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.