Now for the provisions and the partition-walls. Here the order of the work changes according to the diameter of the cylinder. My glass tubes vary greatly in dimensions. The largest have an inner width of a dozen millimetres (Nearly half an inch.--Translator's Note.); the narrowest measure six or seven. (About a quarter of an inch.--Translator's Note.) In the latter, if the bottom suit her, the Osmia sets to work bringing pollen and honey. If the bottom do not suit her, if the sorghum-pith plug with which I have closed the rear-end of the tube be too irregular and badly-joined, the Bee coats it with a little mortar. When this small repair is made, the harvesting begins.
In the wider tubes, the work proceeds quite differently. At the moment when the Osmia disgorges her honey and especially at the moment when, with her hind-tarsi, she rubs the pollen-dust from her ventral brush, she needs a narrow aperture, just big enough to allow of her passage. I imagine that in a straitened gallery the rubbing of her whole body against the sides gives the harvester a support for her brushing-work.
In a spacious cylinder this support fails her; and the Osmia starts with creating one for herself, which she does by narrowing the channel.
Whether it be to facilitate the storing of the victuals or for any other reason, the fact remains that the Osmia housed in a wide tube begins with the partitioning.
Her division is made by a dab of clay placed at right angles to the axis of the cylinder, at a distance from the bottom determined by the ordinary length of a cell. The wad is not a complete round; it is more crescent-shaped, leaving a circular space between it and one side of the tube. Fresh layers are swiftly added to the dab of clay; and soon the tube is divided by a partition which has a circular opening at the side of it, a sort of dog-hole through which the Osmia will proceed to knead the Bee-bread. When the victualling is finished and the egg laid upon the heap, the whole is closed and the filled-up partition becomes the bottom of the next cell. Then the same method is repeated, that is to say, in front of the just completed ceiling a second partition is built, again with a side-passage, which is stouter, owing to its distance from the centre, and better able to withstand the numerous comings and goings of the housewife than a central orifice, deprived of the direct support of the wall, could hope to be. When this partition is ready, the provisioning of the second cell is effected; and so on until the wide cylinder is completely stocked.
The building of this preliminary party-wall, with a narrow, round dog-hole, for a chamber to which the victuals will not be brought until later is not restricted to the Three-horned Osmia; it is also frequently found in the case of the Horned Osmia and of Latreille's Osmia. Nothing could be prettier than the work of the last-named, who goes to the plants for her material and fashions a delicate sheet in which she cuts a graceful arch. The Chinaman partitions his house with paper screens; Latreille's Osmia divides hers with disks of thin green cardboard perforated with a serving-hatch which remains until the room is completely furnished. When we have no glass houses at our disposal, we can see these little architectural refinements in the reeds of the hurdles, if we open them at the right season.
By splitting the bramble-stumps in the course of July, we perceive also that the Three-pronged Osmia notwithstanding her narrow gallery, follows the same practice as Latreille's Osmia, with a difference. She does not build a party-wall, which the diameter of the cylinder would not permit; she confines herself to putting up a frail circular pad of green putty, as though to limit, before any attempt at harvesting, the space to be occupied by the Bee-bread, whose depth could not be calculated afterwards if the insect did not first mark out its confines.
If, in order to see the Osmia's nest as a whole, we split a reed lengthwise, taking care not to disturb its contents; or, better still, if we select for examination the string of cells built in a glass tube, we are forthwith struck by one detail, namely, the uneven distances between the partitions, which are placed almost at right angles to the axis of the cylinder. It is these distances which fix the size of the chambers, which, with a similar base, have different heights and consequently unequal holding-capacities. The bottom partitions, the oldest, are farther apart; those of the front part, near the orifice, are closer together. Moreover, the provisions are plentiful in the loftier cells, whereas they are niggardly and reduced to one-half or even one-third in the cells of lesser height. Let me say at once that the large cells are destined for the females and the small ones for the males.
DISTRIBUTION OF THE SEXES.
Does the insect which stores up provisions proportionate to the needs of the egg which it is about to lay know beforehand the sex of that egg? Or is the truth even more paradoxical? What we have to do is to turn this suspicion into a certainty demonstrated by experiment. And first let us find out how the sexes are arranged.
It is not possible to ascertain the chronological order of a laying, except by going to suitably-chosen species. Fortunately there are a few species in which we do not find this difficulty: these are the Bees who keep to one gallery and build their cells in storeys. Among the number are the different inhabitants of the bramble-stumps, notably the Three-pronged Osmiae, who form an excellent subject for observation, partly because they are of imposing size--bigger than any other bramble-dwellers in my neighbourhood--partly because they are so plentiful.
Let us briefly recall the Osmia's habits. Amid the tangle of a hedge, a bramble-stalk is selected, still standing, but a mere withered stump.
In this the insect digs a more or less deep tunnel, an easy piece of work owing to the abundance of soft pith. Provisions are heaped up right at the bottom of the tunnel and an egg is laid on the surface of the food: that is the first-born of the family. At a height of some twelve millimetres (About half an inch.--Translator's Note.), a partition is fixed. This gives a second storey, which in its turn receives provisions and an egg, the second in order of primogeniture.
And so it goes on, storey by storey, until the cylinder is full. Then the thick plug of the same green material of which the partitions are formed closes the home and keeps out marauders.
In this common cradle, the chronological order of births is perfectly clear. The first-born of the family is at the bottom of the series; the last-born is at the top, near the closed door. The others follow from bottom to top in the same order in which they followed in point of time. The laying is numbered automatically; each cocoon tells us its respective age by the place which it occupies.
A number of eggs bordering on fifteen represents the entire family of an Osmia, and my observations enable me to state that the distribution of the sexes is not governed by any rule. All that I can say in general is that the complete series begins with females and nearly always ends with males. The incomplete series--those which the insect has laid in various places--can teach us nothing in this respect, for they are only fragments starting we know not whence; and it is impossible to tell whether they should be ascribed to the beginning, to the end, or to an intermediate period of the laying. To sum up: in the laying of the Three-pronged Osmia, no order governs the succession of the sexes; only, the series has a marked tendency to begin with females and to finish with males.
The mother occupies herself at the start with the stronger sex, the more necessary, the better-gifted, the female sex, to which she devotes the first flush of her laying and the fullness of her vigour; later, when she is perhaps already at the end of her strength, she bestows what remains of her maternal solicitude upon the weaker sex, the less-gifted, almost negligible male sex. There are, however, other species where this law becomes absolute, constant and regular.
In order to go more deeply into this curious question I installed some hives of a new kind on the sunniest walls of my enclosure. They consisted of stumps of the great reed of the south, open at one end, closed at the other by the natural knot and gathered into a sort of enormous pan-pipe, such as Polyphemus might have employed. The invitation was accepted: Osmiae came in fairly large numbers, to benefit by the queer installation.
Three Osmiae especially (O. Tricornis, Latr., O. cornuta, Latr., O.
Latreillii, Spin.) gave me splendid results, with reed-stumps arranged either against the wall of my garden, as I have just said, or near their customary abode, the huge nests of the Mason-bee of the Sheds.
One of them, the Three-horned Osmia, did better still: as I have described, she built her nests in my study, as plentifully as I could wish.
We will consult this last, who has furnished me with documents beyond my fondest hopes, and begin by asking her of how many eggs her average laying consists. Of the whole heap of colonized tubes in my study, or else out of doors, in the hurdle-reeds and the pan-pipe appliances, the best-filled contains fifteen cells, with a free space above the series, a space showing that the laying is ended, for, if the mother had any more eggs available, she would have lodged them in the room which she leaves unoccupied. This string of fifteen appears to be rare; it was the only one that I found. My attempts at indoor rearing, pursued during two years with glass tubes or reeds, taught me that the Three-horned Osmia is not much addicted to long series. As though to decrease the difficulties of the coming deliverance, she prefers short galleries, in which only a part of the laying is stacked. We must then follow the same mother in her migration from one dwelling to the next if we would obtain a complete census of her family. A spot of colour, dropped on the Bee's thorax with a paint-brush while she is absorbed in closing up the mouth of the tunnel, enables us to recognize the Osmia in her various homes.
In this way, the swarm that resided in my study furnished me, in the first year, with an average of twelve cells. Next year, the summer appeared to be more favourable and the average became rather higher, reaching fifteen. The most numerous laying performed under my eyes, not in a tube, but in a succession of Snail-shells, reached the figure of twenty-six. On the other hand, layings of between eight and ten are not uncommon. Lastly, taking all my records together, the result is that the family of the Osmia fluctuates roundabout fifteen in number.
I have already spoken of the great differences in size apparent in the cells of one and the same series. The partitions, at first widely spaced, draw gradually nearer to one another as they come closer to the aperture, which implies roomy cells at the back and narrow cells in front. The contents of these compartments are no less uneven between one portion and another of the string. Without any exception known to me, the large cells, those with which the series starts, have more abundant provisions than the straitened cells with which the series ends. The heap of honey and pollen in the first is twice or even thrice as large as that in the second. In the last cells, the most recent in date, the victuals are but a pinch of pollen, so niggardly in amount that we wonder what will become of the larva with that meagre ration.
One would think that the Osmia, when nearing the end of the laying, attaches no importance to her last-born, to whom she doles out space and food so sparingly. The first-born receive the benefit of her early enthusiasm: theirs is the well-spread table, theirs the spacious apartments. The work has begun to pall by the time that the last eggs are laid; and the last-comers have to put up with a scurvy portion of food and a tiny corner.
The difference shows itself in another way after the cocoons are spun.
The large cells, those at the back, receive the bulky cocoons; the small ones, those in front, have cocoons only half or a third as big.
Before opening them and ascertaining the sex of the Osmia inside, let us wait for the transformation into the perfect insect, which will take place towards the end of summer. If impatience get the better of us, we can open them at the end of July or in August. The insect is then in the nymphal stage; and it is easy, under this form, to distinguish the two sexes by the length of the antennae, which are larger in the males, and by the glassy protuberances on the forehead, the sign of the future armour of the females. Well, the small cocoons, those in the narrow front cells, with their scanty store of provisions, all belong to males; the big cocoons, those in the spacious and well-stocked cells at the back, all belong to females.
The conclusion is definite: the laying of the Three-horned Osmia consists of two distinct groups, first a group of females and then a group of males.
With my pan-pipe apparatus displayed on the walls of my enclosure and with old hurdle-reeds left lying flat out of doors, I obtained the Horned Osmia in fair quantities. I persuaded Latreille's Osmia to build her nest in reeds, which she did with a zeal which I was far from expecting. All that I had to do was to lay some reed-stumps horizontally within her reach, in the immediate neighbourhood of her usual haunts, namely, the nests of the Mason-bee of the Sheds. Lastly, I succeeded without difficulty in making her build her nests in the privacy of my study, with glass tubes for a house. The result surpassed my hopes.
With both these Osmiae, the division of the gallery is the same as with the Three-horned Osmia. At the back are large cells with plentiful provisions and widely-spaced partitions; in front, small cells, with scanty provisions and partitions close together. Also, the larger cells supplied me with big cocoons and females; the smaller cells gave me little cocoons and males. The conclusion therefore is exactly the same in the case of all three Osmiae.
These conclusions, as my notes show, apply likewise, in every respect, to the various species of Mason-bees; and one clear and simple rule stands out from this collection of facts. Apart from the strange exception of the Three-pronged Osmia, who mixes the sexes without any order, the Bees whom I studied and probably a crowd of others produce first a continuous series of females and then a continuous series of males, the latter with less provisions and smaller cells. This distribution of the sexes agrees with what we have long known of the Hive-bee, who begins her laying with a long sequence of workers, or sterile females, and ends it with a long sequence of males. The analogy continues down to the capacity of the cells and the quantities of provisions. The real females, the Queen-bees, have wax cells incomparably more spacious than the cells of the males and receive a much larger amount of food. Everything therefore demonstrates that we are here in the presence of a general rule.
OPTIONAL DETERMINATION OF THE SEXES.
But does this rule express the whole truth? Is there nothing beyond a laying in two series? Are the Osmiae, the Chalicodomae and the rest of them fatally bound by this distribution of the sexes into two distinct groups, the male group following upon the female group, without any mixing of the two? Is the mother absolutely powerless to make a change in this arrangement, should circumstances require it?
The Three-pronged Osmia already shows us that the problem is far from being solved. In the same bramble-stump, the two sexes occur very irregularly, as though at random. Why this mixture in the series of cocoons of a Bee closely related to the Horned Osmia and the Three-horned Osmia, who stack theirs methodically by separate sexes in the hollow of a reed? What the Bee of the brambles does cannot her kinswomen of the reeds do too? Nothing, so far as I know, explains this fundamental difference in a physiological act of primary importance.
The three Bees belong to the same genus; they resemble one another in general outline, internal structure and habits; and, with this close similarity, we suddenly find a strange dissimilarity.
There is just one thing that might possibly arouse a suspicion of the cause of this irregularity in the Three-pronged Osmia's laying. If I open a bramble-stump in the winter to examine the Osmia's nest, I find it impossible, in the vast majority of cases, to distinguish positively between a female and a male cocoon: the difference in size is so small.
The cells, moreover, have the same capacity: the diameter of the cylinder is the same throughout and the partitions are almost always the same distance apart. If I open it in July, the victualling-period, it is impossible for me to distinguish between the provisions destined for the males and those destined for the females. The measurement of the column of honey gives practically the same depth in all the cells.
We find an equal quantity of space and food for both sexes.
This result makes us foresee what a direct examination of the two sexes in the adult form tells us. The male does not differ materially from the female in respect of size. If he is a trifle smaller, it is scarcely noticeable, whereas, in the Horned Osmia and the Three-horned Osmia, the male is only half or a third the size of the female, as we have seen from the respective bulk of their cocoons. In the Mason-bee of the Walls there is also a difference in size, though less pronounced.
The Three-pronged Osmia has not therefore to trouble about adjusting the dimensions of the dwelling and the quantity of the food to the sex of the egg which she is about to lay; the measure is the same from one end of the series to the other. It does not matter if the sexes alternate without order: one and all will find what they need, whatever their position in the row. The two other Osmiae, with their great disparity in size between the two sexes, have to be careful about the twofold consideration of board and lodging.
The more I thought about this curious question, the more probable it appeared to me that the irregular series of the Three-pronged Osmia and the regular series of the other Osmiae and of the Bees in general were all traceable to a common law. It seemed to me that the arrangement in a succession first of females and then of males did not account for everything. There must be something more. And I was right: that arrangement in series is only a tiny fraction of the reality, which is remarkable in a very different way. This is what I am going to prove by experiment.
The succession first of females and then of males is not, in fact, invariable. Thus, the Chalicodoma, whose nests serve for two or three generations, ALWAYS lays male eggs in the old male cells, which can be recognized by their lesser capacity, and female eggs in the old female cells of more spacious dimensions.
This presence of both sexes at a time, even when there are but two cells free, one spacious and the other small, proves in the plainest fashion that the regular distribution observed in the complete nests of recent production is here replaced by an irregular distribution, harmonizing with the number and holding-capacity of the chambers to be stocked. The Mason-bee has before her, let me suppose, only five vacant cells: two larger and three smaller. The total space at her disposal would do for about a third of the laying. Well, in the two large cells, she puts females; in the three small cells she puts males.
As we find the same sort of thing in all the old nests, we must needs admit that the mother knows the sex of the eggs which she is going to lay, because that egg is placed in a cell of the proper capacity. We can go further, and admit that the mother alters the order of succession of the sexes at her pleasure, because her layings, between one old nest and another, are broken up into small groups of males and females according to the exigencies of space in the actual nest which she happens to be occupying.
Here then is the Chalicodoma, when mistress of an old nest of which she has not the power to alter the arrangement, breaking up her laying into sections comprising both sexes just as required by the conditions imposed upon her. She therefore decides the sex of the egg at will, for, without this prerogative, she could not, in the chambers of the nest which she owes to chance, deposit unerringly the sex for which those chambers were originally built; and this happens however small the number of chambers to be filled.
When the mother herself founds the dwelling, when she lays the first rows of bricks, the females come first and the males at the finish.
But, when she is in the presence of an old nest, of which she is quite unable to alter the general arrangement, how is she to make use of a few vacant rooms, the large and small alike, if the sex of the egg be already irrevocably fixed? She can only do so by abandoning the arrangement in two consecutive rows and accommodating her laying to the varied exigencies of the home. Either she finds it impossible to make an economical use of the old nest, a theory refuted by the evidence, or else she determines at will the sex of the egg which she is about to lay.
The Osmiae themselves will furnish the most conclusive evidence on the latter point. We have seen that these Bees are not generally miners, who themselves dig out the foundation of their cells. They make use of the old structures of others, or else of natural retreats, such as hollow stems, the spirals of empty shells and various hiding-places in walls, clay or wood. Their work is confined to repairs to the house, such as partitions and covers. There are plenty of these retreats; and the insects would always find first-class ones if it thought of going any distance to look for them. But the Osmia is a stay-at-home: she returns to her birthplace and clings to it with a patience extremely difficult to exhaust. It is here, in this little familiar corner, that she prefers to settle her progeny. But then the apartments are few in number and of all shapes and sizes. There are long and short ones, spacious ones and narrow. Short of expatriating herself, a Spartan course, she has to use them all, from first to last, for she has no choice. Guided by these considerations, I embarked on the experiments which I will now describe.
I have said how my study became a populous hive, in which the Three-horned Osmia built her nests in the various appliances which I had prepared for her. Among these appliances, tubes, either of glass or reed, predominated. There were tubes of all lengths and widths. In the long tubes, entire or almost entire layings, with a series of females followed by a series of males, were deposited. As I have already referred to this result, I will not discuss it again. The short tubes were sufficiently varied in length to lodge one or other portion of the total laying. Basing my calculations on the respective lengths of the cocoons of the two sexes, on the thickness of the partitions and the final lid, I shortened some of these to the exact dimensions required for two cocoons only, of different sexes.
Well, these short tubes, whether of glass or reed, were seized upon as eagerly as the long tubes. Moreover, they yielded this splendid result: their contents, only a part of the total laying, always began with female and ended with male cocoons. This order was invariable; what varied was the number of cells in the long tubes and the proportion between the two sorts of cocoons, sometimes males predominating and sometimes females.
When confronted with tubes too small to receive all her family, the Osmia is in the same plight as the Mason-bee in the presence of an old nest. She thereupon acts exactly as the Chalicodoma does. She breaks up her laying, divides it into series as short as the room at her disposal demands; and each series begins with females and ends with males. This breaking up, on the one hand, into sections in all of which both sexes are represented and the division, on the other hand, of the entire laying into just two groups, one female, the other male, when the length of the tube permits, surely provide us with ample evidence of the insect's power to regulate the sex of the egg according to the exigencies of space.
And besides the exigencies of space one might perhaps venture to add those connected with the earlier development of the males. These burst their cocoons a couple of weeks or more before the females; they are the first who hasten to the sweets of the almond-tree. In order to release themselves and emerge into the glad sunlight without disturbing the string of cocoons wherein their sisters are still sleeping, they must occupy the upper end of the row; and this, no doubt, is the reason that makes the Osmia end each of her broken layings with males. Being next to the door, these impatient ones will leave the home without upsetting the shells that are slower in hatching.
I had offered at the same time to the Osmiae in my study some old nests of the Mason-bee of the Shrubs, which are clay spheroids with cylindrical cavities in them. These cavities are formed, as in the old nests of the Mason-bee of the Pebbles, of the cell properly so-called and of the exit-way which the perfect insect cut through the outer coating at the time of its deliverance. The diameter is about 7 millimetres (.273 inch.--Translator's Note.); their depth at the centre of the heap is 23 millimetres (.897 inch.--Translator's Note.) and at the edge averages 14 millimetres. (.546 inch.--Translator's Note.)
The deep central cells receive only the females of the Osmia; sometimes even the two sexes together, with a partition in the middle, the female occupying the lower and the male the upper storey. Lastly, the deeper cavities on the circumference are allotted to females and the shallower to males.
We know that the Three-horned Osmia prefers to haunt the habitations of the Bees who nidify in populous colonies, such as the Mason-bee of the Sheds and the Hairy-footed Anthophora, in whose nests I have noted similar facts.