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A New Upper Carboniferous Stemorthopteran (Insecta) from Ningxia (China)

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A New Upper Carboniferous Stemorthopteran (Insecta) from Ningxia (China) Insect Science (2012) 19, 153–158, DOI 10.1111/j.1744-7917.2011.01468.x ORIGINAL ARTICLE A new Upper Carboniferous stem-orthopteran (Insecta) from Ningxia (China) Olivier Bethoux´ 1,2, Jun-Jie Gu1 and Dong Ren1 1College of Life Sciences, Capital Normal University, Beijing, China, 240 rue d’Aveillans, 38770 La Motte d’Aveillans, France Abstract During the Upper Carboniferous, orthopteran insects (grasshoppers, katydids, and crickets) were represented by numerous species distantly related to crown-orthopterans, such as lobeattid and cnemidolestodean insects. The panorthopterans, including total- orthopterans and their closest relatives, are represented by comparatively rarer species in localities of this period. Here we describe Heterologus duyiwuer sp. nov., an infrequent panorthopteran from the Late Carboniferous locality of Xiaheyan Village (Zhongwei City, Ningxia Hui Autonomous Region, China). The only available specimen is composed of an isolated forewing exhibiting a combination of character states previously unknown, in particular the lack of posterior radius (RP) / anterior Media (MA) connection, late branchings of the media (M) and anterior cubitus (CuA), and a branched posterior branch of the posterior cubitus (CuPb). Based on its unusual branching pattern, the composite stem resulting from the fusion of CuA and CuPaα (second anterior branch of CuP) is assumed to be composed of a branched CuA and a simple CuPaα. Key words Archaeorthoptera, homology, Namurian, stem-orthopterans, Tupo Formation Introduction initially conceptualized as including orthopterans and their closest stem-relatives (Bethoux´ & Nel, 2002a) has The systematics of stem-orthopterans received renewed been altered by Rafael et al. (2008). According to these attention in the last decade, in particular at the species authors this taxon is a synonym of Orthopterida, and en- level. A number of revisions of Upper Carboniferous his- compasses not only total-orthopterans but also the or- torical material led to the identification of a series of ders Grylloblattodea and Mantophasmatodea (among oth- successive stem-representatives of the orthopteran lin- ers). This case exemplifies the excessive plasticity of eage (grasshoppers, katydids and crickets). A new tax- Linnaean-based taxon names, resulting in ambiguity that onomic and nomenclatural framework is currently de- impedes optimal communication (Bethoux,´ 2010). Under veloped under the cladotypic nomenclatural approach the cladotypic nomenclature such volatility is prevented (Bethoux,´ 2007b, 2007c, 2010; see practical introduc- by the association of taxon names together with a defini- tioninBethoux´ & Herd, 2009). Under this scheme the tion. Only those species exhibiting the defining character taxon Archaeorthoptera nom. Bethoux´ and Nel, 2002a, state of a taxon belong to it. With regard to this context, dis.-typ. Bethoux,´ 2007d includes these stem-orthopterans the presumed superiority of this nomenclatural procedure together with crown-orthopterans (Fig. 1). Notice that un- will continue to be tested on the systematics of stem- der the Linnaean approach, the ‘Archaeorthoptera’ taxon, orthopterans. A few species might be sister-representatives of all other Archaeorthoptera (Bethoux,´ 2003, 2006). Two nu- Correspondence: D. Ren, College of Life Sciences, Cap- merically important groups of Upper Carboniferous basal ital Normal University, Beijing 100048, China. email: Archaeorthoptera have been outlined and reported from [email protected] many deposits, namely lobeattid insects (Bethoux,´ 2008a; C 2012 The Authors 153 Journal compilation C Institute of Zoology, Chinese Academy of Sciences 154 O. Bethoux´ et al. daeans are characterized by a highly variable wing vena- tion (Bethoux´ & Nel, 2003). A few other relevant species are known from Mazon Creek, IL, USA (Carpenter, 1944) and the Piesberg locality, Germany (Brauckmann & Herd, 2006), but they are represented by a limited sample, and frequently by incomplete wings only. Herein we describe a new and rare species based on an isolated forewing from the Upper Carboniferous local- ity near Xiaheyan village. Its preservation is exquisite, and its combination of character states unknown to date. Material and methods The material investigated in this contribution is housed at the College of Life Science, Capital Normal University, Beijing (China). Drawings were produced with a Leica MZ7 equipped with a 1.0 planachromatic lens and a draw- ing tube. The specimen was photographed with a Canon EOS 450D coupled with a Canon 50 mm macro lens and a 12 mm elongation tube. The photograph reproduced in Figure 2 is a dry-ethanol composite (see Bethoux´ & Briggs, 2008). The specimen was prepared by O. Bethoux´ in January 2010. Primary homologies for wing venation of total- orthopterans developed by Bethoux´ and Nel (2001, 2002a) are followed. Comments regarding these conjectures Fig. 1 Current phylogenetic framework for stem-orthopterans (Gorochov,2005) and their implications (Rasnitsyn, 2007) (dumasii Brongniart, 1879 and spilopterus Handlirsch, 1911 have been addressed elsewhere (Bethoux, 2007a, 2008b). are assigned to the genera Protophasma Brongniart, 1879 and ´ Cacurgus Handlirsch, 1911 under the Linnaean procedure, The corresponding nomenclature is: ScP, posterior sub- respectively). costa; RA, anterior radius; RP, posterior radius; M, me- dia; CuA, anterior cubitus; CuP, posterior cubitus; CuPa, Bethoux´ & Nel, 2004, 2005; Bethoux´ & Poschmann, anterior branch of CuP; CuPaα, anterior branch of CuPa; 2009; Bethoux´ & Jarzembowski, 2010; Prokop & Ren, CuPaβ, posterior branch of CuPa; CuPb, posterior branch 2007, among others), and cnemidolestodeans (Bethoux,´ of CuP; AA: anterior analis. 2005, among others). These groups abound in the Upper Throughout this contribution we use a nomenclatural Carboniferous locality of Xiaheyan Village (Zhongwei procedure predominantly cladotypic. However, regarding City, Ningxia Hui Autonomous Region, China; O. species names, a binominal name is provided for the new Bethoux´ & J.-J.Gu, pers. obs.) and will be the focus of species by assigning it to a pre-existing genus, so that the future detailed descriptions. The monophyly of these two name is made available under the International Code of groups is uncertain yet. Zoological Nomenclature (ICZN). Genera of previously According to Bethoux´ and Nel (2002a) a more derived described species are indicated at the first mention of the taxon, the panorthopteran insects, includes jumping or- species. Later on Lanham’s species names are used (else- thopterans (saltatorians) and their closest fossil relatives. where referred to as uninominal names; Lanham, 1965; Panorthopterans exhibit, in forewings, a branched anterior and see Dayrat et al., 2004). Under the cladotypic nomen- branch of the posterior cubitus (basal to its fusion with the clature taxa are referred to by names written in italics anterior cubitus), a character state pointed out as a putative with a capital letter. Linnaean taxon names are avoided autapomorphy of the group by these authors. Inference re- and ‘vernacularized’ names are preferred (e.g., orthopter- garding the panorthopteran plesiomorphic character states ans rather than Orthoptera). The choice of the nomen- combination (i.e., ground-plan) is made difficult by the clatural procedure is based on the decision of one of us lack of appropriate data. Among panorthopterans, gerari- (OB; see also Bethoux,´ 2010) and does not imply the C 2012 The Authors Journal compilation C Institute of Zoology, Chinese Academy of Sciences, Insect Science, 19, 153–158 A new stem-orthopteran 155 Fig. 2 Heterologus duyiwuer sp. nov. (holotype CNU-NX1–266, left forewing), drawing and photograph (positive imprint, composite, reversed). support of other authors (JJG, DR) to all aspects of this Description Holotype CNU-NX1–266: incomplete procedure. negative imprint and complete positive imprint of a right forewing, very well preserved; length 27.6 mm, width (op- Systematics posite the end of CuPaα) 10.3 mm; area between anterior wing margin and ScP broad, filled with ScP veinlets, with Taxon Archaeorthoptera nom. Bethoux´ and Nel, 2002a few cross-veins between them; ScP reaching RA; ScP+ dis.-typ. Bethoux,´ 2007d RA with weak distal anterior veinlets (cross-veins?); R Genus Heterologus Carpenter, 1944 simple for 8.8 mm; RP simple for 6.1 mm, with 8 branches Heterologus duyiwuer sp.nov. reaching wing apex, without clear branching pattern; RP Diagnosis ScP reaching RA; RP and M without any preserved as tube basal to its first branching, branches of connection; M weak, concave, simple for a long dis- RP preserved as depressions; M + CuA convex, branching tance; fusion CuA + CuPaα short; CuA branched dis- into M and CuA 7.7 mm distal to wing base; M (diverg- tally; CuPaα and CuPaβ simple; CuPb branched; cross- ing from M + CuA) weakly sclerotized, concave, simple veins rarely reticulated, often forming cells broader than for 8.2 mm, with 3 distal branches, branches of M im- long. printed as depressions; CuA (diverging from M + CuA) C 2012 The Authors Journal compilation C Institute of Zoology, Chinese Academy of Sciences, Insect Science, 19, 153–158 156 O. Bethoux´ et al. 0.9 mm long before its fusion with CuPaα;CuA+ CuPaα raphyletic basal saltatorians) chiefly after the branching short (1.2 mm long; see Comments for homologization of pattern of CuA + CuPaα, with a long distance between CuA+ CuPaα); distal to the divergence of CuPaα (from the first basal branch (itself homologized as CuPaα,see
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