A new, large Permian Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) found in Shanxi Province (China)

2025  47  18 geodiversitas Geodiversitas est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Geodiversitas is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 Directeur De la publication / Publication director : Gilles Bloch, Président du Muséum national d’Histoire naturelle réDacteur en chef / editor-in-chief : Didier Merle réDacteur associé / associate editor: Sylvain Charbonnier assistant De réDaction / assistant editor : Emmanuel Côtez ([email protected]) Mise en page / Page layout : Emmanuel Côtez coMité scientifique / scientific board : Christine Argot (Muséum national d’Histoire naturelle, Paris) Beatrix Azanza (Museo Nacional de Ciencias Naturales, Madrid) Raymond L. 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Geodiversitas est indexé dans / Geodiversitas is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Physical, Chemical, and Earth Sciences® – Scopus® Geodiversitas est distribué en version électronique par / Geodiversitas is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Geodiversitas sont référencés par / Articles and nomenclatural novelties published in Geodiversitas are referenced by: – ZooBank® (http://zoobank.org) 705 GEODIVERSITAS • 2025 • 47 (18) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com Zi Q. XU Nan YANG College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing (China) [email protected] [email protected] (these authors contributed equally to this work and should be considered co-first authors) Nozomu OYAMA Centre de recherche en paléontologie – Paris (CR2P), CNRS, MNHN, Sorbonne Université, Département Origines et Évolution, Muséum national d’Histoire naturelle, case postale 38, 57 rue Cuvier, F-75231 Paris cedex 05 (France) and The Kyushu University Museum, 6-10-1 Hakozaki, Higashi-ku, Fukuoka 812-8581 (Japan) [email protected] Yanqi XU Geological Survey of Jiangsu Province, 700 Zhujiang Road, Nanjing City, Jiangsu (China) [email protected] Liyang DONG Shanxi Natural History Museum, 17 North, West Binhe Road, Taiyuan city, Shanxi (China) [email protected] Dong REN College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, China [email protected] (corresponding author) Olivier BÉTHOUX Centre de recherche en paléontologie – Paris (CR2P), CNRS, MNHN, Sorbonne Université, Département Origines et Évolution, Muséum national d’Histoire naturelle, case postale 38, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] Submitted on 9 September 2024 | accepted on 14 November 2024 | published on 22 October 2025 A new, large Permian Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) found in Shanxi Province (China) urn:lsid:zoobank.org:pub:B0887E1E-FDED-4E6D-8317-9E40C5544022 Xu Z. Q., Yang N., Oyama N., Xu Y., Dong L., Ren D. & Béthoux O. 2025. — A new, large Permian Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) found in Shanxi Province (China). Geodiversitas 47 (18): 705-712. https://doi.org/10.5252/geodiversitas2025v47a18. http://geodiversitas.com/47/18 ABSTRACT The insect family Parelmoidae Rohdendorf, 1962 belongs to the extinct order Diaphanopterodea Handlirsch, 1919, itself belonging to the larger taxon Megasecopteromorpha Béthoux in Yang, 706 GEODIVERSITAS • 2025 • 47 (18) Xu Z. Q. et al. City, Shanxi Province, North China (Fig. 1B). This locality belongs to the Shanxi Fm, composed of deposits of continental-oceanic interaction facies, with the main lithologies being sandstone, mudstone and thin coal seams, including coal seams No. 3 and 4-1 as identified by Kong et al. (1996) for the Shuocheng District section (coal seam No. 3, see p.79, within a set of layers numbered ‘51’; coal seam No. 4-1, see p.80, layer numbered ‘48’), bordered by the Taiyuan Formation and the Xiashihezi Formation (Fig. 1C; and see Yang et al. 2024), all part of the North China Block (NCB). More specifically, the new locality is part of the lower Shanxi Fm. The fossiliferous layer is a grayish, fine-grained siltstone layer (about 0.7 m) situated between a greyish-white layer of coarse-grained sandstone and the coal beds 4. Besides the recovered insect material, it contains few plant fragments (Kong et al. 1996). The North China Block (NCB) is a series of deposits of continental – oceanic interaction facies, which were not simultaneous deposited throughout the area, and therefore the age of the Shanxi Formation on the NCB varies among different studies. Currently, the most accurate age range for the Shanxi Fm in the northern part of the NCB is 298.18 ± 0.32 to 295.346 ± 0.080 Ma, obtained for the Baode County using U-Pb data (Wu et al. 2021; Shen et al. 2022). This point provides a reliable inference on the age of the Zhiyu locality, because it is located in its vicinity. Moreover, the fossil locality is located along a line of sections of very similar age (points 1 to 8 in Shen et al. 2022). We can therefore regard the Zhiyu locality as early to middle Asselian. INTRODUCTION The insect order Diaphanopterodea Handlirsch, 1906, known from Pennsylvanian and Permian strata, is particular among the extinct Rostropalaeoptera Kukalová-Peck in Wootton& Kukalová-Peck, 2000 (also referred to as Palaeodictyopterida Grimaldi& Engel, 2005, as in Prokop & Engel [2019]; and see Yang et al. 2020) for its ability to hold wings backward, along the abdomen, at rest (Carpenter 1992; Rohdendorf 1962), a trait regarded as a convergence with neopteran insects. The family Parelmoidae Rohdendorf, 1962, the most diverse diaphanopterodean family, has been recovered from many renowned Permian localities, in particular from the Cisuralian, such as Obora, Elmo & Midco, and Chekarda, but also from the new, Shuiquan Gully locality, Shanxi Formation (Shanxi Province, China; Yang etal. 2024). Here, we describe a new isolated wing discovered from a new locality also belonging to the Shanxi Formation, near the Zhiyu Village. The new specimen, belonging to a new genus and species within Parelmoidae, is also the largest representative of this family. MATERIAL AND METHODS GeoloGical settinG The new material (specimen CNU-DIA-SS-2024001) was collected from a new locality, namely ‘Zhiyu locality’ (Fig. 1), located near the Zhiyu Village, Shuocheng District, Shuozhou MOTS CLÉS Permien, Cisuralien, Palaeoptera, Chine, Insecte fossile, nervation alaire, translocation de nervure, genre nouveau, espèce nouvelle. KEY WORDS Permian, Cisuralian, Palaeoptera, China, fossil insect, wing venation, vein translocation, new genus, new species. Ren& Béthoux, 2020 . Although generally rare, representatives of this family are a typical component of Permian environments, particularly in the Cisuralian, with occurrence in renowned localities such as Obora, Elmo & Midco, and Chekarda. Here we describe a new Parelmoidae, Grandelmoa jingqii n. gen., n. sp., known from a well-preserved, isolated wing discovered from a new locality, named ‘Zhiyu’ (Shanxi Formation; Shanxi Province, China). The peculiar wing venation pattern of this new taxon is best explained by a translocation (as it is, of MP branches onto CuA), a phenomenon yet rarely documented among palaeopteran taxa. The new species occurs to be, by a small amount, the largest Permian Diaphanopterodea known to date. RÉSUMÉ Un nouveau Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) du Permien, de grande taille, découvert dans la province du Shanxi (Chine). La famille d’insectes Parelmoidae Rohdendorf, 1962 appartient à l’ordre éteint des Diaphanopterodea Handlirsch, 1919 , lui-même appartenant au plus grand taxon Megasecopteromorpha Béthoux in Yang, Ren& Béthoux, 2020. Bien que généralement rares, les représentants de cette famille sont une composante typique des environnements permiens, particulièrement durant le Cisuralien, avec une occurrence dans des localités célèbres comme Obora, Elmo & Midco, et Chekarda. Nous décrivons ici un nouveau Parelmoidae, Grandelmoa jingqii n. gen., n. sp., connu par une aile isolée, bien conservée, découverte dans une nouvelle localité dénommée ‘Zhiyu’ (Formation Shanxi ; Province du Shanxi, Chine). La meilleure explication du patron de nervation alaire particulier du nouveau taxon implique une translocation (dans ce cas, d’une branche de MP sur CuA), un phénomène encore rarement documenté parmi les taxons de paléoptères. La nouvelle espèce est, de peu, le plus grand Diaphanopterodea permien connu à ce jour. 707 A new Permian Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) from China GEODIVERSITAS • 2025 • 47 (18) fig. 1 . — Geographic and stratigraphic information on the Zhiyu locality (red triangle): A, B, location of the collecting site; A, palaeogeographic map of Cisuralian; B, map of Shaanxi Province, China; C, chronological and stratigraphic framework of Pennsylvanian and Cisuralian strata according to Kong et al. (1996; including coal seams numbering), Shen et al. (2020), Shen et al. (2022) and Sun et al. (2022). Scale bar: 100 km (B). Palaeogeographic map of Cisuralian, redrawn by A. Lethiers (CR2P, Paris), from reconstruction by R. Blakey. All Chinese maps data source, Tianditu (http://www.tianditu.gov.cn, 2024/05/08). Silhouettes in C are from PhyloPic, unaltered, courtesy of Jagged Fang, Ben Moon, Felix Vaux, Katie S. Collins, T. Michael Keesey, and Ferran Sayol (CC BY 3.0). N 109°E 110° 111° 112° 113° 114° 115° 116° 35°N 36° 37° 38° 39° 40° Datong Shuozhou Xinzhou Taiyuan Yangquan Lvliang Jinzhong Yuncheng Jincheng Changzhi Linfen Hebei Province Henan Province Shaanxi Province Inner Mongolia Autonomous Region South China North China Paleotethys Siberia Laurasia Gondwana 10 160 20 30 40 50 60 70 80 90 100 110 120 130 140 150 0 (m) Ordovician Carboniferous Permian Majiagou Fm Benxi Fm Taiyuan Fm Shanxi Fm Xiashihezi Fm System Series Formation Lithologic profile Pennsylvanian Cisuralian Siltstone Trilobites Sandstone Fusulinids Mudstone Plants Limestone Gastropods Coal bed Bivalves Conglomerate Brachiopods Marl Insects Al Fe Fe Fe Fe Fe Fe Fe No. 11 No. 9 No. 5 No. 4-1 No. 3 A B C 708 GEODIVERSITAS • 2025 • 47 (18) Xu Z. Q. et al. Documentation of fossil material The new specimen was collected from a locality near Zhiyu village, Shuozhou City, Shanxi Province, China (Kong et al. 1996; Shen et al. 2020; Shen et al. 2022). It is housed at the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University (CNU), Beijing, China, and bears the specimen number CNU-DIA-SS-2024001. The specimen was photographed using a Canon EOS 5D Mark IV digital camera (Canon, Tokyo, Japan) coupled with a Canon MP-E 65 mm macro lens, under both dry and ethanol conditions. Original photographs were optimized using Adobe Photoshop CS6. In addition, Reflectance Transformation Imaging (RTI) files of the complete wing and of the wing base were generated for each side of the specimen, and are accessible from an online Dryad repository (Xu et al. 2025). These files were generated using a Fly Dome automated light dome, ~30 cm diameter and equipped with 54 LEDs arranged in three rows (Fly Dome, Paris, France), driving a Canon EOS 5DS digital camera, itself coupled to a Canon MP-E 65 mm macro lens. Original photographs were then optimized using Adobe Photoshop CS6 prior to RTI processing, itself achieved using the RTI builder software (Cultural Heritage Imaging). In details, the entire set of photographs was used to compute RTI files of complete wing, but only 42 photographs were used to compute RTI files of the wing base, for the Canon MP-E 65 mm macro lens, once deployed, prevented light from LEDs of the upper ring from reaching the specimen surface. The photograph reproduced on Fig. 2 was generated by combining a RTI snapshot under a setting with a low-angle of incidence (of the side ‘b’ of the specimen -its negative imprint) and photographs taken under dry conditions under zenithal light (both sides) (‘RTI-dry-dry’ composite). terminoloGy We follow standard wing venation terminology and abbreviations, as follows: AA Analis anterior; AP Analis posterior; Cu Cubitus; CuA Cubitus anterior; cua-cup first cross-vein in the area between CuA and CuP; CuP Cubitus posterior; M Media; MA Media anterior; MP Media posterior; R Radius; RA Radius anterior; RP Radius posterior; rp-ma first cross-vein to connect RP and MA; ScP Subcosta posterior. SYSTEMATIC PALAEONTOLOGY Class INSECTA Linnaeus, 1758 ROSTROPALAEOPTERA Kukalová-Peck in Wootton & Kukalová-Peck, 2000 MEGASECOPTEROMORPHA Béthoux in Yang et al., 2020 Order DIAPHANOPTERODEA Handlirsch, 1906 Family ParelmoiDae Rohdendorf, 1962 tyPe Genus. — Parelmoa Carpenter, 1947. incluDeD Genera. — Diapha Kukalová-Peck, 1974; Elmodiapha Kukalová-Peck, 1974; Paradiapha Kukalová-Peck, 1974; Permelmoa Prokop & Nel, 2011; Permodiapha Kukalová-Peck, 1974; Permuralia Sinichenkova & Kukalová-Peck, 1997; Protodiapha Kukalová-Peck, 1974; Pseudelmoa Carpenter, 1947; Stenodiapha Kukalová-Peck, 1974; Sinoelmoa Yang, Cui, Xu& Béthoux, in Yang et al. (2024); and Grandelmoa n. gen. Genus Grandelmoa Xu, Yang, Oyama, Ren & Béthoux, n. gen. urn:lsid:zoobank.org:act:64A26BFF-A953-485D-A727-F03038D113D4 tyPe sPecies. — Grandelmoa jingqii Xu, Yang, Oyama, Ren & Béthoux, n.sp. DiaGnosis. — By monotypy, same as for the type species. Gender feminine. etymoloGy. — The genus name derives from the Latin ‘grandis’, referring to the large size of the type species, as the largest individual in the family Parelmoidae, and the genus Elmoa Tillyard, 1937. Grandelmoa jingqii Xu, Yang, Oyama, Ren & Béthoux, n. gen., n.sp. (Fig. 2) urn:lsid:zoobank.org:act:9405F20C-D2EC-4B4C-8777-27E49B8E5E92 tyPe material. — Holotype (by monotypy): China • North China, Shanxi Province, Shuozhou City, Shuocheng District, Zhiyu Village, Zhiyu locality; 39°24’11”N, 112°21’46”E; 26.III.2023; Jingqi Zhang leg.; CNU-DIA-SS-2024001. DiaGnosis. — ScP long, ending beyond the second fork of RP; MA/ MP split distal of the RA/RP split, itself located in a very basal position; cua-cup cross-vein located distally (distal to the RA/RP split) and basal to MA/MP split; CuP forked; anal area well-developed, with 10 terminal branches; two gradate rows of cross-veins, subparallel to the posterior wing margin. etymoloGy. — Named after Jingqi Zhang, the collector of type specimen. The specific epithet is to be treated as a noun in genitive case. tyPe locality anD stratiGraPhy. — Zhiyu locality, between coal seam layers no. 3 and 4-1 (following numbering by Kong et al. 1996); Shanxi Formation; Permian, Cisuralian, Asselian (Kong et al. 1996; Shen et al. 2020; Shen et al. 2022). measurements. — Wing length 22.7 mm, width 7.6 mm. DescriPtion Positive and negative imprints of a complete right wing, possibly a forewing; veins elevation sharply contrasted in the basal half, and attenuated more distally; near wing base, ScP long, ending on RA beyond the second RP fork; stem of R+M convex, with a conspicuous inflexion opposite the point of 709 A new Permian Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) from China GEODIVERSITAS • 2025 • 47 (18) separation of R and M (located about 4.0 mm distal from wing base); RA convex, simple and strong, almost parallel to anterior wing margin; RP with 6 terminal branches, with the two first posterior ones simple; MA/MP split located behind the RA/RP split; MA diverging somewhat obliquely (i.e. M and MP are aligned); rp-ma cross-vein long and oblique, located just distal of the MA/MP split; MA simple; MP forked distally, with 4 branches, 2 of them successively arising from CuA (under the ‘translocated MP branch’ interpretation; see Fig. 2C and Remarks section); short Cu stem, distinct from (R+)M, distinguishable; CuA diverging obliquely from Cu, then running close and parallel to R+M stem for some AP(?) ScP RP MA CuP RA MP CuA AA A-C AP(?) ScP RP MA CuP RA MP CuA AA A B B C fig. 2. — Grandelmoa jingqii n. gen., n. sp., holotype, specimen CNU-DIA-SS-2024001: A, drawing under the ‘branched CuA’ interpretation; B, photograph (‘RTIdry-dry’ composite); C, drawing under the ‘translocated MP branch’ interpretation (large black arrow indicating translocation). Scale bar: 5 mm. 710 GEODIVERSITAS • 2025 • 47 (18) Xu Z. Q. et al. distance, suddenly diverging posteriorly, very close to the R/M split; CuA simple (under the ‘translocated MP branch’ interpretation; see Fig. 2C and Remarks section); CuP forked distally (distal to first point of emergence of MP branches translocated onto CuA); CuA–CuP area narrow until the first cross-vein occurring in this area, which is short and strong, located slightly distal to RA/RP split and basal to MA/MP split; anal area very well-developed, with a total of ten terminal branches (AA with 3 terminal branches; AP(?) with 7 terminal branches); 12 oblique, sigmoidal veinlets evenly dispersed in the area between anterior margin and ScP/RA; two gradate rows of cross-veins paralleled with each other. r emarks A seemingly branched CuA is present in the new specimen (‘branched CuA’ interpretation; see Fig. 2A). However, a branched CuA is extremely rare in Megasecopteromorpha. Based on our literature search, the only known cases are two isolated wings belonging to the Parelmoidae species Stenodiapha moravica Kukalová-Peck, 1974 (see original description). However, a translocation of a branch of MP onto CuA, coupled with a pectinate fusion, is an alternative, plausible interpretation. Vein translocation has been documented in many different taxa of Neoptera, and in particular in orthopterans (Béthoux 2007, 2012) and gryllo blattodeans (Cui etal. 2015). Translocation can also lead to a ‘pectinate fusion’, and one of the best examples is the organisation of the radial system in dictyopterans (Guo et al. 2013). Such transformation is also known in Hemerobiidae Leach, 1815 (neuropterans; among others, see Carpenter [1940], Aspöck et al. [1980]) and some orthopterans (Béthoux 2012). Such transformations are apparently less common among palaeopteran taxa, but it can be considered established in Spilapteridae Brongniart, 1893 (Rostropalaeoptera; Liu et al. 2015; Sinitshenkova 2025). Bearing this in mind, a translocation of a posterior branch of MP, with a partial fusion of its posterior branch with CuA, is a plausible explanation for the morphology observed in the specimen CNU-DIA-SS-2024001 (‘translocated MP branch’ interpretation; Fig. 2C). Moreover, in known Parelmoidae, MP usually has two or three branches, and rarely four. Under the ‘branched CuA’ interpretation (Fig. 2A), the specimen would then have a two-branched MP, which is incongruous, given its large size (relative to other Parelmoidae), very generally implying a relatively higher number of vein branches. Conversely, if the first two anterior branches of the seemingly branched CuA are assigned to MP (assuming translocation and pectinate fusion), the latter vein then has a total of four branches, more consistent with its size. It must be noted that veins elevation in this area of the wing is homogeneous, and therefore does not provide insights on the nature of the corresponding veins. In summary, the ‘translocated MP branch’ interpretation provides a plausible answer to the peculiar configuration observed in the specimen CNU-DIA-SS-2024001. It remains unclear whether a translocation of a MP branch is an unusual or a usual feature of the corresponding species. DISCUSSION systematic Placement According to Yang et al. (2024; and references therein), the combination of the character states ‘long fusion of Cu/CuA with R+M (or, CuA running very close to R+M for some distance)’ and ‘occurrence of a very short and strong cuacup cross-vein’ allows assigning the species to which the new specimen belongs to the families Parelmoidae or Elmoidae Tillyard, 1937, within Diaphanopterodea. Then, two main character states allow distinguishing it from the Elmoidae, namely ‘long ScP, ending on RA beyond the wing mid-length’ and ‘a very well-developed anal area’. Characters relevant to the placement of the species to which the new specimen belongs are summarized in Table 1, showing that it displays a unique combination of traits. Among them, the position of cua-cup cross-vein is worthy of attention. It is located between the RA/RP split and the MA/MP split in the new specimen, a state shared only with Sinoelmoa (Yang et al. 2024; Shuiquan Gully locality; China; Cisuralian, Asselian), and Protodiapha (KukalováPeck 1974; Obora; Czech Republic; Cisuralian, Sakmarian). However, the wing lengths of species belonging to these genera are much smaller than in the new specimen. The new specimen further differs from Sinoelmoa by several traits (Table 1), including the presence of clear veinlets in the area between ScP and the anterior wing margin, ScP remaining strong distally (as opposed to ‘vanishing’), and the presence of a rp-ma cross-vein (instead of a fusion of RP and MA). As for Protodiapha, the new specimen also differs from this genus by well distant RP and MA, and the lack of dark spots on the wing membrane, among other traits (see Table 1). Another relevant trait is the development of the anal area, which is large in many genera of Parelmoidae. However, this area is even more extensively developed in the new specimen. In this trait, it compares, to some extent, with Pseudelmoa (Elmo & Midco; USA; Cisuralian, Artinskian). Nevertheless, the larger size implies that the new material belongs to another taxon. Another prominent trait is the position of the RA/RP split. It is located basally in the genera Parelmoa and Pseudelmoa, but it is even more basal in the new specimen. Finally, as mentioned above, it is not unlikely that a translocation of a branch of MP onto CuA is a usual feature of the species to which belongs the new specimen, in which case it would compose a very unique trait. In summary, it is legitimate to erect a new genus and species for the new material. size asPect A general trend of size reduction has been documented for the Megasecoptera (Nel et al. 2023) and likely affected the entire Rostropalaeoptera, including Diaphanopterodea. It is therefore significant to unravel a large-sized Diaphanopterodea from Permian strata. To date, the largest known Permian member of this order is Eumartynovia raaschi Carpenter, 1947 (family Martynoviidae Tillyard, 1932; Elmo & Midco locality), with a wing length of 22.5 mm. Therefore, with a wing length 711 A new Permian Parelmoidae (Insecta, Megasecopteromorpha, Diaphanopterodea) from China GEODIVERSITAS • 2025 • 47 (18) of 22.7 mm, Grandelmoa jingqii n. gen., n. sp. is the largest known Permian Diaphanopterodea (and also the largest Parelmoidae; with Elmodiapha ovata Kukalová-Peck, 1974, from Obora, being the second largest, with a wing length of 21.4 mm). This discovery, made in a new locality, shows that our documentation of this insect group remains largely incomplete. Data archival statement A Dryad dataset including four RTI files of the new material is available at https://doi.org/10.5061/dryad.2bvq83c09 Acknowledgements We are grateful to N.Sinitshenkova and C. Jouault for their useful comments, and to the editorial board of Geodiversitas for handling peer reviewing and publication process. We are grateful to Zhongshu Liu, who contributed to the fieldwork at the Zhiyu locality; and to Jingqi Zhang, who collected and donated the new specimen. Competing interests The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding This research was supported by grants from National Natural Science Foundation of China [No. 32020103006, 42472001], and Support Project of High-level Teachers in Beijing Municipal Universities [No. BPHR20220114], and from the Japan Society for the Promotion of Science (JSPS) [KAKENHI, No. 24K17161]. Author contributions Conceptualization: DR and OB; Investigation: ZX, NY, YX, NO, LD and OB; Resources: DR; Writing – original draft preparation: NY and OB; writing – review and editing: ZX, NY, NO, DR and OB; visualization: ZX, NY and OB; supervision: DR; project administration: DR; funding acquisition: DR All authors have read and agreed to the published version of the manuscript. REFERENCES asPöck h., asPöck u. & hölzel h. (eds) 1980. — Die Neuropteren Europas: eine zusammenfassende Darstellung der Systematik, Ökologie und Chorologie der Neuropteroidea (Megaloptera, Raphidioptera, Planipennia) Europas. Geocke and Evers, Krefeld, 2volumes, 495+ 355p. B éthoux o. 2007. — Cladotypic taxonomy applied: titanopterans are orthopterans. Arthropod Systematics & Phylogeny 65: 135-156. https://doi.org/10.3897/asp.65.e31674 table 1. — Characters (and their states) relevant for the Parelmoidae systematics at the genus level and for the placement of the new taxon (framed). Taxon authorships are indicate p. 708. Wing length (mm) Coloration Position of cua-cup Well-developped veinlets in the area between ScP and the anterior margin Ending of ScP Relation of RP and MA Diapha 14-18.7 ×basal to the RA/RP split and MA/MP split reaching RA beyond wing mid-length fusion Elmodiapha 18.4-21.4 ×variable reaching RA beyond wing mid-length no fusion, long rp-ma Grandelmoa n. gen. 22.7 × between the RA/RP split and the MA/MP split  remaining strong distally, reaching anterior margin no fusion, long rp-ma Paradiapha 13.4 ×basal to the RA/RP split and MA/MP split reaching RA beyond wing mid-length fusion Parelmoa 12.3-16.1 ×basal to the RA/RP split and MA/MP split reaching RA beyond wing mid-length no fusion, long rp-ma Permelmoa 12.7 × between the RA/RP split and the MA/MP split × remaining strong distally, reaching anterior margin no fusion, long rp-ma Permodiapha 12.9-16.3 dark spots on wing membrane, variable basal to the RA/RP split and MA/MP split reaching RA beyond wing mid-length fusion Permuralia 13.5-18.0 ×basal to the MA/MP split and RA/RP split reaching RA beyond wing mid-length fusion Protodiapha 13.5 dark spots on wing membrane, variable variable reaching RA beyond wing mid-length no fusion, short rp-ma Pseudelmoa 19.6 × opposite the RA/RP split and basal to the MA/MP split  reaching RA beyond wing mid-length no fusion, long rp-ma Sinoelmoa 18.2 (estimated) dark area between RA and the anterior margin between the RA/RP split and the MA/MP split ×vanishing between RA and anterior margin RP briefly touching MA Stenodiapha 15.5-20.2 ×variable reaching RA beyond wing mid-length fusion