The Glow-Worm and Other Beetles

Let us begin by observing precisely at what spot the eggs are laid. A female has just been fertilized before my eyes; she is forthwith placed in a large glass jar, into which I put, at the same time, some clods of earth containing Anthophora-cells. These cells are occupied partly by larvæ and partly by nymphs that are still quite white; some are slightly open and afford a glimpse of their contents. Lastly, in the inner surface of the cork which closes the jar I sink a cylindrical well, a blind alley, of the same diameter as the corridors of the Anthophora. In order that the insect, if it so desire, may enter this artificial corridor, I lay the bottle horizontally.

The female, painfully dragging her big abdomen, perambulates all the nooks and corners of her makeshift dwelling, exploring them with her palpi, which she passes everywhere. After half an hour of groping and careful investigation, she ends by selecting the horizontal gallery dug in the cork. She thrusts her abdomen into this cavity and, with her head hanging outside, begins her laying. Not until thirty-six hours later was the operation completed; and during this incredible lapse of time the patient creature remained absolutely motionless.

The eggs are white, oval and very small. They measure barely two-thirds of a millimetre11 in length. They stick together slightly and are piled in a shapeless heap which might be likened to a good-sized pinch of the unripe seeds of some orchid. As for their number, I will admit that it tried my patience to no purpose. I do not, however, believe that I am exaggerating when I estimate it as at least two thousand. Here are the data on which I base this figure: the laying, as I have said, lasts thirty-six hours; and my frequent visits to the female working in the cavity in the cork convinced me that there was no perceptible interruption in the successive emission of the eggs. Now less than a minute elapses between the arrival of one egg and that of the next; and the number of these eggs cannot therefore be lower than the number of minutes contained in thirty-six hours, or 2160. But the exact number is of no importance: we need only note that it is very large, which implies, for the young larvæ issuing from the eggs, very numerous chances of destruction, since so lavish a supply of germs is necessary to maintain the species in the requisite proportions.

Enlightened by these observations and informed of the shape, the number and the arrangement of the eggs, I searched the galleries of the Anthophoræ for those which the Sitares had laid there and invariably found them gathered in a heap inside the galleries, at a distance of an inch or two from the orifice, which is always open to the outer world. Thus, contrary to what one was to some extent entitled to suppose, the eggs are not laid in the cells of the pioneer Bee; they are simply dumped in a heap inside the entrance to her dwelling. Nay more, the mother does not make any protective structure for them; she takes no pains to shield them from the rigours of winter; she does not even attempt, by stopping for a short distance, as best she can, the entrance-lobby in which she has laid them, to protect them from the thousand enemies that threaten them; for, as long as the frosts of winter have not arrived, these open galleries are trodden by Spiders, by Acari, by Anthrenus-grubs and other plunderers, to whom these eggs, or the young larvæ about to emerge from them, must be a dainty feast. In consequence of the mother's heedlessness, the number of those who escape all these voracious hunters and the inclemencies of the weather must be curiously small. This perhaps explains why she is compelled to make up by her fecundity for her deficient industry.

The hatching occurs a month later, about the end of September or the beginning of October. The season being still propitious, I was led to suppose that the young larvæ must at once make a start and disperse, in order that each might seek to gain access, through some imperceptible fissure, to an Anthophora-cell. This presumption turned out to be entirely at fault. In the boxes in which I had placed the eggs laid by my captives, the young larvæ, little black creatures at most a twenty-fifth of an inch long, did not move away, provided though they were with vigorous legs; they remained higgledy-piggledy with the white skins of the eggs whence they had emerged.

In vain I placed within their reach lumps of earth containing nests of the Anthophora, open cells, larvæ and nymphs of the Bee: nothing was able to tempt them; they persisted in forming, with the egg-skins, a powdery heap of speckled black and white. It was only by drawing the point of a needle through this pinch of living dust that I was able to provoke an active wriggling. Apart from this, all was still. If I forcibly removed a few larvæ from the common heap, they at once hurried back to it, in order to hide themselves among the rest. Perhaps they had less reason to fear the cold when thus collected and sheltered beneath the egg-skins. Whatever may be the motive that impels them to remain thus gathered in a heap, I recognized that none of the means suggested by my imagination succeeded in forcing them to abandon the little spongy mass formed by the skins of the eggs, which were slightly glued together. Lastly, to assure myself that the larvæ, in the free state, do not disperse after they are hatched, I went during the winter to Carpentras and inspected the banks inhabited by the Anthophoræ. There, as in my boxes, I found the larvæ piled into heaps, all mixed up with the skins of the eggs.

CHAPTER III

THE PRIMARY LARVA OF THE SITARES

Nothing new happens before the end of the following April. I shall profit by this long period of repose to tell you more about the young larva, of which I will begin by giving a description. Its length is a twenty-fifth of an inch, or a little less. It is hard as leather, a glossy greenish black, convex above and flat below, long and slender, with a diameter increasing gradually from the head to the hinder extremity of the metathorax, after which it rapidly diminishes. Its head is a trifle longer than it is wide and is slightly dilated at the base; it is pale-red near the mouth and darker about the ocelli.

The labrum forms a segment of a circle; it is reddish, edged with a small number of very short, stiff hairs. The mandibles are powerful, red-brown, curved and sharp; when at rest they meet without crossing. The maxillary palpi are rather long, consisting of two cylindrical sections of equal length, the outer ending in a very short bristle. The jaws and the lower lip are not sufficiently visible to lend themselves to accurate description.

The antennæ consist of two cylindrical segments, equal in length, not very definitely divided; these segments are nearly as long as those of the palpi; the outer is surmounted by a cirrus whose length is as much as thrice that of the head and tapers off until it becomes invisible under a powerful pocket-lens. Behind the base of either antennæ are two ocelli, unequal in size and almost touching.

The thoracic segments are of equal length and increase gradually in width from front to back. The prothorax is wider than the head, but is narrower in front than at the base and is slightly rounded at the sides. The legs are of medium length and fairly robust, ending in a long, powerful, sharp and very mobile claw. On the haunch and thigh of each leg is a long cirrus, like that of the antennæ, almost as long as the whole limb and standing at right angles to the plane of locomotion when the creature moves. There are a few stiff bristles on the legs.

The abdomen has nine segments, of practically equal length, but shorter than those of the thorax and diminishing very rapidly in width toward the last. Fixed below the eighth segment, or rather below the strip of membrane separating this segment and the last, we see two spikes, slightly curved, short, but with strong, sharp, hard points, and placed one to the right and the other to the left of the median line. These two appendages are able, by means of a mechanism recalling, on a smaller scale, that of the Snail's horns, to withdraw into themselves, as a result of the membranous character of their base. They can also retreat under the eighth segment, borne, as they are, by the anal segment, when this last, as it contracts, withdraws into the eighth. Lastly, the ninth or anal segment bears on its hinder edge two long cirri, like those of the legs and the antennæ, curving backwards from tip to base. At the rear of this segment a fleshy nipple appears, more or less prominent; this is the anus. I do not know where the stigmata are placed; they have evaded my investigations, though these were undertaken with the aid of the microscope.

When the larva is at rest, the various segments overlap evenly; and the membranous intervals, corresponding with the articulations, do not show. But, when the larva walks, all the articulations, especially those of the abdominal segments, are distended and end by occupying almost as much space as the horny arches. At the same time the anal segment emerges from the sheath formed by the eighth; the anus, in turn, is stretched into a nipple; and the two points of the penultimate ring rise, at first slowly, and then suddenly stand up with an abrupt motion similar to that of a spring when released. In the end, these two points diverge like the horns of a crescent. Once this complex apparatus is unfolded, the tiny creature is ready to crawl upon the most slippery surface.

The last segment and its anal button are curved at right angles to the axis of the body; and the anus comes and presses upon the surface of locomotion, where it ejects a tiny drop of transparent, treacly fluid, which glues and holds the little creature firmly in position, supported on a sort of tripod formed by the anal button and the two cirri of the last segment. If we are observing the animal's manner of locomotion on a strip of glass, we can hold the strip in a vertical position, or even turn it upside down, or shake it lightly, without causing the larva to become detached and fall, held fast as it is by the glutinous secretion of the anal button.

If it has to proceed along a surface where there is no danger of a fall, the microscopic creature employs another method. It crooks its belly and, when the two spikes of the eighth segment, now fully outspread, have found a firm support by ploughing, so to speak, the surface of locomotion, it bears upon that base and pushes forward by expanding the various abdominal articulations. This forward movement is also assisted by the action of the legs, which are far from remaining inactive. This done, it casts anchor with the powerful claws of its feet; the abdomen contracts; the various segments draw together; and the anus, pulled forward, obtains a fresh purchase, with the aid of the two spikes, before beginning the second of these curious strides.

During these manoeuvres, the cirri of the flanks and thighs drag along the supporting surface and by their length and elasticity appear fitted only to impede progress. But let us not be in a hurry to conclude that we have discovered an inconsistency: the least of creatures is adapted to the conditions amid which it has to live; there is reason to believe that these filaments, far from hampering the pigmy's progress, must, in normal circumstances, be of some assistance to it.

Even the little that we have just learnt shows us that the young Sitaris-larva is not called upon to move on an ordinary surface. The spot, whatever it may be like, where this larva is to live later exposes it to the risk of many dangerous falls, since, in order to prevent them, it is not only equipped with strong and extremely mobile talons and a steel-shod crescent, a sort of ploughshare capable of biting into the most highly polished substance, but is further provided with a viscous liquid, sufficiently tenacious and adhesive to hold it in position without the help of other appliances. In vain I racked my brains to guess what the substance might be, so shifting, so uncertain and so perilous, which the young Sitares are destined to inhabit; and I discovered nothing to explain the necessity for the structure which I have described. Convinced beforehand, by an attentive examination of this structure, that I should witness some peculiar habits, I waited with eager impatience for the return of the warm weather, never doubting that by dint of persevering observation the mystery would be disclosed to me next spring. At last this spring, so fervently desired, arrived; I brought to bear all the patience, all the imagination, all the insight and discernment that I may possess; but, to my utter shame and still greater regret, the secret escaped me. Oh, how painful are those tortures of indecision, when one has to postpone till the following year an investigation which has led to no result!

My observations made during the spring of 1856, although purely negative, nevertheless have an interest of their own, because they prove the inaccuracy of certain suppositions to which the undeniable parasitism of the Sitares naturally inclines us. I will therefore relate them in a few words. At the end of April, the young larvæ, hitherto motionless and concealed in the spongy heap of the egg-skins, emerge from their immobility, scatter and run about in all directions through the boxes and jars in which they have passed the winter. By their hurried gait and their indefatigable evolutions we readily guess that they are seeking something which they lack. What can this something be, unless it be food? For remember that these larvæ were hatched at the end of September and that since then, that is to say, for seven long months, they have taken no nourishment, though they have spent this period in the full enjoyment of their vitality, as I was able to assure myself all through the winter by irritating them, and not in a state of torpor similar to that of the hibernating animals. From the moment of their hatching they are doomed, although full of life, to an absolute abstinence of seven months' duration; and it is natural to suppose, when we see their present excitement, that an imperious hunger sets them bustling in this fashion.

The desired nourishment could only be the contents of the cells of the Anthophora, since we afterwards find the Sitares in these cells. Now these contents are limited to honey or larvæ. It just happens that I have kept some Anthophora-cells occupied by larvæ or nymphs. I place a few of these, some open, some closed, within reach of the young Sitares, as I had already done directly after the hatching. I even slip the Sitares into the cells: I place them on the sides of the larva, a succulent morsel to all appearances; I do all sorts of things to tempt their appetite; and, after exhausting my ingenuity, which continues fruitless, I remain convinced that my famished grubs are seeking neither the larvæ nor nymphs of the Anthophora.

Let us now try honey. We must obviously employ honey prepared by the same species of Anthophora as that at whose cost the Sitares live. But this Bee is not very common in the neighbourhood of Avignon; and my engagements at the college12 do not allow me to absent myself for the purpose of repairing to Carpentras, where she is so abundant. In hunting for cells provisioned with honey I thus lose a good part of the month of May; however, I end by finding some which are newly sealed and which belong to the right Anthophora. I open these cells with the feverish impatience of a sorely-tried longing. All goes well: they are half-full of fluid, dark, nauseating honey, with the Bee's lately-hatched larva floating on the surface. This larva is removed; and taking a thousand precautions, I lay one or more Sitares on the surface of the honey. In other cells I leave the Bee's larva and insert Sitares, placing them sometimes on the honey and sometimes on the inner wall of the cell or simply at the entrance. Lastly, all the cells thus prepared are put in glass tubes, which enable me to observe them readily, without fear of disturbing my famished guests at their meal.

But what am I saying? Their meal? There is no meal! The Sitares, placed at the entrance to a cell, far from seeking to make their way in, leave it and go roaming about the glass tube; those which have been placed on the inner surface of the cells, near the honey, emerge precipitately, half-caught in the glue and tripping at every step; lastly, those which I thought I had favoured the most, by placing them on the honey itself, struggle, become entangled in the sticky mass and perish in it, suffocated. Never did experiment break down so completely! Larvæ, nymphs, cells, honey: I have offered you them all! Then what do you want, you fiendish little creatures?

Tired of all these fruitless attempts, I ended where I ought to have begun: I went to Carpentras. But it was too late: the Anthophora had finished her work; and I did not succeed in seeing anything new. During the course of the year I learnt from Léon Dufour,13 to whom I had spoken of the Sitares, that the tiny creature which he had found on the Andrenæ14 and described under the generic name of Triungulinus, was recognized later by Newport15 as the larva of a Meloe, or Oil-beetle. Now it so happened that I had found a few Oil-beetles in the cells of the same Anthophora that nourishes the Sitares. Could there be a similarity of habits between the two kinds of insects? This idea threw a sudden light for me upon the subject; but I had plenty of time in which to mature my plans: I had another year to wait.

When April came, my Sitaris-larvæ began, as usual, to bestir themselves. The first Bee to appear, an Osmia, is dropped alive into a glass jar containing a few of these larvæ; and after a lapse of some fifteen minutes I inspect them through the pocket-lens. Five Sitares are embedded in the fleece of the thorax. It is done, the problem's solved! The larvæ of the Sitares, like those of the Oil-beetles, cling like grim death to the fleece of their generous host and make him carry them into the cell. Ten times over I repeat the experiment with the various Bees that come to plunder the lilac flowering outside my window and in particular with male Anthophoræ; the result is still the same: the larvæ embed themselves in the hair of the Bees' thorax. But after so many disappointments one becomes distrustful and it is better to go and observe the facts upon the spot; besides, the Easter holidays fall very conveniently and afford me the leisure for my observations.

I will admit that my heart was beating a little faster than usual when I found myself once again standing in front of the perpendicular bank in which the Anthophora nests. What will be the result of the experiment? Will it once more cover me with confusion? The weather is cold and rainy; not a Bee shows herself on the few spring flowers that have come out. Numbers of Anthophoræ cower, numbed and motionless, at the entrance to the galleries. With the tweezers, I extract them one by one from their lurking-places, to examine them under the lens. The first has Sitaris-larvæ on her thorax; so has the second; the third and fourth likewise; and so on, as far as I care to pursue the examination. I change galleries ten times, twenty times; the result is invariable. Then, for me, occurred one of the moments which come to those who, after considering and reconsidering an idea for years and years from every point of view, are at last able to cry: "Eureka!"

On the days that followed, a serene and balmy sky enabled the Anthophoræ to leave their retreats and scatter over the countryside and despoil the flowers. I renewed my examination on those Anthophoræ flying incessantly from one flower to another, whether in the neighbourhood of the places where they were born or at great distances from these places. Some were without Sitaris-larvæ; others, more numerous, had two, three, four, five or more among the hairs of their thorax. At Avignon, where I have not yet seen Sitaris humeralis, the same species of Anthophora, observed at almost the same season, while pillaging the lilac-blossom, was always free of young Sitaris-grubs; at Carpentras, on the contrary, where there is not a single Anthophora-colony without Sitares, nearly three-quarters of the specimens which I examined carried a few of these larvæ in their fleece.

But, on the other hand, if we look for these larvæ in the entrance-lobbies where we found them, a few days ago, piled up in heaps, we no longer see them. Consequently, when the Anthophoræ, having opened their cells, enter the galleries to reach the exit and fly away, or else when the bad weather and the darkness bring them back there for a time, the young Sitaris-larvæ, kept on the alert in these same galleries by the stimulus of instinct, attach themselves to the Bees, wriggling into their fur and clutching it so firmly that they need not fear a fall during the long journeys of the insect which carries them. By thus attaching themselves to the Anthophoræ the young Sitares evidently intend to get themselves carried, at the opportune moment, into the victualled cells.

One might even at first sight believe that they live for some time on the Anthophora's body, just as the ordinary parasites, the various species of Lice, live on the body of the animal that feeds them. But not at all. The young Sitares, embedded in the fleece, at right angles to the Anthophora's body, head inwards, rump outwards, do not stir from the point which they have selected, a point near the Bee's shoulders. We do not see them wandering from spot to spot, exploring the Anthophora's body, seeking the part where the skin is more delicate, as they would certainly do if they were really deriving some nourishment from the juices of the Bee. On the contrary, they are nearly always established on the toughest and hardest part of the Bee's body, on the thorax, a little below the insertion of the wings, or, more rarely, on the head; and they remain absolutely motionless, fixed to the same hair, by means of the mandibles, the feet, the closed crescent of the eighth segment and, lastly, the glue of the anal button. If they chance to be disturbed in this position, they reluctantly repair to another point of the thorax, pushing their way through the insect's fur and in the end fastening on to another hair, as before.

To confirm my conviction that the young Sitaris-grubs do not feed on the Anthophora's body, I have sometimes placed within their reach, in a glass jar, some Bees that have long been dead and are completely dried up. On these dry corpses, fit at most for gnawing, but certainly containing nothing to suck, the Sitaris-larvæ took up their customary position and there remained motionless as on the living insect. They obtain nothing, therefore, from the Anthophora's body; but perhaps they nibble her fleece, even as the Bird-lice nibble the birds' feathers?

To do this, they would require mouth-parts endowed with a certain strength and, in particular, horny and sturdy jaws, whereas their jaws are so fine that a microscopic examination failed to show them to me. The larvæ, it is true, are provided with powerful mandibles; but these finely-pointed mandibles, with their backward curve, though excellent for tugging at food and tearing it to pieces, are useless for grinding it or gnawing it. Lastly, we have a final proof of the passive condition of the Sitaris-larvæ on the body of the Anthophoræ in the fact that the Bees do not appear to be in any way incommoded by their presence, since we do not see them trying to rid themselves of the grubs. Some Anthophoræ which were free from these grubs and some others which were carrying five or six upon their bodies were placed separately in glass jars. When the first disturbance resulting from their captivity was appeased, I could see nothing peculiar about those occupied by the young Sitares. And, if all these arguments were not sufficient, I might add that a creature which has already been able to spend seven months without food and which in a few days' time will proceed to drink a highly-flavoured fluid would be guilty of a singular inconsistency if it were to start nibbling the dry fleece of a Bee. It therefore seems to me undeniable that the young Sitares settle on the Anthophora's body merely to make her carry them into the cells which she will soon be building.

But until then the future parasites must hold tight to the fleece of their hostess, despite her rapid evolutions among the flowers, despite her rubbing against the walls of the galleries when she enters to take shelter and, above all, despite the brushing which she must often give herself with her feet to dust herself and keep spick and span. Hence no doubt the need for that curious apparatus which no standing or moving upon ordinary surfaces could explain, as was said above, when we were wondering what the shifting, swaying, dangerous body might be on which the larva would have to establish itself later. This body is a hair of a Bee who makes a thousand rapid journeys, now diving into her narrow galleries, now forcing her way down the tight throat of a corolla, and who never rests except to brush herself with her feet and remove the specks of dust collected by the down which covers her.