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;