A survey is made of the major features of the venation, articulation, and folding in the hind wings of Coleoptera. The documentation is based upon examination of 108 Coleoptera families and 200 specimens, and shown in 101 published figures. Wing veins and articular sclerites are homologized with elements of the neopteran wing groundplan, resulting in wing vein terminology that differs substantially from that generally used by coleopterists. We tabulate the differences between currently used venational nomenclature and the all-pterygote homologous symbols. The use of the neopteran groundplan, combined with the knowledge of the way in which veins evolved, provides many strong characters linked to the early evolutionary radiation of Coleoptera. The order originated with the development of the apical folding of the hind wings under the elytra executed by the radial and medial loop. The loops, which are very complex venational structures, further diversified in four distinctly different ways which mark the highest (suborder) taxa. The remaining venation and the wing articulation have changed with the loops, which formed additional synapomorphies and autapomorphies at the suborder, superfamily, and sometimes even family and tribe levels. Relationships among the four currently recognized suborders of Coleoptera are reexamined using hind wing characters. The number of wing-related apomorphies are 16 in Coleoptera, seven in Archostemata + Adephaga–Myxophaga, four in Adephaga–Myxophaga, seven in Myxophaga, nine in Archostemata, and five in Polyphaga. The following phylogenetic scheme is suggested: Polyphaga [Archostemata (Adephaga + Myxophaga)]. Venational evidence is given to define two major lineages (the hydrophiloid and the eucinetoid) within the suborder Polyphaga. The unique apical wing folding mechanism of beetles is described. Derived types of wing folding are discussed, based mainly on a survey of recent literature. A sister group relationship between Coleoptera and Strepsiptera is supported by hind wing evidence.

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.

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Abstract. The Dermaptera are a small order of insects, marked by reduced forewings, hindwings with a unique and complicated folding pattern, and by pincer-like cerci. Hindwing characters of 25 extant dermapteran species are documented. The highly derived hindwing venation and articulation is accurately homologized with the other pterygote orders for the first time. The hindwing base of Dermaptera contains phylogenetically informative characters. They are compared with their homologues in fossil dermapteran ancestors, and in Plecoptera, Orthoptera (Caelifera), Dictyoptera (Mantodea, Blattodea, Isoptera), Fulgoromorpha and Megaloptera. A fully homologized character matrix of the pterygote wing complex is offered for the first time. The wing venation of the Coleoptera is re-interpreted and slightly modified. The all-pterygote character analysis suggests the following relationships: Pterygota: Palaeoptera + Neoptera; Neoptera: [Pleconeoptera + Orthoneoptera] + [Blattoneoptera + (Hemineoptera + Endoneoptera)]. Blattoneoptera share at least 15 wing apomorphies with the sistergroup Hemineoptera + Endoneoptera and none with the Orthoneoptera and Pleconeoptera; Blattoneoptera: (Grylloblattodea + (Dermaptera + Dictyoptera)); Dictyoptera: (Mantodea + (Blattodea + Isoptera). Dermaptera share 13 wing apomorphies with the sistergroup Dictyoptera. In order to document the intra-ordinal relation ships of Dermaptera, 18 new characters of venation and articulation are added to an existing data set and analyzed cladistically. The following relationships are suggested (43 characters, tree length 72, CI 0.819 and RI 0.935). Dermaptera: Karschiellidae + ("Diplatyidae" + ("Pygidicranidae" + (AHostethus indicum + (Anisolabididae + ("Labiduridae" + [Forficulidae + (Chelisochidae + Spongiphoridae)]))))). The taxa in quotation marks are probably paraphyletic. Fossil Dermaptera and "Protelytroptera" show that wing-folding characters were already present in Permian ancestors. The evolution of the dermapteran wing-folding mechanism is discussed and the hindwing is presented as a working "origami" model, which will fold as in living earwigs. The functional role of the wing base in wing folding is examined. Characters in orders and other higher taxa are not independent and cannot be analyzed out of context with their groundplans. Higher systematics is dealing with diametrally different problems than species-level systematics. The necessity of using a different methodology for species-level and higher-level phylogenetics is discussed and recommenda tions are made. CONTENTS