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CHAPTER IV
THE PRIMARY LARVA OF THE OIL-BEETLES

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I interrupt the history of the Sitares to speak of the Meloes, those uncouth Beetles, with their clumsy belly and their limp wing-cases yawning over their back like the tails of a fat man's coat that is far too tight for its wearer. The insect is ugly in colouring, which is black, with an occasional blue gleam, and uglier still in shape and gait; and its disgusting method of defence increases the repugnance with which it inspires us. If it judges itself to be in danger, the Meloe resorts to spontaneous bleeding. From its joints a yellowish, oily fluid oozes, which stains your fingers and makes them stink. This is the creature's blood. The English, because of its trick of discharging oily blood when on the defensive, call this insect the Oil-beetle. It would not be a particularly interesting Beetle save for its metamorphoses and the peregrinations of its larva, which are similar in every respect to those of the larva of the Sitares. In their first form, the Oil-beetles are parasites of the Anthophoræ; their tiny grub, when it leaves the egg, has itself carried into the cell by the Bee whose victuals are to form its food.

Observed in the down of various Bees, the queer little creature for a long time baffled the sagacity of the naturalists, who, mistaking its true origin, made it a species of a special family of wingless insects. It was the Bee-louse (Pediculus apis) of Linnæus;17 the Triungulin of the Andrenæ (Triungulinus andrenetarum) of Léon Dufour. They saw in it a parasite, a sort of Louse, living in the fleece of the honey-gatherers. It was reserved for the distinguished English naturalist Newport to show that this supposed Louse was the first state of the Oil-beetles. Some observations of my own will fill a few lacunæ in the English scientist's monograph. I will therefore sketch the evolution of the Oil-beetles, using Newport's work where my own observations are defective. In this way the Sitares and the Meloes, alike in habits and transformations, will be compared; and the comparison will throw a certain light upon the strange metamorphoses of these insects.

The same Mason-bee (Anthophora pilipes) upon whom the Sitares live also feeds a few scarce Meloes (M. cicatricosus) in its cells. A second Anthophora of my district (A. parietina) is more subject to this parasite's invasions. It was also in the nests of an Anthophora, but of a different species (A. retusa), that Newport observed the same Oil-beetle. These three lodgings adopted by Meloe cicatricosus may be of some slight interest, as leading us to suspect that each species of Meloe is apparently the parasite of diverse Bees, a suspicion which will be confirmed when we examine the manner in which the larvæ reach the cell full of honey. The Sitares, though less given to change of lodging, are likewise able to inhabit nests of different species. They are very common in the cells of Anthophora pilipes; but I have found them also, in very small numbers, it is true, in the cells of A. personata.

Despite the presence of Meloe cicatricosus in the dwellings of the Mason-bee, which I so often ransacked in compiling the history of the Sitares, I never saw this insect, at any season of the year, wandering on the perpendicular soil, at the entrance of the corridors, for the purpose of laying its eggs there, as the Sitares do; and I should know nothing of the details of the egg-laying if Godart,18 de Geer19 and, above all, Newport had not informed us that the Oil-beetles lay their eggs in the earth. According to the last-named author, the various Oil-beetles whom he had the opportunity of observing dig, among the roots of a clump of grass, in a dry soil exposed to the sun, a hole a couple of inches deep which they carefully fill up after laying their eggs there in a heap. This laying is repeated three or four times over, at intervals of a few days during the same season. For each batch of eggs the female digs a special hole, which she does not fail to fill up afterwards. This takes place in April and May.

The number of eggs laid in a single batch is really prodigious. In the first batch, which, it is true, is the most prolific of all, Meloe proscarabæus, according to Newport's calculations, produces the astonishing number of 4,218 eggs, which is double the number of eggs laid by a Sitaris. And what must the number be, when we allow for the two or three batches that follow the first! The Sitares, entrusting their eggs to the very corridors through which the Anthophora is bound to pass, spare their larvæ a host of dangers which the larvæ of the Meloe have to run, for these, born far from the dwellings of the Bees, are obliged to make their own way to their hymenopterous foster-parents. The Oil-beetles, therefore, lacking the instinct of the Sitares, are endowed with incomparably greater fecundity. The richness of their ovaries atones for the insufficiency of instinct by proportioning the number of germs in accordance with the risks of destruction. What transcendent harmony is this, which thus holds the scales between the fecundity of the ovaries and the perfection of instinct!

The hatching of the eggs takes place at the end of May or in June, about a month after they are laid. The eggs of the Sitares also are hatched after the same lapse of time. But the Meloe-larvæ, more greatly favoured, are able to set off immediately in search of the Bees that are to feed them; while those of the Sitares, hatched in September, have to wait motionless and in complete abstinence for the emergence of the Anthophoræ the entrance to whose cells they guard. I will not describe the young Meloe-larva, which is sufficiently well known, in particular by the description and the diagram furnished by Newport. To enable the reader to understand what follows, I will confine myself to stating that this primary larva is a sort of little yellow louse, long and slender, found in the spring in the down of different Bees.

How has this tiny creature made its way from the underground lodging where the eggs are hatched to the fleece of a Bee? Newport suspects that the young Oil-beetles, on emerging from their natal burrow, climb upon the neighbouring plants, especially upon the Cichoriceæ, and wait, concealed among the petals, until a few Bees chance to plunder the flower, when they promptly fasten on to their fur and allow themselves to be borne away by them. I have more than Newport's suspicions upon this curious point; my personal observations and experiments are absolutely convincing. I will relate them as the first phase of the history of the Bee-louse. They date back to the 23rd of May, 1858.

A vertical bank on the road from Carpentras to Bédoin is this time the scene of my observations. This bank, baked by the sun, is exploited by numerous swarms of Anthophoræ, who, more industrious than their congeners, are in the habit of building, at the entrance to their corridors, with serpentine fillets of earth, a vestibule, a defensive bastion in the form of an arched cylinder. In a word, they are swarms of A. parietina. A sparse carpet of turf extends from the edge of the road to the foot of the bank. The more comfortably to follow the work of the Bees, in the hope of wresting some secret from them, I had been lying for a few moments upon this turf, in the very heart of the inoffensive swarm, when my clothes were invaded by legions of little yellow lice, running with desperate eagerness through the hairy thickets of the nap of the cloth. In these tiny creatures, with which I was powdered here and there as with yellow dust, I soon recognized an old acquaintance, the young Oil-beetles, whom I now saw for the first time elsewhere than in the Bees' fur or the interior of their cells. I could not lose so excellent an opportunity of learning how these larvæ manage to establish themselves upon the bodies of their foster-parents.

In the grass where, after lying down for a moment, I had caught these lice were a few plants in blossom, of which the most abundant were three composites: Hedypnois polymorpha, Senecio gallicus and Anthemis arvensis. Now it was on a composite, a dandelion, that Newport seemed to remember seeing some young Oil-beetles; and my attention therefore was first of all directed to the plants which I have named. To my great satisfaction, nearly all the flowers of these three plants, especially those of the camomile (Anthemis) were occupied by young Oil-beetles in greater or lesser numbers. On one head of camomile I counted forty of these tiny insects, cowering motionless in the centre of the florets. On the other hand, I could not discover any on the flowers of the poppy or of a wild rocket (Diplotaxis muralis) which grew promiscuously among the plants aforesaid. It seems to me, therefore, that it is only on the composite flowers that the Meloe-larvæ await the Bees' arrival.

In addition to this population encamped upon the heads of the composites and remaining motionless, as though it had achieved its object for the moment, I soon discovered yet another, far more numerous, whose anxious activity betrayed a fruitless search. On the ground, in the grass, numberless little larvæ were running in a great flutter, recalling in some respects the tumultuous disorder of an overturned Ant-hill; others were hurriedly climbing to the tip of a blade of grass and descending with the same haste; others again were plunging into the downy fluff of the withered everlastings, remaining there a moment and quickly reappearing to continue their search. Lastly, with a little attention, I was able to convince myself that within an area of a dozen square yards there was perhaps not a single blade of grass which was not explored by several of these larvæ.

I was evidently witnessing the recent emergence of the young Oil-beetles from their maternal lairs. Part of them had already settled on the groundsel- and camomile-flowers to await the arrival of the Bees; but the majority were still wandering in search of this provisional refuge. It was by this wandering population that I had been invaded when I lay down at the foot of the bank. It was impossible that all these larvæ, the tale of whose alarming thousands I would not venture to define, should form one family and recognize a common mother; despite what Newport has told us of the Oil-beetles' astonishing fecundity, I could not believe this, so great was their multitude.

Though the green carpet was continued for a considerable distance along the side of the road, I could not detect a single Meloe-larva elsewhere than in the few square yards lying in front of the bank inhabited by the Mason-bee. These larvæ therefore could not have come far; to find themselves near the Anthophoræ they had had no long pilgrimage to make, for there was not a sign of the inevitable stragglers and laggards that follow in the wake of a travelling caravan. The burrows in which the eggs were hatched were therefore in that turf opposite the Bees' abode. Thus the Oil-beetles, far from laying their eggs at random, as their wandering life might lead one to suppose, and leaving their young to the task of approaching their future home, are able to recognize the spots haunted by the Anthophoræ and lay their eggs in the near neighbourhood of those spots.

With such a multitude of parasites occupying the composite flowers in close proximity to the Anthophora's nests, it is impossible that the majority of the swarm should not become infested sooner or later. At the time of my observations, a comparatively tiny proportion of the starving legion was waiting on the flowers; the others were still wandering on the ground, where the Anthophoræ very rarely alight; and yet I detected the presence of several Meloe-larvæ in the thoracic down of nearly all the Anthophoræ which I caught and examined.

I have also found them on the bodies of the Melecta- and Coelioxys-bees,20 who are parasitic on the Anthophoræ. Suspending their audacious patrolling before the galleries under construction, these spoilers of the victualled cells alight for an instant on a camomile-flower and lo, the thief is robbed! A tiny, imperceptible louse has slipped into the thick of the downy fur and, at the moment when the parasite, after destroying the Anthophora's egg, is laying her own upon the stolen honey, will creep upon this egg, destroy it in its turn and remain sole mistress of the provisions. The mess of honey amassed by the Anthophora will thus pass through the hands of three owners and remain finally the property of the weakest of the three.

And who shall say whether the Meloe, in its turn, will not be dispossessed by a fresh thief; or even whether it will not, in the state of a drowsy, fat and flabby larva, fall a prey to some marauder who will munch its live entrails? As we meditate upon this deadly, implacable struggle which nature imposes, for their preservation, on these different creatures, which are by turns possessors and dispossessed, devourers and devoured, a painful impression mingles with the wonder aroused by the means employed by each parasite to attain its end; and, forgetting for a moment the tiny world in which these things happen, we are seized with terror at this concatenation of larceny, cunning and brigandage which forms part, alas, of the designs of alma parens rerum!

The young Meloe-larvæ established in the down of the Anthophoræ or in that of the Melecta- and the Coelioxys-bees, their parasites, had adopted an infallible means of sooner or later reaching the desired cell. Was it, so far as they were concerned, a choice dictated by the foresight of instinct, or just simply the result of a lucky chance? The question was soon decided. Various Flies – Drone-flies and Bluebottles (Eristalis tenax and Calliphora vomitoria) – would settle from time to time on the groundsel- or camomile-flowers occupied by the young Meloes and stop for a moment to suck the sweet secretions. On all these Flies, with very few exceptions, I found Meloe-larvæ, motionless in the silky down of the thorax. I may also mention, as infested by these larvæ, an Ammophila (A. hirsuta),21 who victuals her burrows with a caterpillar in early spring, while her kinswomen build their nests in autumn. This Wasp merely grazes, so to speak, the surface of a flower; I catch her; there are Meloes moving about her body. It is clear that neither the Drone-flies nor the Bluebottles, whose larvæ live in putrefying matter, nor yet the Ammophilæ who victual theirs with caterpillars, could ever have carried the larvæ which invaded them into cells filled with honey. These larvæ therefore had gone astray; and instinct, as does not often happen, was here at fault.

Let us now turn our attention to the young Meloes waiting expectant upon the camomile-flowers. There they are, ten, fifteen or more, lodged half-way down the florets of a single blossom or in their interstices; it therefore needs a certain degree of scrutiny to perceive them, their hiding-place being the more effectual in that the amber colour of their bodies merges in the yellow hue of the florets. So long as nothing unusual happens upon the flower, so long as no sudden shock announces the arrival of a strange visitor, the Meloes remain absolutely motionless and give no sign of life. To see them dipping vertically, head downwards, into the florets, one might suppose that they were seeking some sweet liquid, their food; but in that case they ought to pass more frequently from one floret to another, which they do not, except when, after a false alarm, they regain their hiding-places and choose the spot which seems to them the most favourable. This immobility means that the florets of the camomile serve them only as a place of ambush, even as later the Anthophora's body will serve them solely as a vehicle to convey them to the Bee's cell. They take no nourishment, either on the flowers or on the Bees; and, as with the Sitares, their first meal will consist of the Anthophora's egg, which the hooks of their mandibles are intended to rip open.

Their immobility is, as we have said, complete; but nothing is easier than to arouse their suspended activity. Shake a camomile-blossom lightly with a bit of straw: instantly the Meloes leave their hiding-places, come up and scatter in all directions on the white petals of the circumference, running over them from one end to the other with all the speed which the smallness of their size permits. On reaching the extreme end of the petals, they fasten to it either with their caudal appendages, or perhaps with a sticky substance similar to that furnished by the anal button of the Sitares; and, with their bodies hanging outside and their six legs free, they bend about in every direction and stretch as far out as they can, as though striving to touch an object out of their reach. If nothing offers for them to seize upon, after a few vain attempts they regain the centre of the flower and soon resume their immobility.

But, if we place near them any object whatever, they do not fail to catch on to it with surprising agility. A blade of grass, a bit of straw, the handle of my tweezers which I hold out to them: they accept anything in their eagerness to quit the provisional shelter of the flower. It is true that, after finding themselves on these inanimate objects, they soon recognize that they have gone astray, as we see by their bustling movements to and fro and their tendency to go back to the flower if there still be time. Those which have thus giddily flung themselves upon a bit of straw and are allowed to return to their flower do not readily fall a second time into the same trap. There is therefore, in these animated specks, a memory, an experience of things.

After these experiments I tried others with hairy materials imitating more or less closely the down of the Bees, with little pieces of cloth or velvet cut from my clothes, with plugs of cotton wool, with pellets of flock gathered from the everlastings. Upon all these objects, offered with the tweezers, the Meloes flung themselves without any difficulty; but, instead of keeping quiet, as they do on the bodies of the Bees, they soon convinced me, by their restless behaviour, that they found themselves as much out of their element on these furry materials as on the smooth surface of a bit of straw. I ought to have expected this: had I not just seen them wandering without pause upon the everlastings enveloped with cottony flock? If reaching the shelter of a downy surface were enough to make them believe themselves safe in harbour, nearly all would perish, without further attempts, in the down of the plants.

Let us now offer them live insects and, first of all, Anthophoræ. If the Bee, after we have rid her of the parasites which she may be carrying, be taken by the wings and held for a moment in contact with the flower, we invariably find her, after this rapid contact, overrun by Meloes clinging to her hairs. The larvæ nimbly take up their position on the thorax, usually on the shoulders or sides, and once there they remain motionless: the second stage of their strange journey is compassed.

After the Anthophoræ, I tried the first live insects that I was able to procure at once: Drone-flies, Bluebottles, Hive-bees, small Butterflies. All were alike overrun by the Meloes, without hesitation. What is more, there was no attempt made to return to the flowers. As I could not find any Beetles at the moment, I was unable to experiment with them. Newport, experimenting, it is true, under conditions very different from mine, since his observations related to young Meloes held captive in a glass jar, while mine were made in the normal circumstances, Newport, I was saying, saw Meloes fasten to the body of a Malachius and stay there without moving, which inclines me to believe that with Beetles I should have obtained the same results as, for instance, with a Drone-fly. And I did, in fact, at a later date, find some Meloe-larvæ on the body of a big Beetle, the Golden Rose-chafer (Cetonia aurata), an assiduous visitor of the flowers.

After exhausting the insect class, I put within their reach my last resource, a large black Spider. Without hesitation they passed from the flower to the arachnid, made for places near the joints of the legs and settled there without moving. Everything therefore seems to suit their plans for leaving the provisional abode where they are waiting; without distinction of species, genus, or class, they fasten to the first living creature that chance brings within their reach. We now understand how it is that these young larvæ have been observed upon a host of different insects and especially upon the early Flies and Bees pillaging the flowers; we can also understand the need for that prodigious number of eggs laid by a single Oil-beetle, since the vast majority of the larvæ which come out of them will infallibly go astray and will not succeed in reaching the cells of the Anthophoræ. Instinct is at fault here; and fecundity makes up for it.

But instinct recovers its infallibility in another case. The Meloes, as we have seen, pass without difficulty from the flower to the objects within their reach, whatever these may be, smooth or hairy, living or inanimate. This done, they behave very differently, according as they have chanced to invade the body of an insect or some other object. In the first case, on a downy Fly or Butterfly, on a smooth-skinned Spider or Beetle, the larvæ remain motionless after reaching the point which suits them. Their instinctive desire is therefore satisfied. In the second case, in the midst of the nap of cloth or velvet, or the filaments of cotton, or the flock of the everlasting, or, lastly, on the smooth surface of a leaf or a straw, they betray the knowledge of their mistake by their continual coming and going, by their efforts to return to the flower imprudently abandoned.

How then do they recognize the nature of the object to which they have just moved? How is it that this object, whatever the quality of its surface, will sometimes suit them and sometimes not? Do they judge their new lodging by sight? But then no mistake would be possible; the sense of sight would tell them at the outset whether the object within reach was suitable or not; and emigration would or would not take place according to its decision. And then how can we suppose that, buried in the dense thicket of a pellet of cotton-wool or in the fleece of an Anthophora, the imperceptible larva can recognize, by sight, the enormous mass which it is perambulating?

Is it by touch, by some sensation due to the inner vibrations of living flesh? Not so, for the Meloes remain motionless on insect corpses that have dried up completely, on dead Anthophoræ taken from cells at least a year old. I have seen them keep absolutely quiet on fragments of an Anthophora on a thorax long since nibbled and emptied by the Mites. By what sense then can they distinguish the thorax of an Anthophora from a velvety pellet, when sight and touch are out of the question? The sense of smell remains. But in that case what exquisite subtlety must we not take for granted? Moreover, what similarity of smell can we admit between all the insects which, dead or alive, whole or in pieces, fresh or dried, suit the Meloes, while anything else does not suit them? A wretched louse, a living speck, leaves us mightily perplexed as to the sensibility which directs it. Here is yet one more riddle added to all the others.

After the observations which I have described, it remained for me to search the earthen surface inhabited by the Anthophoræ: I should then have followed the Meloe-larva in its transformations. It was certainly cicatricosus whose larvæ I had been studying; it was certainly this insect which ravaged the cells of the Mason-bee, for I found it dead in the old galleries which it had been unable to leave. This opportunity, which did not occur again, promised me an ample harvest. I had to give it all up. My Thursday was drawing to a close; I had to return to Avignon, to resume my lessons on the electrophorus and the Toricellian tube. O happy Thursdays! What glorious opportunities I lost because you were too short!

We will go back a year to continue this history. I collected, under far less favourable conditions, it is true, enough notes to map out the biography of the tiny creature which we have just seen migrating from the camomile-flowers to the Anthophora's back. From what I have said of the Sitaris-larvæ, it is plain that the Meloe-larvæ perched, like the former, on the back of a Bee, have but one aim: to get themselves conveyed by this Bee to the victualled cells. Their object is not to live for a time on the body that carries them.

Were it necessary to prove this, it would be enough to say that we never see these larvæ attempt to pierce the skin of the Bee, or else to nibble at a hair or two, nor do we see them increase in size so long as they are on the Bee's body. To the Meloes, as to the Sitares, the Anthophora serves merely as a vehicle which conveys them to their goal, the victualled cell.

It remains for us to learn how the Meloe leaves the down of the Bee which has carried it, in order to enter the cell. With larvæ collected from the bodies of different Bees, before I was fully acquainted with the tactics of the Sitares, I undertook, as Newport had done before me, certain investigations intended to throw light on this leading point in the Oil-beetle's history. My attempts, based upon those which I had made with the Sitares, resulted in the same failure. The tiny creatures, when brought into contact with Anthophora-larvæ or – nymphs, paid no attention whatever to their prey; others, placed near cells which were open and full of honey, did not enter them, or at most ventured to the edge of the orifice; others, lastly, put inside the cell, on the dry wall or on the surface of the honey, came out again immediately or else got stuck and died. The touch of the honey is as fatal to them as to the young Sitares.

Searches made at various periods in the nests of the Hairy-footed Anthophora had taught me some years earlier that Meloe cicatricosus, like the Sitares, is a parasite of that Bee; indeed I had at different times discovered adult Meloes, dead and shrivelled, in the Bee's cells. On the other hand, I knew from Léon Dufour that the little yellow animal, the Louse found in the Bee's down, had been recognized, thanks to Newport's investigations, as the larva of the Oil-beetle. With these data, rendered still more striking by what I was learning daily on the subject of the Sitares, I went to Carpentras, on the 21st of May, to inspect the nests of the Anthophoræ, then building, as I have described. Though I was almost certain of succeeding, sooner or later, with the Sitares, who were excessively abundant, I had very little hope of the Meloes, which on the contrary are very scarce in the same nests. Circumstances, however, favoured me more than I dared hope and, after six hours' labour, in which the pick played a great part, I became the possessor, by the sweat of my brow, of a considerable number of cells occupied by Sitares and two other cells appropriated by Meloes.

While my enthusiasm had not had time to cool at the sight, momentarily repeated, of a young Sitaris perched upon an Anthophora's egg floating in the centre of the little pool of honey, it might well have burst all restraints on beholding the contents of one of these cells. On the black, liquid honey a wrinkled pellicle is floating; and on this pellicle, motionless, is a yellow louse. The pellicle is the empty envelope of the Anthophora's egg; the louse is a Meloe-larva.

The story of this larva becomes self-evident. The young Meloe leaves the down of the Bee at the moment when the egg is laid; and, since contact with the honey would be fatal to the grub, it must, in order to save itself, adopt the tactics followed by the Sitaris, that is to say, it must allow itself to drop on the surface of the honey with the egg which is in the act of being laid. There, its first task is to devour the egg which serves it for a raft, as is attested by the empty envelope on which it still remains; and it is after this meal, the only one that it takes so long as it retains its present form, that it must commence its long series of transformations and feed upon the honey amassed by the Anthophora. This was the reason of the complete failure both of my attempts and of Newport's to rear the young Meloe-larvæ. Instead of offering them honey, or larvæ, or nymphs, we should have placed them on the eggs recently laid by the Anthophora.

On my return from Carpentras, I meant to try this method, together with that of the Sitares, with which I had been so successful; but, as I had no Meloe-larvæ at my disposal and could not obtain any save by searching for them in the Bees' fleece, the Anthophora-eggs were all discovered to have hatched in the cells which I brought back from my expedition, when I was at last able to find some. This lost experiment is little to be regretted, for, since the Meloes and the Sitares exhibiting the completest similarity not only in habits but also in their method of evolution, there is no doubt whatever that I should have succeeded. I even believe that this method may be attempted with the cells of various Bees, provided that the eggs and the honey do not differ too greatly from the Anthophora's. I should not, for example, count on being successful with the cells of the three-horned Osmia, who shares the Anthophora's quarters: her egg is short and thick; and her honey is yellow, odourless, solid, almost a powder and very faintly flavoured.

17

Carolus Linnæus (Karl von Linné, 1707-1778), the celebrated Swedish botanist and naturalist, founder of the Linnæan system of classification. —Translator's Note.

18

Jean Baptiste Godart (1775-1823), the principal editor of L'Histoire naturelle des lépidoptères de France. —Translator's Note.

19

Baron Karl de Geer (1720-1778), the Swedish entomologist, author of Mémoires pour servir à l'histoire des insectes (1752-1778). —Translator's Note.

20

Cf. The Mason-bees: chaps. viii. and ix. —Translator's Note.

21

For the Wasp known as the Hairy Ammophila, who feeds her young on the Grey Worm, the caterpillar of the Turnip Moth, cf. The Hunting Wasps, chaps. xviii. to xx. —Translator's Note.

The Glow-Worm and Other Beetles

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