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The position of the Palaeozoic Anthracoptilidae has been a major problem of insect systematics for over a century. The previous hypotheses suggested affinities of Anthracoptilidae with the Palaeodictyopteroida, Protorthoptera, Hypoperlida, stem-Mantodea, Paraneoptera, Holometabola, or Eoblattida. Herein we put forward a new hypothesis based mainly on a comprehensive re-evaluation of the wing venation characters and re-examination of the type material of Anthracoptilus perrieri and Mesoptilus dolloi. The Anthracoptilidae are considered as belonging to the paoliid lineage, sister group of the Dictyoptera. In particular this result refutes the presence of Mantodea in the Paleozoic fossil record. The two families Strephocladidae and Strephoneuridae, are considered as junior synonyms of the Anthracoptilidae, while the previous synonymy of the Anthracoptilidae with the Ischnoneuridae is rejected. We consider the Permarrhaphidae, previously considered as synonym of the Anthracoptilidae, (and Permarrhaphus) as Insecta incertae sedis. The following new taxa are proposed: Mesoptilus carpenteri sp. nov. from the early Permian of Wellington Formation in Oklahoma (USA) which extends the range and distribution of the genus; Pseudomesoptilus gen. nov. is designated to include Mesoptilus sellardsi; Strephocladus permianus sp. nov. from the middle Permian of South of France; Westphaloptilus gallicus gen. et sp. nov. from the Bashkirian of the North of France.
New representatives of a stem group Paoliida attributed to family Paoliidae (Insecta: Protoptera) are described from the Upper Carboniferous (Langsettian) sphero−sideritic concretions of the Upper Silesian Coal Basin (USCB) in Poland. Zdenekia silesiensis sp. nov. is based on forewing venation and supplemented by material of isolated hindwing similar in venation pattern. Darekia sanguinea gen. et sp. nov. differs from all other paoliid genera by the presence of a short contact between veins MP and CuA behind the division CuA and CuP. Composition of insect fauna exhibits high abundance of paoliid insects in the early Late Carboniferous ecosystems known also from other European localities such as Hagen Vorhalle in Ruhr Basin (Germany), and South Limbourg (Belgium and the Netherlands). It is the first record of true paoliids from the Polish part of paralic USCB supplementing a single historical record of Stygne roemeri considered as a taxon closely related to Paoliidae. The high abundance of paoliid insects from sphero−sideritic concretions in Sosnowiec and coal deposits previously known from the Czech part of Upper Silesian Coal Basin indicates considerable similarity of both faunas supported as well by their close stratigraphical correlation. Morphology of basal wing parts with remnants of articular sclerites preserved supports neopteran relationships of paoliids. Discovery of the first paoliid immature wing is reported suggesting similar living habitat for larvae and adults.
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