Archaeogerarus schubneli, Nel & Roques, 2021

Nel, André & Roques, Patrick, 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), pp. 879-884 : 880-883

Archaeogerarus schubneli sp. nov.

Fig. 1.

Zoobank LSID: urn:lsid:zoobank.org:act:81243C11­E3B1­4DF4­9561­C0E248E23A41

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, Pas­de­Calais, 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: 2494907A­6BFE­4FCC­97 CA­E615D573272D

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:7D23AD1B­CA20­4BB8­A926­CEE97C49ADF1

Etymology: Named after Jun­Jie 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, Pas­de­Calais, 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 Martins­Neto, Gallego, and Brauckmann, 2007 , Bouleites Lameere, 1917 , Carbonokata Aristov, 2013 , Cnemidolestes Handlirsch, 1906 , Evenkiophlebia Aristov, 2013 , Irajanarkemina Martins­Neto, 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 Martins­Neto, 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

Major clade Family, order Species

? Nugonioneuridae Avionugonioneura jouaulti Nel and Roques, 2021a

Archaeogeraridae Archaeogerarus schubneli gen. et sp. nov.

Avionxixia gui gen. et sp. nov.

Archaeorthoptera

Piesbergopterum avionensis Nel and Roques, 2021c

nec Panorthoptera

Cnemidolestidae Aviocladus pectinatus Prokop, Roques, and Nel, 2014

Aviohapaloptera bethouxi Prokop, Roques, and Nel, 2014

Tococladus sp. ( Coty, Háva, Prokop, Roques, and Nel, 2014)

Aviobiella garrousti Nel and Roques, 2021b

Caloneurodea

Aviogramma gracilis Prokop, Roques, and Nel, 2014

Titanoptera Theiatitan azari Schubnel, Roques, and Nel, 2021

Panorthoptera

Cacurgidae Cacurgus avionensis Schubnel, Roberts, Roques, Garrouste, Desutter­Grandcolas, and Nel, 2019

Eoblattidae Beloatta duquesnei Nel, Garrouste, and Roques, 2020

Family undetermined Aviologus duquesnei Coty, Háva, Prokop, Roques, and Nel, 2014

same characters ( Scudder 1890: 301–302, pl. 17: 9; Martins­Neto 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.