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New Significant Fossil Insects from the Upper Carboniferous of Ningxia in Northern China (Palaeodictyoptera, Archaeorthoptera)

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New Significant Fossil Insects from the Upper Carboniferous of Ningxia in Northern China (Palaeodictyoptera, Archaeorthoptera) Eur. J. Entomol. 104: 267–275, 2007 http://www.eje.cz/scripts/viewabstract.php?abstract=1228 ISSN 1210-5759 New significant fossil insects from the Upper Carboniferous of Ningxia in northern China (Palaeodictyoptera, Archaeorthoptera) JAKUB PROKOP1 and DONG REN2 1Charles University in Prague, Faculty of Science, Department of Zoology, Viniþná 7, CZ-128 44, Praha 2, Czech Republic; e-mail: [email protected] 2College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100037, P.R. China; e-mail: rendong@ mail.cnu.edu.cn Key words. Palaeodictyopteroida, Eugereonoidea, Breyerioidea, Archaeorthoptera, “Protorthoptera”, fam. n., gen. n., sp. n., taxonomy, Late Paleozoic, Upper Carboniferous, Namurian, Ningxia, China Abstract. New palaeodictyopterid Namuroningxia elegans sp. n. (Namuroningxiidae fam. n.) and archaeorthopterid Sinopteron huangheense gen. n., sp. n. (uncertain family assignment) are described from the Tupo Formation (Upper Carboniferous, Namurian, northern China, Ningxia Hui Autonomous Region). Both new taxa exhibit a unique structure of their wing venation pattern. The Namuroningxiidae fam. n. bears a mosaic of characters diagnostic of several “palaeopteran” families throughout Palaeodictyopter- oida and Eupalaeoptera. This fact is highly interesting in reference to the Namurian age known for the occurrence of the first winged insects as well as the palaeogeographical position of the locality far from all sites in Laurussia. The preliminary study of entomo- fauna diversity exhibit frequent occurrence of archaeorthopterid wing fragments (e.g., Sinopteron huangheense) forming up to 40 percent of all insect specimens demonstrate high abundance of this group in the early Late Carboniferous ecosystems. INTRODUCTION genus of dragonfly Shenzhousia Zhang & Hong, 2006 The Namurian is the earliest stage of the Upper Car- (Protodonata: Meganeuridae). boniferous with sudden occurrence of diverse winged Fossils known as “Qilianshan biota” including plants, insects (Pterygota) (e.g., Brauckmann et al., 2003). As a insects, other invertebrates and vertebrates such as fish consequence of the scarce insect fossil record and the are preserved in layers of Tupo Formation in outcrops paucity of rather fragmentary specimens in the Late along Qilianshan Mountains located in the Ningxia, Devonian and Early Carboniferous we cannot estimate Gansu and Inner Mongolia regions (China). Fossil insects the time of origin of the pterygote insect groups (Prokop have been collected in layers of the Tupo Formation in et al., 2005; Engel & Grimaldi, 2004). The Namurian Zhongwei County of southern Ningxia Hui Autonomous insect deposits are currently known from few localities all Region, representing the oldest entomofauna known from located in the Euro-American region, such as the Upper China so far (Hong, 1998). The paralic development of Silesian Coal Basin in the Czech Republic and Poland, Tupo Formation is formed by alternated beds of terrestrial Hagen-Vorhalle, Ruhr area in Germany, Charleroi Coal and marine facies dated to stage C2t (Bashkirian – Mosco- Basin in Belgium, South Limburg coal-field in the Neth- vian) as equivalent of European Namurian B–C (Peng et erlands and Manning Canyon Shale in Utah (USA) al., 2005). The palaeoecological analysis based on fish (Brauckmann et al., 1996; Hennig, 1981; Rasnitsyn & fossils indicates near shore palaeoenvironment with pos- Quicke, 2002). Outside of this area a new insect fauna sible role of secondary transportation (Lu et al., 2002). known as “Qilianshan entomofauna” has been recently Taxa described herein are attributed to the Palaeodicty- discovered from the Namurian C of Qilianshan Moun- opteroida (Namuroningxiidae fam. n., Namuroningxia tains in northern China (Hong, 1998). This locality is the elegans gen. n., sp. n.) and Archaeorthoptera (Sinopteron first Namurian site providing fossil insects from the Far huangheense gen. n., sp. n.), and both could be consid- East. Its distant palaeogeographical situation from the ered as suitable candidates to basal groups within Ptery- other Carboniferous insect localities and occurrence in gota. low latitudes is rather significant for our understanding of MATERIAL AND METHODS insect evolution (Rasnitsyn & Quicke, 2002) (see Fig. 1). The fossil specimens were observed under a stereomicroscope In the first insect systematic studies based on these depos- Olympus SZX-12 in dry state and under ethyl alcohol. The its, Peng et al. (2005) and Zhang et al. (2006) presented venation patterns were drawn directly using a stereomicroscope descriptions of a new family Sinonamuropteridae Peng, with a camera lucida and finally readjusted to the photograph Hong & Zhang, 2005 (all taxa erroneously attributed by scales using image-editing software (Adobe Photoshop). Photo- the authors to Diaphanopterodea, bearing character of graphs were made simultaneously using a digital camera branched CuA typical for Grylloblattida sensu Storoz- Olympus 5050 by single sided cross-light exposure. henko, 2002 after Béthoux, pers. comm.) and a new 267 Genus Namuroningxia gen. n. Type species. Namuroningxia elegans sp. n. by present descrip- tion. Diagnosis. As for the family. Etymology. Composite name after Namurian (stratigraphical stage) and Ningxia (Ningxia Hui Autonomous Region, name in northern China); feminine in gender. Namuroningxia elegans sp. n. (Figs 2–3) Diagnosis. By monotypy, that of the genus. Fig. 1. Palaeogeographical world map of the Upper Carbonif- Description. Holotype specimen CNU-P-NX2006001. erous with approximate position of the locality in Zhongwei Wing large with the broadest part at about mid wing, County, Ningxia Hui Autonomous Region (China) indicated by membrane probably originally hyaline with pattern of black triangle, after Scotese (2005, modified). dark coloration; wing generally covered with coarse irregular network of cross-veins and rather dense small Type material referred to as “CNU” is housed at the Key Lab cells in distal part of radial branches; zigzagged inter- of Insect Evolution & Environmental Changes, the College of calary veins present; posterior wing margin distinctly Life Science, Capital Normal University, Beijing, China (CNU; bearing rather long and dense setation; vein axillary scler- Ren Dong, Curator). We follow the wing venation nomenclature of Kukalová-Peck ites solid and well recognizable; short CP reaching ante- (1991) and Béthoux & Nel (2002a) as elaborated for Archaeor- rior margin close to wing base about 4.4 mm; prominent thoptera. Systematics and divisions follow the concept of short convex ScA reaching anterior margin about twice Palaeodictyopteroida sensu Bechly (1996, 2005) and Archaeor- the distance as CP; concave ScP nearly straight, reaching thoptera sensu Béthoux & Nel (2002a, b). anterior wing margin very distally, about 6 mm from wing apex; two rows of cells in basal part and numerous SYSTEMATICS simple cross-veins in area between C and ScP; RA and Superorder Palaeodictyopteroida Bechly, 1996 RP basally fused into R+ veinal stem, separating from Superfamily Eugereonoidea Sinitshenkova, 2002 each, at about 1/3 wing length, 21.7 mm from wing base; simple RA orientated towards ScP probably reaching Family Namuroningxiidae fam. n. anterior wing margin near apex; RP nearly straight Type genus. Namuroningxia gen. n. divided into five visible branches, all reaching apical area Diagnosis. Based on fore wing venation: wing elon- of wing; first fork of RP about 41 mm from wing base; gated with the broadest part at about mid wing, wing gen- RP area is covered with dense irregular network of small erally covered with coarse irregular network of cells in distal part; stem of M adjacent to radial stem in cross-veins and rather dense small cells (= archedictyon) basal part, M divides into MA and MP about the same in distal part; zigzagged intercalary veins present; poste- level as division RA and RP; MA simple, orientated rior wing margin with long and dense setation; CP towards RP in basal part, ending on posterior wing mar- reaching anterior margin close to wing base; ScA gin; MA and RP connected by short cross-vein, about 5 reaching anterior margin about twice distance as CP; ScP mm from division of MA and MP; MP basally divided nearly straight, reaching anterior wing margin close to dichotomously into two main branches about 28.9 mm wing apex; RA and RP basally fused into R+ veinal stem, from wing base, both secondarily forked reaching poste- bifurcating at about 1/3 wing length; simple RA basally rior wing margin together with five terminal branches; diverged to ScP reaching anterior wing margin near apex; CuA and CuP fused basally into Cu veinal stem, RP divided into five branches; stem of M adjacent to diverging from each other at about 11.3 mm from wing radial stem in basal part, M divided into MA and MP base; CuA simple, orientated towards M in basal part, about the same level as division RA and RP; MA simple; ending on posterior wing margin 44.5 mm from wing MA and RP connected by a short cross-vein (brace); MP base; short prominent oblique cross-vein (brace) between basally divided dichotomously into two main branches, CuA and M stem about 6.5 mm from division of CuA and both secondarily forked reaching posterior wing margin CuP; concave CuP bifurcated basally into two main together with five terminal branches; CuA simple, branches near division CuA and CuP; long anterior strongly diverging from M in basal part; short oblique branch of CuP secondarily divided dichotomously
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