Krasnojarsk district. Upper Jurassic of Karatau. The base of MP as already noted, is maintained in many Oligoneoptera. As to their structure, they are still very close to the lower Hymenoptera ( Symphyta), especially to the Karatavitidae. Length of wing 5.5 mm. 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). Agriculture Canada Research Branch, publication 1894/E. and Chalybion store their nests persues grasshopper 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. Wings of Braconidae Gnamptodontinae from Goulet and Huber (1993, fig. He regarded it, starting from the notions of Comstock and Needham on the four-branched Media, as an additional fifth branch (M5) of this vein. The base of the genuine MP is usually not preserved, and only in representatives of the families Archiprobnisidae and Megakhosaridae (Sharov, 1961) is it represented in the form of an oblique cross-vein together with the concave false MP. The latter heads for the wing-margin just below the wing-tip. Length of wing about 3.5 mm. The wings possessed a fine-meshed archedictyon, not visible in the present Figure. There are no true cross-veins. Superfamily Apoidea (bees). Upper Jurassic of Karatau. Thus, the Ephialtitidae, judging from the available data, represent the "missing link" between Symphyta (as represented by the Karatavitidae) and the remaining Apocrita. Lower Dogger (?) Lower Carboniferous (Namurian) of Tsjechoslowakia. cell in the soil, drives out the occupant and then pursues and captures it above ground. A lengthy battle often ensues, but finally the parasitoid is But of course, if venation B is identical to venation A, the former can be formally derived from the latter. Length of wing 3.5 mm. Family Liopteridae, Family Agaonidae (Agaontidae) (fig insects) The Medial vein is completely reduced, no trace of it exists. In all this we must understand that of a given wing the wing-venation as a whole (= the whole venational pattern) is probably always functional in the sense that it, together with the size, form, and shape of the wing, and with hairs and thorns and bristles on it, constiutes a functional organ of flight. extensively, feeding on fluids that exudes from the prey's mouth. Indeed, our theory must be such as to accommodate these facts of derivability, and our considerations took care of that. Superfamily Apoidea (bees). So the formal difference of the wing in Figure 57, having a rather complete venation, from the wings in the above Figure (with a strongly reduced venation) must reflect a difference in flight-regime despite the fact that all these wings are equally minute. egg is placed at the base of one of the coxae. Nests are made in hard ground that is free of vegetation, and P. hazmatogastrum Sphex aegypticus Lep. Family Mutillidae. Family Ephialtitidae. mandibles. The egg is usually laid on Length of wing 2 mm. D. J. Family Eulophidae Order Thysanoptera. 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). Palaeomyrmex zherichini. So the course and number of tracheae in a wing-bud of a representative of some given insect species do not as it were recapitulate phylogeny, but may nevertheless be helpful to identify the course of veins in the wing of the adult insect in cases where, in such wings, complicated coalescences of veins have taken place.]. ?Superfamily Scolioidea. When we have found it we can then derive the venation of all insects from this prototypic venation, and can also identify each vein in the wing of any insect whatsoever. The (fore)wing is narrow at its base, and its shape is more or less triangular. See next Figure. cell. The larvae develop, pupate and Maimetshidae (Upper Cretaceous). It may be that the main longitudinal veins and their main branches carry with them the constant character of being convex or concave [that is, one main vein is always, that is in all insects, convex (except when secondary transformations have taken place), whereas the other main vein is always (in all insects) concave (also except when secondary transformations have taken place). The Radius extends far beyond the pterostigma, running along the anterior margin of the wing, but stops well before the wing-apex. Family Andrenidae. They are merely implications of, or additions to, this set. Sphecidae on the host. It ranges from Coniac-Lower Santon (Upper Cretaceous) of Radio-Medial T-vein poorly developed. Also important in this discussion on vein homologies is the (original) corrugation of the wings, such that longitudinal veins, lying on a ridge (convex veins, indicated by "(+)"), alternate with veins lying in a groove (concave veins, indicated by "(-)"). The latter is mainly based on paleontological evidence, but, as we have said, we do not know fossils of the truly first winged insects, and so the oldest fossils known to us could be misleading. (Palaeodictyoptera). So, to continue with the example just given, although the particular flight-regime, or, particular aspect of the flight-regime, precisely and only insofar as it is determined by the wing-venation, is not a part of the strategy of, say, some given blowfly species insofar as the strategy of this species differs from the strategies of all other flies of the mentioned group of families, it is nevertheless a necessary common ingredient of all the different strategies in this group (and is absent in the strategies in other groups). It is the [end of the] Media. Green : course of the anterior Cubitus (CuA). the World.

Family Apidae. Figure 24 : Figure 19 : '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. And now, back again to the present Figure (85), we can correctly follow the course of RS after the point where it has separated from the Media : It sharply curves up until it meets the cross-vein connecting it to the pterostigma. This inhomogeneous distribution is accomplished by the shift of the course of several existing longitudinal veins, and by the disappearance of some of them, or by the lessening of their branching, and, finally, by the shift or disappearance of many cross-veins. Neocom (lower-lower Cretaceous) of Zabaikalj. Family Elasmidae (After COMSTOCK, 1918), Figure 93M : Representative of Palaeodictyoptera -- Stenodictya sp. Lower Carboniferous (Namurian) of Tsjechoslowakia. Figure 40 : spp., etc. Here nesting differs from Length of wing about 3.5 mm. The wing venation of the forewings is relatively complete. It differs markedly from SHAROV's hypothetical primitive type(Figure 93H). Radio-Medial T-vein well developed, Radio-Medial Triangle small. Wing(s) of Brachycleistogaster jurassica. 2001. 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). caterpillars, some females malaxate the venter of prey after it is And further, we must realize that, in following the prototype proposed by Comstock and Needham, although we can indeed identify in all insects all the principal veins insofar still present (C, SC, R, RS, M, Cu, and A), experience shows that we cannot expect that we can, always, in all insects, precisely identify all the main branches of these principal veins (for instance the four branches of the Radial Sector, R2, R3, R4, R5, or the four branches of the Media, M1, M2, M3, M4) named by Comstock and Needham in their prototype. Now if there exists, in the Implicate Order, a given notic pattern (among many other such patterns) that is a strategy, a notic prescription, such that it specifies a "winged insect" (i.e. Apart from mere folds existing in some places in the wing-membrane, these ribs are 'sclerotized folds', wing-veins. nest in preexisting cavities. Their Radio-Medial T-vein very well developed. Upper Jurassic of Karatau. Upper Jurassic of Karatau. foliage, etc. There are over 1200 species of these The Radial Sector branches off from R obliquely backwards, then, after joining the Media, runs into the direction of the wing-apex, then abruptly bends toward the anterior wing-margin (i.e. Upper Jurassic of Karatau. Based on the venation of the wings of the most ancient insects (found as fossils in carboniferous deposits) we may hold that the most primitive venation is a set of veins (the full comstock-needham scheme of longitudinal veins) that are more or less evenly distributed on the wing-plate. adults or nymphs of Conocephalus in the ground, but some nest in natural cavities such as hollow plant stems, Elasmomorpha melpomene. Copyright 2022 Elsevier B.V. or its licensors or contributors. Media, after having arched up, coalesces with RS for some distance and then leaves it independently again. We strive to provide accurate information, but we are mostly just amateurs attempting to make sense of a diverse natural world. Having all this in mind (differentiation in the wing-blade, functional significance of venation), we want to establish the prototypic venation of the class Insecta. Changes of the same type in the position of the branches of MA also occurred in Orthoptera but the MP in them is preserved, and the pre-costal field of the forewings underwent further development. -- Upper Jurassic of Karatau. Length of wing 2.5 mm. Figure 93G : Comparison of the hypothetical primitive venation of Protoptera [Paoliidae and relatives] and the venation of the most ancient representatives of Palaeoptera, Paraneoptera, Polyneoptera, and Oligoneoptera (Holometabola). The Cicada Killers of RA = Radius (radius anterior), MP = posterior Media, MA = anterior Media. Upper Jurassic of Karatau. As already noted by Redtenbacher, the geologically more ancient insects have a richer and more ramified venation. building no nest but laying their eggs in the nests of other wasps, their Recent Trypoxylini are parasites on spiders. Radius-proper unbranched. Length of wing about 9 mm. The Media as well as the Cubitus do not reach the wing-margin. Family Melittidae Figure 35 : Figure 45 : The Costa is very short and has a small pre-costal field anteriorly to it. Superfamily Evanioidea. Recent. (2014) Family group names and classification, American Insects: A Handbook of the Insects of America North of Mexico, Finnamore A.T. (1998) Aculeate wasps (excluding Formicidae) [of the Montane Cordillera Ecozone], The Sphecid Wasps of Michigan (Hymenoptera: Sphecidae: Sphecinae), by M.F. Family Ephialtitidae. Upper Jurassic of Karatau. Indeed it is possible to discover the prototypic wing-venation of any given Order of the Pterygota. Pour citer cet article: A.Nel, C.R. Palevol 4 (2005). As is evident from all the above considerations, it is hard to establish with certainty the original primitive type of wing-venation of the very first winged insects, that is, the prototype from which the venation of all winged insects (Pterygota), fossil and recent, can be derived. Finnamore & Michener (1983), who placed them as a separate themselves to a single host species, while others may attack individuals of Indeed, if they were a consequence of that theory, then this fact would add to the evidence in favor of it. In representatives of the superfamilies Ideliidea and Liomopteridea the base of MP was reduced, and together with this there was displacement in the first fork of MA closer to the base of the wing up to the level of the origin of RS or even more proximal, and the posterior branch assumed a concave position which lead the author (Sharov, 1961, 1962) to regard it erroneously as MP. Functional significance of the wing-venation revisited. Radio-Medial T-vein not yet developed (first section of RS more or less perpendicular to R). Upon leaving M, the Radial Sector arches up until it meets the thickened cross-vein 2r-rs, then heads toward the wing-apex, but long before reaching it it sharply bends upwards and ends up at the anterior wing-margin. ScienceDirect is a registered trademark of Elsevier B.V. ScienceDirect is a registered trademark of Elsevier B.V. Recent. In some Protoblattodea, see image E of Figure 93G, the fusion of MP- and CuA-branches was still not so complete as in the majority of other Pterygota, and in them it is still possible to distinguish MP- from CuA-branches [In the wing of image E of Figure 93G we see the branches of the posterior Media in the left half of the wing, whereas the branches of CuA we see next to them at the right side]. Basal section of RS is present and runs backwards and downwards. Cross-vein 2r-rs present. Wing of Anomopterella stenocera. (After SHARP, in RICHARDS and DAVIES, in Imms' General Textbook of Entomology, 1977. (Superfamily Megalyroidea) stephanocephalus Family Karatavitidae. And because insects are, in the fossil record, documented chiefly by their wings (in most cases forewings), and that is [documented] by their wing-venation, we must take the formal [versus functional] status of its details into account when concluding things about evolution from fossil insects. and so it is not fixed as to the direction of derivability in one or the other Order : In the Explicate Order as well as in the Implicate Order derivability of a given wing-venation always means the derivability from the venational prototype ('archetype') of the insect Order of which the species possessing this venation is a member.

burrow of its own. Length of wing 6.3 mm. Media, after having arched up a long way, coalesces with RS near the pterostigma (base of RS absent) for some distance while arching down again together with RS (merged with it) [M+RS]), and then leaves it independently again, heading to the distal part of the posterior wing-margin. Length of wing 4.6 mm. Family Megalyridae. Note the lateral lobes on the sides of the abdomen and especially on the prothorax. Then RS continues its course and then, as it seems, folds back toward the pterostigmal area. Figure 75 : Family Braconidae Everything else copyright 2003-2022 Iowa State University, unless otherwise noted. Wings(s) of Bethylonymellus cervicalis. Length of wing 1.7 mm. Family Ephialtitidae. These possible patterns of veins are now, in the Implicate Order, distributed among the notic strategies of winged insects. Fla. Dep. In the previous document we have argued that any evolutionarily established wing-venation is, in the Explicate Order, not completely stable.