly and I place the Scolia on the opening. The dish pleases my charge; and it would be strange indeed if this were not so, considering that another Scolia-grub, the larva of the Garden Scolia, feeds on the Oryctes. The dish suits it, for before long it has burrowed half-way into the succulent paunch. This time all goes well. Will the rearing be successful? Not a bit of it! On the third day, the Oryctes decomposes and the Scolia dies. Which shall we hold responsible for the failure, myself or the grub? Myself who, perhaps too unskilfully, administered the injection of ammonia, or the grub which, a novice at dissecting a prey differing from its own, did not know how to practise its craft upon a changed victim and began to bite before the proper time?
In my uncertainty, I try again. This time I shall not interfere, so that my clumsiness cannot be to blame. As I described when speaking of the Cetonia-larva, the Oryctes-larva now lies bound, quite alive, on a strip of cork. As usual, I make a small opening in the belly, to entice the grub by means of a bleeding wound and facilitate its access. I obtain the same negative result. In a little while, the Oryctes is a noisome mass on which the nurseling lies poisoned. The failure was foreseen: to the difficulties presented by a prey unknown to my charge was added the commotion caused by the wriggling of an unparalysed animal.
We will try once more, this time with a victim paralysed not by me, an unskilled operator, but by an adept whose ability ranks so high that it is beyond discussion. Chance favours me to perfection: yesterday, in a warm sheltered corner, at the foot of a sandy bank, I discovered three cells of the Languedocian Sphex, each with its Ephippiger and the recently laid egg. This is the game I want, a corpulent prey, of a size suited to the Scolia and, what is more, in splendid condition, artistically paralysed according to rule by a master among masters.
As usual, I install my three Ephippigers in a glass jar, on a bed of mould; I remove the egg of the Sphex and on each victim, after slightly incising the skin of the belly, I place a young Scolia-grub. For three or four days my charges feed upon this game, so novel to them, without any sign of repugnance or hesitation. By the fluctuations of the digestive canal I perceive that the work of nutrition is proceeding as it should; things are happening just as if the dish were a Cetonia-larva. The change of diet, complete though it is, has in no way affected the appetite of the Scolia-grubs. But this prosperous condition does not last long. About the fourth day, a little sooner in one case, a little later in another, the three Ephippigers become putrid and the Scoliae die at the same time.
This result is eloquent. Had I left the egg of the Sphex to hatch, the larva coming out of it would have fed upon the Ephippiger; and for the hundredth time I should have witnessed an incomprehensible spectacle, that of an animal which, devoured piecemeal for nearly a fortnight, grows thin and empty, shrivels up and yet retains to the very end the freshness peculiar to living flesh. Substitute for this Sphex-larva a Scolia-larva of almost the same size; let the dish be the same though the guest is different; and healthy live flesh is promptly replaced by pestilent rotten flesh. That which under the mandibles of the Sphex would for a long while have remained wholesome food promptly becomes a poisonous liquescence under the mandibles of the Scolia.
It is impossible to explain the preservation of the victuals until finally consumed by supposing that the venom injected by the Wasp when she delivers her paralysing stings possesses antiseptic properties. The three Ephippigers were operated on by the Sphex. Able to keep fresh under the mandibles of the Sphex-larvae, why did they promptly go bad under the mandibles of the Scolia-larvae? Any idea of an antiseptic must needs be rejected: a liquid preservative which would act in the first case could not fail to act in the second, as its virtues would not depend on the teeth of the consumer.
Those of you who are versed in the knowledge attaching to this problem, investigate, I beg you, search, sift, see if you can discover the reason why the victuals keep fresh when consumed by a Sphex, whereas they promptly become putrid when consumed by a Scolia. For me, I see only one reason; and I very much doubt whether any one can suggest another.
Both larvae practise a special art of eating, which is determined by the nature of the game. The Sphex, when sitting down to an Ephippiger, the food that has fallen to its lot, knows thoroughly how to consume it and how to preserve, to the very end, the glimmer of life which keeps it fresh; but, if it has to browse upon a Cetonia-grub, whose different structure would confuse its talents as a dissector, it would soon have nothing before it but a heap of putrescence. The Scolia, in its turn, is familiar with the method of eating the Cetonia-grub, its invariable portion; but it does not understand the art of eating the Ephippiger, though the dish is to its taste. Unable to dissect this unknown species of game, its mandibles slash away at random, killing the creature outright as soon as they take their first bites of the deeper tissues of the victim. That is the whole secret.
One more word, on which I shall enlarge in another chapter. I observe that the Scoliae to which I give Ephippigers paralysed by the Sphex keep in excellent condition, despite the change of diet, so long as the provisions retain their freshness. They languish when the game goes high; and they die when putridity supervenes. Their death, therefore, is due not to an unaccustomed diet, but to poisoning by one or other of those terrible toxins which are engendered by animal corruption and which chemistry calls by the name of ptomaines. Therefore, notwithstanding the fatal outcome of my three attempts, I remain persuaded that the unfamiliar method of rearing would have been perfectly successful had the Ephippigers not gone bad, that is, if the Scoliae had known how to eat them according to the rules.
What a delicate and dangerous thing is the art of eating in these carnivorous larvae supplied with a single victim, which they have to spend a fortnight in consuming, on the express condition of not killing it until the very end! Could our physiological science, of which, with good reason, we are so proud, describe, without blundering, the method to be followed in the successive mouthfuls? How has a miserable grub learnt what our knowledge cannot tell us? By habit, the Darwinians will reply, who see in instinct an acquired habit.
Before deciding this serious matter, I will ask you to reflect that the first Wasp, of whatever kind, that thought of feeding her progeny on a Cetonia-grub or on any other large piece of game demanding long preservation could necessarily have left no descendants unless the art of consuming food without causing putrescence had been practised, with all its scrupulous caution, from the first generation onwards. Having as yet learnt nothing by habit or by atavistic transmission, since it was making a first beginning, the nurseling would bite into its provender at random. It would be starving, it would have no respect for its prey. It would carve its joint at random; and we have just seen the fatal consequence of an ill-directed bite. It would perish—I have just proved this in the most positive manner—it would perish, poisoned by its victim, already dead and putrid.
To prosper, it would have, although a novice, to know what was permitted and what forbidden in ransacking the creature's entrails; nor would it be enough for the larva to be approximately in possession of this difficult secret: it would be indispensable that it should possess the secret completely, for a single bite, if delivered before the right moment, would inevitably involve its own demise. The Scoliae of my experiments are not novices, far from it: they are the descendants of carvers that have practised their art since Scoliae first came into the world; nevertheless they all perish from the decomposition of the rations supplied, when I try to feed them on Ephippigers paralysed by the Sphex. Very expert in the method of attacking the Cetonia, they do not know how to set about the business of discreetly consuming a species of game new to them. All that escapes them is a few details, for the trade of an ogre fed on live flesh is familiar to them in its general features; and these unheeded details are enough to turn their food into poison. What, then, happened in the beginning, when the larva bit for the first time into a luscious victim? The inexperienced creature perished; of that there is not a shadow of doubt, unless we admit an absurdity and imagine the larva of antiquity feeding upon those terrible ptomaines which so swiftly kill its descendants to-day.
Nothing will ever make me admit and no unprejudiced mind can admit that what was once food has become a horrible poison. What the larva of antiquity ate was live flesh and not putrescence. Nor can it be admitted that the chances of fortune can have led at the first trial to success in a system of nourishment so full of pit-falls: fortuitous results are preposterous amid so many complications. Either the feeding is strictly methodical at the beginning, in conformity with the organic exigencies of the prey devoured, and the Wasp established her race; or else it was hesitating, without determined rules, and the Wasp left no successor. In the first case we behold innate instinct; in the second acquired habit.
A strange acquisition, truly! An acquisition presumed to be made by an impossible creature; an acquisition supposed to develop in no less impossible successors! Though the snow-ball, slowly rolling, at last becomes an enormous sphere, it is still necessary that the starting-point shall not have been NIL. The big ball implies the little ball, as small as you please. Now, in harking back to the origin of these acquired habits, if I interrogate the possibilities I obtain zero as the only answer. If the animal does not know its trade thoroughly, if it has to acquire something, all the more if it has to acquire everything, it perishes: that is inevitable; without the little snow-ball the big snow-ball cannot be rolled. If it has nothing to acquire, if it knows all that it needs to know, it flourishes and leaves descendants behind it. But then it possesses innate instinct, the instinct which learns nothing and forgets nothing, the instinct which is steadfast throughout time.
The building up of theories has never appealed to me: I suspect them one and all. To argue nebulously upon dubious premises likes me no better. I observe, I experiment and I let the facts speak for themselves. We have just heard these facts. Let each now decide for himself whether instinct is an innate faculty or an acquired habit.
CHAPTER 4. THE CETONIA-LARVA.
The Scolia's feeding-period lasts, on the average, for a dozen days or so. By then the victuals are no more than a crumpled bag, a skin emptied of the last scrap of nutriment. A little earlier, the russet-yellow tint announces the extinction of the last spark of life in the creature that is being devoured. The empty skin is pushed back to make space; the dining-room, a shapeless cavity with crumbling walls, is tidied up a little; and the Scolia-grub sets to work on its cocoon without further delay.
The first courses form a general scaffolding, which finds a support here and there on the earthen walls, and consist of a rough, blood-red fabric. When the larva is merely laid, as required by my investigations, in a hollow made with the finger-tip in the bed of mould, it is not able to spin its cocoon, for want of a ceiling to which to fasten the upper threads of its network. To weave its cocoon, every spinning larva is compelled to isolate itself in a hammock slung in an open-work enclosure, which enables it to distribute its thread uniformly in all directions. If there be no ceiling, the upper part of the cocoon cannot be fashioned, because the worker lacks the necessary points of support. Under these conditions my Scolia-grubs contrive at most to upholster their little pit with a thick down of reddish silk. Discouraged by futile endeavours, some of them die. It is as if they had been killed by the silk which they omit to disgorge because they are unable to make the right use of it. This, if we were not watchful, would be a very frequent cause of failure in our attempts at artificial rearing. But, once the danger has been perceived, the remedy is simple. I make a ceiling over the cavity by laying a short strip of paper above it. If I want to see how matters are progressing, I bend the strip into a semicircle, into a half-cylinder with open ends. Those who wish to play the breeder for themselves will be able to profit by these little practical details.
In twenty-four hours the cocoon is finished; at least, it no longer allows us to see the grub, which is doubtless making the walls of its dwelling still thicker. At first the cocoon is a vivid red; later it changes to a light chestnut-brown. Its form is that of an ellipsoid, with a major axis 26 millimetres in length, while the minor axis measures 11 millimetres. (1.014 x.429 inch.—Translator's Note.) These dimensions, which incidentally are inclined to vary slightly, are those of the female cocoons. In the other sex they are smaller and may measure as little as 17 millimetres in length by 7 millimetres in width. (.663 x.273 inch.—Translator's Note.)
The two ends of the ellipsoid have the same form, so much so that it is only thanks to an individual peculiarity, independent of the shape, that we can tell the cephalic from the anal extremity. The cephalic pole is flexible and yields to the pressure of my tweezers; the anal pole is hard and unyielding. The wrapper is double, as in the cocoons of the Sphex. (Cf. "The Hunting Wasps": chapters 4 to 10 et passim.—Translator's Note.) The outer envelope, consisting of pure silk, is thin, flexible and offers little resistance. It is closely superimposed upon the inner envelope and is easily separated from it everywhere, except at the anal end, where it adheres to the second envelope. The adhesion of the two wrappers at one end and the non-adhesion at the other are the cause of the differences which the tweezers reveal when pinching the two ends of the cocoon.
The inner envelope is firm, elastic, rigid and, to a certain point, brittle. I do not hesitate to look upon it as consisting of a silken tissue which the larva, towards the end of its task, has steeped thoroughly in a sort of varnish prepared not by the silk-glands but by the stomach. The cocoons of the Sphex have already shown us a similar varnish. This product of the chylific ventricle is chestnut-brown. It is this which, saturating the thickness of the tissue, effaces the bright red of the beginning and replaces it by a brown tint. It is this again which, disgorged more profusely at the lower end of the cocoon, glues the two wrappers together at that point.
The perfect insect is hatched at the beginning of July. The emergence takes place without any violent effraction, without any ragged rents. A clean, circular fissure appears at some distance from the top; and the cephalic end is detached all of a piece, as a loose lid might be. It is as though the recluse had only to raise a cover by butting it with her head, so exact is the line of division, at least as regards the inner envelope, the stronger and more important of the two. As for the outer wrapper, its lack of resistance enables it to yield without difficulty when the other gives way.
I cannot quite make out by what knack the Wasp contrives to detach the cap of the inner shell with such accuracy. Is it the art practised by the tailor when cutting his stuff, with mandibles taking the place of scissors? I hardly venture to admit as much: the tissue is so tough and the circle of division so precise. The mandibles are not sharp enough to cut without leaving a ragged edge; and then what geometrical certainty they would need for an operation so perfect that it might well have been performed with the compasses!
I suspect therefore that the Scolia first fashions the outer sac in accordance with the usual method, that is, by distributing the silk uniformly, without any special preparation of one part of the wall more than of another, and that it afterwards changes its method of weaving in order to attend to the main work, the inner shell. In this it apparently imitates the Bembex (Cf. "The Hunting Wasps": chapters 14 to 16.—Translator's Note.), which weaves a sort of eel-trap, whose ample mesh allows it to gather grains of sand outside and encrust them one by one in the silky network, and completes the performance with a cap fitting the entrance to the trap. This provides a circular line of least resistance, along which the casket breaks open afterwards. If the Scolia really works in the same manner, everything is explained: the eel-trap, while still open, enables it to soak with varnish both the inside and the outside of the inner shell, which has to acquire the consistency of parchment; lastly, the cap which completes and closes the structure leaves for the future a circular line capable of splitting easily and neatly.
This is enough on the subject of the Scolia-grub. Let us go back to its provender, of whose remarkable structure we as yet know nothing. In order that it may be consumed with the delicate anatomical discretion imposed by the necessity of having fresh food to the last, the Cetonia-grub must be plunged into a state of absolute immobility: any twitchings on its part—as the experiments which I have undertaken go to prove—would discourage our nibbling larva and impede the work of carving, which has to be effected with so much circumspection. It is not enough for the victim to be unable to move from place to place beneath the soil: in addition to this, the contractible power in its sturdy muscular organism must be suppressed.
In its normal state, this larva, at the very least disturbance, curls itself up, almost as the Hedgehog does; and the two halves of the ventral surface are laid one against the other. You are quite surprised at the strength which the creature displays in keeping itself thus contracted. If you try to unroll it, your fingers encounter a resistance far greater than the size of the animal would have caused you to suspect. To overcome the resistance of this sort of spring coiled upon itself, you have to force it, so much so that you are afraid, if you persist, of seeing the indomitable spiral suddenly burst and shoot forth its entrails.
A similar muscular energy is found in the larvae of the Oryctes (Also known as the Rhinoceros Beetle.—Translator's Note.), the Anoxia (A Beetle akin to the Cockchafer.—Translator's Note.), the Cockchafer. Weighed down by a heavy belly and living underground, where they feed either on leaf-mould or on roots, these larvae all possess the vigorous constitution needed to drag their corpulence through a resisting medium. All of them also roll themselves into a hook which is not straightened without an effort.
Now what would become of the egg and the new-born grub of the Scoliae, fixed under the belly, at the centre of the Cetonia's spiral, or inside the hook of the Oryctes or the Anoxia? They would be crushed between the jaws of the living vice. It is essential that the arc should slacken and the hook unbend, without the least possibility of their returning to a state of tension. Indeed, the well-being of the Scoliae demands something more: those powerful bodies must not retain even the power to quiver, lest they derange a method of feeding which has to be conducted with the greatest caution.
The Cetonia-grub to which the Two-banded Scolia's egg is fastened fulfils the required conditions admirably. It is lying on its back, in the midst of the mould, with its belly fully extended. Long accustomed though I be to this spectacle of victims paralysed by the sting of the Hunting Wasp, I cannot suppress my astonishment at the profound immobility of the prey before my eyes. In the other victims with flexible skins, Caterpillars, Crickets, Mantes, Ephippigers, I perceived at least some pulsations of the abdomen, a few feeble contortions under the stimulus of a needle. There is nothing of the sort here, nothing but absolute inertia, except in the head, where I see, from time to time, the mouth-parts open and close, the palpi give a tremor, the short antennae sway to and fro. A prick with the point of a needle causes no contraction, no matter what the spot pricked. Though I stab it through and through, the creature does not stir, be it ever so little. A corpse is not more inert. Never, since my remotest investigations, have I witnessed so profound a paralysis. I have seen many wonders due to the surgical talent of the Wasp; but to-day's marvel surpasses them all.
I am doubly surprised when I consider the unfavourable conditions under which the Scolia operates. The other paralysers work in the open air, in the full light of day. There is nothing to hinder them. They enjoy full liberty of action in seizing the prey, holding it in position and sacrificing it; they are able to see the victim and to parry its means of defence, to avoid its spears, its pincers. The spot or spots to be attained are within their reach; they drive the dagger in without let or hindrance.
What difficulties, on the other hand, await the Scolia! She hunts underground, in the blackest darkness. Her movements are laboured and uncertain, owing to the mould, which is continually giving way all round her; she cannot keep her eyes on the terrible mandibles, which are capable of cutting her body in two with a single bite. Moreover, the Cetonia-grub, perceiving that the enemy is approaching, assumes its defensive posture, rolls itself up and makes a shield for its only vulnerable part, the ventral surface, with its convex back. No, it cannot be an easy operation to subdue the powerful larva in its underground retreat and to stab with the precision which immediate paralysis requires.
We wish that we might witness the struggle between the two adversaries and see at first hand what happens, but we cannot hope to succeed. It all takes place in the mysterious darkness of the soil; in broad daylight, the attack would not be delivered, for the victim must remain where it is and then and there receive the egg, which is unable to thrive and develop except under the warm cover of vegetable mould. If direct observation is impracticable, we can at least foresee the main outlines of the drama by allowing ourselves to be guided by the warlike manoeuvres of other burrowers.
I picture things thus: digging and rummaging through the heap of mould, guided perhaps by that singular sensibility of the antennae which enables the Hairy Ammophila to discover the Grey Worm (The caterpillar of the Turnip Moth. Cf. "The Hunting Wasps": chapters 18 to 20.—Translator's Note.) underground, the Scolia ends by finding a Cetonia-larva, a good plump one, in the pink of condition, having reached its full growth, just what the grub which is to feed on it requires. Forthwith, the assaulted victim, contracting desperately, rolls itself into a ball. The other seizes it by the skin of the neck. To unroll it is impossible to the insect, for I myself have some trouble in doing so. One single point is accessible to the sting: the under part of the head, or rather of the first segments, which are placed outside the coil, so that the grub's hard cranium makes a rampart for the hinder extremity, which is less well defended. Here the Wasp's sting enters and here only can it enter, within a narrowly circumscribed area. One stab only of the lancet is given at this point, one only because there is no room for more; and this is enough: the larva is absolutely paralysed.
The nervous functions are abolished instantly; the muscular contractions cease; and the animal uncoils like a broken spring. Henceforth motionless, it lies on its back, its ventral surface fully exposed from end to end. On the median line of this surface, towards the rear, near the brown patch due to the alimentary broth contained in the intestine, the Scolia lays her egg and without more ado, leaves everything lying on the actual spot where the murder was committed, in order to go in search of another victim.
This is how the deed must be done: the results prove it emphatically. But then the Cetonia-grub must possess a very exceptional structure in its nervous organization. The larva's violent contraction leaves but a single point of attack open to the sting, the under part of the neck, which is doubtless uncovered when the victim tries to defend itself with its mandibles; and yet a stab in this one point produces the most thorough paralysis that I have ever seen. It is the general rule that larvae possess a centre of innervation for each segment. This is so in particular with the Grey Worm, the sacrificial victim of the Hairy Ammophila. The Wasp is acquainted with this anatomical secret: she stabs the caterpillar again and again, from end to end, segment by segment, ganglion by ganglion. With such an organization the Cetonia-grub, unconquerably coiled upon itself would defy the paralyser's surgical skill.
If the first ganglion were wounded, the others would remain uninjured; and the powerful body, actuated by these last, would lose none of its powers of contraction. Woe then to the egg, to the young grub held fast in its embrace! And how insurmountable would be the difficulties if the Scolia, working in the profound darkness amid the crumbling soil and confronted by a terrible pair of mandibles, had to stab each segment in turn with her sting, with the certainty of method displayed by the Ammophila! The delicate operation is possible in the open air, where nothing stands in the way, in broad daylight, where the sight guides the scalpel, and with a patient which can always be released if it becomes dangerous. But in the dark, underground, amidst the ruins of a ceiling which crumbles in consequence of the conflict and at close quarters with an opponent greatly her superior in strength, how is the Scolia to guide her sting with the accuracy that is essential if the stabs are to be repeated?
So profound a paralysis; the difficulty of vivisection underground; the desperate coiling of the victim: all these things tell me that the Cetonia-grub, as regards its nervous system, must possess a structure peculiar to itself. The whole of the ganglia must be concentrated in a limited area in the first segments, almost under the neck. I see this as clearly as though it had been revealed to me by a post-mortem dissection.
Never was anatomical forecast more fully confirmed by direct examination. After forty-eight hours in benzine, which dissolves the fat and renders the nervous system more plainly visible, the Cetonia-grub is subjected to dissection. Those of my readers who are familiar with these investigations will understand my delight. What a clever school is the Scolia's! It is just as I thought! Admirable! The thoracic and abdominal ganglia are gathered into a single nervous mass, situated within the quadrilateral bounded by the four hinder legs, which legs are very near the head. It is a tiny, dull-white cylinder, about three millimetres long by half a millimetre wide. (.117 x.019 inch.—Translator's Note.) This is the organ which the Scolia's sting must attack in order to secure the paralysis of the whole body, excepting the head, which is provided with special ganglia. From it run numbers of filaments which actuate the feet and the powerful muscular layer which is the creature's essential motor organ. When examined merely through the pocket-lens, this cylinder appears to be slightly furrowed transversely, a proof of its complex structure. Under the microscope, it is seen to be formed by the close juxtaposition, the welding, end to end, of the ganglia, which can be distinguished one from the other by a slight intermediate groove. The bulkiest are the first, the fourth and the tenth, or last; these are all very nearly of equal size. The rest are barely half or even a third as large as those mentioned.
The Interrupted Scolia experiences the same hunting and surgical difficulties when she attacks, in the crumbling, sandy soil, the larvae of the Shaggy Anoxia or of the Morning Anoxia, according to the district; and these difficulties, if they are to be overcome, demand in the victim a concentrated nervous system, like the Cetonia's. Such is my logical conviction before making my examination; such also is the result of direct observation. When subjected to the scalpel, the larva of the Morning Anoxia shows me its centres of innervation for the thorax and the abdomen, gathered into a short cylinder, which, placed very far forward, almost immediately after the head, does not run back beyond the level of the second pair of legs. The vulnerable point is thus easily accessible to the sting, despite the creature's posture of defence, in which it contracts and coils up. In this cylinder I recognize eleven ganglia, one more than in the Cetonia. The first three, or thoracic, ganglia are plainly distinguishable from one another, although they are set very close together; the rest are all in contact. The largest are the three thoracic ganglia and the eleventh.
After ascertaining these facts, I remembered Swammerdam's investigations into the grub of the Monoceros, our Oryctes nasicornis. (Jan Swammerdam (1637-1680), the Dutch naturalist and anatomist.—Translator's Note.) I chanced to possess an abridgement of the "Biblia naturae," the masterly work of the father of insect anatomy. I consulted the venerable volume. It informed me that the learned Dutchman had been struck, long before I was, by an anatomical peculiarity similar to that which the larvae of the Cetoniae and Anoxiae had shown me in their nerve-centres. Having observed in the Silk-worm a nervous system formed of ganglia distinct one from the other, he was quite surprised to find that, in the grub of the Oryctes, the same system was concentrated into a short chain of ganglia in juxtaposition. His was the surprise of the anatomist who, studying the organ qua organ, sees for the first time an unusual conformation. Mine was of a different nature: I was amazed to see the precision with which the paralysis of the victim sacrificed by the Scolia, a paralysis so profound in spite of the difficulties of an underground operation, had guided my forecast as to structure when, anticipating the dissection, I declared in favour of an exceptional concentration of the nervous system. Physiology perceived what anatomy had not yet revealed, at all events to my eyes, for since then, on dipping into my books, I have learnt that these anatomical peculiarities, which were then so new to me, are now within the domain of current science. We know that, in the Scarabaeidae, both the larva and the perfect insect are endowed with a concentrated nervous system.
The Garden Scolia attacks Oryctes nasicornis; the Two-banded Scolia the Cetonia; the Interrupted Scolia the Anoxia. All three operate below ground, under the most unfavourable conditions; and all three have for their victim a larva of one of the Scarabaeidae, which, thanks to the exceptional arrangement of its nerve-centres, lends itself, alone of all larvae, to the Wasp's successful enterprises. In the presence of this underground game, so greatly varied in size and shape and yet so judiciously selected to facilitate paralysis, I do not hesitate to generalize and I accept, as the ration of the other Scoliae, larvae of Lamellicorns whose species will be determined by future observation. Perhaps one of them will be found to give chase to the terrible enemy of my crops, the voracious White Worm, the grub of the Cockchafer; perhaps the Hemorrhoidal Scolia, rivalling in size the Garden Scolia and like her, no doubt, requiring a copious diet, will be entered in the insects' "Who's Who" as the destroyer of the Pine-chafer, that magnificent Beetle, flecked with white upon a black or brown ground, who of an evening, during the summer solstice, browses on the foliage of the fir-trees. Though unable to speak with certainty or precision, I am inclined to look upon these devourers of Scarabaeus-grubs as valiant agricultural auxiliaries.
The Cetonia-larva has figured hitherto only in its quality of a paralysed victim. We will now consider it in its normal state. With its convex back and its almost flat ventral surface, the creature is like a semi-cylinder in shape, fuller in the hinder portion. On the back, each of the segments, except the last, or anal, segment, puckers into three thick pads, bristling with stiff, tawny hairs. The anal segment, much wider than the rest, is rounded at the end and coloured a deep brown by the contents of the intestine, which show through the translucent skin; it bristles with hairs like the other segments, but is level, without pads. On the ventral surface, the segments have no creases; and the hairs, though abundant, are rather less so than on the back. The legs, which are quite well-formed, are short and feeble in comparison with the animal's size. The head has a strong, horny cap for a cranium. The mandibles are powerful, with bevelled tips and three or four teeth on the edge of the bevel.
Its mode of locomotion marks it as an idiosyncratic, exceptional, fantastic creature, having no fellow, that I know of, in the insect world. Though endowed with legs—a trifle short, it is true, but after all as good as those of a host of other larvae—it never uses them for walking. It progresses on its back, always on its back, never otherwise. By means of wriggling movements and the purchase afforded by the dorsal bristles, it makes its way belly upwards, with its legs kicking the empty air. The spectator to whom these topsy-turvy gymnastics are a novelty thinks at first that the creature must have had a fright of some sort and that it is struggling as best it can in the face of danger. He puts it back on its belly; he lays it on its side. Nothing is of any use; it obstinately turns over and resumes its dorsal progress. That is its manner of travelling over a flat surface; it has no other.
This reversal of the usual mode of walking is so peculiar to the Cetonia-larva that it is enough in itself to reveal the grub's identity to the least expert eyes. Dig into the vegetable mould formed by the decayed wood in the hollow trunks of old willow-trees, search at the foot of rotten stumps or in heaps of compost; and, if you come upon a plumpish grub moving along on its back, there is no room for doubt: your discovery is a Cetonia-larva.
This topsy-turvy progress is fairly swift and is not less in speed to that of an equally fat grub travelling on its legs. It would even be greater on a polished surface, where walking on foot is hampered by incessant slips, whereas the numerous hairs of the dorsal pads find the necessary support by multiplying the points of contact. On polished wood, on a sheet of paper and even on a strip of glass, I see my grubs moving from point to point with the same ease as on a surface of garden mould. In the space of one minute, on the wood of my table, they cover a distance of eight inches. The pace is no swifter on a horizontal bed of sifted mould. A strip of glass reduces the distance covered by one half. The slippery surface only half paralyses this strange method of locomotion.
We will now place side by side with the Cetonia-grub the larva of the Morning Anoxia, the prey of the Interrupted Scolia. It is very like the larva of the Common Cockchafer. It is a fat, pot-bellied grub, with a thick, red cap on its head and armed with strong, black mandibles, which are powerful implements for digging and cutting through roots. The legs are sturdy and end in a hooked nail. The creature has a long, heavy, brown paunch. When placed on the table, it lies on its side; it struggles without being able to advance or even to remain on its belly or back. In its usual posture it is curled up into a narrow hook. I have never seen it straighten itself completely; the bulky abdomen prevents it. When placed on a surface of moist sand, the ventripotent creature is no better able to shift its position: curved into a fish-hook, it lies on its side.
To dig into the earth and bury itself, it uses the fore-edge of its head, a sort of weeding-hoe with the two mandibles for points. The legs take part in this work, but far less effectually. In this way it contrives to dig itself a shallow pit. Then, bracing itself against the wall of the pit, with the aid of wriggling movements which are favoured by the short, stiff hairs bristling all over its body, the grub changes its position and plunges into the sand, but still with difficulty.
Apart from a few details, which are of no importance here, we may repeat this sketch of the Anoxia-grub and we shall have, if the size be at least quadrupled, a picture of the larva of Oryctes nasicornis, the monstrous prey of the Garden Scolia. Its general appearance is the same: there is the same exaggeration of the belly; the same hook-like curve; the same incapacity for standing on its legs. And as much may be said of the larva of Scarabaeus pentodon, a fellow-boarder of the Oryctes and the Cetonia.