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 first ant-termite syninclusion in amber with CT-scan analysis of taphonomy.

We describe here a co-occurrence (i.e. a syninclusion) of ants and termites in a piece of Mexican amber (Totolapa deposit, Chiapas), whose importance is two-fold. First, this finding suggests at least a middle Miocene antiquity for the modern, though poorly documented, relationship between Azteca ants and Nasutitermes termites. Second, the presence of a Neivamyrmex army ant documents an in situ raiding behaviour of the same age and within the same community, confirmed by the fact that the army ant is holding one of the termite worker between its mandibles and by the presence of a termite with bitten abdomen. In addition, we present how CT-scan imaging can be an efficient tool to describe the topology of resin flows within amber pieces, and to point out the different states of preservation of the embedded insects. This can help achieving a better understanding of taphonomical processes, and tests ethological and ecological hypotheses in such complex syninclusions.

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.