a new upper carboniferous stem-orthopteran (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 Bethoux1,2, 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 inlocalities of this period. Here we describe Heterologus duyiwuer sp. nov., an infrequentpanorthopteran from the Late Carboniferous locality of Xiaheyan Village (Zhongwei City,Ningxia Hui Autonomous Region, China). The only available specimen is composed ofan isolated forewing exhibiting a combination of character states previously unknown,in particular the lack of posterior radius (RP) / anterior Media (MA) connection, latebranchings of the media (M) and anterior cubitus (CuA), and a branched posterior branchof the posterior cubitus (CuPb). Based on its unusual branching pattern, the compositestem resulting from the fusion of CuA and CuPaα (second anterior branch of CuP) isassumed to be composed of a branched CuA and a simple CuPaα.

Key words Archaeorthoptera, homology, Namurian, stem-orthopterans, Tupo Formation

Introduction

The systematics of stem-orthopterans received renewedattention in the last decade, in particular at the specieslevel. A number of revisions of Upper Carboniferous his-torical material led to the identification of a series ofsuccessive stem-representatives of the orthopteran lin-eage (grasshoppers, katydids and crickets). A new tax-onomic and nomenclatural framework is currently de-veloped under the cladotypic nomenclatural approach(Bethoux, 2007b, 2007c, 2010; see practical introduc-tion in Bethoux & Herd, 2009). Under this scheme thetaxon Archaeorthoptera nom. Bethoux and Nel, 2002a,dis.-typ. Bethoux, 2007d includes these stem-orthopteranstogether with crown-orthopterans (Fig. 1). Notice that un-der the Linnaean approach, the ‘Archaeorthoptera’ taxon,

Correspondence: D. Ren, College of Life Sciences, Cap-ital Normal University, Beijing 100048, China. email:[email protected]

initially conceptualized as including orthopterans andtheir closest stem-relatives (Bethoux & Nel, 2002a) hasbeen altered by Rafael et al. (2008). According to theseauthors this taxon is a synonym of Orthopterida, and en-compasses not only total-orthopterans but also the or-ders Grylloblattodea and Mantophasmatodea (among oth-ers). This case exemplifies the excessive plasticity ofLinnaean-based taxon names, resulting in ambiguity thatimpedes optimal communication (Bethoux, 2010). Underthe cladotypic nomenclature such volatility is preventedby the association of taxon names together with a defini-tion. Only those species exhibiting the defining characterstate of a taxon belong to it. With regard to this context,the presumed superiority of this nomenclatural procedurewill continue to be tested on the systematics of stem-orthopterans.

A few species might be sister-representatives of allother Archaeorthoptera (Bethoux, 2003, 2006). Two nu-merically important groups of Upper Carboniferous basalArchaeorthoptera have been outlined and reported frommany deposits, namely lobeattid insects (Bethoux, 2008a;

C© 2012 The AuthorsJournal compilation C© Institute of Zoology, Chinese Academy of Sciences

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154 O. Bethoux et al.

Fig. 1 Current phylogenetic framework for stem-orthopterans(dumasii Brongniart, 1879 and spilopterus Handlirsch, 1911are assigned to the genera Protophasma Brongniart, 1879 andCacurgus Handlirsch, 1911 under the Linnaean procedure,respectively).

Bethoux & Nel, 2004, 2005; Bethoux & Poschmann,2009; Bethoux & Jarzembowski, 2010; Prokop & Ren,2007, among others), and cnemidolestodeans (Bethoux,2005, among others). These groups abound in the UpperCarboniferous locality of Xiaheyan Village (ZhongweiCity, Ningxia Hui Autonomous Region, China; O.Bethoux & J.-J.Gu, pers. obs.) and will be the focus offuture detailed descriptions. The monophyly of these twogroups is uncertain yet.

According to Bethoux and Nel (2002a) a more derivedtaxon, the panorthopteran insects, includes jumping or-thopterans (saltatorians) and their closest fossil relatives.Panorthopterans exhibit, in forewings, a branched anteriorbranch of the posterior cubitus (basal to its fusion with theanterior cubitus), a character state pointed out as a putativeautapomorphy of the group by these authors. Inference re-garding the panorthopteran plesiomorphic character statescombination (i.e., ground-plan) is made difficult by thelack of appropriate data. Among panorthopterans, gerari-

daeans are characterized by a highly variable wing vena-tion (Bethoux & Nel, 2003). A few other relevant speciesare 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 onan isolated forewing from the Upper Carboniferous local-ity near Xiaheyan village. Its preservation is exquisite,and its combination of character states unknown todate.

Material and methods

The material investigated in this contribution is housed atthe College of Life Science, Capital Normal University,Beijing (China). Drawings were produced with a LeicaMZ7 equipped with a 1.0 planachromatic lens and a draw-ing tube. The specimen was photographed with a CanonEOS 450D coupled with a Canon 50 mm macro lens anda 12 mm elongation tube. The photograph reproducedin Figure 2 is a dry-ethanol composite (see Bethoux &Briggs, 2008). The specimen was prepared by O. Bethouxin January 2010.

Primary homologies for wing venation of total-orthopterans developed by Bethoux and Nel (2001, 2002a)are followed. Comments regarding these conjectures(Gorochov, 2005) and their implications (Rasnitsyn, 2007)have been addressed elsewhere (Bethoux, 2007a, 2008b).The corresponding nomenclature is: ScP, posterior sub-costa; RA, anterior radius; RP, posterior radius; M, me-dia; CuA, anterior cubitus; CuP, posterior cubitus; CuPa,anterior branch of CuP; CuPaα, anterior branch of CuPa;CuPaβ, posterior branch of CuPa; CuPb, posterior branchof CuP; AA: anterior analis.

Throughout this contribution we use a nomenclaturalprocedure predominantly cladotypic. However, regardingspecies names, a binominal name is provided for the newspecies by assigning it to a pre-existing genus, so that thename is made available under the International Code ofZoological Nomenclature (ICZN). Genera of previouslydescribed species are indicated at the first mention of thespecies. Later on Lanham’s species names are used (else-where referred to as uninominal names; Lanham, 1965;and see Dayrat et al., 2004). Under the cladotypic nomen-clature taxa are referred to by names written in italicswith a capital letter. Linnaean taxon names are avoidedand ‘vernacularized’ names are preferred (e.g., orthopter-ans rather than Orthoptera). The choice of the nomen-clatural procedure is based on the decision of one of us(OB; see also Bethoux, 2010) and does not imply the

C© 2012 The AuthorsJournal 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 thisprocedure.

Systematics

Taxon Archaeorthoptera nom. Bethoux and Nel, 2002adis.-typ. Bethoux, 2007d

Genus Heterologus Carpenter, 1944Heterologus duyiwuer sp.nov.

Diagnosis ScP reaching RA; RP and M without anyconnection; M weak, concave, simple for a long dis-tance; fusion CuA + CuPaα short; CuA branched dis-tally; CuPaα and CuPaβ simple; CuPb branched; cross-veins rarely reticulated, often forming cells broader thanlong.

Description Holotype CNU-NX1–266: incompletenegative imprint and complete positive imprint of a rightforewing, very well preserved; length 27.6 mm, width (op-posite the end of CuPaα) 10.3 mm; area between anteriorwing margin and ScP broad, filled with ScP veinlets, withfew cross-veins between them; ScP reaching RA; ScP+RA with weak distal anterior veinlets (cross-veins?); Rsimple for 8.8 mm; RP simple for 6.1 mm, with 8 branchesreaching wing apex, without clear branching pattern; RPpreserved as tube basal to its first branching, branches ofRP preserved as depressions; M + CuA convex, branchinginto M and CuA 7.7 mm distal to wing base; M (diverg-ing from M + CuA) weakly sclerotized, concave, simplefor 8.2 mm, with 3 distal branches, branches of M im-printed as depressions; CuA (diverging from M + CuA)



0.9 mm long before its fusion with CuPaα; CuA + CuPaαshort (1.2 mm long; see Comments for homologization ofCuA+ CuPaα); distal to the divergence of CuPaα (fromCuA+ CuPaα), CuA simple for 5.5 mm, posteriorly pecti-nate, with 4 branches reaching posterior wing margin;imprint of CuA turns from a tube-like structure to adepression basal to the first branching of CuA; CuPbranched 3.2 mm distal to wing base; CuPa simple for4.0 mm; CuPaα 1.3 mm long before fusion with CuA;CuPaα and CuPaβ simple; CuPb simple for 7.3 mm, pos-teriorly pectinate, with 3 branches reaching posterior wingmargin; CuPaα, CuPaβ and CuPb preserved as tubularimprints in their basal part, then as depressions; first AAvein branched distally; 2 additional convex AA veins oc-cur; cross-veins convex, strong, especially in the middleof their course, straight, rarely reticulated, often formingcells broader than long; in areas between CuA and CuPaβ,the most distal cross-vein (convex) reaches the posteriormargin obliquely, forming a main-vein-like fork pattern(main vein concave); wing overall dark, darker colorationalong cross-veins (observed in the middle part).

Material examined Holotype CNU-NX1–266, com-posed of negative (fragment) and positive (complete) im-prints of an isolated left forewing.

Type locality and horizon Tupo Formation (NingxiaHui Autonomous region, China); Namurian, UpperCarboniferous (Lu et al., 2002).

Etymology ‘In a class by oneself’ [duyiwuer] inChinese.

Discussion

The new material belongs to a species that can be as-signed to Archaeorthoptera as it exhibits the definingcharacter state of this taxon, namely in forewings, a fu-sion of CuA (diverging from M + CuA) with a branchof CuP (Bethoux, 2007d). This species is considered asa panorthopteran insect as it exhibits one of the diagnos-tic character states of the group pointed out by Bethouxand Nel (2002a), namely the branching of CuPa basal tothe fusion of its anterior branch (CuPaα) with CuA (thetaxon name ‘Panorthoptera’ is not yet associated with adefinition under cladotypic nomenclature).

Among panorthopterans, the new material differs fromspecies of geraridaeans after its more distal branching ofM, the lack of strong and oblique anterior branches ofRA, and the lack of connection of RP with the anteriorbranch of M (see Bethoux and Nel, 2003; among oth-ers). It differs from Late Palaeozoic oedischioideans (pa-

raphyletic basal saltatorians) chiefly after the branchingpattern of CuA + CuPaα, with a long distance betweenthe first basal branch (itself homologized as CuPaα, seebelow) and the next fork (giving rise to branches of CuA,see below). In oedischioideans, whichever conjecture ofprimary homology is followed for CuA + CuPaα (CuAsimple + branched CuPaα vs. CuA branched + simpleCuPaα vs. CuA branched + CuPaα branched), branchesare emitted at a regular pace (Bethoux and Nel, 2002b;Bethoux et al., 2002; Carpenter, 1992; Gorochov, 1987;Sharov, 1968, 1971, among others). In addition all stem-saltatorians exhibit a first branching of M near the point ofdivergence of CuA from M + CuA, while M is simple fora long distance in the specimen CNU-NX1–266. Lastly,most of these basal saltatorians exhibit a connection ofRP with an anterior branch of M.

The species to which the specimen CNU-NX1–266belongs differs from trecwithiensis Kukalova-Peck &Brauckmann, 1992 (genus Osnogerarus Kukalova-Peck& Brauckmann, 1992; see Brauckmann & Herd, 2006:fig. 3) after its simple CuPaβ (branched in trecwithien-sis), the lack of connection between RP and the anteriorbranch of M, a lower number of branches of M, the lack ofstrong branches from ScP and the cross-venation (denselyreticulated in trecwithiensis).

After its late branching of M and its cross-venation, thenew material resembles langfordorum Carpenter, 1944(genus Heterologus Carpenter, 1944). However, langfor-dorum has a simple CuPb, while this vein is branched inthe specimen CNU-NX1–266. In addition langfordorumhas more numerous branches of RP, M and CuA + CuPaα.The erection of a new species is therefore justified.

The species is provisionally assigned to the genusHeterologus Carpenter, 1944 in order to make the speciesname available under the ICZN. The genus was composedof langfordorum Carpenter, 1944 only. As newly delim-ited, the genus is probably paraphyletic but this does notprevent the name Heterologus duyiwuer to be valid underthe ICZN. Notice that under the cladotypic nomenclat-ural procedure the species can be referred to as duyiwerBethoux, Gu & Ren, 2011, or Archaeorthoptera duyiwuerBethoux, Gu & Ren, 2011.

Except for ScP, RA and the most basal AA veins, im-prints of all main veins turn suddenly from a tube-like toa depression-like preservation, from the forewing base tothe apex and posterior wing margin. The origin of thisswitch is unknown, but clearly, vein elevation cannot beconsidered as relevant for discussion regarding vein ho-mologies in the distal part.

The branching pattern of CuA + CuPaα in duyiwueris not usual. In lobeattid insects and oedischioideans,among others, CuA + CuPaα emit posterior branches at a


A new stem-orthopteran 157

regular pace (Bethoux, 2008a; Bethoux & Nel, 2004;Sharov, 1968, among others). In duyiwuer a basal branchdiverges near the fusion of CuA with CuPaα, while othersbranch off more distally. This pattern suggests that thefirst posterior branch is CuPaα, while distal branches be-long to CuA. The composite vein CuA + CuPaα wouldtherefore be composed of a branched CuA and a simpleCuPaα.

Conclusion

Recent progress in the systematics of Upper Carbonifer-ous Archaeorthoptera allowed us to identify a species ofphylogenetic importance among the large sample assem-bled by DR’s the research team. Indeed the new materialallowed a new primary homology for the branches ofCuA + CuPaα to be conjectured in a stem-orthoptean.In addition the new species exhibits a combination ofcharacter states previously unknown. It will likely bea key taxon in future phylogenetic analysis of basalArchaeorthoptera.

Acknowledgements

We thank an anonymous reviewer for constructive com-ments. The first author was a recipient of an Endeavourpostdoctoral Award at the time of writing. This researchis supported by the National Natural Science Founda-tion of China (No. 40872022, 31071964, 31172143), Na-tional Basic Research Program of China (973 Program)(2012CB821906), Nature Science Foundation of Beijing(No. 5082002), the Scientific Research Key Program(KZ200910028005) and PHR Project of Beijing Munici-pal Commission of Education (20090509, 201107120).

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Accepted May 26, 2011


a new upper carboniferous stem-orthopteran (insecta) from ningxia (china)

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