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I mark a plentiful number of my guests with a variety of dots on the thorax, which enables me to follow any one Osmia from the beginning to the end of her laying. The tubes and their respective holes are numbered; a list, always lying open on my desk, enables me to note from day to day, sometimes from hour to hour, what happens in each tube and particularly the actions of the Osmiae whose backs bear distinguishing marks. As soon as one tube is filled, I replace it by another. Moreover, I have scattered in front of either hive a few handfuls of empty Snail-shells, specially chosen for the object which I have in view. Reasons which I will explain later led me to prefer the shells of Helix caespitum. Each of the shells, as and when stocked, received the date of the laying and the alphabetical sign corresponding with the Osmia to whom it belonged. In this way, I spent five or six weeks in continual observation. To succeed in an enquiry, the first and foremost condition is patience. This condition I fulfilled; and it was rewarded with the success which I was justified in expecting.
The tubes employed are of two kinds. The first, which are cylindrical and of the same width throughout, will be of use for confirming the facts observed in the first year of my experiments in indoor rearing. The others, the majority, consist of two cylinders which are of very different diameters, set end to end. The front cylinder, the one which projects a little way outside the hive and forms the entrance-hole, varies in width between 8 and 12 millimetres. (Between.312 to .468 inch.—Translator's Note.) The second, the back one, contained entirely within my packing-case, is closed at its far end and is 5 to 6 millimetres (.195 to.234 inch.—Translator's Note.) in diameter. Each of the two parts of the double-galleried tunnel, one narrow and one wide, measures at most a decimetre (3.9 inches.—Translator's Note.) in length. I thought it advisable to have these short tubes, as the Osmia is thus compelled to select different lodgings, each of them being insufficient in itself to accommodate the total laying. In this way I shall obtain a greater variety in the distribution of the sexes. Lastly, at the mouth of each tube, which projects slightly outside the case, there is a little paper tongue, forming a sort of perch on which the Osmia alights on her arrival and giving easy access to the house. With these facilities, the swarm colonized fifty-two double-galleried tubes, thirty-seven cylindrical tubes, seventy-eight Snail-shells and a few old nests of the Mason-bee of the Shrubs. From this rich mine of material I will take what I want to prove my case.
Every series, even when incomplete, begins with females and ends with males. To this rule I have not yet found an exception, at least in galleries of normal diameter. In each new abode, the mother busies herself first of all with the more important sex. Bearing this point in mind, would it be possible for me, by manoeuvring, to obtain an inversion of this order and make the laying begin with males? I think so, from the results already ascertained and the irresistible conclusions to be drawn from them. The double-galleried tubes are installed in order to put my conjectures to the proof.
The back gallery, 5 or 6 millimetres (.195 to.234 inch.—Translator's Note.) wide, is too narrow to serve as a lodging for normally developed females. If, therefore, the Osmia, who is very economical of her space, wishes to occupy them, she will be obliged to establish males there. And her laying must necessarily begin here, because this corner is the rear-most part of the tube. The foremost gallery is wide, with an entrance-door on the front of the hive. Here, finding the conditions to which she is accustomed, the mother will go on with her laying in the order which she prefers.
Let us now see what has happened. Of the fifty-two double galleried tubes, about a third did not have their narrow passage colonized. The Osmia closed its aperture communicating with the large passage; and the latter alone received the eggs. This waste of space was inevitable. The female Osmiae, though nearly always larger than the males, present marked differences among one another: some are bigger, some are smaller. I had to adjust the width of the narrow galleries to Bees of average dimensions. It may happen therefore that a gallery is too small to admit the large-sized mothers to whom chance allots it. When the Osmia is unable to enter the tube, obviously she will not colonize it. She then closes the entrance to this space which she cannot use and does her laying beyond it, in the wide tube. Had I tried to avoid these useless apparatus by choosing tubes of larger calibre, I should have encountered another drawback: the medium-sized mothers, finding themselves almost comfortable, would have decided to lodge females there. I had to be prepared for it: as each mother selected her house at will and as I was unable to interfere in her choice, a narrow tube would be colonized or not, according as the Osmia who owned it was or was not able to make her way inside.
There remain some forty pairs of tubes with both galleries colonized. In these there are two things to take into consideration. The narrow rear tubes of 5 or 5 1/2 millimetres (.195 to.214 inch.—Translator's Note.)—and these are the most numerous—contain males and males only, but in short series, between one and five. The mother is here so much hampered in her work that they are rarely occupied from end to end; the Osmia seems in a hurry to leave them and to go and colonize the front tube, whose ample space will leave her the liberty of movement necessary for her operations. The other rear tubes, the minority, whose diameter is about 6 millimetres (.234 inch.—Translator's Note.), contain sometimes only females and sometimes females at the back and males towards the opening. One can see that a tube a trifle wider and a mother slightly smaller would account for this difference in the results. Nevertheless, as the necessary space for a female is barely provided in this case, we see that the mother avoids as far as she can a two-sex arrangement beginning with males and that she adopts it only in the last extremity. Finally, whatever the contents of the small tube may be, those of the large one, following upon it, never vary and consist of females at the back and males in front.
Though incomplete, because of circumstances very difficult to control, the result of the experiment is none the less very striking. Twenty-five apparatus contain only males in their narrow gallery, in numbers varying from a minimum of one to a maximum of five. After these comes the colony of the large gallery, beginning with females and ending with males. And the layings in these apparatus do not always belong to late summer or even to the intermediate period: a few small tubes contain the earliest eggs of the Osmiae. A couple of Osmiae, more forward than the others, set to work on the 23rd of April. Both of them started their laying by placing males in the narrow tubes. The meagre supply of provisions was enough in itself to show the sex, which proved later to be in accordance with my anticipations. We see then that, by my artifices, the whole swarm starts with the converse of the normal order. This inversion is continued, at no matter what period, from the beginning to the end of the operations. The series which, according to rule, would begin with females now begins with males. Once the larger gallery is reached, the laying is pursued in the usual order.
We have advanced one step and that no small one: we have seen that the Osmia, when circumstances require it, is capable of reversing the sequence of the sexes. Would it be possible, provided that the tube were long enough, to obtain a complete inversion, in which the entire series of the males should occupy the narrow gallery at the back and the entire series of the females the roomy gallery in front? I think not; and I will tell you why.
Long and narrow cylinders are by no means to the Osmia's taste, not because of their narrowness but because of their length. Remember that for each load of honey brought the worker is obliged to move backwards twice. She enters, head first, to begin by disgorging the honey-syrup from her crop. Unable to turn in a passage which she blocks entirely, she goes out backwards, crawling rather than walking, a laborious performance on the polished surface of the glass and a performance which, with any other surface, would still be very awkward, as the wings are bound to rub against the wall with their free end and are liable to get rumpled or bent. She goes out backwards, reaches the outside, turns round and goes in again, but this time the opposite way, so as to brush off the load of pollen from her abdomen on to the heap. If the gallery is at all long, this crawling backwards becomes troublesome after a time; and the Osmia soon abandons a passage that is too small to allow of free movement. I have said that the narrow tubes of my apparatus are, for the most part, only very incompletely colonized. The Bee, after lodging a small number of males in them, hastens to leave them. In the wide front gallery, she can stay where she is and still be able to turn round easily for her different manipulations; she will avoid those two long journeys backwards, which are so exhausting and so bad for her wings.
Another reason no doubt prompts her not to make too great a use of the narrow passage, in which she would establish males, followed by females in the part where the gallery widens. The males have to leave their cells a couple of weeks or more before the females. If they occupy the back of the house, they will die prisoners or else they will overturn everything on their way out. This risk is avoided by the order which the Osmia adopts.
In my tubes with their unusual arrangement, the mother might well find the dilemma perplexing: there is the narrowness of the space at her disposal and there is the emergence later on. In the narrow tubes, the width is insufficient for the females; on the other hand, if she lodges males there, they are liable to perish, since they will be prevented from issuing at the proper moment. This would perhaps explain the mother's hesitation and her obstinacy in settling females in some of my apparatus which looked as if they could suit none but males.
A suspicion occurs to me, a suspicion aroused by my attentive examination of the narrow tubes. All, whatever the number of their inmates, are carefully plugged at the opening, just as separate tubes would be. It might therefore be the case that the narrow gallery at the back was looked upon by the Osmia not as the prolongation of the large front gallery, but as an independent tube. The facility with which the worker turns as soon as she reaches the wide tube, her liberty of action, which is now as great as in a doorway communicating with the outer air, might well be misleading and cause the Osmia to treat the narrow passage at the back as though the wide passage in front did not exist. This would account for the placing of the female in the large tube above the males in the small tube, an arrangement contrary to her custom.
I will not undertake to decide whether the mother really appreciates the danger of my snares, or whether she makes a mistake in considering only the space at her disposal and beginning with males. At any rate, I perceive in her a tendency to deviate as little as possible from the order which safeguards the emergence of the two sexes. This tendency is demonstrated by her repugnance to colonizing my narrow tubes with long series of males. However, so far as we are concerned, it does not matter much what passes at such times in the Osmia's little brain. Enough for us to know that she dislikes narrow and long tubes, not because they are narrow, but because they are at the same time long.
And, in fact, she does very well with a short tube of the same diameter. Such are the cells in the old nests of the Mason-bee of the Shrubs and the empty shells of the Garden Snail. With the short tube, the two disadvantages of the long tube are avoided. She has very little of that crawling backwards to do when she has a Snail-shell for the home of her eggs and scarcely any when the home is the cell of the Mason-bee. Moreover, as the stack of cocoons numbers two or three at most, the deliverance will be exempt from the difficulties attached to a long series. To persuade the Osmia to nidify in a single tube long enough to receive the whole of her laying and at the same time narrow enough to leave her only just the possibility of admittance appears to me a project without the slightest chance of success: the Bee would stubbornly refuse such a dwelling or would content herself with entrusting only a very small portion of her eggs to it. On the other hand, with narrow but short cavities, success, without being easy, seems to me at least quite possible. Guided by these considerations, I embarked upon the most arduous part of my problem: to obtain the complete or almost complete permutation of one sex with the other; to produce a laying consisting only of males by offering the mother a series of lodgings suited only to males.
Let us in the first place consult the old nests of the Mason-bee of the Shrubs. I have said that these mortar spheroids, pierced all over with little cylindrical cavities, are adopted pretty eagerly by the Three-horned Osmia, who colonizes them before my eyes with females in the deep cells and males in the shallow cells. That is how things go when the old nest remains in its natural state. With a grater, however, I scrape the outside of another nest so as to reduce the depth of the cavities to some ten millimetres. (About two-fifths of an inch.—Translator's Note.) This leaves in each cell just room for one cocoon, surmounted by the closing stopper. Of the fourteen cavities in the nests, I leave two intact, measuring fifteen millimetres in depth. (.585 inch.—Translator's Note.) Nothing could be more striking than the result of this experiment, made in the first year of my home rearing. The twelve cavities whose depth had been reduced all received males; the two cavities left untouched received females.
A year passes and I repeat the experiment with a nest of fifteen cells; but this time all the cells are reduced to the minimum depth with the grater. Well, the fifteen cells, from first to last, are occupied by males. It must be quite understood that, in each case, all the offspring belonged to one mother, marked with her distinguishing spot and kept in sight as long as her laying lasted. He would indeed be difficult to please who refused to bow before the results of these two experiments. If, however, he is not yet convinced, here is something to remove his last doubts.
The Three-horned Osmia often settles her family in old shells, especially those of the Common Snail (Helix aspersa), who is so common under the stone-heaps and in the crevices of the little unmortared walls that support our terraces. In this species, the spiral is wide open, so that the Osmia, penetrating as far down as the helical passage permits, finds, immediately above the point which is too narrow to pass, the space necessary for the cell of a female. This cell is succeeded by others, wider still, always for females, arranged in a line in the same way as in a straight tube. In the last whorl of the spiral, the diameter would be too great for a single row. Then longitudinal partitions are added to the transverse partitions, the whole resulting in cells of unequal dimensions in which males predominate, mixed with a few females in the lower storeys. The sequence of the sexes is therefore what it would be in a straight tube and especially in a tube with a wide bore, where the partitioning is complicated by subdivisions on the same level. A single Snail-shell contains room for six or eight cells. A large, rough earthen stopper finishes the nest at the entrance to the shell.
As a dwelling of this sort could show us nothing new, I chose for my swarm the Garden Snail (Helix caespitum), whose shell, shaped like a small, swollen Ammonite, widens by slow degrees, the diameter of the usable portion, right up to the mouth, being hardly greater than that required by a male Osmia-cocoon. Moreover, the widest part, in which a female might find room, has to receive a thick stopping-plug, below which there will often be a free space. Under all these conditions, the house will hardly suit any but males arranged one after the other.
The collection of shells placed at the foot of each hive includes specimens of different sizes. The smallest are 18 millimetres (.7 inch.—Translator's Note.) in diameter and the largest 24 millimetres (.936 inch.—Translator's Note.) There is room for two cocoons, or three at most, according to their dimensions.
Now these shells were used by my visitors without any hesitation, perhaps even with more eagerness than the glass tubes, whose slippery sides might easily be a little annoying to the Bee. Some of them were occupied on the first few days of the laying; and the Osmia who had started with a home of this sort would pass next to a second Snail-shell, in the immediate neighbourhood of the first, to a third, a fourth and others still, always close to one another, until her ovaries were emptied. The whole family of one mother would thus be lodged in Snail-shells which were duly marked with the date of the laying and a description of the worker. The faithful adherents of the Snail-shell were in the minority. The greater number left the tubes to come to the shells and then went back from the shells to the tubes. All, after filling the spiral staircase with two or three cells, closed the house with a thick earthen stopper on a level with the opening. It was a long and troublesome task, in which the Osmia displayed all her patience as a mother and all her talents as a plasterer. There were even some who, scrupulous to excess, carefully cemented the umbilicus, a hole which seemed to inspire them with distrust as being able to give access to the interior of the dwelling. It was a dangerous-looking cavity, which for the greater safety of the family it was prudent to block up.
When the pupae are sufficiently matured, I proceed to examine these elegant abodes. The contents fill me with joy: they fulfil my anticipations to the letter. The great, the very great majority of the cocoons turn out to be males; here and there, in the bigger cells, a few rare females appear. The smallness of the space has almost done away with the sixty-eight Snail-shells colonized. But, of this total number, I must use only those series which received an entire laying and were occupied by the same Osmia from the beginning to the end of the egg-season. Here are a few examples, taken from among the most conclusive.
From the 6th of May, when she started operations, to the 25th of May, the date at which her laying ceased, the Osmia occupied seven Snail-shells in succession. Her family consists of fourteen cocoons, a number very near the average; and, of these fourteen cocoons, twelve belong to males and only two to females. These occupy the seventh and thirteenth places in chronological order.
Another, between the 9th and 27th of May, stocked six Snail-shells with a family of thirteen, including ten males and three females. The places occupied by the latter in the series were numbers 3, 4 and 5.
A third, between the 2nd and 29th of May, colonized eleven Snail-shells, a prodigious task. This industrious one was also exceedingly prolific. She supplied me with a family of twenty-six, the largest which I have ever obtained from one Osmia. Well, this abnormal progeny consisted of twenty-five males and one female, one alone, occupying place 17.
There is no need to go on, after this magnificent example, especially as the other series would all, without exception, give us the same result. Two facts are immediately obvious. The Osmia is able to reverse the order of her laying and to start with a more or less long series of males before producing any females. In the first case, the first female appears as number 7; in the third, as number 17. There is something better still; and this is the proposition which I was particularly anxious to prove: the female sex can be permuted with the male sex and can be permuted to the point of disappearing altogether. We see this especially in the third case, where the presence of a solitary female in a family of twenty-six is due to the somewhat larger diameter of the corresponding Snail-shell and also, no doubt, to some mistake on the mother's part, for the female cocoon, in a series of two, occupies the upper storey, the one next to the orifice, an arrangement which the Osmia appears to me to dislike.
This result throws so much light on one of the darkest corners of biology that I must attempt to corroborate it by means of even more conclusive experiments. I propose next year to give the Osmiae nothing but Snail-shells for a lodging, picked out one by one, and rigorously to deprive the swarm of any other retreat in which the laying could be effected. Under these conditions, I ought to obtain nothing but males, or nearly, for the whole swarm.
There would still remain the inverse permutation: to obtain only females and no males, or very few. The first permutation makes the second seem very probable, although I cannot as yet conceive a means of realizing it. The only condition which I can regulate is the dimensions of the home. When the rooms are small, the males abound and the females tend to disappear. With generous quarters, the converse would not take place. I should obtain females and afterwards an equal number of males, confined in small cells which, in case of need, would be bounded by numerous partitions. The factor of space does not enter into the question here. What artifice can we then employ to provoke this second permutation? So far, I can think of nothing that is worth attempting.
It is time to conclude. Leading a retired life, in the solitude of a village, having quite enough to do with patiently and obscurely ploughing my humble furrow, I know little about modern scientific views. In my young days I had a passionate longing for books and found it difficult to procure them; to-day, when I could almost have them if I wanted, I am ceasing to wish for them. It is what usually happens as life goes on. I do not therefore know what may have been done in the direction whither this study of the sexes has led us. If I am stating propositions that are really new or at least more comprehensive than the propositions already known, my words will perhaps sound heretical. No matter: as a simple translator of facts, I do not hesitate to make my statement, being fully persuaded that time will turn my heresy into orthodoxy. I will therefore recapitulate my conclusions.
Bees lay their eggs in series of first females and then males, when the two sexes are of different sizes and demand an unequal quantity of nourishment. When the two sexes are alike in size, the same sequence may occur, but less regularly.
This dual arrangement disappears when the place chosen for the nest is not large enough to contain the entire laying. We then see broken layings, beginning with females and ending with males.
The egg, as it issues from the ovary, has not yet a fixed sex. The final impress that produces the sex is given at the moment of laying or a little before.
So as to be able to give each larva the amount of space and food that suits it according as it is male or female, the mother can choose the sex of the egg which she is about to lay. To meet the conditions of the building, which is often the work of another or else a natural retreat that admits of little or no alteration, she lays either a male egg or a female egg as she pleases. The distribution of the sexes depends upon herself. Should circumstances require it, the order of the laying can be reversed and begin with males; lastly, the entire laying can contain only one sex.
The same privilege is possessed by the predatory Hymenoptera, the Wasps, at least by those in whom the two sexes are of a different size and consequently require an amount of nourishment that is larger in the one case than in the other. The mother must know the sex of the egg which she is going to lay; she must be able to choose the sex of that egg so that each larva may obtain its proper portion of food.
Generally speaking, when the sexes are of different sizes, every insect that collects food and prepares or selects a dwelling for its offspring must be able to choose the sex of the egg in order to satisfy without mistake the conditions imposed upon it.
The question remains how this optional assessment of the sexes is effected. I know absolutely nothing about it. If I should ever learn anything about this delicate point, I shall owe it to some happy chance for which I must wait, or rather watch, patiently. Towards the end of my investigations, I heard of a German theory which relates to the Hive-bee and comes from Dzierzon, the apiarist. (Johann Dzierzon, author of "Theorie und Praxis des neuen Bienenfreundes."—Translator's Note.) If I understand it aright, according to the very incomplete documents which I have before me, the egg, as it issues from the ovary, is said already to possess a sex, which is always the same; it is originally male; and it becomes female by fertilization. The males are supposed to proceed from non-fertilized eggs, the females from fertilized eggs. The Queen-bee would thus lay female eggs or male eggs according as she fertilized them or not while they were passing into her oviduct.
Coming from Germany, this theory cannot but inspire me with profound distrust. As it has been given acceptance, with rash precipitancy, in standard works, I will overcome my reluctance to devoting my attention to Teutonic ideas and will submit it not to the test of argument, which can always be met by an opposite argument, but to the unanswerable test of facts.
For this optional fertilization, determining the sex, the mother's organism requires a seminal reservoir which distils its drop of sperm upon the egg contained in the oviduct and thus gives it a feminine character, or else leaves it its original character, the male character, by refusing it that baptism. This reservoir exists in the Hive-bee. Do we find a similar organ in the other Hymenoptera, whether honey-gatherers or hunters? The anatomical treatises are either silent on this point or, without further enquiry, apply to the order as a whole the data provided by the Hive-bee, however much she differs from the mass of Hymenoptera owing to her social habits, her sterile workers and especially her tremendous fertility, extending over so long a period.
I at first doubted the universal presence of this spermatic receptacle, having failed to find it under my scalpel in my former investigations into the anatomy of the Sphex-wasps and some other game-hunters. But this organ is so delicate and so small that it very easily escapes the eye, especially when our attention is not specially directed in search of it; and, even when we are looking for it and it only, we do not always succeed in discovering it. We have to find a globule attaining in many cases hardly as much as a millimetre (About one-fiftieth of an inch.—Translator's Note.) in diameter, a globule headed amidst a tangle of air-ducts and fatty patches, of which it shares the colour, a dull white. Then again, the merest slip of the forceps is enough to destroy it. My first investigations, therefore, which concerned the reproductive apparatus as a whole, might very well have allowed it to pass unperceived.
In order to know the rights of the matter once and for all, as the anatomical treatises taught me nothing, I once more fixed my microscope on its stand and rearranged my old dissecting-tank, an ordinary tumbler with a cork disk covered with black satin. This time, not without a certain strain on my eyes, which are already growing tired, I succeeded in finding the said organ in the Bembex-wasps, the Halicti (Cf. Chapters 12 to 14 of the present volume.—Translator's Note.), the Carpenter-bees, the Bumble-bees, the Andrenae (A species of Burrowing Bees.—Translator's Note.) and the Megachiles. (Or Leaf-cutting Bees. Cf. Chapter 8 of the present volume.—Translator's Note.) I failed in the case of the Osmiae, the Chalicodomae and the Anthophorae. Is the organ really absent? Or was there want of skill on my part? I lean towards want of skill and admit that all the game-hunting and honey-gathering Hymenoptera possess a seminal receptacle, which can be recognized by its contents, a quantity of spiral spermatozoids whirling and twisting on the slide of the microscope.
This organ once accepted, the German theory becomes applicable to all the Bees and all the Wasps. When copulating, the female receives the seminal fluid and holds it stored in her receptacle. From that moment, the two procreating elements are present in the mother at one and the same time: the female element, the ovule; and the male element, the spermatozoid. At the egg-layer's will, the receptacle bestows a tiny drop of its contents upon the matured ovule, when it reaches the oviduct, and you have a female egg; or else it withholds its spermatozoids and you have an egg that remains male, as it was at first. I readily admit it: the theory is very simple, lucid and seductive. But is it correct? That is another question.
One might begin by reproaching it with making a singular exception to one of the most general rules. Which of us, casting his eyes over the whole zoological progression, would dare to assert that the egg is originally male and that it becomes female by fertilization? Do not the two sexes both call for the assistance of the fertilizing element? If there be one undoubted truth, it is certainly that. We are, it is true, told very curious things about the Hive-bee. I will not discuss them: this Bee stands too far outside the ordinary limits; and then the facts asserted are far from being accepted by everybody. But the non-social Bees and the predatory insects have nothing special about their laying. Then why should they escape the common rule, which requires that every living creature, male as well as female, should come from a fertilized ovule? In its most solemn act, that of procreation, life is one and uniform; what it does here it does there and there and everywhere. What! The sporule of a scrap of moss requires an antherozoid before it is fit to germinate; and the ovule of a Scolia, that proud huntress, can dispense with the equivalent in order to hatch and produce a male? These new-fangled theories seem to me to have very little value.
One might also bring forward the case of the Three-pronged Osmia, who distributes the two sexes without any order in the hollow of her reed. What singular whim is the mother obeying when, without decisive motive, she opens her seminal phial at haphazard to anoint a female egg, or else keeps it closed, also at haphazard, to allow a male egg to pass unfertilized? I could imagine impregnation being given or withheld for periods of some duration; but I cannot understand impregnation and non-impregnation following upon each other anyhow, in any sort of order, or rather with no order it all. The mother has just fertilized an egg. Why should she refuse to fertilize the next, when neither the provisions nor the lodgings differ in the smallest respect from the previous provisions and lodgings? These capricious alternations, so unreasonable and so exceedingly erratic, are scarcely appropriate to an act of such importance.
But I promised not to argue and I find myself arguing. My reasoning is too fine for dull wits. I will pass on and come to the brutal fact, the real sledge-hammer blow.
Towards the end of the Bee's operations, in the first week of June, the last acts of the Three-horned Osmia become so exceptionally interesting that I made her the object of redoubled observation. The swarm at this time is greatly reduced in numbers. I have still some thirty laggards, who continue very busy, though their work is in vain. I see some very conscientiously stopping up the entrance to a tube or a Snail-shell in which they have laid nothing at all. Others are closing the home after only building a few partitions, or even mere attempts at partitions. Some are placing at the back of a new gallery a pinch of pollen which will benefit nobody and then shutting up the house with an earthen stopper as thick, as carefully made as though the safety of a family depended on it. Born a worker, the Osmia must die working. When her ovaries are exhausted, she spends the remainder of her strength on useless works: partitions, plugs, pollen-heaps, all destined to be left unemployed. The little animal machine cannot bring itself to be inactive even when there is nothing more to be done. It goes on working so that its last vibrations of energy may be used up in fruitless labour. I commend these aberrations to the staunch supporters of reasoning-powers in the animal.
Before coming to these useless tasks, my laggards have laid their last eggs, of which I know the exact cells, the exact dates. These eggs, as far as the microscopes can tell, differ in no respect from the others, the older ones. They have the same dimensions, the same shape, the same glossiness, the same look of freshness. Nor are their provisions in any way peculiar, being very well suited to the males, who conclude the laying. And yet these last eggs do not hatch: they wrinkle, fade and wither on the pile of food. In one case, I count three or four sterile eggs among the last lot laid; in another, I find two or only one. Elsewhere in the swarm, fertile eggs have been laid right up to the end.
Those sterile eggs, stricken with death at the moment of their birth, are too numerous to be ignored. Why do they not hatch like the other eggs, which outwardly they resemble in every respect? They have received the same attention from the mother and the same portion of food. The searching microscope shows me nothing in them to explain the fatal ending.
To the unprejudiced mind, the answer is obvious. Those eggs do not hatch because they have not been fertilized. Any animal or vegetable egg that had not received the life-giving impregnation would perish in the same way. No other answer is possible. It is no use talking of the distant period of the laying: eggs of the same period laid by other mothers, eggs of the same date and likewise the final ones of a laying, are perfectly fertile. Once more, they do not hatch because they were not fertilized.
And why were they not fertilized? Because the seminal receptacle, so tiny, so difficult to see that it sometimes escaped me despite all my scrutiny, had exhausted its contents. The mothers in whom this receptacle retained a remnant of sperm to the end had their last eggs as fertile as the first; the others, whose seminal reservoir was exhausted too soon, had their last-born stricken with death. All this seems to me as clear as daylight.
If the unfertilized eggs perish without hatching, those which hatch and produce males are therefore fertilized; and the German theory falls to the ground.
Then what explanation shall I give of the wonderful facts which I have set forth? Why, none, absolutely none. I do not explain facts, I relate them. Growing daily more sceptical of the interpretations suggested to me and more hesitating as to those which I may have to suggest myself, the more I observe and experiment, the more clearly I see rising out of the black mists of possibility an enormous note of interrogation.
Dear insects, my study of you has sustained me and continues to sustain me in my heaviest trials. I must take leave of you for to-day. The ranks are thinning around me and the long hopes have fled. Shall I be able to speak to you again? (This is the closing paragraph of Volume 3 of the "Souvenirs entomologiques," of which the author has lived to publish seven more volumes, containing over 2,500 pages and nearly 850,000 words.—Translator's Note.)
CHAPTER 6. INSTINCT AND DISCERNMENT.
The Pelopaeus (A Mason-wasp forming the subject of essays which have not yet been published in English.—Translator's Note.) gives us a very poor idea of her intellect when she plasters up the spot in the wall where the nest which I have removed used to stand, when she persists in cramming her cell with Spiders for the benefit of an egg no longer there and when she dutifully closes a cell which my forceps has left empty, extracting alike germ and provisions. The Mason-bees (Cf. "The Mason-bees": chapter 7.—Translator's Note.), the caterpillar of the Great Peacock Moth (Cf. "Social Life in the Insect World" by J.H. Fabre, translated by Bernard Miall: chapter 14.—Translator's Note.) and many others, when subjected to similar tests, are guilty of the same illogical behaviour: they continue, in the normal order, their series of industrious actions, though an accident has now rendered them all useless. Just like millstones unable to cease revolving though there be no corn left to grind, let them once be given the compelling power and they will continue to perform their task despite its futility. Are they then machines? Far be it from me to think anything so foolish.
It is impossible to make definite progress on the shifting sands of contradictory facts: each step in our interpretation may find us embogged. And yet these facts speak so loudly that I do not hesitate to translate their evidence as I understand it. In insect mentality, we have to distinguish two very different domains. One of these is INSTINCT properly so called, the unconscious impulse that presides over the most wonderful part of what the creature achieves. Where experience and imitation are of absolutely no avail, instinct lays down its inflexible law. It is instinct and instinct alone that makes the mother build for a family which she will never see; that counsels the storing of provisions for the unknown offspring; that directs the sting towards the nerve-centres of the prey and skilfully paralyses it, so that the game may keep good; that instigates, in fine, a host of actions wherein shrewd reason and consummate science would have their part, were the creature acting through discernment.
This faculty is perfect of its kind from the outset, otherwise the insect would have no posterity. Time adds nothing to it and takes nothing from it. Such as it was for a definite species, such it is to-day and such it will remain, perhaps the most settled zoological characteristic of them all. It is not free nor conscious in its practice, any more than is the faculty of the stomach for digestion or that of the heart for pulsation. The phases of its operations are predetermined, necessarily entailed one by another; they suggest a system of clock-work wherein one wheel set in motion brings about the movement of the next. This is the mechanical side of the insect, the fatum, the only thing which is able to explain the monstrous illogicality of a Pelopaeus when misled by my artifices. Is the Lamb when it first grips the teat a free and conscious agent, capable of improvement in its difficult art of taking nourishment? The insect is no more capable of improvement in its art, more difficult still, of giving nourishment.
But, with its hide-bound science ignorant of itself, pure insect, if it stood alone, would leave the insect unarmed in the perpetual conflict of circumstances. No two moments in time are identical; though the background remain the same, the details change; the unexpected rises on every side. In this bewildering confusion, a guide is needed to seek, accept, refuse and select; to show preference for this and indifference to that; to turn to account, in short, anything useful that occasion may offer. This guide the insect undoubtedly possesses, to a very manifest degree. It is the second province of its mentality. Here it is conscious and capable of improvement by experience. I dare not speak of this rudimentary faculty as intelligence, which is too exalted a title: I will call it DISCERNMENT. The insect, in exercising its highest gifts, discerns, differentiates between one thing and another, within the sphere of its business, of course; and that is about all.
As long as we confound acts of pure instinct and acts of discernment under the same head, we shall fall back into those endless discussions which embitter controversy without bringing us one step nearer to the solution of the problem. Is the insect conscious of what it does? Yes and no. No, if its action is in the province of instinct; yes, if the action is in that of discernment. Are the habits of an insect capable of modification? No, decidedly not, if the habit in question belongs to the province of instinct; yes, if it belongs to that of discernment. Let us state this fundamental distinction more precisely by the aid of a few examples.
The Pelopaeus builds her cells with earth already softened, with mud. Here we have instinct, the unalterable characteristic of the worker. She has always built in this way and always will. The passing ages will never teach her, neither the struggle for life nor the law of selection will ever induce her to imitate the Mason-bee and collect dry dust for her mortar. This mud nest needs a shelter against the rain. The hiding-place under a stone suffices at first. But should she find something better, the potter takes possession of that something better and instals herself in the home of man. (The Pelopaeus builds in the fire-places of houses.—Translator's Note.) There we have discernment, the source of some sort of capacity for improvement.
The Pelopaeus supplies her larvae with provisions in the form of Spiders. There you have instinct. The climate, the longitude or latitude, the changing seasons, the abundance or scarcity of game introduce no modification into this diet, though the larva shows itself satisfied with other fare provided by myself. Its forebears were brought up on Spiders; their descendants consumed similar food; and their posterity again will know no other. Not a single circumstance, however favourable, will ever persuade the Pelopaeus that young Crickets, for instance, are as good as Spiders and that her family would accept them gladly. Instinct binds her down to the national diet.
But, should the Epeira (The Weaving or Garden Spider. Cf. "The Life of the Spider" by J. Henri Fabre translated by Alexander Teixeira de Mattos; chapters 9 to 14 and appendix.—Translator's Note.), the favourite prey, be lacking, must the Pelopaeus therefore give up foraging? She will stock her warehouses all the same, because any Spider suits her. There you have discernment, whose elasticity makes up, in certain circumstances, for the too-great rigidity of instinct. Amid the innumerable variety of game, the huntress is able to discern between what is Spider and what is not; and, in this way, she is always prepared to supply her family, without quitting the domain of her instinct.
The Hairy Ammophila gives her larva a single caterpillar, a large one, paralysed by as many pricks of her sting as it has nervous centres in its thorax and abdomen. Her surgical skill in subduing the monster is instinct displayed in a form which makes short work of any inclination to see in it an acquired habit. In an art that can leave no one to practise it in the future unless that one be perfect at the outset, of what avail are happy chances, atavistic tendencies, the mellowing hand of time? But the grey caterpillar, sacrificed one day, may be succeeded on another day by a green, yellow or striped caterpillar. There you have discernment, which is quite capable of recognizing the regulation prey under very diverse garbs.
The Megachiles build their honey-jars with disks cut out of leaves; certain Anthidia make felted cotton wallets; others fashion pots out of resin. There you have instinct. Will any rash mind ever conceive the singular idea that the Leaf-cutter might very well have started working in cotton, that the cotton-wool-worker once thought or will one day think of cutting disks out of the leaves of the lilac- and the rose-tree, that the resin-kneader began with clay? Who would dare to indulge in any such theories? Each Bee has her art, her medium, to which she strictly confines herself. The first has her leaves; the second her wadding; the third her resin. None of these guilds has ever changed trades with another; and none ever will. There you have instinct, keeping the workers to their specialities. There are no innovations in their workshops, no recipes resulting from experiment, no ingenious devices, no progress from indifferent to good, from good to excellent. To-day's method is the facsimile of yesterday's; and to-morrow will know no other.
But, though the manufacturing-process is invariable, the raw material is subject to change. The plant that supplies the cotton differs in species according to the locality; the bush out of whose leaves the pieces will be cut is not the same in the various fields of operation; the tree that provides the resinous putty may be a pine, a cypress, a juniper, a cedar or a spruce, all very different in appearance. What will guide the insect in its gleaning? Discernment.
These, I think, are sufficient details of the fundamental distinction to be drawn in the insect's mentality; the distinction, that is, between instinct and discernment. If people confuse these two provinces, as they nearly always do, any understanding becomes impossible; the last glimmer of light disappears behind the clouds of interminable discussions. From an industrial point of view, let us look upon the insect as a worker thoroughly versed from birth in a craft whose essential principles never vary; let us grant that unconscious worker a gleam of intelligence which will permit it to extricate itself from the inevitable conflict of attendant circumstances; and I think that we shall have come as near to the truth as the state of our knowledge will allow for the moment.
Having thus assigned a due share both to instinct and the aberrations of instinct when the course of its different phases is disturbed, let us see what discernment is able to do in the selection of a site for the nest and materials for building it; and, leaving the Pelopaeus, upon whom it is useless to dwell any longer, let us consider other examples, picked from among those richest in variations.
The Mason-bee of the Sheds (Chalicodoma rufitarsis, PEREZ) well deserves the name which I have felt justified in giving her from her habits: she settles in numerous colonies in our sheds, on the lower surface of the tiles, where she builds huge nests which endanger the solidity of the roof. Nowhere does the insect display a greater zeal for work than in one of these colossal cities, an estate which is constantly increasing as it passes down from one generation to another; nowhere does it find a better workshop for the exercise of its industry. Here it has plenty of room: a quiet resting-place, sheltered from damp and from excess of heat or cold.
But the spacious domain under the tiles is not within the reach of all: sheds with free access and the proper sunny aspect are pretty rare. These sites fall only to the favoured of fortune. Where will the others take up their quarters? More or less everywhere. Without leaving the house in which I live, I can enumerate stone, wood, glass, metal, paint and mortar as forming the foundation of the nests. The green-house with its furnace heat in the summer and its bright light, equalling that outside, is fairly well-frequented. The Mason-bee hardly ever fails to build there each year, in squads of a few dozen apiece, now on the glass panes, now on the iron bars of the framework. Other little swarms settle in the window embrasures, under the projecting ledge of the front door or in the cranny between the wall and an open shutter. Others again, being perhaps of a morose disposition, flee society and prefer to work in solitude, one in the inside of a lock or of a pipe intended to carry the rain-water from the leads; another in the mouldings of the doors and windows or in the crude ornamentation of the stone-work. In short, the house is made use of all round, provided that the shelter be an out-of-door one; for observe that the enterprising invader, unlike the Pelopaeus, never penetrates inside our dwellings. The case of the conservatory is an exception more apparent than real: the glass building, standing wide open throughout the summer, is to the Mason-bee but a shed a little lighter than the others. There is nothing here to arouse the distrust with which anything indoors or shut up inspires her. To build on the threshold of an outer door, or to usurp its lock, a hiding-place to her fancy, is all that she allows herself; to go any farther is an adventure repugnant to her taste.
Lastly, in the case of all these dwellings, the Mason-bee is man's free tenant; her industry makes use of the products of our own industry. Can she have no other establishments? She has, beyond a doubt; she possesses some constructed on the ancient plan. On a stone the size of a man's fist, protected by the shelter of a hedge, sometimes even on a pebble in the open air, I see her building now groups of cells as large as a walnut, now domes emulating in size, shape and solidity those of her rival, the Mason-bee of the Walls.
The stone support is the most frequent, though not the only one. I have found nests, but sparsely inhabited it is true, on the trunks of trees, in the seams of the rough bark of oaks. Among those whose support was a living plant, I will mention two that stand out above all the others. The first was built in the lobe of a torch-thistle as thick as my leg; the second rested on a stalk of the opuntia, the Indian fig. Had the fierce armour of these two stout cactuses attracted the attention of the insect, which looked upon their tufts of spikes as furnishing a system of defence for its nest? Perhaps so. In any case, the attempt was not imitated; I never saw another installation of the kind. There is one definite conclusion to be drawn from my two discoveries. Despite the oddity of their structure, which is unparalleled among the local flora, the two American importations did not compel the insect to go through an apprenticeship of groping and hesitation. The one which found itself in the presence of those novel growths, and which was perhaps the first of its race to do so, took possession of their lobes and stalks just as it would have done of a familiar site. From the start, the fleshy plants from the New World suited it as well as the trunk of a native tree.
The Mason-bee of the Pebbles (Chalicodoma parietina) has none of this elasticity in the choice of a site. In her case, the smooth stone of the parched uplands is the almost invariable foundation of her structures. Elsewhere, under a less clement sky, she prefers the support of a wall, which protects the nest against the prolonged snows. Lastly, the Mason-bee of the Shrubs (Chalicodoma rufescens, PEREZ) fixes her ball of clay to a twig of any ligneous plant, from the thyme, the rock-rose and the heath to the oak, the elm and the pine. The list of the sites that suit her would almost form a complete catalogue of the ligneous flora.
The variety of places wherein the insect instals itself, so eloquent of the part played by discernment in their selection, becomes still more remarkable when it is accompanied by a corresponding variety in the architecture of the cells. This is more particularly the case with the Three-horned Osmia, who, as she uses clayey materials very easily affected by the rain, requires, like the Pelopaeus, a dry shelter for her cells, a shelter which she finds ready-made and uses just as it is, after a few touches by way of sweeping and cleansing. The homes which I see her adopt are especially the shells of Snails that have died under the stone-heaps and in the low, unmortared walls which support the cultivated earth of the hills in shelves or terraces. The use of Snail-shells is accompanied by the no less active use of the old cells of both the Mason-bee of the Sheds and of certain Anthophorae (A. pilipes, A. parietina and A. personata).
We must not forget the reed, which is highly appreciated when—a rare find—it appears under the requisite conditions. In its natural state, the plant with the mighty hollow cylinders is of no possible use to the Osmia, who knows nothing of the art of perforating a woody wall. The gallery of an internode has to be wide open before the insect can take possession of it. Also, the clean-cut stump must be horizontal, otherwise the rain would soften the fragile edifice of clay and soon lay it low; also, the stump must not be lying on the ground and must be kept at some distance from the dampness of the soil. We see therefore that, without the intervention of man, involuntary in the vast majority of cases and deliberate only on the experimenter's part, the Osmia would hardly ever find a reed-stump suited to the installation of her family. It is to her a casual acquisition, a home unknown to her race before men took it into their heads to cut reeds and make them into hurdles for drying figs in the sun.
How did the work of man's pruning-knife bring about the abandonment of the natural lodging? How was the spiral staircase of the Snail-shell replaced by the cylindrical gallery of the reed? Was the change from one kind of house to another effected by gradual transitions, by attempts made, abandoned, resumed, becoming more and more definite in their results as generation succeeded generation? Or did the Osmia, finding the cut reed that answered her requirements, instal herself there straightway, scorning her ancient dwelling, the Snail-shell? These questions called for a reply; and they have received one. Let us describe how things happened.
Near Serignan are some great quarries of coarse limestone, characteristic of the miocene formation of the Rhone valley. These have been worked for many generations. The ancient public buildings of Orange, notably the colossal frontage of the theatre whither all the intellectual world once flocked to hear Sophocles' "Oedipus Tyrannus," derive most of their material from these quarries. Other evidence confirms what the similarity of the hewn stone tells us. Among the rubbish that fills up the spaces between the tiers of seats, they occasionally discover the Marseilles obol, a bit of silver stamped with the four-spoked wheel, or a few bronze coins bearing the effigy of Augustus or Tiberius. Scattered also here and there among the monuments of antiquity are heaps of refuse, accumulations of broken stones in which various Hymenoptera, including the Three-horned Osmia in particular, take possession of the dead Snail-shell.
The quarries form part of an extensive plateau which is so arid as to be nearly deserted. In these conditions, the Osmia, at all times faithful to her birth-place, has little or no need to emigrate from her heap of stones and leave the shell for another dwelling which she would have to go and seek at a distance. Since there are heaps of stone there, she probably has no other dwelling than the Snail-shell. Nothing tells us that the present-day generations are not descended in the direct line from the generations contemporary with the quarryman who lost his as or his obol at this spot. All the circumstances seem to point to it: the Osmia of the quarries is an inveterate user of Snail-shells; so far as heredity is concerned, she knows nothing whatever of reeds. Well, we must place her in the presence of these new lodgings.
I collect during the winter about two dozen well-stocked Snail-shells and instal them in a quiet corner of my study, as I did at the time of my enquiries into the distribution of the sexes. The little hive with its front pierced with forty holes has bits of reed fitted to it. At the foot of the five rows of cylinders I place the inhabited shells and with these I mix a few small stones, the better to imitate the natural conditions. I add an assortment of empty Snail-shells, after carefully cleaning the interior so as to make the Osmia's stay more pleasant. When the time comes for nest-building, the stay-at-home insect will have, close beside the house of its birth, a choice of two habitations: the cylinder, a novelty unknown to its race; and the spiral staircase, the ancient ancestral home.
The nests were finished at the end of May and the Osmiae began to answer my list of questions. Some, the great majority, settled exclusively in the reeds; the others remained faithful to the Snail-shell or else entrusted their eggs partly to the spirals and partly to the cylinders. With the first, who were the pioneers of cylindrical architecture, there was no hesitation that I could perceive: after exploring the stump of reed for a time and recognizing it as serviceable, the insect instals itself there and, an expert from the first touch, without apprenticeship, without groping, without any tendencies bequeathed by the long practice of its predecessors, builds its straight row of cells on a very different plan from that demanded by the spiral cavity of the shell which increases in size as it goes on.
The slow school of the ages, the gradual acquisitions of the past, the legacies of heredity count for nothing therefore in the Osmia's education. Without any novitiate on its own part or that of its forebears, the insect is versed straight away in the calling which it has to pursue; it possesses, inseparable from its nature, the qualities demanded by its craft: some which are invariable and belong to the domain of instinct; others, flexible, belonging to the province of discernment. To divide a free lodging into chambers by means of mud partitions; to fill those chambers with a heap of pollen-flour, with a few sups of honey in the central part where the egg is to lie; in short, to prepare board and lodging for the unknown, for a family which the mothers have never seen in the past and will never see in the future: this, in its essential features, is the function of the Osmia's instinct. Here, everything is harmoniously, inflexibly, permanently preordained; the insect has but to follow its blind impulse to attain the goal. But the free lodging offered by chance varies exceedingly in hygienic conditions, in shape and in capacity. Instinct, which does not choose, which does not contrive, would, if it were alone, leave the insect's existence in peril. To help her out of her predicament, in these complex circumstances, the Osmia possesses her little stock of discernment, which distinguishes between the dry and the wet, the solid and the fragile, the sheltered and the exposed; which recognizes the worth or the worthlessness of a site and knows how to sprinkle it with cells according to the size and shape of the space at disposal. Here, slight industrial variations are necessary and inevitable; and the insect excels in them without any apprenticeship, as the experiment with the native Osmia of the quarries has just proved.
Animal resources have a certain elasticity, within narrow limits. What we learn from the animals' industry at a given moment is not always the full measure of their skill. They possess latent powers held in reserve for certain emergencies. Long generations can succeed one another without employing them; but, should some circumstance require it, suddenly those powers burst forth, free of any previous attempts, even as the spark potentially contained in the flint flashes forth independently of all preceding gleams. Could one who knew nothing of the Sparrow but her nest under the eaves suspect the ball-shaped nest at the top of a tree? Would one who knew nothing of the Osmia save her home in the Snail-shell expect to see her accept as her dwelling a stump of reed, a paper funnel, a glass tube? My neighbour the Sparrow, impulsively taking it into her head to leave the roof for the plane-tree, the Osmia of the quarries, rejecting her natal cabin, the spiral of the shell, for my cylinder, alike show us how sudden and spontaneous are the industrial variations of animals.
CHAPTER 7. ECONOMY OF ENERGY.
What stimulus does the insect obey when it employs the reserve powers that slumber in its race? Of what use are its industrial variations? The Osmia will yield us her secret with no great difficulty. Let us examine her work in a cylindrical habitation. I have described in full detail, in the foregoing pages, the structure of her nests when the dwelling adopted is a reed-stump or any other cylinder; and I will content myself here with recapitulating the essential features of that nest-building.
We must first distinguish three classes of reeds according to their diameter: the small, the medium-sized and the large. I call small those whose narrow width just allows the Osmia to go about her household duties without discomfort. She must be able to turn where she stands in order to brush her abdomen and rub off its load of pollen, after disgorging the honey in the centre of the heap of flour already collected. If the width of the tube does not admit of this operation, if the insect is obliged to go out and then come in again backwards in order to place itself in a favourable posture for the discharge of the pollen, then the reed is too narrow and the Osmia is rather reluctant to accept it. The middle-sized reeds and a fortiori the large ones leave the victualler entire liberty of action; but the former do not exceed the width of a cell, a width agreeing with the bulk of the future cocoon, whereas the latter, with their excessive diameter, require more than one chamber on the same floor.
When free to choose, the Osmia settles by preference in the small reeds. Here, the work of building is reduced to its simplest expression and consists in dividing the tube by means of earthen partitions into a straight row of cells. Against the partition forming the back wall of the preceding cell the mother places first a heap of honey and pollen; next, when the portion is seen to be enough, she lays an egg in the centre of it. Then and then only she resumes her plasterer's work and marks out the length of the new cell with a mud partition. This partition in its turn serves as the rear-wall of another chamber, which is first victualled and then closed; and so on until the cylinder is sufficiently colonized and receives a thick terminal stopper at its orifice. In a word, the chief characteristic of this method of nest-building, the roughest of all, is that the partition in front is not undertaken so long as the victualling is still incomplete, or, in other words, that the provisions and the egg are deposited before the Bee sets to work on the partition.
At first sight, this latter detail hardly deserves attention: is it not right to fill the pot before we put a lid on? The Osmia who owns a medium-sized reed is not at all of this opinion; and other plasterers share her views, as we shall see when we watch the Odynerus building her nest. (A genus of Mason-wasps, the essays on which have not yet been translated into English.—Translator's Note.) Here we have an excellent illustration of one of those latent powers held in reserve for exceptional occasions and suddenly brought into play, although often very far removed from the insect's regular methods. If the reed, without being of inordinate width from the point of view of the cocoon, is nevertheless too spacious to afford the Bee a suitable purchase against the wall at the moment when she is disgorging honey and brushing off her load of pollen; the Osmia altogether changes the order of her work; she sets up the partition first and then does the victualling.
All round the inside of the tube she places a ring of mud, which, as the result of her constant visits to the mortar, ends by becoming a complete diaphragm minus an orifice at the side, a sort of round dog-hole, just large enough for the insect to pass through. When the cell is thus marked out and almost wholly closed, the Osmia attends to the storing of her provisions and the laying of her eggs. Steadying herself against the margin of the hole at one time with her fore-legs and at another with her hind-legs, she is able to empty her crop and to brush her abdomen; by pressing against it, she obtains a foothold for her little efforts in these various operations. When the tube was narrow, the outer wall supplied this foothold and the earthen partition was postponed until the heap of provisions was completed and surmounted by the egg; but in the present case the passage is too wide and would leave the insect floundering helplessly in space, so the partition with its serving-hatch takes precedence of the victuals. This method is a little more expensive than the other, first in materials, because of the diameter of the reed, and secondly in time, if only because of the dog-hole, a delicate piece of mortar-work which is too soft at first and cannot be used until it has dried and become harder. Therefore the Osmia, who is sparing of her time and strength, accepts medium-sized reeds only when there are no small ones available.
The large tubes she will use only in grave emergencies and I am unable to state exactly what these exceptional circumstances are. Perhaps she decides to make use of those roomy dwellings when the eggs have to be laid at once and there is no other shelter in the neighbourhood. While my cylinder-hives gave me plenty of well-filled reeds of the first and second class, they provided me with but half-a-dozen at most of the third, notwithstanding my precaution to furnish the apparatus with a varied assortment.
The Osmia's repugnance to big cylinders is quite justified. The work in fact is longer and more costly when the tubes are wide. An inspection of a nest constructed under these conditions is enough to convince us. It now consists not of a string of chambers obtained by simple transverse partitions, but of a confused heap of clumsy, many-sided compartments, standing back to back, with a tendency to group themselves in storeys without succeeding in doing so, because any regular arrangement would mean that the ceilings possessed a span which it is not in the builder's power to achieve. The edifice is not a geometrical masterpiece and it is even less satisfactory from the point of view of economy. In the previous constructions, the sides of the reed supplied the greater part of the walls and the work was limited to one partition for each cell. Here, except at the actual periphery, where the tube itself supplies a foundation, everything has to be obtained by sheer building: the floor, the ceiling, the walls of the many-sided compartment are one and all made of mortar. The structure is almost as costly in materials as that of the Chalicodoma or the Pelopaeus.
It must be pretty difficult, too, when one thinks of its irregularity. Fitting as best she can the projecting angles of the new cell into the recessed corners of the cell already built, the Osmia runs up walls more or less curved, upright or slanting, which intersect one another at various points, so that each compartment requires a new and complicated plan of construction, which is very different from the circular-partition style of architecture, with its row of parallel dividing-disks. Moreover, in this composite arrangement, the size of the recesses left available by the earlier work to some extent decides the assessment of the sexes, for, according to the dimensions of those recesses, the walls erected take in now a larger space, the home of a female, and now a smaller space, the home of a male. Roomy quarters therefore have a double drawback for the Osmia: they greatly increase the outlay in materials; and also they establish in the lower layers, among the females, males who, because of their earlier hatching, would be much better placed near the mouth of the nest. I am convinced of it: if the Osmia refuses big reeds and accepts them only in the last resort, when there are no others, it is because she objects to additional labour and to the mixture of the sexes.
The Snail-shell, then, is but an indifferent home for her, which she is quite ready to abandon should a better offer. Its expanding cavity represents an average between the favourite small cylinder and the unpopular large cylinder, which is accepted only when there is no other obtainable. The first whorls of the spiral are too narrow to be of use to the Osmia, but the middle ones have the right diameter for cocoons arranged in single file. Here things happen as in a first-class reed, for the helical curve in no way affects the method of structure employed for a rectilinear series of cells. Circular partitions are erected at the required distances, with or without a serving-hatch, according to the diameter. These mark out the first cells, one after the other, which are reserved solely for the females. Then comes the last whorl, which is much too wide for a single row of cells; and here we once more find, exactly as in a wide reed, a costly profusion of masonry, an irregular arrangement of the cells and a mixture of the sexes.
Having said so much, let us go back to the Osmia of the quarries. Why, when I offer them simultaneously Snail-shells and reeds of a suitable size, do the old frequenters of the shells prefer the reeds, which in all probability have never before been utilized by their race? Most of them scorn the ancestral dwelling and enthusiastically accept my reeds. Some, it is true, take up their quarters in the Snail-shell; but even among these a goodly number refuse my new shells and return to their birth-place, the old Snail-shell, in order to utilize the family property, without much labour, at the cost of a few repairs. Whence, I ask, comes this general preference for the cylinder, never used hitherto? The answer can be only this: of two lodgings at her disposal the Osmia selects the one that provides a comfortable home at a minimum outlay. She economizes her strength when restoring an old nest; she economizes it when replacing the Snail-shell by the reed.
Can animal industry, like our own, obey the law of economy, the sovran law that governs our industrial machine even as it governs, at least to all appearances, the sublime machine of the universe? Let us go deeper into the question and bring other workers into evidence, those especially who, better equipped perhaps and at any rate better fitted for hard work, attack the difficulties of their trade boldly and look down upon alien establishments with scorn. Of this number are the Chalicodomae, the Mason-bees proper.
The Mason-bee of the Pebbles does not make up her mind to build a brand-new dome unless there be a dearth of old and not quite dilapidated nests. The mothers, sisters apparently and heirs-at-law to the domain, dispute fiercely for the ancestral abode. The first who, by sheer brute force, takes possession of the dome, perches upon it and, for long hours, watches events while polishing her wings. If some claimant puts in an appearance, forthwith the other turns her out with a volley of blows. In this way the old nests are employed so long as they have not become uninhabitable hovels.
Without being equally jealous of the maternal inheritance, the Mason-bee of the Sheds eagerly uses the cells whence her generation issued. The work in the huge city under the eaves begins thus: the old cells, of which, by the way, the good-natured owner yields a portion to Latreille's Osmia and to the Three-horned Osmia alike, are first made clean and wholesome and cleared of broken plaster and then provisioned and shut. When all the accessible chambers are occupied, the actual building begins with a new stratum of cells upon the former edifice, which becomes more and more massive from year to year.
The Mason-bee of the Shrubs, with her spherical nests hardly larger than walnuts, puzzled me at first. Does she use the old buildings or does she abandon them for good? To-day perplexity makes way for certainty: she uses them very readily. I have several times surprised her lodging her family in the empty rooms of a nest where she was doubtless born herself. Like her kinswoman of the Pebbles, she returns to the native dwelling and fights for its possession. Also, like the dome-builder, she is an anchorite and prefers to cultivate the lean inheritance alone. Sometimes, however, the nest is of exceptional size and harbours a crowd of occupants, who live in peace, each attending to her business, as in the colossal hives in the sheds. Should the colony be at all numerous and the estate descend to two or three generations in succession, with a fresh layer of masonry each year, the normal walnut-sized nest becomes a ball as large as a man's two fists. I have gathered on a pine-tree a nest of the Mason-bee of the Shrubs that weighed a kilogram (2.205 pounds avoirdupois.—Translator's Note.) and was the size of a child's head. A twig hardly thicker than a straw served as its support. The casual sight of that lump swinging over the spot on which I had sat down made me think of the mishap that befell Garo. (The hero of La Fontaine's fable, "Le Gland et la Citrouille," who wondered why acorns grew on such tall trees and pumpkins on such low vines, until he fell asleep under one of the latter and a pumpkin dropped upon his nose.—Translator's Note.) If such nests were plentiful in the trees, any one seeking the shade would run a serious risk of having his head smashed.
After the Masons, the Carpenters. Among the guild of wood-workers, the most powerful is the Carpenter-bee (Xylocopa violacea (Cf. "The Life of the Spider": chapter 1.—Translator's Note.)), a very large Bee of formidable appearance, clad in black velvet with violet-coloured wings. The mother gives her larvae as a dwelling a cylindrical gallery which she digs in rotten wood. Useless timber lying exposed to the air, vine-poles, large logs of fire-wood seasoning out of doors, heaped up in front of the farmhouse porch, stumps of trees, vine-stocks and big branches of all kinds are her favourite building-yards. A solitary and industrious worker, she bores, bit by bit, circular passages the width of one's thumb, as clear-cut as though they were made with an auger. A heap of saw-dust accumulates on the ground and bears witness to the severity of the task. Usually, the same aperture is the entrance to two or three parallel corridors. With several galleries there is accommodation for the entire laying, though each gallery is quite short; and the Bee thus avoids those long series which always create difficulties when the moment of hatching arrives. The laggards and the insects eager to emerge are less likely to get in each other's way.
After obtaining the dwelling, the Carpenter-bee behaves like the Osmia who is in possession of a reed. Provisions are collected, the egg is laid and the chamber is walled in front with a saw-dust partition. The work is pursued in this way until the two or three passages composing the house are completely stocked. Heaping up provisions and erecting partitions are an invariable feature of the Xylocopa's programme; no circumstance can release the mother from the duty of providing for the future of her family, in the matter both of ready-prepared food and of separate compartments for the rearing of each larva. It is only in the boring of the galleries, the most laborious part of the work, that economy can occasionally be exercised by a piece of luck. Well, is the powerful Carpenter, all unheeding of fatigue, able to take advantage of such fortunate occasions? Does she know how to make use of houses which she has not tunnelled herself? Why, yes: a free lodging suits her just as much as it does the various Mason-bees. She knows as well as they the economic advantages of an old nest that is still in good condition: she settles down, as far as possible, in her predecessors' galleries, after freshening up the sides with a superficial scraping. And she does better still. She readily accepts lodgings which have never known a drill, no matter whose. The stout reeds used in the trellis-work that supports the vines are valuable discoveries, providing as they do sumptuous galleries free of cost. No preliminary work or next to none is required with these. Indeed, the insect does not even trouble to make a side-opening, which would enable it to occupy the cavity contained within two nodes; it prefers the opening at the end cut by man's pruning-knife. If the next partition be too near to give a chamber of sufficient length, the Xylocopa destroys it, which is easy work, not to be compared with the labour of cutting an entrance through the side. In this way, a spacious gallery, following on the short vestibule made by the pruning-knife, is obtained with the least possible expenditure of energy.
Guided by what was happening on the trellises, I offered the black Bee the hospitality of my reed-hives. From the very beginning, the insect gladly welcomed my advances; each spring, I see it inspect my rows of cylinders, pick out the best ones and instal itself there. Its work, reduced to a minimum by my intervention, is limited to the partitions, the materials for which are obtained by scraping the inner sides of the reed.
As first-rate joiners, next to the Carpenter-bees come the Lithurgi, of whom my district possesses two species: L. cornutus, FAB., and L. chrysurus, BOY. By what aberration of nomenclature was the name of Lithurgus, a worker in stone, given to insects which work solely in wood? I have caught the first, the stronger of the two, digging galleries in a large block of oak that served as an arch for a stable-door; I have always found the second, who is more widely distributed, settling in dead wood—mulberry, cherry, almond, poplar—that was still standing. Her work is exactly the same as the Xylocopa's, on a smaller scale. A single entrance-hole gives access to three or four parallel galleries, assembled in a serried group; and these galleries are subdivided into cells by means of saw-dust partitions. Following the example of the big Carpenter-bee, Lithurgus chrysurus knows how to avoid the laborious work of boring, when occasion offers: I find her cocoons lodged almost as often in old dormitories as in new ones. She too has the tendency to economize her strength by turning the work of her predecessors to account. I do not despair of seeing her adopt the reed if, one day, when I possess a large enough colony, I decide to try this experiment on her. I will say nothing about L. cornutus, whom I only once surprised at her carpentering.
The Anthophorae, those children of the precipitous earthy banks, show the same thrifty spirit as the other members of the mining corporation. Three species, A. parietina, A. personata and A. pilipes, dig long corridors leading to the cells, which are scattered here and there and one by one. These passages remain open at all seasons of the year. When spring comes, the new colony uses them just as they are, provided that they are well preserved in the clayey mass baked by the sun; it increases their length if necessary, runs out a few more branches, but does not decide to start boring in new ground until the old city, which, with its many labyrinths, resembles some monstrous sponge, is too much undermined for safety. The oval niches, the cells that open on those corridors, are also profitably employed. The Anthophora restores their entrance, which has been destroyed by the insect's recent emergence; she smooths their walls with a fresh coat of whitewash, after which the lodging is fit to receive the heap of honey and the egg. When the old cells, insufficient in number and moreover partly inhabited by diverse intruders, are all occupied, the boring of new cells begins, in the extended sections of the galleries, and the rest of the eggs are housed. In this way, the swarm is settled at a minimum of expense.
To conclude this brief account, let us change the zoological setting and, as we have already spoken of the Sparrow, see what he can do as a builder. The simplest form of his nest is the great round ball of straw, dead leaves and feathers, in the fork of a few branches. It is costly in material, but can be set up anywhere, when the hole in the wall or the shelter of a tile are lacking. What reasons induced him to give up the spherical edifice? To all seeming, the same reasons that led the Osmia to abandon the Snail-shell's spiral, which requires a fatiguing expenditure of clay, in favour of the economical cylinder of the reed. By making his home in a hole in the wall, the Sparrow escapes the greater part of his work. Here, the dome that serves as a protection from the rain and the thick walls that offer resistance to the wind both become superfluous. A mere mattress is sufficient; the cavity in the wall provides the rest. The saving is great; and the Sparrow appreciates it quite as much as the Osmia.
This does not mean that the primitive art has disappeared, lost through neglect; it remains an ineffaceable characteristic of the species, ever ready to declare itself should circumstances demand it. The generations of to-day are as much endowed with it as the generations of yore; without apprenticeship, without the example of others, they have within themselves, in the potential state, the industrial aptitude of their ancestors. If aroused by the stimulus of necessity, this aptitude will pass suddenly from inaction to action. When, therefore, the Sparrow still from time to time indulges in spherical building, this is not progress on his part, as is sometimes contended; it is, on the contrary, a retrogression, a return to the ancient customs, so prodigal of labour. He is behaving like the Osmia who, in default of a reed, makes shift with a Snail-shell, which is more difficult to utilize but easier to find. The cylinder and the hole in the wall stand for progress; the spiral of the Snail-shell and the ball-shaped nest represent the starting-point.
I have, I think, sufficiently illustrated the inference which is borne out by the whole mass of analogous facts. Animal industry manifests a tendency to achieve the essential with a minimum of expenditure; after its own fashion, the insect bears witness to the economy of energy. On the one hand, instinct imposes upon it a craft that is unchangeable in its fundamental features; on the other hand, it is left a certain latitude in the details, so as to take advantage of favourable circumstances and attain the object aimed at with the least possible expenditure of time, materials and work, the three elements of mechanical labour. The problem in higher geometry solved by the Hive-bee is only a particular case—true, a magnificent case,—of this general law of economy which seems to govern the whole animal world. The wax cells, with their maximum capacity as against a minimum wall-space, are the equivalent, with the superaddition of a marvellous scientific skill, of the Osmia's compartments in which the stonework is reduced to a minimum through the selection of a reed. The artificer in mud and the artificer in wax obey the same tendency: they economize. Do they know what they are doing? Who would venture to suggest it in the case of the Bee grappling with her transcendental problem? The others, pursuing their rustic art, are no wiser. With all of them, there is no calculation, no premeditation, but simply blind obedience to the law of general harmony.
CHAPTER 8. THE LEAF-CUTTERS.
It is not enough that animal industry should be able, to a certain extent, to adapt itself to casual exigencies when choosing the site of a nest; if the race is to thrive, something else is required, something which hide-bound instinct is unable to provide. The Chaffinch, for instance, introduces a great quantity of lichen into the outer layer of his nest. This is his method of strengthening the edifice and making a stout framework in which to place first the bottom mattress of moss, fine straw and rootlets and then the soft bed of feathers, wool and down. But, should the time-honoured lichen be lacking, will the bird refrain from building its nest? Will it forgo the delight of hatching its brood because it has not the wherewithal to settle its family in the orthodox fashion?
No, the chaffinch is not perplexed by so small a matter; he is an expert in materials, he understands botanical equivalents. In the absence of the branches of the evernias, he picks the long beards of the usneas, the wartlike rosettes of the parmelias, the membranes of the stictises torn away in shreds; if he can find nothing better, he makes shift with the bushy tufts of the cladonias. As a practical lichenologist, when one species is rare or lacking in the neighbourhood, he is able to fall back on others, varying greatly in shape, colour and texture. And, if the impossible happened and lichen failed entirely, I credit the Chaffinch with sufficient talent to be able to dispense with it and to build the foundations of his nest with some coarse moss or other.
What the worker in lichens tells us the other weavers of textile materials confirm. Each has his favourite flora, which hardly ever varies when the plant is easily accessible and which can be supplemented by plenty of others when it is not. The bird's botany would be worth examining; it would be interesting to draw up the industrial herbal of each species. In this connection, I will quote just one instance, so as not to stray too far from the subject in hand.
The Red-backed Shrike (Lanius collurio), the commonest variety in my district, is noteworthy because of his savage mania for forked gibbets, the thorns in the hedgerows whereon he impales the voluminous contents of his game-bag—little half-fledged birds, small Lizards, Grasshoppers, caterpillars, Beetles—and leaves them to get high. To this passion for the gallows, which has passed unnoticed by the country-folk, at least in my part, he adds another, an innocent botanical passion, which is so much in evidence that everybody, down to the youngest bird's-nester, knows all about it. His nest, a massive structure, is made of hardly any other materials than a greyish and very fluffy plant, which is found everywhere among the corn. This is the Filago spathulata of the botanists; and the bird also makes use, though less frequently, of the Filago germanica, or common cotton-rose. Both are known in Provencal by the name herbo dou tarnagas, or Shrike-herb. This popular designation tells us plainly how faithful the bird is to its plant. To have struck the agricultural labourer, a very indifferent observer, the Shrike's choice of materials must be remarkably persistent.
Have we here a taste that is exclusive? Not in the least. Though cotton-roses of all species are plentiful on level ground, they become scarce and impossible to find on the parched hills. The bird, on its side, is not given to journeys of exploration and takes what it finds to suit it in the neighbourhood of its tree or hedge. But on arid ground, the Micropus erectus, or upright micropus, abounds and is a satisfactory substitute for the Filago so far as its tiny, cottony leaves and its little fluffy balls of flowers are concerned. True, it is short and does not lend itself well to weaver's work. A few long sprigs of another cottony plant, the Helichrysum staechas, or wild everlasting, inserted here and there, will give body to the structure. Thus does the Shrike manage when hard up for his favourite materials: keeping to the same botanical family, he is able to find and employ substitutes among the fine cotton-clad stalks.
He is even able to leave the family of the Compositae and to go gleaning more or less everywhere. Here is the result of my botanizings at the expense of his nests. We must distinguish between two genera in the Shrike's rough classification: the cottony plants and the smooth plants. Among the first, my notes mention the following: Convolvulus cantabrica, or flax-leaved bindweed; Lotus symmetricus, or bird's-foot trefoil; Teucrium polium, or poly; and the flowery heads of the Phragmites communis, or common reed. Among the second are these: Medicago lupulina, or nonesuch; Trifolium repens, or white clover; Lathyrus pratensis, or meadow lathyrus; Capsella bursa pastoris, or shepherd's purse; Vicia peregrina, or broad-podded vetch; Convolvulus arvensis, or small bindweed; Pterotheca nemausensis, a sort of hawkweed; and Poa pratensis, or smooth-stalked meadow-grass. When it is downy, the plant forms almost the whole nest, as is the case with the flax-leaved bindweed; when smooth, it forms only the framework, destined to support a crumbling mass of micropus, as is the case with the small bindweed. When making this collection, which I am far from giving as the birds' complete herbarium, I was struck by a wholly unexpected detail: of the various plants, I found only the heads still in bud; moreover, all the sprigs, though dry, possessed the green colouring of the growing plant, a sign of swift desiccation in the sun. Save in a few cases, therefore, the Shrike does not collect the dead and withered remains: it is from the growing plants that he reaps his harvest, mowing them down with his beak and leaving the sheaves to dry in the sun before using them. I caught him one day hopping about and pecking at the twigs of a Biscayan bindweed. He was getting in his hay, strewing the ground with it.
The evidence of the Shrike, confirmed by that of all the other workers—weavers, basket-makers or woodcutters—whom we may care to call as witnesses, shows us what a large part must be assigned to discernment in the bird's choice of materials for its nest. Is the insect as highly gifted? When it works with vegetable matter, is it exclusive in its tastes? Does it know only one definite plant, its special province? Or has it, for employment in its manufactures, a varied flora, in which its discernment exercises a free choice? For answers to these questions we may look, above all, to the Leaf-cutting Bees, the Megachiles. Reaumur has told the story of their industry in detail; and I refer the reader who wishes for further particulars to the master's Memoirs.
The man who knows how to use his eyes in his garden will observe, some day or other, a number of curious holes in the leaves of his lilac- and rose-trees, some of them round, some oval, as if idle but skilful hands had been at work with the pinking-iron. In some places, there is scarcely anything but the veins of the leaves left. The author of the mischief is a grey-clad Bee, a Megachile. For scissors, she has her mandibles; for compasses, producing now an oval and anon a circle, she has her eye and the pivot of her body. The pieces cut out are made into thimble-shaped wallets, destined to contain the honey and the egg: the larger, oval pieces supply the floor and sides; the smaller, round pieces are reserved for the lid. A row of these thimbles, placed one on top of the other, up to a dozen or more, though often there are less: that is, roughly, the structure of the Leaf-cutter's nest.
When taken out of the recess in which the mother has manufactured it, the cylinder of cells seems to be an indivisible whole, a sort of tunnel obtained by lining with leaves some gallery dug underground. The real thing does not correspond with its appearance: under the least pressure of the fingers, the cylinder breaks up into equal sections, which are so many compartments independent of their neighbours as regards both floor and lid. This spontaneous break up shows us how the work is done. The method agrees with those adopted by the other Bees. Instead of a general scabbard of leaves, afterwards subdivided into compartments by transverse partitions, the Megachile constructs a string of separate wallets, each of which is finished before the next is begun.
A structure of this sort needs a sheath to keep the pieces in place while giving them the proper shape. The bag of leaves, in fact, as turned out by the worker, lacks stability; its numerous pieces, not glued together, but simply placed one after the other, come apart and give way as soon as they lose the support of the tunnel that keeps them united. Later, when it spins its cocoon, the larva infuses a little of its fluid silk into the gaps and solders the pieces to one another, especially the inner ones, so much so that the insecure bag in due course becomes a solid casket whose component parts it is no longer possible to separate entirely.
The protective sheath, which is also a framework, is not the work of the mother. Like the great majority of the Osmiae, the Megachiles do not understand the art of making themselves a home straight away: they want a borrowed lodging, which may vary considerably in character. The deserted galleries of the Anthophorae, the burrows of the fat Earth-worms, the tunnels bored in the trunks of trees by the larva of the Cerambyx-beetle (The Capricorn, the essay on which has not yet been published in English.—Translator's Note.), the ruined dwellings of the Mason-bee of the Pebbles, the Snail-shell nests of the Three-horned Osmia, reed-stumps, when these are handy, and crevices in the walls are all so many homes for the Leaf-cutters, who choose this or that establishment according to the tastes of their particular genus.
For the sake of clearness, let us cease generalizing and direct our attention to a definite species. I first selected the White-girdled Leaf-cutter (Megachile albocincta, PEREZ), not on account of any exceptional peculiarities, but solely because this is the Bee most often mentioned in my notes. Her customary dwelling is the tunnel of an Earth-worm opening on some clay bank. Whether perpendicular or slanting, this tunnel runs down to an indefinite depth, where the climate would be too damp for the Bee. Besides, when the time comes for the hatching of the adult insect, its emergence would be fraught with peril if it had to climb up from a deep pit through crumbling rubbish. The Leaf-cutter, therefore, uses only the front portion of the Worm's gallery, two decimetres at most. (7.8 inches.—Translator's Note.) What is to be done with the rest of the tunnel? It is an ascending shaft, tempting to an enemy; and some underground ravager might come this way and destroy the nest by attacking the row of cells at the back.
The danger is foreseen. Before fashioning her first honey-bag, the Bee blocks the passage with a strong barricade composed of the only materials used in the Leaf-cutter's guild. Fragments of leaves are piled up in no particular order, but in sufficient quantities to make a serious obstacle. It is not unusual to find in the leafy rampart some dozens of pieces rolled into screws and fitting into one another like a stack of cylindrical wafers. For this work of fortification, artistic refinement seems superfluous; at any rate, the pieces of leaves are for the most part irregular. You can see that the insect has cut them out hurriedly, unmethodically and on a different pattern from that of the pieces intended for the cells.
I am struck with another detail in the barricade. Its constituents are taken from stout, thick, strong-veined leaves. I recognize young vine-leaves, pale-coloured and velvety; the leaves of the whitish rock-rose (Cistus albidus), lined with a hairy felt; those of the holm-oak, selected among the young and bristly ones; those of the hawthorn, smooth but tough; those of the cultivated reed, the only one of the Monocotyledones exploited, as far as I know, by the Megachiles. In the construction of cells, on the other hand, I see smooth leaves predominating, notably those of the wild briar and of the common acacia, the robinia. It would appear, therefore, that the insect distinguishes between two kinds of materials, without being an absolute purist and sternly excluding any sort of blending. The very much indented leaves, whose projections can be completely removed with a dexterous snip of the scissors, generally furnish the various layers of the barricade; the little robinia-leaves, with their fine texture and their unbroken edges, are better suited to the more delicate work of the cells. |
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