Introduction

More than 15 y ago, earthworks east of Niedermoschel, a village located about 45 km southwest of Mainz (Rhineland-Palatinate, Germany), prompted collecting at a site which has been known to yield fossils since the beginning of the last century (Reis 1913). The exposed sediments belong to the Niedermoschel bed (Meisenheim Formation, ‘Lower Rotliegend’; Schindler 1997) and are thus latest Late Carboniferous to earliest Early Permian (Gzhelian/Asselian boundary) in age (Königer et al. 2002). They represent the profundal fades of a large lake and yield an aquatic fauna consisting of vertebrates and crustaceans, such as ostracodes, conchostracans, and syncarids. The abundance of terrestrial plants and insects has been attributed to a strong deltaic input (Poschmann & Schindler 1997).

The Niedermoschel insects are preserved with remains of organic material still present, but this has probably been altered by contact metamorphism (Montenari & Uhl 2005). The insects are usually preserved as paired or isolated wings, or fragments thereof. Completely preserved large wings (> 2 cm) are rare. Animals with body and wings preserved are rare exceptions, and the number of such fossils decreases with increasing size (Hörnschemeyer 1998). Hörnschemeyer (1998) briefly outlined the taxonomic composition of the insect fauna, which is dominated by lemmatophoridaeans, miomopterans, and blattinopsidaeans. The latter group has been revised by Hörnschemeyer & Stapf (2001).

Herein we provide the description of a large-sized archaeorthopteran from this Lagerstätte. Archaeorthoptera is a taxon including orthopterans, stem orthopterans, and a number of extinct groups, such as caloneurodeans and cnemidolestodeans (Béthoux 2007d: Appendix 2). Such insects form only a minor part of the Niedermoschel assemblage.

Methods

The fossil was photographed under alcohol immersion in both normal and polarized light using a digital camera, and a unique composite photograph was produced. Drawings were made with both the help of enlarged photographs and a SteREO Discovery V8 stereomicroscope equipped with a pair of W-PL 10x/23 eye pieces, a Plan Apo S 1.0x FWD objective, and a camera lucida. Figures were processed and arranged with image-editing software (Adobe Photoshop).

We use the cladotypic nomenclatural procedure elaborated by Béthoux (2007b, 2007c, in press) for taxa other than species, and follow suggestions of Dayrat et al. (2004; and references therein) for species names [see introduction and application of this approach in JOR by Béthoux & Herd (2009)]. All formal species and taxon names are written in italics. Capitals indicate a taxon name. In other words, a capitalized name written in italics does not refer to a genus but to a taxon at any level of the phylogeny. The choice of the taxonomic procedure is based on the decision of one of us (OB) and does not imply the support of the other author (MP) for all aspects of this procedure. An appendix compliant with the ICZN is provided, so as to allow the mention of the new species under a Linnaean framework. Although the paper does not consistently apply rules of the ICZN, therefore jeopardizing the availability of newly erected names, the need for experimentation with alternative nomenclatural approaches (Béthoux 2009) requires some flexibility.

It must be emphasized that uninominal species names are applicable under a nomenclatural approach using ranks and onomatophore-based taxon conceptualization (i.e., as under the Linnaean approach). In addition, regarding taxon names used in this paper, Archaeorthoptera/Archaeorthopera refers to a taxon which is above the family-group names under a Linnaean hierarchy, and therefore is not regulated by the ICZN. Owing to its definition and delimitation, the taxon name Miamia is valid both as a cladotypic taxon name and as an ICZN-compliant generic name (Béthoux 2008a). In summary, concerns regarding the recourse to an alternative nomenclatural approach are, in this case, not fully justified. On the other hand, the comments subsection benefits from the avoidance of numerous monotypic generic names, which are by definition synonyms of specific epithets, therefore supernumerary (Béthoux in press). In addition, assignment of the new species to the taxon Archaeorthoptera is objectively ascertained, owing to the occurrence of the defining character state of this taxon in the new species (Béthoux in press).

Fig. 1.

Species niedermoschelensis sp. nov., holotype (PE2008/5007-LS), drawing (remains of the left hind wing omitted; with tentative interpretation) and photograph (negative imprint, composite); see text for wing venation abbreviations; other abbreviations: le, lateral eye; md, mandible; msl, mesothoracic leg; mst, mesothorax; mtt, metathorax; pl, prothoracic leg; pt, prothorax; lfw: left forewing; rfw: right forewing: rhw: right hind wing; ^* indicates CuPb. See Plates.

We use the wing venation nomenclature elaborated by Béthoux & Nel (2002) for Archaeorthoptera, itself based on that of orthopterans (Béthoux & Nel 2001) and on the serial insect wing venation pattern (Lameere 1922, 1923). Corresponding abbreviations are repeated herein for convenience: ScP, posterior subcosta; R, radius; RA, anterior radius; RP, posterior radius; M, media; MA, anterior media; MP, posterior media; CuA, anterior cubitus; CuP, posterior cubitus; CuPa, anterior branch of CuP; CuPb, posterior branch of CuP; AA, anterior analis; AA1, first anterior Analis; AA2, second anterior Analis. A vein (or vein sector) is said to be convex if located on an elevation, and concave if located in a depression. Criticisms expressed by Gorokhov (2005) regarding this homologization hypothesis are addressed in Béthoux (2007a). Subsequent comments by Rasnitsyn (2007) are addressed in Béthoux (2008b).

Systematic Palaeontology

Among lobeattids, only the taxon Miamia nom. Dana, 1864, dis.typ. Béthoux, 2008a has been defined (Béthoux 2008a). The new species does not exhibit the defining character state of this taxon, viz. a fusion of CuPa with M + CuA (i.e., prior to the divergence of CuA from M + CuA). The monophyly of the lobeattid group is not demonstrated (Béthoux 2005, 2008a), and the group is not defined. Therefore, the only possible taxonomic assignment of niedermoschelensis sp. nov. is to Archaeorthoptera.

The species niedermoschelensis sp. nov. differs from other lobeattids such as species assigned to Miamia, and the species elongata Brongniart, 1893: 433, silvatica Laurentiaux & Laurentiaux-Vieira, 1980, and schneideri Béthoux, 2005 by its comparatively long CuA in the forewings [see Béthoux (2005, 2008a), Béthoux & Nel (2004)]. Its comparatively small forewing size allows niedermoschelensis sp. nov. to be distinguished from cubitalis Handlirsch, 1911 and mazonus Béthoux, 2005 (Béthoux 2005). In addition the loose forewing crossvenation distinguishes it from elongata, cubitalis, and mazonus. Regarding huangheense Prokop & Ren, 2007 and liuae Liu et al., 2009 (which are most likely synonyms), niedermoschelensis sp. nov. lacks a connection of the anterior branch of MA with RP (or of MA for liuae), and well-individualized branches of ScP, diagnostic of the former species [in Prokop & Ren (2007), according to the text and their fig. 4, the anterior branch of MA is fully fused with RP in huangheense, contra their fig. 5]. Although the available material is incomplete, the erection of a new species is justified.

Discussion

Lobeattid insects exhibit a homogenous morphology from which no Permian groups seem to derive. They appear as a constant component of Late Carboniferous insect faunas and have been reported from the Tupo Formation (Namurian B-C, China, Liu et al. 2009, Prokop & Ren 2007), Mazon Creek (Westphalian D, Béthoux 2005, Béthoux 2008a), the North Coal basin of Pas-de-Calais (Westphalian C, France, Béthoux 2008a), and Commentry (Stephanian B-C, France, Béthoux & Nel 2004). The occurrence of the species niedermoschelensis sp. nov. near the Gzhelian/Asselian boundary constitutes the latest record of the group (assuming that it is monophyletic).

Paradoxically, niedermoschelensis sp. nov. exhibits a long free part of CuA, which is most probably a plesiomorphy within Archaeorthoptera: most lobeattid species have a short CuA (e.g., schneideri, elongata; it is nonexistent in Miamia; see references above), while huangheense, the earliest lobeattid, and putative outgroups of lobeattids [e.g., dumasii Brongniart, 1879, macroptera Van Beneden & Coemans, 1867, palmiformis Bolton, 1922; see Béthoux 2003; Béthoux & Nel 2004, 2005] exhibit a long one. The species niedermoschelensis sp. nov. may therefore occupy a very basal position with respect to other lobeattids, together with huangheense.

Conclusion

The importance of the various character-state combinations exhibited by lobeattid insects will have to be evaluated based on a numerical cladistic analysis involving Palaeozoic Archaeorthoptera as a whole, taking into account recent revisions and newly described material. The character-state combinations of huangheense and niedermoschelensis sp. nov. might be important for supporting the monophyly of lobeattids, or of a subgroup within lobeattids.