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The most primitive known wing articulation and wing folding in the Pterygota occurs in the extinct Palaeoptera the Diaphanopterodea. The oldest Diaphanopterodea Namurodiapha sippelorum n.gen., n.sp. (Namurodiaphidae n.fam.) is described from the early Late Carboniferous (Namurian B) of Hagen-Vorhalle, Ruhr area, West Germany. Unlike other Palaeoptera, the Diaphanopterodea show similar apomorphic venational features and parallel evolutionary trends in wing evolution, as in the unrelated Neoptera. This is explained (i) by convergence caused by the wing folding present in both Neoptera and Diaphanopterodea but absent in the rest of the Palaeoptera, and (ii) by a progressive diminution in size which affected the diaphanopterid and neopterous wing venation in a similar way. The wing articulation and the evolution of wing folding in the earliest known Palaeoptera and Neoptera are examined.

Gigantic as well as very large mayflies from the middle Upper Carboniferous (Westphalian) strata of Europe and North America are described: the adult and nymph of Bojophlebia prokopi n. gen., n. sp. (Bojophlebiidae n. fam.) and the nymphs of Lithoneura piecko n. sp. and Lithoneura clayesi n. sp. (Syntonopteridae). Evolution of ephemerid wing venation during 300 million years is summarized. Autapomorphic, apomorphic, and plesiomorphic character states of venation are categorized. Venational nomenclature of Recent Ephemerida is emended based on its evolutionary changes. Evidence that wing veins occurred primitively as a pair of fluted sectors is documented in Carboniferous mayflies in the costa, subcosta, radius, anal, and jugal. Ephemeroids and odonatoids are sister groups that share the veinal anal brace AA fused with CuP at an area important for flight. Ancestral Odonatoephemerida are the sister group of the extinct haustellate Paleoptera. The Carboniferous nymphs bear three pairs of almost homonomous thoracic wings and, on the abdomen, nine pairs of legs and nine pairs of tracheal gills (wing homologues). This proves that abdominal legs have been totally reduced in Recent Ephemerida except for the claspers (gonopods) and that tracheal gills are not flattened legs. The metamorphic instar probably originated in relatively young instars. Insectan cerci developed from segmented, arched, functional legs of abdominal segment 11, which were still present in this primitive condition in Carboniferous Monura.

The oldest known odonatoid wings are described from the Namurian of Argentina: Eugeropteron lunatum Riek n.g. et sp. and Geropteron arcuatum Riek n.g. and sp. (Meganisoptera: Meganeurina: Eugeropteridae n. fam.). The wings are generalized and support a reinterpretation of the venation of living Odonata as being fully homologous to that of other pterygotes and closely related to Ephemeropteroidea, but different from Neoptera. Therefore, Paleoptera is a valid phylogenetic unit, and Odonatoidea and Ephemeropteroidea are sister groups.

The most primitive known pterygote terga, wing articulation, wings, and upper leg segments with exites, occur in gigantic Upper Carboniferous Paleodictyoptera, Homoiopteridae. Fossil features are used as clues for reinterpreting some structures connected with flight in modern Pterygota. Brief comparisons with Paleozoic Diaphanopterodea, Permothemistida, Ephemeroptera, Protodonata, and with living Ephemeroptera, Odonata, and Neoptera are given. The wing articulation of all Pterygota is derived from a common ancestral ground plan based upon features present in fossils. The ancestral wings were articulated by a closely packed band of multiple sclerites which were hinged to eight lateral tergal lobes, and aligned with eight pairs of wing veins. The axillaria of Neoptera and axillary plates of Paleoptera are composite sclerites, which originated by fusion of several sclerites of the original band. Articular patterns of Paleoptera and Neoptera evolved differently and show (i) the presence or absence of a gap at the cubital level, (ii) the presence or absence of a turning–pivoting composite third axillary sclerite (3Ax), and (iii) a different composition of all composite sclerites. Gliding and wing folding adaptations within the articular band are discussed. A new fossil-based interpretation of veinal stems, veinal sectors, and of their fluting near the wing base is offered. An underlying symmetry of thoracic tergal sulci, articular sclerites, and wing venation seems to point to a nearly symmetrical, nonflying pro-wing engaged in up-and-down movement. Evidence of articulation in Paleozoic nymphal wings and evolution of metamorphic instars are examined. Pitfalls of paleoentomological work are discussed. Criteria for major divisions of Pterygota are reassessed. It is hypothesized that the wing originated from the first segment (epicoxa) of the euarthropodan upper leg and its exite. An epicoxal podomere became incorporated into the body wall and broke up into an articular ring of dorsal and ventral sclerites, and an epicoxal exite flattened and became a pro-wing. The pro-wing originally operated on a row of pivots from the epicoxa and subcoxa (pleuron) and became mobilized by epicoxal leg musculature.