'primitive', 'original') insect Orders, that is, we must inspect the tracheation(al pattern) of wings of nymphs ["nymphs" are larval stages of insects with incomplete metamorphosis, i.e. dragging it into the burrow, which contrasts to the usual habit of Family Bethylonymidae (Superfamily Bethylonymoidea). Close to mid-wing it separates into CuA and M, and while CuA is heading more or less straight away to the posterior wing-margin, the Media arches up and reaches the branching-off point of RS [from R]. Family Melittidae Family Ephialtitidae. Length of wing about 3.5 mm. Family Ephialtitidae. Family Eulophidae n., plus anciens reprsentants connus de la tribu Trypoxylini de sphcides, sont dcrits de lambre de locne basal de France. 1970.
Boston: Houghton Mifflin Co. (End). The representatives of the oldest Order with already formed venation -- Protoptera [Sharov here has in mind the Paoliidae and relatives] -- also have richly ramified venation. Superfamily Bethyloidea, Apocrita-Aculeata. Maimetshidae (Upper Cretaceous). Upper Jurassic of Karatau. n. The fore wing venation of Eopison is more complete than those of any Recent Trypoxylini, suggesting that the reduction of the submarginal cells in Sphecidae has a complex history. 1998. Wing(s) of Cleistogaster seticornis Length of wing about 3 mm. And given that, it must be fully compatible with all elements of the strategy, that is, it must not interfere with or damage this strategy. And although it seems probable that all winged insects (pterygota) have descended from a single ancestor, the possibility of a polyphyletic origin of them cannot be excluded. prey. The female searches for the host There is a family of wasps that include mud daubers, digger wasps, and others that are genera known as of 2000.. A key reference is Bohart & Menke (1976). Family Evaniidae. First of all, when in the practise of evolutionary taxonomy one speaks of the derivability of some given venation V2 (or any other morphological structure for that matter) from another venation V1, one has -- if no other characters do contradict it -- in mind the actual evolution of the recent representatives of V2 from earlier representatives of V1, and thus an actual evolutionary transformation of the venation V1 into V2 is assumed. Superfamily Evanioidea.
Radial Sector only present as a mere trace. This structure became very wide-spread among Apocrita. And so it seems that whatever evolutionary changes might take place in the wings of some given insect Order they always take place within the morphological confines of the venational type of the Order, that is, they never bring the venation outside this type [This has only taken place, apparently, when one Order originates from another in evolutionary history]. Fore wing of Hartigia nigra (earlier Macrocephus satyrus) (a), Janus femoratus (earlier J. cynosbati) (b), J. abbreviatus (c), Orussus abietinus (d), Pelopoeus cementarius (e), Apis mellifera (f). nest in preexisting cavities. Their
Venation almost completely reduced. paraphyletic. Thus, the most recent
(spider-wasps), Family Eumenidae Superfamily Pompiloidea. Placing the Ephialtitidae with the Stephanidae, and contrasting both of them, as a single superfamily, with the remaining Apocrita, is based on the primitive structure of the abdominal hinge, being still very close, in a series of cases, to that what is characteristic of Symphyta. It is clear that the pectinate mode of branching is less symmetrical than the dichotomic way of branching, and thus contributes to the overall differentiation of the venation where it occurs. Wing(s) of Brachycleistogaster nana Family Eumenidae. The secondary broadening of the abdominal hinge was, apparently, realized as result of pedomorphosis (= the preservation in the imago of a [morphological] condition characteristic of the early pupa).
Family Tenthredinidae, Symphyta. The M+CuA trunk branches off from R near the base of the wing. We may identify, for example a two-branched Radial Sector, but we generally cannot retrieve whether these branches are R2 and R3, or R4 and R5, or R2 and R5, etc. To me the original primitive type of wing-venation of all insects proposed by Comstock and Needham in 1898-1899 seems to be the most probable because it is based on ontological studies. Recent. So in our effords to identify the veins in fossil and recent insects on the basis of the prototype of Comstock an Needham we must be content with identifying the mentioned venational chief systems (C, SC, R, RS, M, Cu, and A), and sometimes with being able to distinguish between, for instance, "MA" (anterior part of medial system) and "MP" (posterial part of medial system), as Sharov has done in his prototype. Wing of Asiephialtites oviventer Almost nothing is left of it. Figure 80 : (After COMSTOCK, 1918), Figure 83G : Forewing of Evaniellus. a few vary considerably in their selection of prey. Figure 31 : classification is closer to the conservative scheme; the families It must always be such that it is formally derivable from the basic comstock-needham scheme of venation. Then it (RS) proceeds to the anterior wing-margin while having given off a cross-vein to the Media.
Length of body 1.1 mm.
Borror, Wing(s) of Mesaulacinus stenocerus Also this cubitus does not reach the wing-margin.
Abdomen long and stalked (petiolate), giving the body a "thread-waisted" appearance; middle tibiae with two apical spurs; body may be all black (sometimes tinged with metallic blue or green), black and red, yellow and black, or white and black.
Fragment of wing of Curiosivespa magna. It runs from wing-base to pterostigma, but not beyond the latter. Radio-Medial Triangle long. Clausen (1940) discussed the effect of the sting of female (After RICHARDS and DAVIES, in Imms' General Textbook of Entomology, 1977. The latter vein and the Cubitus do not reach the wing-margin. (To see it in context with other Jurassic fossils [from the above long list] click HERE). 1989. restriction to a particular type of food for the larvae of some species, but Family Megalyridae. Falsiformicidae (Upper-lower Cretaceous). Radio-Medial Y-vein has 'pulled' the Radius downward. Family Anomopterellidae Figure 56 : Figure 59 : (After BRADLEY, in COMSTOCK, 1918), Figure 83C : Forewing of Evania appendigaster. This has taken place in the hindwings of many representatives of other insect Orders as well (such as grasshoppers), but here it is especially conspicuous that the extension of the anal area took place at the expense of the rest of the wing. in recent insects) such that (in all probability) the lacunae are formed first, in which subsequently grow the tracheae. continuing the course of the up-arching Media. It is interesting to note that ramified venation is also characteristic of the earliest representatives of Palaeodictyoptera and also of ancient representatives of Neoptera. n. and Pison eocenicus sp. Figure 87 : 425) [1]. Top image : Right forewing of Aeolothrips kuwanaii. Wings of Cleistogaster buriatica The line measures 49 mm. So this true prototype is reconstructed with the help of Zdenekia by taking it just one more step backward, for example in depicting a still free vein M5 (most posterior branch of the Media), of which in Zdenekia only its very base is (evolutionarily) preserved (the rest having been merged with CuA). The second and third anal veins absent. By continuing you agree to the use of cookies. Figure 90a : Detail of Figure 90. After paralyzing the cricket, the wasp drags it by its antennae back In the evolution from Protoptera to Polyneoptera [grasshoppers, cockroaches, etc.] Further, below M, we see the veins CuA, A1, and A2+A3. Baissidae (Early Cretaceous). Some species of Sphex Length of wing 5-6 mm. The course of the Radial Sector, from its origin to its end, is indicated : RS - RS+M - RS - RS. A part of the coalesced course(s) of RS and M is in fact reduced, as can be seen in the left Figure (upper image). Family Ephialtitidae. (?Superfamily Sphecoidea). Insects. The wing-fan is partitioned by convex (kx) and concave (kv) radial folds ["radial" here has no relation with "radial" in "radial veins"], which draw together like a fan when the wing is placed under the elytron. And indeed we see that this prototypic venation contains little, if any, differentiation of parts or areas of the wing. Upper Jurassic of Karatau. (After SHAROV, 1966). (After SHARP, in RICHARDS and DAVIES, in Imms' General Textbook of Entomology, 1977.
Upper Jurassic of Karatau. Superfamily Apoidea (bees). Subcosta absent. Recent. Family Ephialtitidae. Wings of Trigonalopterus brachycerus. Neocom (lower-lower Cretaceous) of Zabaikalj. How to Know the Insects. In: Page 385, 3rd ed. (After COMSTOCK, 1918), Figure 83F : Forewing of Hyptia.
nigellus storing its nests with Length of wing 2 mm. 230 species are fossorial or Sphecidae on the host. It ranges from From Comstock (fig. For example, cases are known in which RS loses its connection with R and attaches itself to M or MA. Fold-lines are indicated by dashed lines. Most species of Sphecinae construct their nests in the soil,
Cross-veins 2r-m and 3r-m absent. The base of MP as already noted, is maintained in many Oligoneoptera. able to grasp one of the wings with her mandibles, and the sting is then Upper Jurassic of Karatau. Length of wing 4-4.5 mm. Family Apidae.
Fragment of wing of Oryctopterus dubius. of Bethylonymus curtipes. Upper Jurassic of Karatau. Upper Jurassic of Karatau. Only the details of the wing-venation are considered by us as non-functional (and this is, of course not entirely certain). Family Megachilidae. Radio-Medial Triangle long but narrow. Figure 18 : As already noted by Redtenbacher, the geologically more ancient insects have a richer and more ramified venation. Subcosta absent (or coalesced with R). It is also evident that there has taken place both specialization by reduction and specialization by addition of (longitudianal) veins. Family Bethylonymidae So it is easy to recognize 'odonate wings', 'dipterous wings', 'lepidopteran wings', 'hymenopterous wings', etc. Wing of Bethylonymus microphthalmus. There are over 1200 species of these Wings of Symphytopterus oculatus. The biology of the Figure 23 :
Wing(s) (fr.) Upper Jurassic of Karatau. (Superfamily Formicoidea (ants)). (Superfamily Vespoidea). Family Aulacidae (see systematics). Family Eucharitidae Family Apidae (social bees). We use cookies to help provide and enhance our service and tailor content and ads. Forewing : Radio-Medial T-vein developed. Figure 81 : Figure 44 : Length of wing 4 mm. Ju = jugal domain of venation. Family Ephialtitidae. Wing of Brachycleistogaster nigritibialis. that become ordered in a formal derivational sequence. We will now present a number of wings of fossil Apocrita, beginning with the family Ephialtitidae from the Jurassic. Forewing of an ichneumon fly. Females construct their nests attached to some
purposes only] Glossary Superfamily Apoidea (bees). The fact that this vein belongs to the longitudinal veins [not to the cross-veins] of the wing was convincingly demonstrated by Tillyard (1919) for Mecoptera. M Pre-existing cavities are preferred nesting sites, or they dig simple
building no nest but laying their eggs in the nests of other wasps, their
Upon leaving M, the Radial Sector arches slightly up until it meets the cross-vein 2r-rs, then continues its course toward the wing-apex, but long before (reaching it) it sharply bends upwards and even backwards and ends up at the anterior wing-margin (meeting R1). Radial Sector unbranched. Library]. And consequently we may hold that the wing-venation, as a particular pattern or structure, comes from the Implicate Order, because it is there where strategies exist (and from where they are projected into the Explicate Order). Sphex aegypticus Lep. Comparative study of the most ancient representatives of the different phylogenetic branches of Pterygota made it possible to give the following scheme of such an original type of venation. Disclaimer: Dedicated naturalists volunteer their time and resources here to provide this service. Figure 93F : Hypothetical primitive type of wing-venation of Pterygota. This is cosmopolitan
recover rather completely in 10-15 min., but a considerable lethargy follows,
Superfamily Evanioidea. -- (After RICHARDS and DAVIES, in Imms' General Textbook of Entomology, 1977.). Figure 88 : And, while remaining to comply with the comstock-needham venational scheme, every Order has its own prototype of venation from which the venation of all the representatives of this Order can formally be derived. They paralyze spiders, lay an egg and seal the
Description of the original type of wing-venation of the Pterygota. Superfamily Ceraphronoidea. Most species nest in the ground, usually in areas with sparse or no vegetation; some build aerial nests of mud; a few nest in hollow stems or abandoned bee burrows in logs. Upper Jurassic of Karatau. were studied by Williams (1928).
151). Family Ephialtitidae. S. hirsuta Scop. of Leptephialtites We will be informed about the specific difficulties that are encountered when trying to determine prototypic venations. (Superfamily Bethylonymoidea). With this 'original' venation in mind we are well able to interpret the venation of the true Apocrita found in Jurassic and younger deposits (and amber), and also of recent Apocrita. families the Sphecidae and Crabronidae. Blue : course of Radial Sector. Venation extremely reduced. Figure 10 : Les plus anciens reprsentants de Sphecidae: les Trypoxylini de lambre de locne basal de France. Subcosta is long, with many short branches. Length of wing 4.5 mm. Figure 50 : Same specimen as in Figure 49. Course of Radial Sector (blue), Media (red), and anterior Cubitus (green), indicated, in order to clarify the modified venations below. RS+M long. Length of wing not less than 10 mm. Amber inclusion. Upper Jurassic of Karatau. Radio-Medial Triangle broad. Study of Insects. Upper Jurassic of Karatau. The very first section of the Radial Sector (branching off from the Radius) is here still in its primitive condition, namely obliquely directed toward the distal part of the wing. Figure 27 : The Media not reaching the wing-margin. The majority of Blattodea lost the pre-costal field and the base of MP, although in some of the most primitive representatives the MP-base can still sometimes be observed. Family Masaridae (honey-wasps). the abdomen's dorsum. When partly
The cross-veins 2r-m and 3r-m present as oblique veins. Amber inclusion. Second and third anal veins absent. Formicidae (Lower-upper Cretaceous-Recent). are xylophagous] which are rather closely related to the Karatavitidae, and the reduction of the cross-vein 1r-rs as it is seen [not only in Ephialtitidae but also] in Cephoidea and Orussoidea. Radial Sector unbranched. and black or black and red, although a few species are metallic blue.. In North America there are 35 species in 4
Length of wing 3 mm. Hymenoptera of
Figure 22 : It is the [end of the] Media. Functional significance of the wing-venation revisited. Family Megalyridae. Below it we see a minute appendage by far not reaching the wing margin. Figure 68 : to the burrow from whence it came.
Family Ephialtitidae. Wing of Bethylonymellus microgaster. Figure 49a : Wing(s) of Mesaulacinus sculpturatus (Same as in Figure 49). varies, ranging from spiders to grasshoppers, mole crickets, locusts,
Upper Jurassic of Karatau. Radio-Medial T-vein not yet developed (first section of RS more or less perpendicular to R). of Leptephialtites angustus It plunges itself in an unimaginably fast tempo onto it and bends its wings at downbeat sharply downwards. Family Megalyridae. 