A further strange Archaeorthoptera from the Moscovian of Nord-Pas-de-Calais (France) (Insecta, Polyneoptera).

The new Archaeorthoptera Duquesnia gallica gen. et sp. nov. Nel Roques, is described from the upper Carboniferous of Northern France. It shows several putative synapomorphies with the three genera Contracladus Dvok et al., 2021 (Pennsylvanian of Germany), Nugonioneura (lower Permian of USA), and Avionugonioneura from the Moscovian of Avion (France).

Sound vs. light: wing-based communication in Carboniferous insects.

Acoustic communication is well-known in insects since the Mesozoic, but earlier evidence of this behavior is rare. Titanoptera, an 'orthopteroid' Permian-Triassic order, is one of the few candidates for Paleozoic intersex calling interactions: some specimens had highly specialized broadened zones on the forewings, which are currently considered-despite inconclusive evidence-as 'resonators' of a stridulatory apparatus. Here we argue that the stridulatory apparatus hypothesis is unlikely because the Titanoptera lack a stridulatory file on their bodies, legs or wings. Instead, comparing these broadened zones with similar structures in extant locusts, flies, and fossil damselflies, we find evidence that the Titanoptera used their wings to produce flashes of light and/or crepitated sounds. Moreover, we describe the first Carboniferous (~310 Mya) Titanoptera, which exhibits such specialized zones, thus corresponding to the oldest record of wing communication in insects. Whether these communication systems were used to attract sexual partners and/or escape predators remain to be demonstrated.

New archaeorthopteran insects from the Late Carboniferous of the Nord and Pas-de-Calais basins in northern France (Insecta: Cnemidolestodea, Panorthoptera).

New polyneopteran insects are described from Pennsylvanian (Bashkirian, Moscovian) compressed fossils from the North of France (Insecta: Archaeorthoptera). Discovery of wing apex with distinct venation, e.g., apical fusion of RA with RP, numerous parallel posterior branches of RP with transversal crossveins, can be assigned to cf. Tococladus sp. (Cnemidolestodea: Tococladidae). It represents the second record of Cnemidolestodea from the Avion locality apart from Aviocladus pectinatus Prokop et al., 2014. Bruaylogus magnificus gen. et sp. nov., based on forewing venation, is attributed to Panorthoptera nec Orthoptera having some distinct characters for the placement either close to Oedischiidae or a more basal position possibly with affinities to genus Heterologus. Aviologus duquesnei gen. et sp. nov., based on forewing venation, differs from Oedischiidae by the presence of basal fork of M far from point of separation between M and Cu and fusion of MA with first posterior branch of RP. Aviologus share a long stem of M and simple CuPaß with Heterologus duyiwuer and H. langfordorum, but both differ in well separated median and radial veins. These new fossils demonstrate that the archaeorthopterid insect fauna from the North of France was rather diverse with links to late Carboniferous and early Permian assemblages in Euramerica such as the Mazon Creek, Carbondale Formation or Elmo, Wellington Formation (Illinois, Kansas, USA) entomofaunas.

The earliest known holometabolous insects.

The Eumetabola (Endopterygota (also known as Holometabola) plus Paraneoptera) have the highest number of species of any clade, and greatly contribute to animal species biodiversity. The palaeoecological circumstances that favoured their emergence and success remain an intriguing question. Recent molecular phylogenetic analyses have suggested a wide range of dates for the initial appearance of the Holometabola, from the Middle Devonian epoch (391 million years (Myr) ago) to the Late Pennsylvanian epoch (311 Myr ago), and Hemiptera (310 Myr ago). Palaeoenvironments greatly changed over these periods, with global cooling and increasing complexity of green forests. The Pennsylvanian-period crown-eumetabolan fossil record remains notably incomplete, particularly as several fossils have been erroneously considered to be stem Holometabola (Supplementary Information); the earliest definitive beetles are from the start of the Permian period. The emergence of the hymenopterids, sister group to other Holometabola, is dated between 350 and 309 Myr ago, incongruent with their current earliest record (Middle Triassic epoch). Here we describe five fossils--a Gzhelian-age stem coleopterid, a holometabolous larva of uncertain ordinal affinity, a stem hymenopterid, and early Hemiptera and Psocodea, all from the Moscovian age--and reveal a notable penecontemporaneous breadth of early eumetabolan insects. These discoveries are more congruent with current hypotheses of clade divergence. Eumetabola experienced episodes of diversification during the Bashkirian-Moscovian and the Kasimovian-Gzhelian ages. This cladogenetic activity is perhaps related to notable episodes of drying resulting from glaciations, leading to the eventual demise in Euramerica of coal-swamp ecosystems, evidenced by floral turnover during this interval. These ancient species were of very small size, living in the shadow of Palaeozoic-era 'giant' insects. Although these discoveries reveal unexpected Pennsylvanian eumetabolan diversity, the lineage radiated more successfully only after the mass extinctions at the end of the Permian period, giving rise to the familiar crown groups of their respective clades.

Traits and evolution of wing venation pattern in paraneopteran insects.

Two different patterns of wing venation are currently supposed to be present in each of the three orders of Paraneoptera. This is unlikely compared with the situation in other insects where only one pattern exists per order. We propose for all Paraneoptera a new and unique interpretation of wing venation pattern, assuming that the convex cubitus anterior gets fused with the common stem of median and radial veins at or very near to wing base, after separation from concave cubitus posterior, and re-emerges more distally from R + M stem. Thereafter, the vein between concave cubitus posterior and CuA is a specialized crossvein called "cua-cup," proximally concave and distally convex. We show that despite some variations, that is, cua-cup can vary from absent to hypertrophic; CuA can re-emerge together with M or not, or even completely disappear, this new interpretation explains all situations among all fossil and recent paraneopteran lineages. We propose that the characters "CuA fused in a common stem with R and M"and "presence of specialized crossvein cua-cup" are venation apomorphies that support the monophyly of the Paraneoptera. In the light of these characters, we reinterpret several Palaeozoic and early Mesozoic fossils that were ascribed to Paraneoptera, and confirm the attribution of several to this superorder as well as possible attribution of Zygopsocidae (Zygopsocus permianus Tillyard, 1935) as oldest Psocodea. We discuss the situation in extinct Hypoperlida and Miomoptera, suggesting that both orders could well be polyphyletic, with taxa related to Archaeorthoptera, Paraneoptera, or even Holometabola. The Carboniferous Protoprosbolidae is resurrected and retransferred into the Paraneoptera. The genus Lithoscytina is restored. The miomopteran Eodelopterum priscum Schmidt, 1962 is newly revised and considered as a fern pinnule. In addition, the new paraneopteran Bruayaphis oudardi gen. nov. et sp. nov. is described fromthe Upper Carboniferous of France (see Supporting Information).