A new family and two new genera from Avion, Northern France, confirm the high Moscovian (late Carboniferous) diversity of the insect superorder Archaeorthoptera

ANDRÉ NEL and PATRICK ROQUES

Nel, A. and Roques, P. 2021. A new family and two new genera from Avion, Northern France, confirm the high Moscovian (late Carboniferous) diversity of the insect superorder Archaeorthoptera. Acta Palaeontologica Polonica 66 (4): 879–884.

The new archaeorthopteran family Archaeogeraridae, based on Archaeogerarus schubneli gen. et sp. nov., is described from the Moscovian of Avion. It has several unique wing venation characters allowing to separate it from the other groups of this superorder, viz. a vein RP separating from RA very close to wing base and closely parallel to it; M shortly fused with RP; CuA+CuPa simple and straight; and CuPb branched. This last character is convergently present in a few Palaeozoic Panorthoptera, a feature of great interest for a future phylogenetic analysis of the whole superorder. Avionxixia gui gen. et sp. nov., second Cnemidolestidae from the Moscovian of Avion, is described and illustrated. It shares numerous characters with the Chinese Namurian genus Xixia and the European (Germany and France) Moscovian genus Piesbergopterum, suggesting possible phylogenetic affinities between these three genera. It is mainly separated from these two genera by the costal area much narrower than the subcostal one and the presence of only one posterior branch of the anterior branch of MP+CuA+CuPa. This new taxon confirms the high diversity of the Cnemidolestidae during the late Carboniferous.

Key words: Insecta, Polyneoptera , Cnemidolestodea, Archaeogeraridae, forewing venation convergences, Carboni­ferous, France.

André Nel [anel@mnhn.fr, ORCID: https://orcid.org/0000-0002-4241-7651], Institut de Systématique, Évolution, Biodiversité (ISYEB) Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, CP50, 57 rue Cuvier 75005 Paris, France.

Patrick Roques [patrick.roques93@wanadoo.fr, ORCID: https://orcid.org/0000-0002-8721-9763], 22 Chemin des Processions, 93360 Neuilly Plaisance, France.

Received 2 September 2021, accepted 6 October 2021, available online 3 December 2021.

Copyright © 2021 A. Nel and P. Roques. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Introduction

The superorder Archaeorthoptera, now-a-day reduced to the sole Orthoptera, witnesses an impressive diversification during the Palaeozoic and the early Mesozoic, with the orders Caloneurodea, Titanoptera, Cnemidolestodea, stem Orthoptera and a complex series of families and genera belonging to the Archaeorthoptera nec Panorthoptera (Béthoux and Nel 2002). Some historical outcrops have given numerous fossils of this superorder (e.g., Mazon Creek, USA; Com­mentry, France), showing that it was probably among the most diverse clade during the late Carboniferous, together with the two other polyneopteran groups Dictyoptera and “Grylloblattodea”. But, if the Dictyoptera were clearly very abundant, their morphological disparity was clearly lower than that of the Archaeorthoptera. That of the “Grylloblattodea” was very high too, but the monophyly of this group of taxa remains to be verified, unlike that of the Archaeorthoptera, well supported by a series of wing venation synapomorphies. Thanks to the efforts of one of us (PR), the Moscovian outcrop of Avion (Northern France) has given recently an impressive series of fossil insects, among which were found the oldest representatives of the Acercaria and Holometabola (Nel et al. 2013), but also numerous Archaeorthoptera of several orders and families. Nevertheless, all these new genera and species are based on single fossils, suggesting that the diversity of the original entomofauna was much higher than what has been recovered.

The present descriptions of two new fossils representing a new family, and two new genera and species support this assumption. They are of great interest for a better knowledge of the Moscovian diversity and morphological disparity of the Archaeorthoptera.

Institutional abbreviations.—MNHN, Muséum National d’Histoire Naturelle, Paris, France.

Other abbreviations.—A, anal vein; C, costa; CuA, cubitus anterior; CuP, cubitus posterior; CuPa, anterior branch of CuP; CuPaa, anterior branch of CuPa; CuPab, posterior branch of CuPa; CuPb, posterior branch of CuP; MA, median anterior vein; MP, median posterior vein; PCu, postcubital vein; RA, radius anterior; RP, radius posterior; ScP, subcostal posterior.

Nomenclatural acts.—This published work and the nomenclatural acts it contains, have been registered in ZooBank: urn:lsid:zoobank.org:pub:7D23AD1B-CA20-4BB8-A926-CEE97C49ADF1

Material and methods

The fossils were found by one of us (PR) in the slag heap of Avion. They were examined and drawn under a Nikon SMZ1500, and photographed with an AmScope camera MU900.

We follow the wing venation terminology of Béthoux and Nel (2002) modified by Schubnel et al. (2020), and the classification of Béthoux (2005) for the Cnemidolestodea.

Systematic palaeontology

Superorder Archaeorthoptera Béthoux and Nel, 2002

Order undetermined

Family Archaeogeraridae nov.

Zoobank LSID: urn:lsid:zoobank.org:act:5283533C-1641-4366-A77D- 5CADECB66536

Type genus: Archaeogerarus gen. nov.; see below.

Diagnosis.—As for the type genus by monotypy.

Genus Archaeogerarus nov.

Zoobank LSID: urn:lsid:zoobank.org:act:95D9C644-779C-440E-BBCC-71D6846D3D40

Type species: Archaeogerarus schubneli sp. nov.; see below.

Etymology: From Ancient Greek ἀρχαῖος (arkhaîos), primitive, and the genus Gerarus for its similarity in the forewing vein CuPb (gender masculine).

Diagnosis.—As for the type species by monotypy.

Archaeogerarus schubneli sp. nov.

Fig. 1.

Zoobank LSID: urn:lsid:zoobank.org:act:81243C11-E3B1-4DF4-9561- C0E248E23A41

Etymology: In honour of Thomas Schubnel, for his impressive scientific activity and the very useful interactions we have in the study of extant and fossil insect wing venation.

Holotype: MNHN.F. A70991, part and counterpart of the basal half of a forewing, with polarity of veins well visible.

Type locality: Terril N°7, Avion, Pas-de-Calais, France.

Type horizon: Moscovian (Westphalian C/D equivalent to Bolsovian/Asturian), middle Pennsylvanian, Carboniferous.

Material.—Type material only.

Diagnosis.—Forewing characters only. Base of RP opposite point of separation of M and CuA; RP closely parallel with RA; M shortly fused with RP and separating again distally (autapomorphy); CuA+CuPa simple (autapomorphy); CuPa very short; CuPb (+ first branch of PCu) with elongate distal branches; second branch of PCu simple; first anal vein with elongate branches; second anal vein elongate and simple. Further characters: ScP ending into C; CuP with three branches; M with four branches.

Description.—Based on forewing venation. Length of preserved part 17.0 mm, maximum width at midwing 7.5 mm; original coloration of wing membrane not preserved or hyaline; concave ScP distally slightly zigzagged, running parallel with costal margin, not ending into RA; costal area with numerous simple crossveins, 0.6 mm wide, slightly narrower than subcostal area, 0.8 mm wide; stem of R diverging from M+CuA near base of wing; division of RA and RP very basal, opposite point of separation of M and CuA, less than 1.0 mm distal of point of separation of R and M+CuA; strongly convex RA simple; concave RP very closely parallel with RA and with at least two posterior branches; neutral vein M diverging from M+CuA very close to its base, parallel to R, shortly fused with RP, and separating again 0.2 mm distally; M divided into MA and MP 4.5 mm distal to its separation from RP; both MA and MP with a distal fork; concave vein CuPa very short, ending into convex CuA 2.0 mm behind divergence of M from CuA; convex CuA+CuPa straight and simple (even if there are several oblique crossveins between it and MP superficially looking like anterior branches of CuA+CuPa); elongate concave CuP basally strongly curved towards R+M+CuA, CuPb with three long branches; a convex vein (anterior branch of PCu) separating from PCu at its base and ending into CuP; anal area with simple convex vein PCu 9.8 mm long, running parallel to CuP and CuPb; a broad area between CuP/CuPb and PCu; first anal vein with three branches; a second simple anal vein; no anal loop.

Remarks.—Archaeogerarus gen. nov. can be clearly attri­buted to the Archaeorthoptera as it shares the main diagnostic character of this superorder, namely the basal fusion of CuA with M and subsequent connection with the concave anterior branch of CuP as CuPa (Béthoux and Nel 2002). The absence of a division of CuPa into two branches CuPaa and CuPab is a symplesiomorphy excluding it from the Panorthoptera sensu Béthoux and Nel (2002). The most remarkable characters of Archaeogerarus gen. nov. is the vein RP closely parallel to RA and branching from R opposite the point of separation of M and CuA, and the vein M directly ending into RP to separate again just distally. A RP very long and closely parallel to RA is present in some Anthracoptilidae (in Palaeozoic order Paoliida), e.g., Mesoptilus dolloi Lameere, 1917 (Guan et al. 2015); but these have a completely different pattern of the median and cubital veins at wing base, with a stem Cu from which the convex CuA and the concave CuP distally emerge. Such a vein RP elongate and closely parallel to RA, is also present in few Archaeorthoptera, viz. Eoblatta robusta (Brongniart, 1893), Beloatta duquesnei Nel, Garrouste, and Roques, 2020, Ctenoptilus elongatus (Brongniart, 1893), Ischnoneura oustaleti Brongniart, 1893, Ischnoptera diaphanes Béthoux and Nel, 2005, Kitshuga ryzhkovae Aristov, 2012 (Béthoux and Nel 2005; Schubnel et al. 2019). All these taxa strongly differ from Archaeogerarus gen. nov. in the CuA+CuPa with numerous strong branches vs. simple in Archaeogerarus schubneli gen. nov., M not fused with RP near their bases, and CuPb simple vs. with branches in Archaeogerarus schubneli gen. nov.

In the other polyneopteran orders, the vein CuP is simple, suggesting that that a branched CuPb is a specialized structure of some Archaeorthoptera. It is encountered in very few Archaeorthopteran Panorthoptera, viz. Gerarus bruesi Meunier, 1909, Owadpteron dareki Dvořák, Pecha­rová, Krzemiński, and Prokop, 2019, and Nace­ko­mia rossae Richardson, 1956 (Béthoux and Nel 2002; Dvořák et al. 2019). But these have a branched CuPa into a CuPaa and a CuPab (as Panorthoptera), and a branched CuA+CuPaa, unlike Archaeogerarus schubneli gen. nov. Archaeogerarus gen. nov. is the first Archaeorthoptera not belonging to the Panorthoptera with a branched CuPb. This character is a putative convergence between it and the panorthopteran genera Gerarus, Owadpteron, and Nacekomia. Notice that the pattern of PCu and of the anal veins of Archaeogerarus schubneli gen. nov. is similar to that of Nacekomia rossae, these veins being very long and parallel.

The presence of this unique combination of characters plus some extremely particular structures supports the attribution of Archaeogerarus schubneli gen. et sp. nov. to a new family of Archaeorthoptera.

Stratigraphic and geographic range.—Type locality and horizon only.

Order Cnemidolestodea Handlirsch, 1937
(sensu Béthoux 2005)

Family Cnemidolestidae Handlirsch, 1906

Genus Avionxixia nov.

Zoobank LSID: uurn:lsid:zoobank.org:act:2494907A-6BFE-4FCC-97 CA-E615D573272D

Type species: Avionxixia gui sp. nov.; see below.

Etymology: Named after the type locality Avion and the genus Xixia.

Diagnosis.—As for the type species by monotypy.

Avionxixia gui sp. nov.

Fig. 2.

Zoobank LSID: urn:lsid:zoobank.org:pub:7D23AD1B-CA20-4BB8-A926-CEE97C49ADF1

Etymology: Named after Jun-Jie Gu, for his work on the Cnemidolestidae.

Holotype: MNHN.F. A70990, part and counterpart of a nearly complete forewing, with only apex missing.

Type locality: Terril N°7, Avion, Pas-de-Calais, France.

Type horizon: Moscovian (Westphalian C/D equivalent to Bolsovian/Asturian, middle Pennsylvanian, Carboniferous).

Material.—Type material only.

Diagnosis.—Forewing characters only. Only one posterior branches of anterior branch of MP+CuA+CuPa; veinlet between MP+CuA+CuPa and posterior branch of MA strongly zigzagged and very short; area between MA and anterior branch of MP+CuA+CuPa quite short, less than three times as long as wide; anterior branch of MA not touching RP; costal area distinctly narrower than subcostal area. Forewing only 12.1 mm long.

Description.—Based on forewing venation. Estimated total wing length 12.1 mm, maximum width at midwing 4.1 mm; bending of costal margin in about basal third; original coloration of wing membrane not preserved or hyaline; concave ScP slightly curved, running parallel with costal margin, ending to RA behind midwing; costal area with numerous simple crossveins, 0.3 mm wide, narrower than subcostal area, 0.6 mm wide; stem of R diverging from M+CuA near base of wing; division of RA and RP proximal of midwing, 1.5 mm basal to connection of ScP on RA; strongly convex RA simple ending on costal margin well basal to wing apex; numerous oblique crossveins present in space between RA and costal wing margin; concave RP posteriorly pectinate ending with four branches covering whole distal part of wing; neutral vein MA diverging from M+CuA and further running parallel to stem of R/RP; MA deeply forked into two simple branches, anterior one MA1 not connected to RP and posterior one MA2 nearly straight; concave vein CuPa ending into MP+CuA shortly behind divergence of MA from MP+CuA; MP+CuA+CuPa emitting a posterior branch apically subdivided into two branches; and an anterior branch parallel to MA, emitting only one posterior branch, distally forked, a strongly zigzagged veinlet between MP+CuA+CuPa and MA, closing a short and broad area between it and MA (0.8 mm wide, 2.0 mm long), crossed by irregular veinlets; areas between branches of RP, MA and MP+CuA+CuPa regularly connected by simple and oblique crossveins; concave CuP basally dividing into CuPa and CuPb, simple CuPb running parallel to MP+CuA+CuPa towards posterior wing margin; anal area with convex vein PCu running parallel to CuP and CuPb; first anal vein distally connected to PCu, forming an elongate “anal loop”, 2.5 mm long; a second short anal vein.

Remarks.—Avionxixia gen. nov. can be attributed to the Archaeorthoptera as it shares the main diagnostic character of this superorder, namely the basal fusion of CuA with M and subsequent connection with the anterior branch of CuP as CuPa (Béthoux and Nel 2002). Furthermore, it displays characters typical of the order Cnemidolestodea, such as ScP terminating on RA, CuPa ending into MP+CuA, MP+CuA+CuPa emitting an anterior branch parallel to MA, with a specialised veinlet between it and MA, thus defining a large area between it and MA crossed by irregular veinlets (Béthoux 2005; Gu et al. 2014: figs. 1, 2). Aristov (2014), using a different diagnosis and wing venation nomenclature for the Cnemidolestodea, proposed a key to families.

Avionxixia gen. nov. falls in the family Cnemidolestidae Handlirsch, 1906 (sensu Aristov 2014: 10) because of the cha­racter “MP weak, ending on CuA or MA, or absent”. Indeed, in Avionxixia gen. nov., MP is clearly basally fused with CuA.

The family Cnemidolestidae comprises the following genera, after Aristov (2014), Gu et al. (2014), and Dvořák et al. (2021): Aetophlebia Scudder, 1885, Amphibolia­cri­dites Langiaux and Parriat, 1974, Anarkemina Aristov, 2014, Argen­tinonarkemina Martins-Neto, Gallego, and Brauck­mann, 2007, Bouleites Lameere, 1917, Carbonokata Aristov, 2013, Cnemidolestes Handlirsch, 1906, Evenkio­phle­bia Ari­stov, 2013, Irajanarkemina Martins-Neto, Gal­lego, and Brauckmann, 2007, Ischnoneura Brongniart, 1893, Longzhua Gu, Béthoux, and Ren, 2011, Narkema Hand­lirsch, 1911, Narkemina Martynov, 1930, Narke­minopsis Whalley, 1979, Narkeminuta Aristov, 2013, Narke­mulla Aristov, 2013, Para­narkemina Pinto and Ornellas, 1980, Piesbergopterum Dvo­řák, Pecharová, Leipner, Nel, and Prokop, 2021, Proto­di­amphipnoa Brongniart, 1885, Tshunoptera Aristov, 2013, Velizphlebia Martins-Neto, Gal­lego, and Brauckmann, 2007, and Xixia Gu, Béthoux, and Ren, 2014.

Narkema (N. taeniatum Handlirsch, 1911, N. alternatum Cockerell, 1924) shares with Avionxixia gen. nov. the presence of only two posterior branches of anterior branch of MP+CuA+CuPa, but it has a very long stem of RP, a very long stem of anterior branch of MP+CuA+CuPa, and posterior branch of MP+CuA+CuPa branched (Handlirsch 1911: fig. 28; Cockerell 1924, 1927).

Irajanarkemina, Velizphlebia, Argentinonarkemina, and Paranarkemina have few or no posterior branch of the anterior branch of MP+CuA+CuPa, but they all have a longer stem of RP and a narrower area between MA and MP+CuA+CuPa than in Avionxixia gen. nov. Aetophlebia and Longzhua also differ from Avionxixia gen. nov. in the same characters (Scudder 1890: 301–302, pl. 17: 9; Martins-Neto et al. 2007: figs. 2, 7; Gu et al. 2011: fig 3). Avionxixia gen. nov. differs from Narkeminopsis in much broader area between MA and MP+CuA+CuPa (Whalley 1979: fig. 3; Brauckmann and Herd 2006: fig. 9; Béthoux and Nel 2005: fig. 20; Aristov 2013: fig. 1b).

Piesbergopterum (Piesbergopterum punctatum Dvořák, Pecharová, Leipner, Nel, and Prokop, 2021, Piesbergopterum schubneli Nel and Roques, 2021c), and Xixia (Xixia huban Gu, Béthoux, and Ren, 2014) share with Avionxixia gen. nov. the presence of few posterior branches of anterior branch of MP+CuA+CuPa (two or three at most) (Gu et al. 2014; Dvořák et al. 2021; Nel and Roques 2021c). The specialized veinlet between MP+CuA+CuPa and posterior branch of MA is short and simple in Piesbergopterum, vs. elongate and more or less sigmoidal with crossvein(s) branching on it in Xixia. In Avionxixia gen. nov., this veinlet is strongly zigzagged and very short. Piesbergopterum has a posterior branch of MP+CuA+CuPa with more than two branches, unlike Avionxixia gen. nov. and Xixia. Lastly, the area between MA and anterior branch of MP+CuA+CuPa is quite short in Avionxixia gen. nov., less than three times as long as wide, while it is three times (or more) as long as wide in Xixia and Piesbergopterum. The anterior branch of MA in Avionxixia gen. nov. is not touching RP, unlike in Piesbergopterum. The costal area of Avionxixia gen. nov. is distinctly narrower than the subcostal area, unlike in Piesbergopterum and Xixia. Lastly the forewing of Avionxixia gen. nov. is only 12.1 mm long, instead of 20–21 mm in the two species of Piesbergopterum, and 17.9–21.8 mm in Xixia huban.

All the other cnemidolestid genera have three or more posterior branches of the anterior branch of MP+CuA+CuPa, which is itself quite elongate, unlike Avionxixia gen. nov.

It is worth to note that apart from Piesbergopterum avionensis there are two other cnemidolestodeans known from Avion: Avionxixia gui gen. nov. differs strongly both from Aviocladus pectinatus Prokop, Roques, and Nel, 2014 and Aviohapaloptera bethouxi Prokop, Roques, and Nel, 2014 in the presence of a strong anterior branch of MP+CuA+CuPa and the different shape of CuA+CuPa (Prokop et al. 2014).

Stratigraphic and geographic range.—Type locality and horizon only.

Concluding remarks

With these two new taxa, the Avion locality shows an impressive diversity of Archaeorthoptera, with no less than 13 ge­nera and 13 species (Table 1; 10% of the ca. 121 Carboniferous archaeorthopteran species, after the Fossilworks Database http://fossilworks.org accessed 30/08/2021). It confirms that this superorder underwent an important diversification during the late Carboniferous, with a high disparity of wing venations. Unfortunately no body structure of these insects has been found in Avion, which is also rarely encountered in the other Carboniferous outcrops. Thus their biology remains unknown in main part. The discovery of Archaeogerarus schubneli gen. et sp. nov. with a highly specialized branched vein CuPb, convergently present in a few Panorthoptera, is especially interesting for a future phylogenetic analysis of the whole superorder Archaeorthoptera.

Acknowledgements

We thank a lot Eric Damay (Society Eiffage Route Nord Est, Avion, France) for the kind authorization to collect fossil insects in the terril of Avion. We also sincerely thank two anonymous referees, and Thomas Schubnel (MNHN) for their useful remarks on the first version of the paper.

References

Aristov, D.S. 2012. Composition and distribution of the family Cacurgidae (Insecta; Grylloblattida). Paleontological Journal 46: 250–257. Crossref

Aristov, DS. 2013. New and little-known Eoblattida (Insecta) from the Paleozoic of Russia. Paleontological Journal 47: 272–282. Crossref

Aristov, D.S. 2014. Classification of the order Cnemidolestida (Insecta: Perlidea) with descriptions of new taxa. Far Eastern Entomologist 277: 1–46.

Béthoux, O. 2005. Cnemidolestodea (Insecta): an ancient order reinstated. Journal of Systematic Palaeontology 3: 403–408. Crossref

Béthoux, O. and Nel, A. 2002. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1–88. Crossref

Béthoux, O. and Nel, A. 2005. Some Palaeozoic “Protorthoptera” are “ancestral” orthopteroids: major wing braces as clues to a new split among the “Protorthoptera”. Journal of Systematic Palaeontology 2 (4): 1–25. Crossref

Brauckmann, C. and Herd, K.J. 2006. Insekten-Funde aus dem Westfalium D (Ober-Karbon) des Piesberges bei Osnabruck (Deutschland). Teil 2: Neoptera. Osnabrücker Naturwissenschaftliche Mitteilungen 30/31: 19–65.

Brongniart, C. 1885. Les insectes fossiles des terrains primaires. Coup d’œil rapide sur la faune entomologique des terrains paléozoïques. Bulletin de la Société des Amis des Sciences Naturelles de Rouen 21: 50–68. Crossref

Brongniart, C. 1893. Recherches pour servir à l’histoire des insectes fossiles des temps primaires précédées d’une étude sur la nervation des ailes des insectes. Bulletin de la Société d’Industrie Minérale de Saint-Etienne (3) 7: 1–491. Crossref

Cockerell, T.D.A. 1924. Fossil insects. Entomological News 35: 28–30.

Cockerell, T.D.A. 1927. The Carboniferous insects of Maryland. Annals and Magazine of Natural History (9) 19: 385–416. Crossref

Coty, D., Háva, J., Prokop, J., Roques, P. and Nel, A. 2014. New archaeorthopteran insects from the late Carboniferous of the Nord and Pas-de-Calais basins in northern France (Insecta: Cnemidolestodea, Panorthoptera). Zootaxa 3878: 462–470. Crossref

Dvořák, T., Pecharová, M., Krzemiński, W., and Prokop, J. 2019. New archa­eorthopteran insects from the Carboniferous of Poland: insights into tangled taxonomy. Acta Palaeontologica Polonica 64 (4): 787–796. Crossref

Dvořák, T., Pecharová, M., Leipner, A., Nel, A., and Prokop, J. 2021. New archaeorthopteran insects from the Pennsylvanian of Piesberg reveal unexpected mosaic of morphological traits and colouration pattern of the tegmina. Historical Biology [published online, https://doi.org/10.1080/08912963.2020.1867127]. Crossref

Gu, J.-J., Béthoux, O., and Ren, D. 2011. Longzhua loculata n. gen. n. sp., one of the most completely documented Pennsylvanian Archaeorthoptera (Insecta; Ningxia, China). Journal of Paleontology 85: 303–314. Crossref

Gu, J.-J., Béthoux, O., and Ren, D. 2014. A new cnemidolestodean stem-­orthopteran insect from the late Carboniferous of China. Acta Palaeontologica Polonica 59: 689–696.

Guan, Z.-Y., Prokop, J., Roques, P., Lapeyrie, J., and Nel, A. 2015. Revision of the enigmatic family Anthracoptilidae enlightens the evolution of Palaeozoic stem-dictyopterans. Acta Palaeontologica Polonica 61: 71–87. Crossref

Handlirsch, A. 1906. Die fossilen Insekten und die Phylogenie der rezenten Formen. Ein Handbuch für Paläontologen und Zoologen. 1430 pp. Wilhelm Engelman, Leipzig.

Handlirsch, A. 1911. New Paleozoic insects from the vicinity of Mazon Creek, Illinois. American Journal of Sciences 31: 297–326, 353–377. Crossref

Handlirsch, A. 1937. Neue Untersuchungen über die fossilen Insekten. Annalen des Naturhistorischen Museums in Wien 48:1–140.

Lameere, A. 1917. Révision sommaire des insectes fossiles du Stéphanien de Commentry. Bulletin du Muséum National d’Histoire Naturelle, Paris 23: 141–200.

Langiaux, J. and Parriat, H. 1974. Faune entomologique du bassin de Blanzy-Montceau. «La Physiophile», Société d’Etude des Sciences Naturelles et Historique de Montceau-les-Mines 81: 62–74.

Martins-Neto, R.G., Gallego, O.F., Brauckmann, C., and Cruz, J.L. 2007. A review of the South American Palaeozoic entomofauna. Part I: the Ischnoneuroidea and Cacurgoidea, with description of new taxa. African Invertebrates 48: 87–101.

Martynov, A.V. 1930. Palaeozoic insects from the Kuznetzk Basin [in Russian with English summary]. Izvestiâ Glavnogo Geologo-Razvedočnogo Upravleniâ Moskva 49: 1221–1248.

Meunier, F. 1909. Nouvelles recherches sur les insectes du terrain houiller de Commentry, Allier. Annales de Paléontologie 4: 125–152.

Nel, A. and Roques, P. 2021a. A new strange Archaeorthoptera from the Moscovian of Avion (France) (Insecta, Polyneoptera). Historical Bio­logy [published online https://doi.org/10.1080/08912963.2021.1978082]. Crossref

Nel, A. and Roques, P. 2021b. The second Caloneurodea from the Moscovian of Avion, France (Insecta, Archaeorthoptera). Palaeoentomology 4: 320–322. Crossref

Nel, A. and Roques, P. 2021c. The second species of the cnemidolestid genus Piesbergopterum from the upper Carboniferous of Avion, Northern France (Archaeorthoptera: Cnemidolestidae). Palaeoentomology 4: 323–325. Crossref

Nel, A., Garrouste, R., and Roques, P. 2020. The first representative of the archaeorthopteran family Eoblattidae in the Konservat-Lagerstätte of Avion (France) (Insecta: Polyneoptera). Palaeoentomology 3: 552–555. Crossref

Nel, A., Roques, P., Nel, P., Prokin, A.A., Bourgoin, T., Prokop, J., Szwedo, J., Azar, D., Desutter-Grandcolas, L., Wappler, T., Garrouste, R., Coty, D., Huang, D., Engel, M., and Kirejtshuk, A.G. 2013. The earliest known holometabolous insects. Nature 503: 257–261. Crossref

Pinto, I.D. and Ornellas, L. 1981. A new Upper Carboniferous paraplecopteran insect from Argentina. Anais do Congresso Latino-Americano de Pale­ontologia 2 (1): 107–111.

Prokop, J., Roques, P., and Nel, A. 2014. New non-holometabolous insects from Pennsylvanian of Avion locality in Pas-de-Calais, France (Insecta: “Exopterygota”). Alcheringa 38: 155–169. Crossref

Richardson, E.S. Jr. 1956. Pennsylvanian invertebrates of the Mazon Creek area, Illinois. Insects. Fieldiana, Geology 12 (1–4): 15–56. Crossref

Schubnel, T., Desutter-Grandcolas, L., Legendre, F., Prokop, J., Mazurier, A., Garrouste, R., Grandcolas, P., and Nel, A. 2020. To be or not to be: postcubital vein in insects revealed by microtomography. Systematic Entomology 45: 327–336. Crossref

Schubnel, T., Legendre, F., Roques, P., Garrouste, P., Cornette, R., Perreau, M., Perreau, N., Desutter-Grandcolas, L., and Nel, A. 2021. Sound vs. light: wing-based communication in Carboniferous insects. Communications Biology 4: 1–11. Crossref

Schubnel, T., Roberts, D., Roques, P., Garrouste, R., Desutter-Grandcolas, L., and Nel, A. 2019. Moscovian fossils shed light on the enigmatic polyneopteran families Cacurgidae and Eoblattidae (Insecta: “Eoblattida”, Archaeorthoptera). Journal of Systematic Palaeontology 18: 499–511. Crossref

Scudder, S.H. 1885. Palaeodictyoptera: or the affinities and classification of Paleozoic Hexapoda. Memoirs of the Boston Society of Natural History 3 (12): 319–351.

Scudder, S.H. 1890. The fossil insects of North America (with notes on some European species). 1. The Pretertiary insects. Report of the United States Geological Survey of the Territories 13: 1–453. Crossref

Whalley, P.E.S. 1979. New species of Protorthoptera and Protodonata (Insecta) from the upper Carboniferous of Britain, with a comment on the origin of wings. Bulletin of the British Museum of Natural History, (Geology) 32: 85–90.

Acta Palaeontol. Pol. 66 (4): 879–884, 2021

https://doi.org/10.4202/app.00940.2021