CHAPTER IV
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;1 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.
1 Carolus Linnæus (Karl von Linné, 1707-1778), the celebrated Swedish botanist and naturalist, founder of the Linnæan system of classification.—Translator's Note.
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,2 de Geer3 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.
2 Jean Baptiste Godart (1775-1823), the principal editor of L'Histoire naturelle des lépidoptères de France.—Translator's Note.
3 Baron Karl de Geer (1720-1778), the Swedish entomologist, author of Mémoires pour servir à l'histoire des insectes (1752-1778).—Translator's Note.
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,4 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.
4 Cf. The Mason-bees: chaps. viii. and ix.—Translator's Note.
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),5 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.
5 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.
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.
CHAPTER V
By a Machiavellian stratagem the primary larva of the Oil-beetle or the Sitaris has penetrated the Anthophora's cell; it has settled on the egg, which is its first food and its life-raft in one. What becomes of it once the egg is exhausted?
Let us, to begin with, go back to the larva of the Sitaris. By the end of a week the Anthophora's egg has been drained dry by the parasite and is reduced to the envelope, a shallow skiff which preserves the tiny creature from the deadly contact of the honey. It is on this skiff that the first transformation takes place, whereafter the larva, which is now organized to live in a glutinous environment, drops off the raft into the pool of honey and leaves its empty skin, split along the back, clinging to the pellicle of the egg. At this stage we see floating motionless on the honey a milk-white atom, oval, flat and a twelfth of an inch long. This is the larva of the Sitaris in its new form. With the aid of a lens we can distinguish the fluctuations of the digestive canal, which is gorging itself with honey; and along the circumference of the flat, elliptical back we perceive a double row of breathing-pores which, thanks to their position, cannot be choked by the viscous liquid. Before describing the larva in detail we will wait for it to attain its full development, which cannot take long, for the provisions are rapidly diminishing.
The rapidity however is not to be compared with that with which the gluttonous larvæ of the Anthophora consume their food. Thus, on visiting the dwellings of the Anthophoræ for the last time, on the 25th of June, I found that the Bee's larvæ had all finished their rations and attained their full development, whereas those of the Sitares, still immersed in the honey, were, for the most part, only half the size which they must finally attain. This is yet another reason why the Sitares should destroy an egg which, were it to develop, would produce a voracious larva, capable of starving them in a very short time. When rearing the larvæ myself in test-tubes, I have found that the Sitares take thirty-five to forty days to finish their mess of honey and that the larvæ of the Anthophora spend less than a fortnight over the same meal.
It is in the first half of July that the Sitaris-grubs reach their full dimensions. At this period the cell usurped by the parasite contains nothing beyond a full-fed larva and, in a corner, a heap of reddish droppings. This larva is soft and white, about half an inch in length and a quarter of an inch wide at its broadest part. Seen from above as it floats on the honey, it is elliptical in form, tapering gradually towards the front and more suddenly towards the rear. Its ventral surface is highly convex; its dorsal surface, on the contrary, is almost flat. When the larva is floating on the liquid honey, it is as it were steadied by the excessive development of the ventral surface immersed in the honey, which enables it to acquire an equilibrium that is of the greatest importance to its welfare. In fact, the breathing-holes, arranged without means of protection on either edge of the almost flat back, are level with the viscous liquid and would be choked by that sticky glue at the least false movement, if a suitably ballasted hold did not prevent the larva from heeling over. Never was corpulent abdomen of greater use: thanks to this plumpness of the belly the larva is protected from asphyxia.
Its segments number thirteen, including the head. This head is pale, soft, like the rest of the body, and very small compared with the rest of the creature. The antennæ are excessively short and consist of two cylindrical joints. I have vainly looked for the eyes with a powerful magnifying-glass. In its former state, the larva, subject to strange migrations, obviously needs the sense of sight and is provided with four ocelli. In its present state, of what use would eyes be to it at the bottom of a clay cell, where the most absolute darkness prevails?
The labrum is prominent, is not distinctly divided from the head, is curved in front and edged with pale and very fine bristles. The mandibles are small, reddish toward the tips, blunt and hollowed out spoonwise on the inner side. Below the mandibles is a fleshy part crowned with two very tiny nipples. This is the lower lip with its two palpi. It is flanked right and left by two other parts, likewise fleshy, adhering closely to the lip and bearing at the tip a rudimentary palp consisting of two or three very tiny joints. These two parts are the future jaws. All this apparatus of lips and jaws is completely immobile and in a rudimentary condition which is difficult to describe. They are budding organs, still faint and embryonic. The labrum and the complicated lamina formed by the lip and the jaws leave between them a narrow slit in which the mandibles work.
The legs are merely vestiges, for, though they consist of three tiny cylindrical joints, they are barely a fiftieth of an inch in length. The creature is unable to make use of them, not only in the liquid honey upon which it lives, but even on a solid surface. If we take the larva from the cell and place it on a hard substance, to observe it more readily, we see that the inordinate protuberance of the abdomen, by lifting the thorax from the ground, prevents the legs from finding a support. Lying on its side, the only possible position because of its conformation, the larva remains motionless or only makes a few lazy, wriggling movements of the abdomen, without ever stirring its feeble limbs, which for that matter could not assist it in any way. In short, the tiny creature of the first stage, so active and alert, is succeeded by a ventripotent grub, deprived of movement by its very obesity. Who would recognize in this clumsy, flabby, blind, hideously pot-bellied creature, with nothing but a sort of stumps for legs, the elegant pigmy of but a little while back, armour-clad, slender and provided with highly perfected organs for performing its perilous journeys?
Lastly, we count nine pairs of stigmata: one pair on the mesothorax and the rest on the first eight segments of the abdomen. The last pair, that on the eighth abdominal segment, consists of stigmata so small that to detect them we have to gather their position by that in the succeeding states of the larva and to pass a very patient magnifying-glass along the direction of the other pairs. These are as yet but vestigial stigmata. The others are fairly large, with pale, round, flat edges.
If in its first form the Sitaris-larva is organized for action, to obtain possession of the coveted cell, in its second form it is organized solely to digest the provisions acquired. Let us take a glance at its internal structure and in particular at its digestive apparatus. Here is a strange thing: this apparatus, in which the hoard of honey amassed by the Anthophora is to be engulfed, is similar in every respect to that of the adult Sitaris, who possibly never takes food. We find in both the same very short oesophagus, the same chylific ventricle, empty in the perfect insect, distended in the larva with an abundant orange-coloured pulp; in both the same gall-bladders, four in number, connected with the rectum by one of their extremities. Like the perfect insect, the larva is devoid of salivary glands or any other similar apparatus. Its nervous system comprises eleven ganglia, not counting the oesophageal collar, whereas in the perfect insect there are only seven: three for the thorax, of which the last two are contiguous, and four for the abdomen.
When its rations are finished the larva remains a few days in a motionless condition, ejecting from time to time a few reddish droppings until the digestive canal is completely cleared of its orange-coloured pulp. Then the creature contracts itself, huddles itself together; and before long we see coming detached from its body a transparent, slightly crumpled and extremely fine pellicle, forming a closed bag, in which the successive transformations will take place henceforth. On this epidermal bag, this sort of transparent leather bottle, formed by the larva's skin detached all of a piece, without a slit of any kind, we can distinguish the several well-preserved external organs: the head, with its antennæ, mandibles, paws and palpi; the thoracic segments, with their vestiges of legs; the abdomen, with its chain of breathing-holes still connected one to another by tracheal threads.
Then beneath this pellicle, which is so delicate that it can hardly bear the most cautious touch, we see a soft, white mass taking shape, a mass which in a few hours acquires a firm, horny consistency and a vivid yellow hue. The transformation is now complete. Let us tear the fine gauze bag enclosing the organism which has just come into being and direct our investigation to this third form of the Sitaris-larva.
It is an inert, segmented body, with an oval outline, a horny consistency, just like that of pupæ and chrysalids, and a bright-yellow colour, which we can best describe by likening it to that of a lemon-drop. Its upper surface forms a double inclined plane with a very blunt ridge; its lower surface is at first flat, but, as the result of evaporation, becomes more concave daily, leaving a projecting rim all around its oval outline. Lastly, its two extremities or poles are slightly flattened. The major axis of the lower surface averages half an inch in length and the minor axis a quarter of an inch.
At the cephalic pole of this body is a sort of mask, modelled roughly on the head of the larva, and at the opposite pole a small circular disk deeply wrinkled at the centre. The three segments that come after the head bear each a pair of very minute knobs, hardly visible without the lens: these are, to the legs of the larva in its previous form, what the cephalic mask is to the head of the same larva. They are not organs, but indications, landmarks placed at the points where these organs will appear later. On either side we count nine stigmata, set as before on the mesothorax and the first eight abdominal segments. The first eight breathing-holes are dark brown and stand out plainly against the yellow colour of the body. They consist of small, shiny, conical knobs, perforated at the top with a round hole. The ninth stigma, though fashioned like the others, is ever so much smaller; it cannot be distinguished without the lens.
The anomaly, already so manifest in the change from the first form to the second, becomes even more so here; and we do not know what name to give to an organism without a standard of comparison, not only in the order of Beetles, but in the whole class of insects. While, on the one hand, this organism offers many points of resemblance to the pupæ of the Flies in its horny consistency, in the complete immobility of its various segments, in the all but absolute absence of relief which would enable one to distinguish the parts of the perfect insect; while, on the other hand, it approximates to the chrysalids, because the creature, to attain this condition, has to shed its skin, as the caterpillars do, it differs from the pupa because it has for covering not the surface skin, which has become horny, but rather one of the inner skins of the larva; and it differs from the chrysalids by the absence of mouldings which in the latter betray the appendages of the perfect insect. Lastly, it differs yet more profoundly from the pupa and the chrysalis because from both these organisms the perfect insect springs straightway, whereas that which follows what we are considering is simply a larva like that which went before. I shall suggest, to denote this curious organism, the term pseudochrysalis; and I shall reserve the names primary larva, secondary larva and tertiary larva to denote, in a couple of words, each of the three forms under which the Sitares possess all the characteristics of larvæ.
Although the Sitaris, on assuming the form of the pseudochrysalis, is transfigured outwardly to the point of baffling the science of entomological phases, this is not so inwardly. I have at every season of the year examined the viscera of the pseudochrysalids, which generally remain stationary for a whole year, and I have never observed other forms among their organs than those which we find in the secondary larva. The nervous system has undergone no change. The digestive apparatus is absolutely void and, because of its emptiness, appears only as a thin cord, sunk, lost amid the adipose sacs. The stercoral intestine has more substance; its outlines are better defined. The four gall-bladders are always perfectly distinct. The adipose tissue is more abundant than ever: it forms by itself the whole contents of the pseudochrysalis, for in the matter of volume the insignificant threads of the nervous system and the digestive apparatus count for nothing. It is the reserve upon which life must draw for its future labours.
A few Sitares remain hardly a month in the pseudochrysalis stage. The other phases are achieved in the course of August; and at the beginning of September the insect attains the perfect state. But as a rule the development is slower; the pseudochrysalis goes through the winter; and it is not, at the earliest, until June in the second year that the final transformations take place. Let us pass in silence over this long period of repose, during which the Sitaris, in the form of a pseudochrysalis, slumbers at the bottom of its cell, in a sleep as lethargic as that of a germ in its egg, and come to the months of June and July in the following year, the period of what we might call a second hatching.
The pseudochrysalis is still enclosed in the delicate pouch formed of the skin of the secondary larva. Outside, nothing fresh has happened; but important changes have taken place inside. I have said that the pseudochrysalis displayed an upper surface arched like a hog's back and a lower surface at first flat and then more and more concave. The sides of the double inclined plane of the upper or dorsal surface also share in this depression occasioned by the evaporation of the fluid constituents; and a time comes when these sides are so depressed that a section of the pseudochrysalis through a plane perpendicular to its axis would be represented by a curvilinear triangle with blunted corners and inwardly convex sides. This is the appearance displayed by the pseudochrysalis during the winter and spring.
But in June it has lost this withered appearance; it represents a perfect balloon, an ellipsoid of which the sections perpendicular to the major axis are circles. Something has also come to pass of greater importance than this expansion, which may be compared with that which we obtain by blowing into a wrinkled bladder. The horny integuments of the pseudochrysalis have become detached from their contents, all of a piece, without a break, just as happened the year before with the skin of the secondary larva; and they thus form a fresh vesicular envelope, free from any adhesion to the contents and itself enclosed in the pouch formed of the secondary larva's skin. Of these two bags without outlet, one of which is enclosed within the other, the outer is transparent, flexible, colourless and extremely delicate; the second is brittle, almost as delicate as the first, but much less translucent because of its yellow colouring, which makes it resemble a thin flake of amber. On this second sac are found the stigmatic warts, the thoracic studs and so forth, which we noted on the pseudochrysalis. Lastly, within its cavity we catch a glimpse of something the shape of which at once recalls to mind the secondary larva.
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.1 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.
1 .787 inch.—Translator's Note.
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.2 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.
2 Later classifiers place both in the family of the Cantharidæ.—Translator's Note.
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.3 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.
3 .393 x .156 inch.—Translator's Note.
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.