Organic Evolution in terms of the Implicate and Explicate Orders.


Hymenoptera (wasps, bees, ants) (Sequel)

The evolutionary diversification in the Order Hymenoptera in terms of Strategies (Sequel).

Evolutionary establisment of the Bees (Apoidea)

Secondary-apoine Phase of the Vespoid Type

For a further understanding of the evolution of nesting instincts of the masarids (Masaridae), [nesting instincts] still more approaching to the apoine type, the observations of Ferton on the habits of Ceramius  tuberculifer Gauss., about which we partly already have spoken earlier, are very instructive. In the Pyreneans (Monlouis) this masarid populates in colonies [aggregations] in hard clayey soil, covered with scarce grassy vegetation, and always in the neighborhood of water. The nearness of water turns out to be necessary to these wasps in order to soften with the help of droplets the hard earth in which they dig. In the second half of July, in which the observations were carried out, the work [of the wasps] was in full swing. Nearby the colony, counting many hundreds of individuals, a small basin of clear water was the place of their constant visiting. The number of individuals bringing-in water[-loads] simultaneously was on average 7-8, but flights-to and flights-from were observed to take place constantly. From the first portions of material obtained from digging out the burrow, an entrance-tube was constructed. It is built in determined order and renewed every time when damaged or destroyed. This tube can resist heavy rains lasting for several hours.

About the meaning and further destiny of this tube the investigator says nothing. Will it, perhaps, merge into the final closure of the nest, or is it, finally, stock-material for the construction of cells?

The cells of C. tuberculifer, lying horizontally in the depth of the burrow, are unified into one whole group. Their outer surface being rough but of regular form, shows individual pieces of cement of which it is made. But on the inside the walls of the cell are polished.
Before preparing provisions, this species deposits an egg in the cell, but the egg is not suspended on a thread as in the nests of Odynerus and Eumenes, but is provisionally deposited on the floor of the cell. The honey-dough having a firm consistency is formed as a pie recalling the shape of a short-necked retort with a widened apical neck. See next Figure.

Figure 1 :  Cell [bottom image] and provisions [upper image] prepared by the masarid Ceramius  tuberculifer Gauss.
(After FERTON, 1901, in MALYSHEV, 1966)

The next Figure depicts a species of the genus Ceramius :

Figure 1a :  Ceramius  lusitanicus.  Family Pompilidae, spider-wasps.  ca. 15 mm.
(After CHINERY, in Nieuwe Insektengids, 1987)

The mother-wasp sticks the egg, laid earlier, onto this projection, with which the just hatched larva starts its feeding. The successive loads of provisions, brought to the cell by the mother, are not mixed up and remain separate from each other, the one following upon the other in a horizontal direction. Having finished provisioning of the cell, the mother closes it with mineral cement. Later, the whole group of constructed and provisioned cells is closed with a common cover of cement.
If one opens a finished cell of this masarid, while not having seen the female-builder, and not knowing the order in which the provisioning had proceeded, then one might easily come to the conclusion that it was built by one or another bee, and certainly not by a wasp. Only the method of work in provisioning the cell (oviposition before preparing provisions, with subsequent transposition of the egg onto the provisions), totally not typical of bees, would have forced one to abandon such a conclusion, at least until clarification of the true nature of the female-builder.
So, only the order of acts in provisioning the cell distinguishes the nesting-instincts of the masarids of the type C. tuberculifer (or C. lusitanicus Klug., as they are described by Ferton) from purely apoine instincts. And this difference is not that essential, as to not reckon the masarids, as to their nesting behavior, also to the bees, but [admittedly] special bees that still have preserved a characteristic feature of their vespoid ancestors -- depositing of the egg before preparing provisions.
The just considered habits of the masarids (provisionally laying the egg in an empty cell, preparing, after that, specially kneadable honey-provisions, and then transposing the egg onto these provisions) are the basic biological features of the present second[ary]-apoine phase of the vespoid type. Insofar as it could be clarified, with this phase the evolution of the instincts of masarids came to an end. However, in the light of what has been expounded, we may easily think of a further change of them toward typical apoine habits. This may take place through a certain simplification :  by way of cancellation of the act of provisional oviposition in the empty cell, and connected with this, the subsequent transposition of the egg onto earlier prepared provisions. Accordingly, in this supposed new phase the egg must be laid only after the preparation and formation of the honey-bun, either directly onto it, or as before, on the wall of the cell nearby the provisions. The latter supposition is more probable. So far, then, these typically apoine instincts are not demonstrated in masarids, and all of them, as far as known, lay the egg before preparing provisions.
To what has been said we must add that the nesting-behavior of a series of masarids (Jugurtia, Trimeria, Quartinia, and others) are still almost or totally unknown.

There exists an indication (Williams, 1927), based on very succinct and incomplete data, according to which the small masarid Euparagia  scutellaris Cress. stocks its cell with larvae of snout-beetles. Meanwhile, in the complex natural conditions, it is, without opening up the cells-to-be-provisioned, not possible to hold that precisely the mentioned masarid, and not some other wasp has hunted in the neighborhood after snout-beetle larvae.
Similar assertions have been made for
Pseudomasaris, allegedly provisioning their cells with caterpillars. Today it is definitely known that also these masarids do not form an exception, they also provision their cells with honey-food.

In the considered behavioral features of the solitary plaited-winged wasps (Eumenidae and Masaridae) we should see the demonstration of an unusual evolutionary plasticity of them. On the basis of this plasticity, as it was already said earlier, lies the cancellation of certain acts from the chain of serial acts, and the subsequent coupling together of the remaining ones. Such a kind of transformation of instincts, not having demanded essential structural changes, in many cases turned out to be very effective as to their results. Such a simple method of transformation of maternal instincts in wasps is undoubtedly expressed in the acceleration of their changes. This also already follows from the very diversity of maternal instincts in the individual wasp-genera Ammophila, Odynerus, Synagris, and many others, dealt with earlier.
While until now we considered chiefly the rearing-instincts of wasps, it is now necessary to adress our attention to the development of their building-activity. The first stages of it were given earlier. In its simplest form it was expressed in adapting an existing ready-made cavity in behalf of a cell, and especially in the ability to construct an opening in the cells. Working in conditions of insufficient moisture (on the sunny side of clayey slopes, and so on), the wasps had, and, evidently, quickly, to acquire the ability to bring water to the construction site, in order to moisten with it the dry building-material and prepare out of it a plastic mass. This also gave them the possibility to use the latter not only for closing the openings of the cells, but also for covering with it the whole nest cavity, as for instance in  Ancistrocerus  callosus Thoms. See next Figure.

Figure 2 :  Plastic [mouldable] clay, prepared by the wasp  Ancistrocerus  callosus Thoms., covers, from the inside, the walls of the cells from all sides.  (After MALYSHEV, 1966)

As a result, the ability to built roofs of the cells was transformed into the ability to model cells, as this already appeared among odyners (Odynerus  laevipes Shuk.) and especially in masarids.

Having now considered apoine behavioral elements in certain vespoid wasps, we will, in the next document turn to sphecoid wasps, and see where, in them, apoine behavioral elements may be found.

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