The Glow-Worm and Other Beetles

And indeed, if we tear the double envelope which protects this mystery, we recognize, not without astonishment, that we have before our eyes a new larva similar to the secondary. After one of the strangest transformations, the creature has gone back to its second form. To describe the new larva is unnecessary, for it differs from the former in only a few slight details. In both there is the same head, with its various appendages barely outlined; the same vestiges of legs, the same stumps transparent as crystal. The tertiary larva differs from the secondary only by its abdomen, which is less fat, owing to the absolute emptiness of the digestive apparatus; by a double chain of fleshy cushions extending along each side; by the rim of the stigmata, crystalline and slightly projecting, but less so than in the pseudochrysalis; by the ninth pair of breathing-holes, hitherto rudimentary but now almost as large as the rest; lastly by the mandibles ending in a very sharp point. Evicted from its twofold sheath, the tertiary larva makes only very lazy movements of contraction and dilation, without being able to advance, without even being able to maintain its normal position, because of the weakness of its legs. It usually remains motionless, lying on its side, or else displays its drowsy activity merely by feeble, wormlike movements.

By dint of these alternate contractions and dilations, indolent though they be, the larva nevertheless contrives to turn right round in the sort of shell with which the pseudochrysalidal integuments provide it, when by accident it finds itself placed head downwards; and this operation is all the more difficult inasmuch as the larva almost exactly fills the cavity of the shell. The creature contracts, bends its head under its belly and slides its front half over its hinder half by wormlike movements so slow that the lens can hardly detect them. In less than a quarter of an hour the larva, at first turned upside down, finds itself again head uppermost. I admire this gymnastic feat, but have some difficulty in understanding it, so small is the space which the larva, when at rest in its cell, leaves unoccupied, compared with that which we should be justified in expecting from the possibility of such a reversal. The larva does not long enjoy the privilege which enables it to resume inside its cell, when this is moved from its original position, the attitude which it prefers, that is to say, with its head up.

Two days, at most, after its first appearance it relapses into an inertia as complete as that of the pseudochrysalis. On removing it from its amber shell, we see that its faculty of contracting or dilating at will is so completely paralysed that the stimulus of a needle is unable to provoke it, though the integuments have retained all their flexibility and though no perceptible change has occurred in the organization. The irritability, therefore, which in the pseudochrysalis is suspended for a whole year, reawakens for a moment, to relapse instantly into the deepest torpor. This torpor will be partly dispelled only at the moment of the passing into the nymphal stage, to return immediately afterwards and last until the insect attains the perfect state.

Further, on holding larvæ of the third form, or nymphs enclosed in their cells, in an inverted position, in glass tubes, we never see them regain an erect position, however long we continue the experiment. The perfect insect itself, during the time that it is enclosed in the shell, cannot regain it, for lack of the requisite flexibility. This total absence of movement in the tertiary larva, when a few days old, and also in the nymph, together with the smallness of the space left free in the shell, would necessarily lead to the conviction, if we had not witnessed the first moments of the tertiary larva, that it is absolutely impossible for the creature to turn right round.

And now see to what curious inferences this lack of observations made at the due moment may lead us. We collect some pseudochrysalids and heap them in a glass jar in all possible positions. The favourable season arrives; and with very legitimate astonishment we find that, in a large number of shells, the larva or nymph occupies an inverted position, that is to say, the head is turned towards the anal extremity of the shell. In vain we watch these reversed bodies for any indications of movement; in vain we place the shells in every imaginable position, to see if the creature will turn round; in vain, once more, we ask ourselves where the free space is which this turning would demand. The illusion is complete: I have been taken in by it myself; and for two years I indulged in the wildest conjectures to account for this lack of correspondence between the shell and its contents, to explain, in short, a fact which is inexplicable once the propitious moment has passed.

On the natural site, in the cells of the Anthophora, this apparent anomaly never occurs, because the secondary larva, when on the point of transformation into the pseudochrysalis, is always careful to place its head uppermost, according as the axis of the cell more or less nearly approaches the vertical. But, when the pseudochrysalids are placed higgledy-piggledy in a box or jar, all those which are upside down will later contain inverted larvæ or nymphs.

After four changes of form so profound as those which I have described, one might reasonably expect to find some modifications of the internal organization. Nevertheless, nothing is changed; the nervous system is the same in the tertiary larva as in the earlier phases; the reproductive organs do not yet show; and there is no need to mention the digestive apparatus, which remains invariable even in the perfect insect.

The duration of the tertiary larva is a bare four or five weeks, which is also about the duration of the second. In July, when the secondary larva passes into the pseudochrysalid stage, the tertiary larva passes into the nymphal stage, still inside the double vesicular envelope. Its skin splits along the back in front; and with the assistance of a few feeble contractions, which reappear at this juncture, it is thrust behind in the shape of a little ball. There is therefore nothing here that differs from what happens in the other Beetles.

Nor does the nymph which succeeds this tertiary larva present any peculiarity: it is the perfect insect in swaddling-bands, yellowish white, with its various external members, clear as crystal, displayed under the abdomen. A few weeks elapse, during which the nymph partly dons the livery of the adult state; and, in about a month, the insect moults for a last time, in the usual manner, in order to attain its final form. The wing-cases are now of a uniform yellowish white, as are the wings, the abdomen and the greater part of the legs; very nearly all the rest of the body is of a glossy black. In the space of twenty-four hours, the wing-cases assume their half-black, half-russet colouring; the wings grow darker; and the legs finish turning black. This done, the adult organism is completed. However, the Sitaris remains still a fortnight in the intact shell, ejecting at intervals white droppings of uric acid, which it pushes back together with the shreds of its last two sloughs, those of the tertiary larva and of the nymph. Lastly, about the middle of August, it tears the double bag that contains it, pierces the lid of the Anthophora's cell, enters a corridor and appears outside in quest of the other sex.

I have told how, while digging in search of the Sitaris, I found two cells belonging to Meloe cicatricosus. One contained an Anthophora's egg; with this egg was a yellow Louse, the primary larva of the Meloe. The history of this tiny creature we know. The second cell also was full of honey. On the sticky liquid floated a little white larva, about a sixth of an inch in length and very different from the other little white larvæ belonging to Sitares. The rapid fluctuations of the abdomen showed that it was eagerly drinking the strong-scented nectar collected by the Bee. This larva was the young Meloe in the second period of its development.

I was not able to preserve these two precious cells, which I had opened wide to examine the contents. On my return from Carpentras, I found that their honey had been spilt by the motion of the carriage and that their inhabitants were dead. On the 25th of June, a fresh visit to the nests of the Anthophoræ furnished me with two larvæ like the foregoing, but much larger. One of them was on the point of finishing its store of honey, the other still had nearly half left. The first was put in a place of safety with a thousand precautions, the second was at once immersed in alcohol.

These larvæ are blind, soft, fleshy, yellowish white, covered with a fine down visible only under the lens, curved into a fish-hook like the larvæ of the Lamellicorns, to which they bear a certain resemblance in their general configuration. The segments, including the head, number thirteen, of which nine are provided with breathing-holes with a pale, oval rim. These are the mesothorax and the first eight abdominal segments. As in the Sitaris-larvæ, the last pair of stigmata, that of the eighth segment of the abdomen, is less developed than the rest.

The head is horny, of a light brown colour. The epistoma is edged with brown. The labrum is prominent, white and trapezoidal. The mandibles are black, strong, short, obtuse, only slightly curved, sharp-edged and furnished each with a broad tooth on the inner side. The maxillary and labial palpi are brown and shaped like very small studs with two or three joints to them. The antennæ, inserted just at the base of the mandibles, are brown, and consist of three sections: the first is thick and globular; the two others are much smaller in diameter and cylindrical. The legs are short, but fairly strong, able to serve the creature for crawling or digging; they end in a strong black claw. The length of the larva when fully developed is one inch.

As far as I can judge from the dissection of the specimen preserved in alcohol, whose viscera were affected by being kept too long in that liquid, the nervous system consists of eleven ganglia, not counting the oesophageal collar; and the digestive apparatus does not differ perceptibly from that of an adult Oil-beetle.

The larger of the two larvæ of the 25th of June, placed in a test-tube with what remained of its provisions, assumed a new form during the first week of the following month. Its skin split along the front dorsal half and, after being pushed half back, left partly uncovered a pseudochrysalis bearing the closest analogy with that of the Sitares. Newport did not see the larva of the Oil-beetle in its second form, that which it displays when it is eating the mess of honey hoarded by the Bees, but he did see its moulted skin half-covering the pseudochrysalis which I have just mentioned. From the sturdy mandibles and the legs armed with a powerful claw which he observed on this moulted skin, Newport assumed that, instead of remaining in the same Anthophora-cell, the larva, which is capable of burrowing, passes from one cell to another in search of additional nourishment. This suspicion seems to me to be well-founded, for the size which the larva finally attains exceeds the proportions which the small quantity of honey enclosed in a single cell would lead us to expect.

Let us go back to the pseudochrysalis. It is, as in the Sitares, an inert body, of a horny consistency, amber-coloured and divided into thirteen segments, including the head. Its length is 20 millimetres.[22 - .787 inch. —Translator's Note.] It is slightly curved into an arc, highly convex on the dorsal surface, almost flat on the ventral surface and edged with a projecting fillet which marks the division between the two. The head is only a sort of mask on which certain features are vaguely carved in still relief, corresponding with the future parts of the head. On the thoracic segments are three pairs of tubercles, corresponding with the legs of the recent larva and the future insect. Lastly, there are nine pairs of stigmata, one pair on the mesothorax and the eight following pairs on the first eight segments of the abdomen. The last pair is rather smaller than the rest, a peculiarity which we have already noted in the larva which precedes the pseudochrysalis.

On comparing the pseudochrysalids of the Oil-beetles and Sitares, we observe a most striking similarity between the two. The same structure occurs in both, down to the smallest details. We find on either side the same cephalic masks, the same tubercles occupying the place of the legs, the same distribution and the same number of stigmata and, lastly, the same colour, the same rigidity of the integuments. The only points of difference are in the general appearance, which is not the same in the two pseudochrysalids, and in the covering formed by the cast skin of the late larva. In the Sitares, in fact, this cast skin constitutes a closed bag, a pouch completely enveloping the pseudochrysalis; in the Oil-beetles, on the contrary, it is split down the back and pushed to the rear and, consequently, only half-covers the pseudochrysalis.

The post-mortem examination of the only pseudochrysalis in my possession showed me that, similarly to that which happens in the Sitares, no change occurred in the organization of the viscera, notwithstanding the profound transformations which take place externally. In the midst of innumerable little sacs of adipose tissue is buried a thin thread in which we easily recognize the essential features of the digestive apparatus, both of the preceding larval form and of the perfect insect. As for the medullary cord of the abdomen, it consists, as in the larva, of eight ganglia. In the perfect insect it comprises only four.

I could not say positively how long the Oil-beetle remains in the pseudochrysalid form; but, if we consider the very complete analogy between the evolution of the Oil-beetles and that of the Sitares, there is reason to believe that a few pseudochrysalids complete their transformation in the same year, while others, in greater numbers, remain stationary for a whole year and do not attain the state of the perfect insect until the following spring. This is also the opinion expressed by Newport.

Be this as it may, I found at the end of August one of these pseudochrysalids which had already attained the nymphal stage. It is with the help of this precious capture that I shall be able to finish the story of the Oil-beetle's development. The horny integuments of the pseudochrysalis are split along a fissure which includes the whole ventral surface and the whole of the head and runs up the back of the thorax. This cast skin, which is stiff and keeps its shape, is half-enclosed, as was the pseudochrysalis, in the skin shed by the secondary larva. Lastly, through the fissure, which divides it almost in two, a Meloe-nymph half-emerges; so that, to all appearances, the pseudochrysalis has been followed immediately by the nymph, which does not happen with the Sitares, which pass from the first of these two states to the second only by assuming an intermediary form closely resembling that of the larva which eats the store of honey.

But these appearances are deceptive, for, on removing the nymph from the split sheath formed by the integuments of the pseudochrysalis, we find, at the bottom of this sheath, a third cast skin, the last of those which the creature has so far rejected. This skin is even now adhering to the nymph by a few tracheal filaments. If we soften it in water, we easily recognize that it possesses an organization almost identical with that which preceded the pseudochrysalis. In the latter case only, the mandibles and the legs are not so robust. Thus, after passing through the pseudochrysalid stage, the Oil-beetles for some time resume the preceding form, almost without modification.

The nymph comes next. It presents no peculiarities. The only nymph that I have reared attained the perfect insect state at the end of September. Under ordinary conditions would the adult Oil-beetle have emerged from her cell at this period? I do not think so, since the pairing and egg-laying do not take place until the beginning of spring. She would no doubt have spent the autumn and the winter in the Anthophora's dwelling, only leaving it in the spring following. It is even probable that, as a rule, the development is even slower and that the Oil-beetles, like the Sitares, for the most part spend the cold season in the pseudochrysalid state, a state well-adapted to the winter torpor, and do not achieve their numerous forms until the return of the warm weather.

The Sitares and Meloes belong to the same family, that of the Meloidæ.[23 - Later classifiers place both in the family of the Cantharidæ. —Translator's Note.] Their strange transformations must probably extend throughout the group; indeed, I had the good fortune to discover a third example, which I have not hitherto been able to study in all its details after twenty-five years of investigation. On six occasions, no oftener, during this long period I have set eyes on the pseudochrysalis which I am about to describe. Thrice I obtained it from old Chalicodoma-nests built upon a stone, nests which I at first attributed to the Chalicodoma of the Walls and which I now refer with greater probability to the Chalicodoma of the Sheds. I once extracted it from the galleries bored by some wood-eating larva in the trunk of a dead wild pear-tree, galleries afterwards utilized for the cells of an Osmia, I do not know which. Lastly, I found a pair of them in between the row of cocoons of the Three-pronged Osmia (O. tridentata, DUF.), who provides a home for her larvæ in a channel dug in the dry bramble stems. The insect in question therefore is a parasite of the Osmiæ. When I extract it from the old Chalicodoma-nests, I have to attribute it not to this Bee but to one of the Osmiæ (O. tricornis and O. Latreillii) who, when making their nests, utilize the old galleries of the Mason-bee.

The most nearly complete instances that I have seen furnishes me with the following data: the pseudochrysalis is very closely enveloped in the skin of the secondary larva, a skin consisting of fine transparent pellicle, without any rent whatever. This is the pouch of the Sitaris, save that it lies in immediate contact with the body enclosed. On this jacket we distinguish three pairs of tiny legs, reduced to short vestiges, to stumps. The head is in place, showing quite perceptibly the fine mandibles and the other parts of the mouth. There is no trace of eyes. Each side has a white edging of shrivelled tracheæ, running from one stigmatic orifice to another.

Next comes the pseudochrysalis, horny, currant-red, cylindrical, cone-shaped at both ends, slightly convex on the dorsal surface and concave on the ventral surface. It is covered with delicate, prominent spots, sprinkled very close together; it takes a lens to show them. It is 1 centimetre long and 4 millimetres wide.[24 - .393 x .156 inch. —Translator's Note.] We can distinguish a large knob of a head, on which the mouth is vaguely outlined; three pairs of little shiny brown specks, which are the hardly perceptible vestiges of the legs; and on each side a row of eight black specks, which are the stigmatic orifices. The first speck stands by itself, in front; the seven others, divided from the first by an empty space, form a continuous row. Lastly, at the opposite end is a little pit, the sign of the anal pore.

Of the six pseudochrysalids which a lucky accident placed at my disposal, four were dead; the other two were furnished by Zonitis mutica. This justified my forecast, which from the first, with analogy for my guide, made me attribute these curious organizations to the genus Zonitis. The meloidal parasite of the Osmiæ, therefore, is recognized. We have still to make the acquaintance of the primary larva, which gets itself carried by the Osmia into the cell full of honey, and the tertiary larva, the one which, at a given moment, must be found contained in the pseudochrysalis, a larva which will be succeeded by the nymph.

Let us recapitulate the strange metamorphoses which I have sketched. Every Beetle-larva, before attaining the nymphal stage, undergoes a greater or smaller number of moults, of changes of skin; but these moults, which are intended to favour the development of the larva by ridding it of covering that has become too tight for it, in no way alter its external shape. After any moult that it may have undergone, the larva retains the same characteristics. If it begin by being tough, it will not become tender; if it be equipped with legs, it will not be deprived of them later; if it be provided with ocelli, it will not become blind. It is true that the diet of these non-variable larvæ remains the same throughout their duration, as do the conditions under which they are destined to live.

But suppose that this diet varies, that the environment in which they are called upon to live changes, that the circumstances accompanying their development are liable to great changes: it then becomes evident that the moult may and even must adapt the organization of the larva to these new conditions of existence. The primary larva of the Sitaris lives on the body of the Anthophora. Its perilous peregrinations demand agility of movement, long-sighted eyes and masterly balancing-appliances; it has, in fact, a slender shape, ocelli, legs and special organs adapted to averting a fall. Once inside the Bee's cell, it has to destroy the egg; its sharp mandibles, curved into hooks, will fulfil this office. This done, there is a change of diet: after the Anthophora's egg the larva proceeds to consume the ration of honey. The environment in which it has to live also changes: instead of balancing itself on a hair of the Anthophora, it has now to float on a sticky fluid; instead of living in broad daylight, it has to remain plunged in the profoundest darkness. Its sharp mandibles must therefore become hollowed into a spoon that they may scoop up the honey; its legs, its cirri, its balancing-appliances must disappear as useless and even harmful, since all these organs can only involve the larva in serious danger, by causing it to stick in the honey; its slender shape, its horny integuments, its ocelli, being no longer necessary in a dark cell where movement is impossible, where there are no rough encounters to be feared, may likewise give place to complete blindness, to soft integuments, to a heavy, slothful form. This transfiguration, which everything shows to be indispensable to the life of the larva, is effected by a simple moult.

We do not so plainly perceive the necessity of the subsequent forms, which are so abnormal that nothing like them is known in all the rest of the insect class. The larva which is fed on honey first adopts a false chrysalid appearance and afterwards goes back to its earlier form, though the necessity for these transformations escapes us entirely. Here I am obliged to record the facts and to leave the task of interpreting them to the future. The larva of the Meloidæ, therefore, undergo four moults before attaining the nymphal state; and after each moult their characteristics alter most profoundly. During all these external changes, the internal organization remains unchangingly the same; and it is only at the moment of the nymph's appearance that the nervous system becomes concentrated and that the reproductive organs are developed, absolutely as in the other Beetles.

Thus, to the ordinary metamorphoses which make a Beetle pass successively through the stages of larva, nymph and perfect insect, the Meloidæ add others which repeatedly transform the larva's exterior, without introducing any modification of its viscera. This mode of development, which preludes the customary entomological forms by the multiple transfigurations of the larva, certainly deserves a special name: I suggest that of hypermetamorphosis.

Let us now recapitulate the more prominent facts of this essay.

The Sitares, the Meloes, the Zonites and apparently other Meloidæ, possibly all of them, are in their earliest infancy parasites of the harvesting Bees.

The larva of the Meloidæ, before reaching the nymphal state, passes through four forms, which I call the primary larva, the secondary larva, the pseudochrysalis and the tertiary larva. The passage from one of these forms to the next is effected by a simple moult, without any changes in the viscera.

The primary larva is leathery and settles on the Bee's body. Its object is to get itself carried into a cell filled with honey. On reaching the cell, it devours the Bee's egg; and its part is played.

The secondary larva is soft and differs completely from the primary larva in its external characteristics. It feeds upon the honey contained in the usurped cell.

The pseudochrysalis is a body deprived of all movement and clad in horny integuments which may be compared with those of the pupæ and chrysalids. On these integuments we see a cephalic mask without distinct or movable parts, six tubercles indicating the legs and nine pairs of breathing-holes. In the Sitares the pseudochrysalis is enclosed in a sort of sealed pouch and in the Zonites in a tight-fitting bag formed of the skin of the secondary larva. In the Meloes it is simply half-sheathed in the split skin of the secondary larva.

The tertiary larva reproduces almost exactly the peculiarities of the second; it is enclosed, in the Sitares and probably also the Zonites, in a double vesicular envelope formed of the skin of the secondary larva and the slough of the pseudochrysalis. In the Meloes, it is half-enclosed in the split integuments of the pseudochrysalis, even as these, in their turn, are half-enclosed in the skin of the secondary larva.

From the tertiary larva onwards the metamorphoses follow their habitual course, that is to say, this larva becomes a nymph; and this nymph the perfect insect.

CHAPTER VI

CEROCOMÆ, MYLABRES AND ZONITES

All has not been told concerning the Meloidæ, those strange parasites, some of which, the Sitares and the Oil-beetles, attach themselves, like the tiniest of Lice, to the fleece of various Bees to get themselves carried into the cell where they will destroy the egg and afterwards feed upon the ration of honey. A most unexpected discovery, made a few hundred yards from my door, has warned me once again how dangerous it is to generalize. To take it for granted, as the mass of data hitherto collected seemed to justify us in doing, that all the Meloidæ of our country usurp the stores of honey accumulated by the Bees, was surely a most judicious and natural generalization. Many have accepted it without hesitation; and I for my part was one of them. For on what are we to base our conviction when we imagine that we are stating a law? We think to take our stand upon the general; and we plunge into the quicksands of error. And behold, the law of the Meloidæ has to be struck off the statutes, a fate common to many others, as this chapter will prove.

On the 16th of July, 1883, I was digging, with my son Émile, in the sandy heap where, a few days earlier, I had been observing the labours and the surgery of the Mantis-killing Tachytes. My purpose was to collect a few cocoons of this Digger-wasp. The cocoons were turning up in abundance under my pocket-trowel, when Émile presented me with an unknown object. Absorbed in my task of collection, I slipped the find into my box without examining it further than with a rapid glance. We left the spot. Half-way home, the ardour of my search became assuaged; and a thought of the problematical object, so negligently dropped into the box among the cocoons, flashed across my mind.

"Hullo!" I said to myself. "Suppose it were that? Why not? But, no, yes, it is that; that's just what it is!"

Then, suddenly turning to Émile, who was rather surprised by this soliloquy:

"My boy," I said, "you have had a magnificent find. It's a pseudochrysalis of the Meloidæ. It's a document of incalculable value; you've struck a fresh vein in the extraordinary records of these creatures. Let us look at it closely and at once."

The thing was taken from the box, dusted by blowing on it and carefully examined. I really had before my eyes the pseudochrysalis of some Meloid. Its shape was unfamiliar to me. No matter: I was an old hand and could not mistake its source. Everything assured me that I was on the track of an insect that rivalled the Sitares and the Oil-beetles in the strangeness of its transformations; and, what was a still more precious fact, its occurrence amid the burrows of the Mantis-killer told me that its habits would be wholly different.

"It's very hot, my poor Émile; we are both of us pretty done. Never mind: let's go back to our sand-hill and dig and have another search. I must have the larva that comes before the pseudochrysalis; I must, if possible, have the insect that comes out of it."

Success responded amply to our zeal. We found a goodly number of pseudochrysalids. More often still, we unearthed larvæ which were busy eating the Mantes, the rations of the Tachytes. Are these really the larvæ that turn into the pseudochrysalids? It seems very probable, but there is room for doubt. Rearing them at home will dispel the mists of probability and replace them by the light of certainty. But that is all: I have not a vestige of the perfect insect to inform me of the nature of the parasite. The future, let us hope, will fill this gap. Such was the result of the first trench opened in the heap of sand. Later searches enriched my harvest a little, without furnishing me with fresh data.

Let us now proceed to examine my double find. And first of all the pseudochrysalis, which put me on the alert. It is a motionless, rigid body, of a waxen yellow, smooth, shiny, curved like a fish-hook towards the head, which is inflected. Under a very powerful magnifying-glass the surface is seen to be strewn with very tiny points which are slightly raised and shinier than the surface. There are thirteen segments, including the head. The dorsal surface is convex, the ventral surface flat. A blunt ridge divides the two surfaces. The three thoracic segments bear each a pair of tiny conical nipples, of a deep rusty red, signs of the future legs. The stigmata are very distinct, appearing as specks of a deeper red than the rest of the integuments. There is one pair, the largest, on the second segment of the thorax, almost on the line dividing it from the first segment. Then follow eight pairs, one on each segment of the abdomen except the last, making in all nine pairs of stigmata. The last pair, that of the eighth abdominal segment, is the smallest.

The anal extremity displays no peculiarity. The cephalic mask comprises eight cone-shaped tubercles, dark red like the tubercles of the legs. Six of these are arranged in two lateral rows; the others are between the two rows. In each row of three nipples, the one in the middle is the largest; it no doubt corresponds with the mandibles. The length of this organism varies greatly, fluctuating between 8 and 15 millimetres.[25 - .312 to .585 inch. —Translator's Note.] Its width is from 3 to 4 millimetres.[26 - .117 to .156 inch. —Translator's Note.]