Patrick Roques

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). J. Morphol., 2012.

The earliest holometabolous insect from the Carboniferous: a “crucial” innovation with delayed success (Insecta Protomeropina Protomeropidae)

Abstract Insects dominate Earth by their diversity, and the most are Holometabola. Therefore, the holometabolous development characterised by a pupal stage between larvae and adult seems to be linked with the extensive radiation of insects. Holometaboly is suspected to appear in the carboniferous period, however until now fossils have not brought univocal evidence. The discovery in the Carboniferous (Early Langsettian, circa 310 mya, Bashkirian Stage) of France of the earliest Holometabola attributed to the Permian amphiesmenopteran or antliophoran family Protomeropidae brings the first irrefutable evidence that holometaboly existed in the Carboniferous. Given the climatic data of France at this period, this discovery contradicts the traditional scenarii of a relation between the acquisition of endopterygote pupal stage and climatic global cooling during Late Carboniferous and Early Permian. This example illustrates the hypothesis that a new, apparently more efficient, biological innovation is not always sufficient to guarantee the ‘evolutionary success’ of the concerned clade. Ecological opportunities have to be considered as well for this innovation success.

From Carboniferous to Recent: wing venation enlightens evolution of thysanopteran lineage

Recent Thysanoptera are characterized by two pairs of slender wings fringed by long hairs and with reduced venation. Fossils presenting more complete venation have been used in earlier studies to link Thysanoptera to ‘Zoropsocinae’, which form with ‘Lophioneurinae’ the ‘Lophioneuridae’, an extinct family of the superorder Thripida. On the basis of one new Carboniferous fossil described herein, Westphalothripides oudardi sp. nov. (Westphalothripidesidae fam. nov.), as well as new Cretaceous fossils, we revise the current interpretation of venation in Thripida and propose new cladistic analyses, which divide this superorder into three clades: Panthysanoptera nov. (including ‘Zoropsocidae’ stat. rest. and Thysanoptera), Lophioneurida (including ‘Lophioneurinae’ and Moundthripidae), and Westphalothripidesidae. We confirm that the Thysanoptera belong to the superorder Thripida but show that ‘Lophioneuridae’ are paraphyletic. Thanks to the venation observed in fossils, we describe a new character of the wing in some Recent species which allows them to be integrated in phylogenetic studies. Results show two clades inside Thysanoptera: Aeolothripidae + Melanthripidae + Merothripidae, and Thripidae + Stenurothripidae + Fauriellidae. Phlaeothripidae could not be studied due to the absence of the diagnostic wing veins in this family. We discuss the appearance of Thripida within Paraneoptera and particularly the importance of one apomorphy, the reduction of the right mandible as a putative adaptation to spore and pollen piercing.

New non-holometabolous insects from the Pennsylvanian of the Avion locality in Pas-de-Calais, France (Insecta: 'Exopterygota')

Prokop, J., Roques, P. & Nel, A., 2014. New non-holometabolous insects from the Pennsylvanian of the Avion locality in Pas-de-Calais, France (Insecta: ‘Exopterygota’). Alcheringa 38, 155–169. ISSN 0311–5518. New palaeopteran and neopteran insects are described from Moscovian impression fossils of the Pas-de-Calais Basin in northern France. Spilaptera splendens sp. nov. (Palaeodictyoptera) is based on new forewing venation characters including a well-developed anal bar in the form of a cuticular reinforcement extending from the anal area to the radial stem, a feature previously unknown in this genus. This character was also confirmed after re-examination of S. libelluloides. The discovery of a wing part with a large anal fan supports the presence of a gigantic palaeodictyopteran that can be assigned to Homoiopteridae. Based on a petiolate wing, Jacquesoudardia magnifica gen. et sp. nov. represents a small damselfly-like odonatopteran that was probably predatory on small arthropods and coexisted with larger meganeurid species such as Gallotupus oudardi from the same locality. The discovery of Paolia sp. (Paoliidae), known previously from the Bashkirian of Indiana (USA) and South Limbourg (the Netherlands), extends the distribution and stratigraphic range of the genus. Characterized by a short ScP, Aviogramma gracilis gen. et sp. nov. is the second member of Caloneurodea from the Pas-de-Calais Basin after Pruvostiella lecomtei (Pruvost). Aviocladus pectinatus gen. et sp. nov. and Aviohapaloptera bethouxi gen. et sp. nov. are both assigned to Archaeorthoptera. Aviocladus pectinatus, a member of Cnemidolestodea, preserves a venation pattern that allows restoration of Tococladus to Archaeorthoptera. Aviohapaloptera has strongly similar venation to Hapaloptera, known from Mazon Creek (USA), in bearing a pectinate CuPb, CuA+CuPa with few posterior branches, a narrow costal area and radial posterior (RP) with few branches. In addition, we restore Narkeminidae stat. rest. and separate it from Proedischiidae. These new fossils demonstrate that the insect fauna from the Pas-de-Calais Basin was highly diverse with links to Bashkirian/Moscovian assemblages in Euramerica, such as the Mazon Creek (Illinois, USA) entomofauna. Jakub Prokop [jprokop@natur.cuni.cz], Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44, Praha 2, Czech Republic. Patrick Roques [patrick.roques93@wanadoo.fr] allée des Myosotis, Neuilly sur Marne, F-93330, France. André Nel [anel@mnhn.fr] CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 rue Buffon, F-75005 Paris, France. Received 15.7.2013, revised 10.9.2013, accepted 23.9.2013.

A new genus and species of Breyeriidae and wings of immature stages from the Upper Carboniferous, Nord-Pas-de-Calais, France (Insecta: Palaeodictyoptera)

A new palaeodictyopterid, Aviobreyeria gracilis gen.n., sp.n., is described from an outcrop of Upper Carboniferous (Westphalian C/D) deposits at Avion in the Nord-Pas-de-Calais department (France). This new taxon is placed in the family Breyeriidae based on a comparison of the pattern of venation in its forewing venation with that of other breyeriids and possibly related genera within the Breyerioidea. In addition, wings of different immature instars of palaeodictyopterids from the same strata with a similar pattern of venation but differing in having an enlarged RP and a broader anal area (supposed hind wings) were also tentatively attributed to Breyeriidae gen. et sp. indet.. These wings have a roughly identifiable venation and exhibit the gradual development of wings through different ontogenetic stages and their position on the thorax, which is sparsely documented for Paleozoic insects. Discovery of immature and adult stages of palaeodictyopterids in the same layers also indicates they lived in similar habitats.

Revision of the Enigmatic Insect Family Anthracoptilidae Enlightens the Evolution of Palaeozoic Stem-Dictyopterans

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 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 CuPaβ 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.

A new, extraordinary ‘damselfly-like’ Odonatoptera from the Pennsylvanian of the Avion locality in Pas-de-Calais, France (Insecta: ‘Exopterygota’)

Nel, A., Roques, P., Prokop, J., & Garrouste, R., 11 September 2018. A new, extraordinary ‘damselfly-like’ Odonatoptera from the Pennsylvanian of the Avion locality in Pas-de-Calais, France (Insecta: ‘Exopterygota’). Alcheringa 43, 241–245. ISSN 0311-5518. Enigmaptera magnifica gen. et sp. nov., type genus and species of the new odonatopteran family Enigmapteridae, is described from the Moscovian of Avion (northern France). It is the sister group of the major clade Neodonatoptera, placed together in the new clade Paneodonatoptera. Its wing venation has characters never found in other Odonatoptera. It is a further case of convergent wing petiolation in this superorder. Enigmaptera magnifica, like the protozygopteran Jacquesoudardia magnifica from the same outcrop, probably lived like the extant damselflies along the shores of lakes and rivers, hunting the small insects found in the same deposits. These discoveries show that very small insects were significant elements of the entomofaunal diversity and trophic chains of the Late Carboniferous ecosystems. Romain Garrouste [garroust@mnhn.fr] Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE, 57 rue Cuvier, CP 50, 75005 Paris, France; Patrick Roques [patrick.roques93@wanadoo.fr] Allée des Myosotis, Neuilly sur Marne, F-93330, France.; Jakub Prokop [jprokop@natur.cuni.cz] Charles University, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44, Praha 2, Czech Republic.