nomos LXII

Organic Evolution in terms of the Implicate and Explicate Orders.

Part LVIII

Hymenoptera (wasps, bees, ants) (Sequel)

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

Familial Ectoparasitic (Hemi-Formicoid) Phase

As a result of the given analysis it became clear that the highly developed form of familial life of hymenoptera, coming close to the formicoid (ant) way of life, appeared only in two [evolutionary] directions : Among chalcidoid terebrants of the family Eulophidae and among bethyloid wasps of the group Scleroderminae. Not in any other hymenoptera is such a characteristic form of familial life encountered. The simularity in the way of life and habits between these mentioned groups is remarkable. Although, based on morphological data we can openly say that the chalcidoids stand farther away from the ants, the features of their life and behavior on this high stage of [evolutionary] development is very indicative : They cast definite light on that particular path which led the chalcidoids to the highest stage of familial life in all terebrants and made them into, we could say, look-alikes of the scleroderms, and together with this brought these or others close to the ants.

Among the mentioned chalcidoids on the foremost place here do step in only Melittobia, and of them first of all Melittobia acasta WALK. This eulophid chiefly attacks preys that live in special hide-aways. She namely parasitizes on fully-grown, having concluded feeding, larvae (more rarely on prepupa and pupae) of solitary wasps and bees, nesting in dry, well sunlit places, for example in overhanging reed roofs, in clay walls and cracks, or in skillfully by themselves modelled nests in open places. These are chiefly constructions by Odynerus, Trypoxylon, Pelopeus, and Agenia [I hope that I have spelled the latter two names correctly], of wasps, or nests of Megachile and Antophora [Also here I hope that I have spelled the last name correctly], of bees.
In order to penetrate into such special shelters, well protected at all sides, except the entrance opening that is also closed later by the constructrix, the melittobia has grown accustomed to a special method : She penetrates into the nest of the wasp or bee at an appropriate time, when the cell is still open and the provisioning continues. The minute terebrant, having a length of not more than 1-1.25 mm, is easily hidden among the roughness of the walls and in the aggregation of provisions. As a result, the constructrix, not having destroyed the melittobia, locks it up in one room together with her just created offspring now destined to die.
Now finding itself in the cell of a wasp or bee, a cell with provisions and an egg, the melittobia patiently waits, apparently not eating anything as long as the emerged host larva grows and consumes its provisions. In line with the progression of growth of the [host] larva, the melittobia stays with it proportionally longer, for a long time palpating it with the antennae. Awaiting for the right moment, the melittobia, with the help of its very short ovipositor, begins to apply stings at the prey's body. For this she holds the ovipositor vertically to the integument of the prey, holds itself to it with all of her legs, presses with her abdomen and makes drilling movements with all of her body, or simply raises and lowers her abdomen. Directly after this, she holds her mouth at the skin of the prey at the places of the stings and licks at them, evidently consuming the blood oozing out. As a result of this her abdomen gets significantly bigger. See next Figure.

Figure 1 : Two females of Melittobia acasta WALK. on a fully-grown larva of the leaf-cutter bee Megachile bombycina L.
(After MALYSHEV, 1966)

On the spot of the sting, inflicted by the melittobia, after a day or two a dark brown, and later almost black, stain appears from which the destitute larva can never separate itself. Normally there is a various number of such stains, often very much of them, and they are distributed over different places of the body of the prey. The larva that is attacked by the melittobia may for much (hundreds or thousands times) surpass in size the melittobia itself, but this doesn't change things. The stings of the melittobia give off a paralyzing effect on the prey : It already does not twine a cocoon anymore, becomes completely immobile and flaccid. Its skin more and more wrinkles, and the size markedly diminishes. But the larva does not directly die. In it life for long weakly smoulders. In artificial conditions in the course of many months it may remain soft and fresh, but in natural conditions events go faster.
As soon as the prey becomes calm, the chalcidoid starts ovipositing onto it and alternately administers stings and oviposits. The eggs are elongated, with one end narrowed, and so minute that for the unaided eye, at least as lying on the body of the host, they are invisible. They are placed onto the integument of the prey without any order, sometimes 2-3 in a row, sometimes individually. They are not attached with the help of anything, and are displaced at the slightest touch. On a single host larva the melittobia lays many of them. The larger the prey, the more eggs are laid onto it. Thus onto the small larva of Trypoxylon figulus L. 30-50 eggs are being laid, whereas onto the larger larva of the leaf-cutter bee Megachile bombycina RAD. up to 100 and more. Oviposition is repeatedly carried out in the course of several days, such that when onto the same prey the last eggs have been laid, the larvae, having hatched from the first eggs, may conclude their development and even transform into pupae, and develop into adult melittobias.
Having done the work in one cell [of the nest of the bee or wasp], the melittobia gnaws through the wall of the cell with its minute jaws, and pentrates the adjacent cell where the host larva has finished at this moment a cocoon, and now is resting in it. When the cocoon lies firmly against the separating wall, then gnawing through the latter, the melittobia at the same time gnaws through the wall of the cocoon, and in this way directly penetrates to the larva. But in the case in which the cocoon does not lie against the separating wall, and its walls moreover stand also far from the body of the larvae that had spun that cocoon, then the melittobia, working with its jaws, penetrates inside the cocoon. But things become yet again different when the delicate wall of the cocoon directly lies against the body of larva having it spun. In this case, remaining outside the cocoon, the melittobia is entering it only with her ovipositor. Having concluded the work on the second prey, as with the first one, the melittobia penetrates into the third cell [of the nest], and after it also into the next one, usually until it has infected the whole nest, whereby a single female may deposit generally more than 1000 eggs.
The changes in the habits of the Melittobia stand out still more when she, instead of the host larva, finds in the cell preudo-cocoons (puparia) of parasitic flies (Pachyophtalmus signatus MG. and others). She also does not penetrate into them [that is, she does not gnaw-through herself into it], but infects them with the ovipositor from the outside. In this way, from an ectoparasite, ovipositing onto the surface of the prey, she now becomes, at least from the viewpoint of instinct, an endoparasite. On the other hand, having penetrated into a cell of a wasp, where, because of one reason or another, remained only not-consumed provisions (for instance leaf-roller caterpillars), the melittobia may deposit eggs onto them, beit in a very limited number.
When a melittobia is placed in a test tube together with its usual prey, then it may be observed that the melittobia lays eggs not only onto the prey placed before it, but not seldom leaves them also on the glass walls of the test tube. Moreover, crawling on the transparent walls of the tube the melittobia leaves behind in large quantities also its excrements in the form of semitransparent brushstrokes, which in natural conditions are hard to discern on the walls of a cell [of a bee or wasp nest infected by it]. After two days, from the eggs, deposited by the melittobia, the larvae emerge, being similar to the usual legless grubs of hymenoptera. The borders between the segments initially being sharp, become faint in the course of time. The larvae suck the prey, not inflicting on it a noticeable wound and not leaving behind any traces at the spot of feeding. They do not move from one place to another, and are only able to weakly bent their body. But when they are deliberately displaced they also at the new place [on the body of the prey] successfully pick up the interrupted feeding. Soon the integument of them and their prey become lightly moistened which aids in adhering the larvae to the prey's body, although generally [this adherence is] very weak. In 8-10 days the melittobia larvae conclude their feeding and, having fallen off the prey's body, remain where they are now. Soon they give off excrements, after which they quickly pupate, not making a cocoon. In still eight more days winged adults appear. In this way, a new generation of melittobia in favorable conditions develops very fast, in 18-20 days, when their mother has still all of her power and successfully continues her activities in the same [bee or wasp] nest, and sometimes even in the same cell.
In Melittobia acasta sexual dimorphism is sharply expressed. The female has normally developed wings, facetted eyes and articulate antennae with spherical segments of the whip. But the male has strongly shortened wings. Instead of complex eyes it has only single ones, one at each side, and its antennae are peculiar.
Usually the males do not leave the room (cell, cocoon, puparium) in which they emerged. The first adult [winged] male slowly tosses and turns itself amidst the heap of pupae, taking with its feet one after another. To the secondly emerged male it is hostile. One constantly tries to keep itself away and hide, and the other hunts for it. In not longer than 24 hours one of the males has its abdomen and feet bitten off, and often being without a head. The one that was happily left over in one piece, as absolute sovereign, proceeds inspecting the pupae. But then a third male appears. And again we see pursuit and struggle with the fatal outcome. Similar events are repeated also subsequently until females appear. For the rest, at this time usually only one male is left. The females, even without it, emerge in a larger number than males do (6-10 times more of them). The male, left over in one piece, persistently pursues females. Mating is brief. In normal circumstances, apparently all females become fertilized. The females display a remarkably strong attraction to light, and the majority of them, after having emerged, leave the nest. Only a few sometimes remain in the same nest where they were born, and lay eggs together with their mother onto the still fresh remains of the prey.
From an experiment it was revealed that M. acasta may reproduce also parthenogenetically, but in this case she lays only 4-5 eggs, from which [only] males emerge. The parthenogenetic [i.e. unfertilized] females display a special attraction to their single offspring : They almost in no instance leave the developing larvae, touching them with their antennae and feet. This affection increases when the larvae turn into pupae. This interest of the unfertilized female is especially expressed when there is only one larva left with her. It is only too clear that precisely to it and to the developed pupa all the work of the mother is concentrated. Only after fertilization by the one male nursed with such exclusive care, the melittobia begins the normal deposition of a great number of eggs, from which again only a few give rise to males. In the course of the year in a moderate climate 4-5 generations of M. acasta develop. Fully-grown larvae hibernate, not having given off excrements, and more seldom pupae (MALYSHEV, 1911).

The investigation cited, -- one of the first works of the author [Malyshev] -- was published not in the form of an individual article, but in a general sketch of the biology of [species of] Odynerus. Later authors have confirmed these data and partly supplemented them.

The second species, Melittobia chalybii ASHM., parasitizing in cocoons of solitary wasps from Trypoxylon and Pelopeus [hoping this latter name is correctly spelled], is, as to its behavior, very close to the previous species. But it went still further in its specialization and, in addition, displays polymorphism : Two forms of females are known, and two of males. See next Figure.

Figure 2 : Melittobia chalybii, an ubiquitous chalcid parasite of many kinds of wasp larvae.
1 and 3 (left) are males, 2 and 4 (right) females. The upper two figures (1 and 2) are of the "normal" form, the lower two (3 and 4) of the rapid-developing form in which the males are blind, the females robust and short-winged.
(After R.G. SCHMIEDER, 1933, in EVANS and EBERHARD, 1973)

One form of females has a brown color with spots, non-functional wings, more delicate integument and weaker delimited sclerites. The abdomen is large and inflated from birth. Development from egg to adult takes in this form all in all 14 days, whereby the life of the adult female maximally lasts for 30 days. She usually does not take food and begins to carry eggs on the day of emergence from the pupa, whereby in toto 40-60 eggs are being laid. Unfertilized individuals carry no more than 10 eggs, but also these largely perish.
The second form of females are dark brown with normally developed wings. Development from egg to adult takes about 90 days. The adult individuals live 60-75 days. Oviposition begins at the 11th-12th day after fertilization. Up to 500-600 eggs may be laid.
The first form is purely productive, whereas the second also serves the distribution of the parasite. Normally, on each prey two consecutive broods develop. Of them the first consists of both forms. The individuals of the first form rapidly reach maturity and generate offspring which transform into the second form which develops alongside with the other individuals of the first brood.
An experiment revealed that the second form may only be derived from the prey after first having developed on it a number of larvae of the first form. It is assumed that the food of the larvae of the first form chiefly consists of blood, while the larvae of the second form chiefly consume the more compact parts of the prey. From this it follows that precisely on trophic [= nutritive] conditions depends the appearance of this or that form, and that of the eggs only one sort is laid.
Of the offspring of fertilized females usually only 3 percent are males, but also this number goes down as a result of the mutual struggle. The males are represented also by two forms, chiefly differing in the construction of eyes and wings.
Winter is spent by M. chalybii in the stage of fully-grown larva of the first form.

About Melittobia chalybii something additional can be given, taken from Evans and EBERHARD, 1973, pp. 223 :

Several species of chalcid wasps are also known to be wasp parasites. Since these are minute insects, many indiviuals may develop within [probably better : on] a single wasp larva. The best known of these chalcids is Melittobia chalybii, a wasp little more than 1 millimeter long, over 500 of which may develop at the expense of a single wasp larva. This parasite has been reared from a wide variety of mud-daubers as well as twig-nesting wasps and bees. In the laboratory it will attack the larvae of social wasps as well as ground-nesting solitary wasps, but those are probably not often attacked in the field (In fact, in the laboratory Melittobia attacks and develops successfully on insects as diverse as cockroaches and beetles, and it is often a serious laboratory pest).
The males of Melittobia chalybii are short-lived and are greatly outnumbered by the females. They have short, nonfunctional wings and generally fertilize the females as soon as they molt to the adult stage. The males are reported to be "belligerent" and to "engage" in mortal combat with one another. According to one author, even a dead male, or a part of one, "will be fiercely pounced upon by another male, and dragged around and thrown about with a great show of anger, like a terrier with a rat." Fertilized females leave the cell by boring through the cap. They evidently seek out fresh wasp nest-cells by walking and hopping. Although their wings are fully developed, they seem to fly little if at all.
Having found a suitable host, the female remains with it for the remainder of her life of two or three months. She pierces the integument frequently and feeds at the exuding blood. After a few days, when her eggs have matured, she begins to lay several eggs a day on the host. Her first few (twelve to twenty) offspring develop rapidly and produce another generation of adults within two to three weeks. These adults differ in several respects from their parents : The males are blind and have even shorter wings, and the females have crumpled wings and stout abdomens (see Figure 2 above). These females are able to lay eggs immediately and at a more rapid pace than their mother, but they live only a few days. The eggs laid by these females -- and the remainder of the offspring of the mother -- undergo a slow development, and after about 90 days (or after a winter diapause) give rise to offspring of the "normal" type. R.G. Schmieder, who was the first to elucidate this unusual life cycle, believed that the larvae of these rapid-developing, short-lived individuals fed on the blood of the host, while the others fed on less nutritious tissues (experimentally, the eggs of either form were equally capable of producing offspring of either form).
This instance of dimorphism resulting from the nature of the larval food is in some ways suggestive of caste determination in social insects. In this case, the two forms together serve to build up a population of several hundred offspring able to utilize fully the larva of the host wasp.

(Continuing with Malyshev again)
Along precisely which way the melittobias arrived at the habit to infect larvae of wasps and bees in their cells, can easily be understood if we take into account that they eagerly oviposit in cocoons and puparia of various parasites encountered in the same cells. Earlier it was already noted that precisely in cocoons and puparia favorable conditions are created for the transition from one form of parasitism to another, including for the transition of terebrant-ectoparasites into hyperparasitism as well as into internal parasitism, and in the present case into parasitism in cells of wasps and bees. As to the members of the family Eulophidae, to which Melittobia belongs, they are, in all their diversity in their parasitic habits, in their overwhelming majority familial parasites. This characteristic feature of them reverberates on the predatory as well as on the ectoparasitic Phases of their evolution.
Some examples, illustrating the appearance of initial hemi-familial traits of life of eulophids, were already given earlier. Recall the habits of Cirrospilus ovisugosus C. and M., Microplectron fuscipennis ZETT., Euplectrus bicolor SWED., Cratotechus longicornis THOMS., and Eulophus viridulus THOMS., which all belong to the family Eulophidae.
On the highest stage reached among among terebrants, the melittobias -- attacking, with respect to them huge, preys, [preys] well isolated from the external environment and almost or totally immobile, with very delicate integument -- obtained the possibility not only of repeatedly ovipositing onto the same prey, but also of feeding on it together with their young. This eventually also led them to the formation of a true typical family of hymenoptera, although still weakly organized, but surely already polymorphic. From this we must assume that the melittobias on their line of evolution absolutely did not go through a solitary phase. Their ancestors were already in their oo-phagous [egg-eating] phase hemi-familial parasites.

Let us now turn to the bethylid S c l e r o d e r m a. [as the most probable ancestor of the ants].
In the south of Europe we encounter half a dozen species of this genus, and in the Balkans some of them are seen also in houses. See next Figure.

Figure 3 : Female of Scleroderma domestica HIEF. (From BERLAND, 1928, in MALYSHEV, 1966)

Substantial data on the biology of the scleroderms are known, although only concerning exotic species, especially many of them on the Hawaiian Islands, and, partly, also one species, S. macrogaster ASH., from Texas.
Although, as has been said, the scleroderma is in many respects a kind of look-alike of melittobia, there exists also a number of essential differences in its characteristics. Here we should note first of all that scleroderms chiefly attack larvae of beetles or -- more seldom -- of butterflies, living in dead wood of trees and shrubs.
Despite their small size, sometimes all in all only for two times larger than the minute melittobias, the female scleroderms are very energetic. When needed, they gnaw, with the help of their firm jaws, with extraordinary tenacity, themselves a way through wood bark, rotting wood, or through plant remains gnawed-up in wood by other insects. Having in this way reached its prey, for instance a larva of one or another member of the family of long-horned beetles, Cerambycidae, or one of the tree-gnawers, Bostrychidae, from time to time being for many, even for thousands of times larger than it, the scleroderm climbs on the back of the prey, grabs it at its skin with its jaws and stings it. Often, but not always, it stings at the region of the mandibular muscles of the prey, eliminating the danger of being bitten. The latter, for that matter, takes place seldom, even in the case when the prey is first hit at some other region of its body.
Holding itself on the prey, biting and stinging it, the scleroderm moves from place to place until the muscles of the prey, grabbed by its jaws, do not show any contraction anymore. Thus, gradually, in the course of 1 to 4 days, the scleroderm applies an uncountable number of stings without any determined consecutiveness and distribution of them over the body of the prey. As a result the larva is completely paralyzed. When the prey becomes immobile, the scleroderm feeds on its sap during several days. For this, the wasp lets prickle the prey's skin at various places, and then, from the wounds, earlier inflicted by the sting, small droplets exude, which the scleroderm licks off. Its abdomen quickly inflates as a result of the growth of the ovaries, and the scleroderm starts laying eggs, sparkling-white, elongated, rather large in relation to its size. The eggs are layed without any preparation of the places on the body of the prey, and are not scattered one by one, but in groups or packets of 10-40 together. The process of oviposition usually takes 2-3 days, after which there is an interruption until a new portion has matured. One female may deposit in this way up to 6 packets of eggs with intervals from 5 to 7 or more days between individual ovipositions. Apparently only few females lay more than 150 eggs during the course of their life which lasts up to three months.
From the eggs, deposited by the scleroderm, after 3-4 days the larvae hatch, being similar to the usual legless grubs of hymenoptera. In the beginning they feed, lying on the surface of the prey. But soon, having almost reached full growth, they gnaw through the skin of the host-larva and sink into it their head and the anterior segments. The rest of their body remains outside and projects sideways, as a result of which the prey looks bristled like a porcupine. In the body of fully-grown larvae white spots shine through as a result of the aggregation of a large quantity of urate crystals in fat formations. The mother-scleroderm, having laid eggs, does not leave the prey, but remains at it with her developing offspring.

Its a pity that this important moment in the life activity of the scleroderms is little cleared up by the investigator. On the basis of what has been expounded we may nevertheless think that the mother-scleroderm, at least in natural conditions, having deposited its whole relatively small stock of eggs onto one and the same huge prey, will not be inclined to look for a new [i.e. second] prey, but will stay definitively in the same hiding place together with its developing offspring. Precisely here lies the difference of them [i.e. the scleroderms] from the melittobias, which are almost ten times as fecund than the scleroderms, and which therefore do not need to limit themselves to just one prey but may attack new preys that are present in the same row -- in adjacent cells of the [wasp or bee] nest.

The mother-scleroderm often stays with her larvae and sometimes licks them, holding them between the forelegs. She also continues to drink blood of the prey, exuding near the heads of her larvae, sunken into the wounds in the skin of the prey. When the prey begins to dry out and decay, the mother-scleroderm eats the eggs laid by her and [eats] her own larvae.
It is observed that several female-scleroderms peacefully live together and in some case may jointly attack a very large larva of a long-horned beetle (Xystocera globosa OLIV.). Not messing each other up, they deposit eggs onto their common prey, and the offspring of some of these females feed and attain maturity without mutual impediment. An experiment revealed that females of even different species of Scleroderma and their larvae may peacefully live [together] and develop on one and the same prey.
In 5 days after hatching the larvae of the scleroderm conclude feeding. They lie down near the wrinkled and almost completely consumed, by them, prey and then take to prepare snow-white cocoons. This work takes two days whereby the cocoons are not constructed isolated from each other but in the form of a beautiful compact mass. Two weeks after the construction of the cocoons, and all in all a month after oviposition, the adult scleroderms emerge. They are polymorph, namely represented by two forms of females and two forms of males.
In different species of Scleroderma the numerical proportion of these forms is not the same. Thus, in S. immigrans BRID. the winged female form makes out 1/3 of the total number of females, whereas 99 percent of the males are winged, and the rest wingless. In another species, on the other hand, S. macrogaster ASHM., winged females are rare, as are also wingless males. The conditions of appearance of all these forms are not cleared up, as is also not cleared up the possible differences in their life and behavior.

See what has been said about polymorphism in Melittobia chalybii ASHM. earlier.

The males emerge from the cocoons first, and directly gnaw for themselves entrances into the cocoons of the females, where also mating takes place. But they may also fertilize mature females that have left their cocoon, unifying sometimes with one and the same individual several times with short intervals. One and the same female may also mate consecutively with a number of males.

Judging from Melittobia acasta, in the data given by Wheeler there may be some imprecision of the observations as a result of the difficulties of determining whether in each individual case actual fertilization of the scleroderm has taken place or merely an attempt to it.

In artificial conditions it is observed that the mother-scleroderm, living for a long time, may mate with one of her sons and greedily paralyzes another beetle larva, producing second offspring, and again may mate with one of her grandsons.
Females of scleroderma are hostile toward the feeding of the males on the prey, and as soon the time of oviposition has arrived chase the males away. Generally, after mating the males do not live long. In some cases when a male has remained with a female and with the, by her, paralyzed prey, it turns out to be decapitated.
When unfertilized females are taken in isolation and offered prey, they greedily paralyze it but oviposition is markedly delayed. The eggs and larvae of them develop normally but produce only males.

The great similarity in a number of basic features of life and behavior between the chalcidoid Melittobia and the bethyloid Scleroderma leads one to think of their having gone through similar phases of evolution. Indeed, melittobia as well as scleroderma look for a prey hidden in such conditions that fully guarantee a successful development of their multiple offspring. In this situation there is no need for them to displace the prey. In addition, the size of the prey, surpassing that of the attacking individual not just a mere six times (as it is the maximum in Bethylus), but hundreds or even thousands times, precludes for them the very possibility to transport it. And then the number of eggs laid by them onto one and the same prey, being many tens, totally does not correspond to that limited number of eggs (normally 1-6, seldom 8) laid by the other group of bethyloids. And the eggs are not deposited individually at rigorously determined places on the prey's body, but are distributed in groups or packets without any kind of order. If we, moreover, take into account that also the cocoons in scleroderms are not twined isolated but in compact masses, then it becomes clear that the scleroderms, as well as the melittobias, in effect absolutely do not live solitarily.
All this points to the fact that Scleroderma took its beginning not from some solitarily developing bethyloid, but went through, as did melittobia, in its development, the hemi-familial ectoparasitic Phase. Then, in this phase, the mother-bethyloid [not yet a scleroderm], attacking not too large a prey, insufficient for feeding her and her larvae, could not stay with her developing offspring and inevitably had to leave them after oviposition. It is very likely that a similar hemi-familial phase of evolution will be encountered also today among the many, not yet studied more closely, species of Scleroderma, in their attacking small larvae of beetles and butterflies [When the prey would be large, the mother could stay, and the hemi-familial mode [of living] would turn into a familial mode. But a familial way of life is not yet a social way of life, because in the latter there is division of labor also among similar individuals.]

With respect to the european Scleroderma domestica LATR. it is known only that it is obtained from pieces of a tree, infested by small xylophagous beetles (Anobium and others).

Accordingly, it is cleared up that the development of scleroderms went independently of solitary bethyloids. The lines of evolution of the one and the other originated, evidently, from one ancient oo-phagous phase that gave rise to also the Dryinidae, about which we have spoken earlier. But while the line of evolution of the Dryinidae and of solitary bethyloids went in the direction of the development of typically solitary vespine life, and the line of the bethyoids of the type Bethylus, laying a small number of eggs on determined places on the prey's body, turned out to be merely a blind side-branch, -- the line of the scleroderms developed further and, passing over to the next Phase, set the beginning of the remarkable bloom of hymenopterous life.

Noëtics of the origin of sociality in insects, especially in ants.

As we, in our noëtic theory of evolution, locate the main lines or pathways of organic evolution, not in the Explicate Order, but in the Implicate Order (noëtic space), we conjecture that also the evolutionary origin of sociality in insects must be traced back to certain noëtic reactions and derivations taking place in noëtic space. These noëtic events, like all other such events, do not take place in time and space, and therefore, the results are always 'already there' in the Implicate Order. Generally, the transition of strategy from s o l i t a r y, or from merely familial, to s o c i a l [way of life] in insects, especially in ants, as it takes place in the Implicate Order (noëtic space) might be visualized as follows : In solitary insects, and also in familial insects, each species represents a single complete strategy. Other living organic species, figuring in such a strategy as food (plant), prey (animal), or host (animal) are, as to the formal possibility of their existence in the Explicate Order, independent of their consumers or enemies : that is, their strategies do not presuppose those of their consumers or enemies (phytophags, predators, parasites), but may contain certain adaptations, independently whether these enemies actually exist (in the Explicate Order) or not [they can exist without their enemies], whereas the strategies of these consumers do definitively presuppose these [species of] food plants, preys, or hosts [they cannot exist without them]. So these consumer strategies can only be p r o j e c t e d (into the Explicate Order) when these plants, preys, or hosts, do actually exist there. Upon projection of the consumer strategy [i.e. the strategy of some given consumer] it appears in the Explicate Order as dual individuals, that is, male-female individuals [each such an individual is represented by two physical individuals, a male and a female], and they one-sidedly depend on the mentioned food plants, preys, or hosts [but mutually depend upon each other like the partners of a symbiosis, implying that they must be projected together.]. Matters become fundamentally different when the dependency is not one-sided but m u t u a l. Thus, in cases of symbiosis of different organic species (as for instance in lichens) the strategy of one implies that of the other, while that of the latter at the same time implies that of the former, meaning that each strategy by itself is incomplete, and cannot therefore be projected into the Explicate Order. So b o t h strategies must be projected together. In the case of social insects, such as ants, each caste (soldier, worker, male, queen(s)) represents a strategy, but these strategies are mutually dependent, causing them to be, each for themselves, incomplete. They must be projected together, and then in the Explicate Order they supplement each other, and all of them together then represent the c o m p l e t e strategy of the species. And because the castes belong to the same (taxonomic) species the joint projection of their corresponding strategies results, in the Explicate Order, in a h i g h e r - l e v e l individual -- the ant colony. And it is these higher-level individuals (ant colonies) that represent -- in the Explicate Order -- the species and its (single) strategy. This 'individual' does perform all the duties (all the functions) simultaneously : foraging, guarding the nest, extending the nest, feeding the larvae, laying the eggs, etc. And this indeed is the essence of social (insect) life. The individualization and segregation of incomplete substrategies within a single original complete insect strategy -- disintegration and subsequent re-integration -- is possible only in certain pre-existing specific strategies of solitary (or merely familial) insects that are in some way pre-adapted to it. It is assumed that in such an original strategy (as being a noëtic description in the Implicate Order) certain noëtic reactions take place between parts of the original strategy, resulting in the disintegration and subsequent re-integration of this original strategy. Perhaps re-integration only takes place in the Explicate Order upon projection. So, in summarizing things, the original strategy must already contain the functions (or at least rudiments of them) that are also present in the super-individual of social insects. Internal noëtic reactions result, as has been said, in the disintegration of the original strategy, meaning that these functions are detached from one another, but in all this each one of them remains connected with the rest (or most of it) of the strategy [which rest is thus multiplied]. In this way different separated (but related) strategies result, which, however, imply one another mutually, and can therefore only project together, and will then be re-integrated such that within the super-individual the mentioned functions are distributed among the lower-level individuals (the usual, physical, individuals), and are kept in this way physically separated from each other. In this way different castes appear in the insects concerned. We assume, with MALYSHEV, that ants have evolved from scleroderm-like ancestors. Indeed -- as we will discuss in the next document -- in the familial life of the scleroderms (as well as of the melittobias), we can see a number of essential features (pre-adaptations) bringing their type of family close to that of the ants, namely :

Presence of a habitation (nest) suitable for the life of the family. Presence in that habitation of a stock of provisions in the form of a huge prey, suitable for the feeding of the mother-foundress herself as well as for her offspring. Oviposition in groups or packets onto this prey without any attachment of the eggs to the place of oviposition. A close contact of the long-living mother with her rapidly developing offspring. A, by this, guaranteed encounter of several (2 or even 3) generations in their adult state in that same habitation. Presence of polymorphism and especially the presence in such insects of two forms of fertile females.

These are the first prerequisites of insect (especially ant) sociality. A strategy from which, in the Implicate Order, may be derived a strategy of, albeit still primitive, social insects such as ants must at least already contain these elements. We will see that in the melittobias a further element is missing, an element which is present in scleroderms. So these are the true ancestors of the ants.

With all this (biology and noëtics), we conclude our exposition of the Familial Ectoparasitic (Hemi-Formicoid) Phase of hymenopterous evolution.
In the next document we will, still on our way to the origin of ants, expound the Primary-Ant (Proformicoid) Phase.

e-mail :