nomos L

Organic Evolution in terms of the Implicate and Explicate Orders.

Part XLVI

Hymenoptera (wasps, bees, ants) (Sequel)

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

Further REMARK on noëtics
The Implicate Order consists of immaterial noëtic patterns. These patterns are the result of noëtic reactions in the Implicate Order, that is, first of all of reactions and interactions between elementary noëtic entities, then [not temporarily meant] between these and the products, and then between the products themselves (and between the products resulting from noëtic reactions between 'earlier' products). Although we see these reactions as more or less analogues of chemical reactions, there is, in noëtic reactions no exhaustion of reactants (because exhaustion has to do with quantities, that is here : number of 'available' individuals, and these are not present in the Implicate Order). There are also no spatial limitations to reactants to react with each other : They are 'always' and 'everywhere' present in the Implicate Order. We assume that the resulting patterns remain 'existing' in the Implicate Order, albeit that here the act of 'existing' is nothing more than the act of being-a-pattern. And a noëtic entity that is qua complexity beyond the mere elementary noëtic entities is only a p a t t e r n when it is internally logically consistent and when it has a definite identity that distinguishes it from other such patterns. In this way all patterns will 'exist' in the Implicate Order as a result of the mentioned noëtic reactions. To such reactions we not only reckon those that involve two or more noëtic reactants, but also those that involve in fact only one reactant. Such a one-reactant noëtic reaction we call a (noëtic) d e r i v a t i o n. And they are supposed to be very common in the Implicate Order. All immaterial patterns strive -- as explained in the REMARKS preceding the content of the previous document -- for ontological completion, which they can only achieve by being p r o j e c t e d from the Implicate Order into the Explicate Order. Of the 'existing' noëtic patterns, most of the c o m p l e x patterns will never be projected because their content can never (as being materialized) exist in the Explicate Order. They are simply irrelevant to existence in this Order. Only those noëtic patterns that turn out to be true s t r a t e g i e s to exist and persist in the Explicate Order will be projected. In fact we can see the earlier described n o ë t i c t r a j e c t o r y as a (non-temporal) succession of formal d e r i v a t i o n s of one noëtic pattern from the other, that is, a sequence of one-reactant noëtic reactions. This noëtic tractory can repeatedly be branched, and although many branches of it will represent successively derived patterns that never will be projected into the Explicate Order because they do not represent existential strategies, many others will be so projected because they represent true existential strategies. And precisely these patterns are the 'essences' of organisms (seen as substances in the metaphysical sense), that is, they are their true, complete and specific contents, which we already earlier had called 'strategies'. And so only those branches of the overall noëtic trajectory will be projected into the Explicate Order that consist of successive existential strategies. It must be emphasized that the whole process of the advancing and branching noëtic trajectory DOES NOT TAKE PLACE IN TIME, it just consists of a branching series of formal derivations (analogous to mathematical or logical derivations) in the Implicate Order. So we c a n n o t say, for instance, that

"although the formally derived patterns in branch 'A' of the noëtic trajectory represent indeed true existential strategies, they cannot yet be projected into the Explicate Order because the ecological situation there is still not yet so structured that it can already receive the projected strategy-pattern. Only if it will finally have become so ecologically structured it can receive that pattern".

The course and the branching of the noëtic trajectory (in the Implicate Order), while it visits one noëtic-stability-field-of-some-noëtic-pattern (in the Implicate Order) after another, that is, when successively one noëtic pattern is formally derived from another, resulting in a branched sequence of derived patterns, is ABSOLUTELY TIMELESS. But upon projection of precisely that c o n t i n u o u s series of derived e x i s t e n t i a l s t r a t e g i e s, not interrupted by derived patterns that do not represent existential strategies, this series will be seen in the Explicate Order as an evolution of strategies -- and therefore of organisms -- IN SPACE AND TIME. That is, the unfolding of the sequence of derived immaterial strategy patterns along the space and time dimensions of the Explicate Order will result in a series of events that we call evolution, that is, concrete evolution, in which ecological situations and changes will play a role. And every strategy involved here is represented by an organic species (or higher-order taxon), itself represented by individual locally existing living organisms. And it is only the Explicate Order in which every thing is spatial, temporal, individual, and local. And indeed the Explicate Order is pure in this respect : There do not exist immaterial, or, as one may say, noëtic, entities anywhere in the Explicate Order. And this means that also 'minds' do not exist in the Explicate Order. So concrete organic evolution, as we see it in the Explicate Order, is the physical reflection of the corresponding (branched) sequence of formal derivations in the Implicate Order.
( End of remark on noëtics)

In Part XXXI, where we began the exposition of the evolution of the Hymenoptera, we had given an overview of this large insect Order, by presenting a diagram outlining -- as it is proposed by its authors -- the chief evolutionary courses in it, resulting in the evolutionary establishment of the larger hymenopterous groups. To see the place of the 'true wasps' (which are about to be expounded) within the Order (these true wasps being represented by all the aculeates except the bees and the ants), it is perhaps instructive to repeat the mentioned overview :

Figure above : Dendrogram showing the probable evolution of the major groups of Hymenoptera. For a magnification click (twice) on the image (and get back by the return button of the browser). (After EVANS and EBERHARD, 1973, The Wasps).
In this dendrogram the Hymenoptera are presented as consisting of two Suborders, the Symphyta and Apocrita. The latter suborder then consists of the Terebrantia and the Aculeata (In our own exposition we consider the Hymenoptera as consisting of three Suborders, the Symphyta, the Terebrantia, and the Aculeata). With respect to the feeding habit of the larvae, the Order originally consisted of phytophags (suborder Symphyta), but soon gave rise to feeders on animal food (arthropods), the terebrants and aculeates. But some representatives of the latter two groups returned to feeding on plants and their products, as is expressed in the diagram.

The evolutionary establishment of
the True Wasps
( Vespiformia sensu lato )

The question of the origin of the true wasps.

The third suborder of the Hymenoptera -- sting-bearers (Aculeata) -- is a well-defined group. However, when it becomes familiar to us in a more detailed way the distinguishing features between the Terebrantia and the Aculeata almost disappear.
If one takes into account the fact that three superfamilies are contained in the suborder Aculeata, namely the wasps, bees, and ants, among which the wasps undoubtedly represent the basic group, then the question of the origin of the Aculeata in fact boils down to the question of the origin of the wasps, or, more completely expressed, [the question] of which ways and under which circumstances these or those archaic forms -- ancestors of the Hymenoptera with a petiolate abdomen (Apocrita, to which also belong the terebrants) -- could have obtained vespine (= wasp-like) features and evolved in the direction of the Aculeata.
According to HANDLIRSCH, 1908, the groups, usually and rightly so unified as the Sting-bearing Hymenoptera (Aculeata), and which we may, on the basis of the structure of the thorax, divide into Vespiformia and Sphegiformia, undoubtedly separated off from the Terebrantia. Here, the starting point of the Aculeata, according to him, was a form still winged in both sexes, not yet constructing any independent accommodation [for the larva], and, consequently, having been in its larval stage an ectoparasite almost similar to what is still the case today in many Scoliidae and Pompilidae [both true wasps]. Later we will see, that although the vespiforms s.l. (= true wasps) originated together with the terebrants, their lines of evolution [of vespiforms and terbrants] branched off from a wider base [that is, they did not arise from a single point].
As chief distinguishing characters, separating the terebrants from the true wasps, and through them also from the other aculeates, systematists come up with in fact two of them.
The first is : The presence of a two-fold trochanter (at least on the hind legs) in the Terebrantia, and a simple, one-segmented, one in the true wasps.

When at the base of the femur, adjoining the trochanter, a constriction is visible, the trochanter is called a two-fold trochanter. Without this constriction it is called a simple trochanter. [The trochanter is a small segment connecting the femur of the leg with the 'hip' (coxa)]

As to the second distinguishing character : In the Terebrantia the to-be-laid egg is led along the ovipositor all the way down to the precize place of oviposition, whereas in the Aculeata it exits at the very base of the sting, which latter organ is homologous with the ovipositor.
Apart from being only little indicative and often badly distinguishable, the first of these two features is also not always true. Thus, for instance, there are forms (Pelecinidae) which in virtue of a series of characters undoubtedly belong to the Terebrantia, but which nevertheless possess a one-segmented trochanter. Or, the other way around, in the Rhopalosomatidae, belonging to the true wasps, the trochanter is two-segmented. So the first [alleged distinguishing] character cannot, at least in the doubtful cases, held to be decisive.
As regards the second character, connected with egg-laying, in practise it often turnes out to be impossible to determine its presence, especially when one has to do with dead and dry specimens.
According to BOUVIER, 1919, the sting of the wasps does not function in leading the eggs anymore, [this] as a result of displacement of the genital opening toward the outside, and [thus] not located at the base of a tubular organ [anymore] [meant is evidently : not located at the end of the tube, i.e. not at the apical 'beginning' ('base') of it, but at the proximal beginning of it] as in the Terebrantia. This displacement also had to evoke deep-seated changes of the behavior of the insects having undergone this. However, the question, when and under what circumstances, such a displacement took place, is not asked.
The insufficient definiteness of the mentioned distinguishing characters urged MUESEBECK, 1951, and other writers to abandon the division of the Hymenoptera under discussion into Terebrantia and Aculeata, and to confine themselves only to establish one single series of superfamilies, a series that is common to both. However, such an establishment does not contribute to a clarification of the chief evolutionary directions in the Hymenoptera. When working from the evolutionary point of view, it is very important first of all to focus on the vespine features -- i.e. features typical of true wasps -- [already] present in the behavior of these or those terebrants, and with it also to focus on the conditions under which these [vespine] features here [among certain terebrants] appear.

Elements of vespine life in terebrants

Most significant to the transformation of maternal instincts of terebrants into vespine instincts was one particular character, having appeared in [certain] terebrants as it were unexpectedly, and subsequently having been applied most widely. This is the ability to paralyze the prey. The origin of this ability must be located at least at that particular moment when the terebrants began to place their eggs at a prey that lives in the open [that is, unconcealed], although the proper elements to this were possesed by them also already in earlier times : secretion from the appendant sex glands and the stinging device itself, the ovipositor. While the incarcerated concealed prey, almost lacking the ability to move, is itself powerless to struggle successfully with terebrants, this cannot at all be said of free-living preys. It is known, for instance, how a caterpillar usually defends itself against the attack by a terebrant on it : It tries to ward it off by a violent movement of its body, to clasp it by its jaws, to sprinkle it with fluid coming from its mouth, to run away, or even to fall off from the plant. And this activity of the prey is especially dangerous to the terebrant when the egg is not simply inserted under the prey's skin, as in the case of internal parasitism, but must be placed on it such that the prey cannot reach it with its jaws, nor the ectoparasitic larva emerging from the egg. To such terebrants the ability to paralyze the prey, if only partially and transient, apparently turned out to be a necessity from the very beginning. And it is indeed observed in similar circumstances.
As a clear example of this we can give the description of I. SHEVIREV of the hunt of the terebrant Paniscus ocellaris Thoms. (an ichneumonoid) after the caterpillar of the winter night owl Agrotis segetum Schiff. (see first next Figure) :

Figure 1 : Female of the terebrant Paniscus ocellaris paralyzes a caterpillar of a night owl. (After SHEVIREV, 1912, in MALYSHEV, 1966)

" Upon detecting the female terebrant, the caterpillar makes some violent movements and speedily runs away, but the female [terebrant] hunts it down, and rising on its feet such that its thorax is directed to the prey, bends toward it the tip of its abdomen with the stretched-out borer and stings it at one of the last segments of the body. Promptly the caterpillar's movements slow down, it stops running and more and more quietly swings its lightly elevated trunk. The female mounts on it and stands on all feet holding its head toward the tail end of the caterpillar, and then attaches the egg behind its second thoracic segment. Now the female withdraws the ovipositor and walks away from the caterpillar, while the small egg, tightly attached by a stalk inserted under the skin, remains on the prey, of which the paralyzed state has already subsided, and which rolls and bends on the ground as if it were attempting to free itself from, or crush in virtue of its weight, the nasty larva" (SHEVIREV, 1912, p. 81).
From this example it is clear that the terebrant directs its paralyzing sting not precisely to the central nervous system of the prey. It does apply it to a special place, though, namely to that section of the caterpillar's body where after it the terebrant will install its egg [This does not precisely fit the above report of SHEVIREV, in which it is stated that the paralyzing sting and the oviposition take place at different sites of the caterpillar's body]. Moreover, it is clear that the effect of paralyzing in this case presents itself immediately and increases gradually. On the other hand, it goes by quickly and the stiffening of the prey stops. Such a kind of effect, without infection of the nervous system [at all], if even only transient, is hardly possible [So when indeed not the central nervous system is hit, some other parts of it must have been reached by the poison, and so (transiently) affected].
Another species of Paniscus -- P. cristatus Thoms -- in similar cases can do without paralyzing. Instead, it acts with remarkable courage and with almost inapprehensible rapidity. Also with such a hellish rapidity Tryphon incestus Holmg. goes its way on the pseudo-cater pillar of a saw-fly.
Temporarily, at the moment of oviposition, also the above mentioned Polysphincta paralyze their spiders [that is, having them paralyzed only for a short time], whereby P. eximia Schm. directs its sting to the mouth of the prey. There exists an indication that those species of true ichneumonoids (Ichneumonidae) and of cryptids (Cryptinae), of which the larvae develop as external parasites on hosts that live concealed at one place (in a cocoon, rolled-up leaves, and the like), paralyze their preys usually for ever. In such a case another result has been obtained : The paralyzed prey cannot undergo further transformation anymore and, consequently, remains at that stage in which it was infected by the terebrant.
The fact of certain terebrants having acquired more or less the ability to paralyze the prey turned out for them to be the source of peculiar behavior. Until precisely the moment of laying the egg onto the prey, that lives inside a plant [and thus automatically being more or less protected], or, living freely [not concealed] but nevertheless able to protect itself, for the adult terebrant there either could not be established a direct contact with the prey, or it would last only an instant. Now, on the other hand, when the ability to paralyze the prey had had a certain development, for the female terebrant it became possible to stay for a certain time on the body of the prey infected by it, and could make contact with that particular point where it had layed its egg. And here it began to lick up blood (hemolymph) of the prey oozing out of the wound. This subsequently gave rise, at least in some specialized terebrants, to a special inclination and method of feeding on the prey infected by them. In these conditions the act of feeding from the wound on the prey's body, resulting from the sting, got separated from the act of oviposition. Now not only the pearcing of the sting would not in anyway take place intrinsically coupled with oviposition, and now being exclusively only for paralyzing the prey, or for the feeding of the female terebrant, but now also oviposition may take place already without any stinging act at all : it may take place directly onto the body of the prey. This of course was a new, very significant step toward the development of vespine behavior.
In the in this way originated relationships we see, in addition, how the ovipositor of [some of the] the terebrants had lost its original function as being an organ to lead the egg to the oviposition site, and aquired a new function -- the origin of the sting. This transformation of the ovipositor of the terebrants into the sting of the true wasps was determined by yet another circumstance. Whereas in those primitive terebrants, such as the Pimpla's (Pimplinae, Ichneumonidae), parasitizing on larvae living concealed in wood, the egg, at the time of its being laid, goes through inside the ovipositor for all its length, now, for instance in Paniscus, the egg-to-lay goes already outside the ovipositor, along its underside. Inside the latter only moves the posterior appendage of the egg, with which it is going to be attached to the integument of the prey. And now we only have to assume the disappearance or non-development of the mentiond appendage at the posterior end of the egg, to understand that oviposition began to take place directly from the base of the ovipositor, or now already the sting. And it is precisely this what is seen in the sting-bearing Hymenoptera, the Aculeata.
In the presently discussed phase of evolution [where elements of vespine life appear in certain terebrants], in the terebrants another remarkable feature appeared, connected with the moment of oviposition. This is the ability of the female to instinctively regulate the gender of the egg-to-be-laid by it, that is, letting it depend on (self-controlled) circumstances. Thus, Pimpla instigator F. and other species, according to the investigations of SEVIREV (1913), PICARD, 1921, and others, can, on encountering larger hosts, lay eggs from which develop females, and in the other case eggs giving rise to males.

If the female applies sperm from the seed purse to the egg to be laid, then a female individual will develop from it. In the other case (when sperm is not applied) a male will develop.

This truly remarkable phenomenon of the instinct in terebrants became widely distributed in the higher forms, where the gender of the to-be-born individual is determined, as was wonderfully shown by FABRE, by the size of the chamber assigned to this individual, and by the amount of provisions beforehand prepared for it, and by other circumstances.

Having now seen the appearance of vespine features already in some terebrants, we will now (next document) turn to the evolutionary development (diversification) of the True Wasps, starting with the Semi-vespine (bethyloid) Phase.

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