Foss Rec 20 253ndash257 2017httpsdoiorg105194fr-20-253-2017copy Author(s) 2017 This work is distributed underthe Creative Commons Attribution 30 License

A new rare and distinctive species of Panorthoptera (InsectaArchaeorthoptera) from the Upper Carboniferous ofXiaheyan (Ningxia China)Jun-Jie Gu1 Olivier Beacutethoux24 and Dong Ren3

1College of Biological Sciences and Engineering North Minzu University 204 Wenchangbeijie Xixia DistrictYinchuan 750021 China2Sorbonne Universiteacutes UPMC (Paris 6) MNHN CNRS UMR 7207 Centre de recherche sur la Paleacuteobiodiversiteacute et lesPaleacuteoenvironnements (CR2P) Paris France3College of Life Sciences Capital Normal University 105 Xisanhuanbeilu Haidian District Beijing 100048 China4Museacuteum National drsquoHistoire Naturelle 57 rue Cuvier CP38 75005 Paris France

Correspondence to Dong Ren (rendongmailcnueducn)

Received 9 May 2017 ndash Revised 4 September 2017 ndash Accepted 22 September 2017 ndash Published 7 November 2017

Abstract The Xiaheyan locality is providing abundant ma-terial on one of the earliest insect faunas The most commonspecies are comparatively remote relatives of Orthoptera(grasshoppers crickets and katydids) belonging to the Ar-chaeorthoptera nec Panorthoptera (the latter including crownOrthoptera) We establish the genus Sinogerarus gen novto accommodate a new rare species S pectinatus specnov which can be conclusively assigned to PanorthopteraIt therefore represents the second occurrence of the group atthis locality The new species displays a distinctive combina-tion of character states adding to the knowledge of a numberof Upper Carboniferous Panorthoptera whose relationshipsare poorly understood

1 Introduction

The Xiaheyan locality (Ningxia China) is a Late Carbonifer-ous outcrop providing insects in very large abundance ie atleast an order of magnitude above known contemporaneouslocalities Several of the described species are represented bytens of specimens including some with both forewings pre-served attached to the pterothorax This exceptional materialallowed solid inferences on wing venation intra-specific vari-ability in various insect lineages (eg Beacutethoux et al 20112012b Cui et al 2011 Gu et al 2011 Li et al 2013Pecharovaacute et al 2015) Moreover both males and females

were identified for two species (Du et al 2017 Pecharovaacuteet al 2015) one of them a relative of grasshoppers cricketsand katydids (Orthoptera) displaying a significant female-biased sexual size dimorphism Such data are essential tobetter delimit early insect species a chronic issue and in par-ticular for those species with no relevant extant counterparts(Beacutethoux 2009)

The current overview of the insect fauna (or at least thetaphocoenosis) suggests that it is dominated by stem lin-eages of Orthoptera Most of the species described to dateare comparatively remote relatives of Orthoptera (Archae-orthoptera nec Panorthoptera) Panorthoptera a taxon includ-ing species more closely related to crown Orthoptera are rep-resented by a single species Heterologus duyiwuer BeacutethouxGu and Ren 2012a with a single forewing described Hereinwe describe the second species of Panorthoptera from thisfauna also documented by a single forewing which displaysa unique combination of character states

2 Material and methods

The studied specimen is housed at the Key Lab of Insect Evo-lution and Environmental Changes College of Life SciencesCapital Normal University Beijing China (CNU) It wascollected from the locality near Xiaheyan village Zhong-wei City Ningxia Hui Autonomous Region China The ex-act layer from which it was collected was not documented

Published by Copernicus Publications on behalf of the Museum fuumlr Naturkunde Berlin

254 J-J Gu et al A new Panorthoptera from Xiaheyan

Previous studies suggested a Namurian BCearly Bashkirianage for the whole locality (Lu et al 2002 Zhang et al2013) but ongoing studies (Truumlmper et al 2017) are reveal-ing that the locality is probably younger A Moscovian age ismore likely for the higher insect-bearing horizons

A draft drawing was produced using a Leica MZ125 dis-secting microscope equipped with a drawing tube (LeicaWetzlar Germany) It was finalized using Adobe IllustratorCS6 (Adobe Systems San Jose CA USA) The photographwas taken using a digital camera Canon 5D Mark III (CanonTokyo Japan) coupled to a Canon MP-E 65 mm macro lensequipped with a polarizing filter The resulting photographwas optimized using Adobe Photoshop CS6

We follow the serial insect wing venation ground pat-tern (Lameere 1922 1923) The corresponding wing vena-tion nomenclature is repeated for convenience ScP posteriorsubcosta R radius RA anterior radius RP posterior radiusM media MA anterior media MP posterior media Cucubitus CuA anterior cubitus (indicated by an asterisk inFig 1) CuP posterior cubitus CuPa anterior branch of CuPCuPaα anterior branch of CuPa CuPaβ posterior branch ofCuPa CuPb posterior branch of CuP AA anterior analisBased on this ground pattern we follow topographic homol-ogy conjectures proposed by Beacutethoux and Nel (2002 2001)for total Orthoptera (see Du et al 2017 for references onalternative proposals and the associated discussion)

3 Description of new taxa

Taxon Archaeorthoptera Bethoux and Nel 2002

Taxon Panorthoptera Crampton 1928 (sensu Beacutethouxand Nel 2002)

Sinogerarus Gu Beacutethoux and Ren gen nov

(urnlsidzoobankorgactD42326AA-4A4A-4222-A2B2-7591FA91CF78) (The registration date is 27September 2017)

Type species Sinogerarus pectinatus spec nov

Gender Masculine

Etymology A combination of ldquoSinordquo referring to thegeographic origin of the holotype and Gerarus a genusname used for species sharing similarities with the new one

Diagnosis By monotypy as for the type species

Sinogerarus pectinatus Gu Beacutethoux and Ren spec nov

(urnlsidzoobankorgact008B9E61-F8FC-429F-A319-FD5914220387) (The registration date is 27September 2017)

(Fig 1)

Material Holotype only CNU-NX1-343 deposited at theCNU

Etymology The species epithet derives from the Latinldquopectinatusrdquo referring to the regular branching patterns ofRP MP and CuA + CuPaα all posteriorly pectinate

Diagnosis Forewing RP regularly posteriorly pecti-nate lack of connection of RP with MA (or the anteriorbranch of M) M (emerging from M + CuA) simple for along distance (first fork at ca the second fifth of M length)MP regularly posteriorly pectinate CuA + CuPaα straightregularly posteriorly pectinate CuPaβ simple CuPb forkeddistally cross-venation reticulated in most areas withloosely organized pseudo-veins in the distal part of the areasbetween RP branches

Description Holotype specimen (CNU-NX1-343) Iso-lated forewing veins elevation not evident (as a consequenceit cannot be determined whether it is a right or a left wing)base most of anterior wing margin and apical area missingwing apparently disrupted just basal of the first RP fork(putative correspondences indicated by double-headedarrows in Fig 1) preserved length 236 mm estimatedlength about 32 mm width about 8 mm area between ScPand RRA narrow with straight and distinct cross-veins RAsimple in preserved parts RP posteriorly pectinate with fivebranches preserved all simple (as preserved) M simple for80 mm before it forks into MA and MP (just basal of thefirst RP fork) MA simple MP posteriorly pectinate withthree branches reaching posterior wing margin all simplefree CuA (emerging from M + CuA) 10 mm long fusingwith CuPaα CuA + CuPaα straight simple for 24 mmthen posteriorly pectinate with a total of six branchesreaching the posterior wing margin all simple CuPa forked(into CuPaα and CuPaβ) shortly before the origin of freeCuA area between M and CuA + CuPaα broad with up tofour rows of cells CuPb forked distally with two terminalbranches most areas of preserved forewing covered by anetwork of cross-veins loosely organized in pseudo-veins inthe distal part of the areas between RP branches

Locality and horizon Xiaheyan village Zhongwei CityNingxia China Tupo Formation Namurian BC UpperCarboniferous

Discussion The new specimen can be assigned to Ar-chaeorthoptera as it exhibits the defining character state ofthis taxon (formally defined in Beacutethoux 2007) namely afusion of CuA (diverging from M + CuA) with the anteriorbranch of CuP The occurrence of a fork of CuPa basalto the fusion with CuA (itself emerging from M + CuA)indicates that within Archaeorthoptera it belongs to theclade Panorthoptera (not formally defined see diagnosisin Beacutethoux and Nel 2002) This clade includes Orthoptera

Foss Rec 20 253ndash257 2017 wwwfoss-recnet202532017

J-J Gu et al A new Panorthoptera from Xiaheyan 255

Figure 1 Sinogerarus pectinatus gen and spec nov holotype (CNU-NX1-343) habitus (a) interpretative drawing and (b) photograph(flipped horizontally) See text for symbols and abbreviations

and their closest fossil relatives such as ldquogeraridaeansrdquoand ldquooedischioideansrdquo whose reciprocal relationships arenot evident (the supposed ldquohemipteroidrdquo affinities of theformer were challenged by Beacutethoux and Briggs 2008)and a number of distinctive species such as H duyiwuerand Bruaylogus magnificus Coty Haacuteva Prokop Roquesand Nel 2014 The Caloneurodea an extinct Palaeozoicorder are also considered close relatives of Orthoptera(Beacutethoux et al 2004) As argued below the new specimenshares similarities with many of these taxa but cannot beconclusively affiliated to any

Most ldquogeraridaeansrdquo and all ldquooedischioideansrdquo display afork of M located shortly after the point of divergence of Mfrom M + CuA (Beacutethoux and Nel 2003 Carpenter 1992Gorochov 1995 Kukalovaacute-Peck and Brauckmann 1992Sharov 1968 1971 Zessin 2009) unlike the new specimenin which M is simple for ca two-fifths of its length Amongldquogeraridaeansrdquo Gerarus fischeri (Brongniart 1885) (revisedin Beacutethoux and Nel 2003) and Osnogerarus trecwithiensisKukalovaacute-Peck and Brauckmann 1992 (revised in Brauck-mann and Herd 2006 ndash including a drawing by O Beacutethoux)both display a comparatively distal first fork of M Yet thenew specimen can be distinguished from both species (andmost ldquooedischioideansrdquo) owing to the lack of ldquoconnectionrdquo(actual fusion or connection via a short cross-vein) betweenRP and MA Moreover cross-venation is denser in G fish-

eri and O trecwithiensis and does not tend to form inter-calary pseudo-veins in the apical area Finally the forkedCuPb precludes an assignment of the new specimen to theldquooedischioideansrdquo in which this vein is always simple

The very regular branching pattern of RP and the apparentdistal fork of M are reminiscent of the Caloneurodea but (1)according to Beacutethoux et al (2004) the apparent distal forkof M in Caloneurodea is actually the first fork of MA thefirst fork of M being super-imposed with the MCuA fork(there is no evidence of such fusion in the new specimen) and(2) in Caloneurodea CuA + CuPaα (actually MP + CuA +

CuPaα) is simple (if with an anterior branch it then corre-sponds to MP CuA + CuPaα is richly branched in the newspecimen) Note that Spiculum mendicum Brauckmann andHerd 2006 is a candidate Caloneurodea Affinities with thisorder can be excluded

Among the ldquounplacedrdquo Panorthoptera species B magnifi-cus has a fusion of CuPaα located opposite the point of diver-gence of M and CuA In other words there is no free part ofCuA emerging from M + CuA unlike in the new specimen(asterisk in Fig 1) Several other features (cross-venationbranching pattern of RP etc) allow the new specimen to bedifferentiated from B magnificus After its late branching ofM the new specimen resembles Heterologus langfordorumCarpenter 1944 but this species does not display the regularbranching pattern of RP MP and CuA + CuPaα it also has a

wwwfoss-recnet202532017 Foss Rec 20 253ndash257 2017

256 J-J Gu et al A new Panorthoptera from Xiaheyan

simple CuPb (forked distally in the new specimen) The newspecimen maybe shares the most numerous similarities withH duyiwuer (from the same locality provisionally assignedto the genus Heterologus in the original description) includ-ing the distal first fork of M and the forked CuPb The mostconspicuous difference regards the cross-venation (reticula-tion occurs only in the distal-most areas in the forewing inH duyiwuer) Also branches of CuA + CuPaα are emittedregularly in the new specimen while in H duyiwuer there isa long distance between the point of divergence of the firstand second posterior branches Based on the available datathe new specimen has a forewing more elongate than that ofH duyiwuer A number of additional species which are pu-tative stem Orthoptera (see Brauckmann and Herd 2006) aredocumented based on very fragmentary material and there-fore will not be discussed We conclude that erecting a newgenus and a new species is justified

Sinogerarus pectinatus displays a combination of charac-ter states whose polarities are difficult to assess due to (1)the scarcity of non-ldquogeraridaeanrdquo and non-ldquooedischioideanrdquoPanorthoptera material and (2) the lack of identified immedi-ate sister group of Panorthoptera Even though it remains per-plexing in some respects the newly described species addsto the knowledge of a poorly documented set of Upper Car-boniferous insects

Data availability No data sets were used in this article

Competing interests The authors declare that they have no conflictof interest

Acknowledgements We thank the anonymous reviewers for theiruseful comments We thank the numerous students who overthe years contributed to fieldwork at Xiaheyan This research issupported by the National Natural Science Foundation of China(nos 31360525 31730087 41688103) Program for ChangjiangScholars and Innovative Research Team in University (IRT17R75)

Edited by Johannes MuumlllerReviewed by two anonymous referees

References

Beacutethoux O Cladotypic taxonomy applied titanopterans are or-thopterans Arth Syst Phyl 65 135ndash156 2007

Beacutethoux O Gaps and nodes between fossil and extant in-sects Syst Ent 34 599ndash609 httpsdoiorg101111j1365-3113200900484x 2009

Beacutethoux O and Briggs D E G How Gerarus lost its head stem-group Orthoptera and Paraneoptera revisited Syst Ent 33 529ndash547 httpsdoiorg101111j1365-3113200800419x 2008

Beacutethoux O and Nel A Venation pattern of OrthopteraJ Orthopt Res 10 195ndash198 httpsdoiorg1016651082-6467(2001)010[0195VPOO]20CO2 2001

Beacutethoux O and Nel A Venation pattern and revision of Or-thoptera sensu nov and sister groups Phylogeny of Palaeo-zoic and Mesozoic Orthoptera sensu nov Zootaxa 96 1ndash88httpsdoiorg1011646zootaxa9611 2002

Beacutethoux O and Nel A Wing venation morphology and vari-ability of Gerarus fischeri (Brongniart 1885) sensu Burnham(Panorthoptera Upper Carboniferous Commentry France) withinferences on flight performances Org Div Evol 3 173ndash183httpsdoiorg1010781439-6092-00070 2003

Beacutethoux O Nel A and Lapeyrie J The extinct order Caloneu-rodea (Insecta Pterygota Panorthoptera) wing venation sys-tematics and phylogenetic relationships Ann Zool 54 289ndash318 2004

Beacutethoux O Cui Y Kondratieff B Stark B and Ren D At lasta Pennsylvanian stem-stonefly (Plecoptera) discovered BMCEvol Biol 11 248 httpsdoiorg1011861471-2148-11-2482011

Beacutethoux O Gu J-j and Ren D A new Upper Carboniferousstem-orthopteran (Insecta) from Ningxia (China) Insect Sci19 153ndash158 httpsdoiorg101111j1744-7917201101468x2012a

Beacutethoux O Gu J Yue Y and Ren D Miamia maimai n spa new Pennsylvanian stem-orthopteran insect and a case studyon the application of cladotypic nomenclature Foss Rec 15103ndash113 httpsdoiorg101002mmng201200008 2012b

Brauckmann C and Herd K J Insekten-Funde aus dem Westfal-ium D (Ober-Karbon) des Piesberges bei Osnabruumlck (Deutsch-land) Teil 2 Neoptera Osnabruumlcker nat Mitt 3031 (for 2005)19ndash65 2006

Brongniart C Les Insectes fossiles des terrains primaires Coupdrsquooeil rapide sur la faune entomologique des terrains paleacuteo-zoiumlques Bull Soc Amis Sci nat Rouen 1885 50ndash68httpsdoiorg105962bhltitle36372 1885

Carpenter F M Carboniferous insects from the vicinity of MazonCreek Illinois Illinois State Mus Sci Pap 3 1ndash20 1944

Carpenter F M Superclass Hexapoda In Treatise on InvertebratePaleontology edited by Kaesler R L The Geological Societyof America and the University of Kansas Boulder 1992

Coty D Haacuteva J Prokop J Roques P and Nel A Newarchaeorthopteran insects from the Late Carboniferous of theNord and Pas-de-Calais basins in northern France (InsectaCnemidolestodea Panorthoptera) Zootaxa 3878 462ndash470httpsdoiorg1011646zootaxa387854 2014

Crampton G C The grouping of the insect orders and their linesof descent The Entomologist 61 82ndash85 1928

Cui Y Beacutethoux O and Ren D Intraindividual variabil-ity in Sinonamuropteridae forewing venation (GrylloblattidaLate Carboniferous) taxonomic and nomenclatural implica-tions Syst Ent 36 44ndash56 httpsdoiorg101111j1365-3113201000545x 2011

Du S Beacutethoux O Gu J and Ren D Protomiamia yangi gen etsp nov (Early Pennsylvanian Xiaheyan China) a sexually di-morphic Palaeozoic stem-Orthoptera J Syst Pal 15 193ndash204httpsdoiorg1010801477201920161154899 2017

Foss Rec 20 253ndash257 2017 wwwfoss-recnet202532017

254 J-J Gu et al A new Panorthoptera from Xiaheyan

Previous studies suggested a Namurian BCearly Bashkirianage for the whole locality (Lu et al 2002 Zhang et al2013) but ongoing studies (Truumlmper et al 2017) are reveal-ing that the locality is probably younger A Moscovian age ismore likely for the higher insect-bearing horizons

A draft drawing was produced using a Leica MZ125 dis-secting microscope equipped with a drawing tube (LeicaWetzlar Germany) It was finalized using Adobe IllustratorCS6 (Adobe Systems San Jose CA USA) The photographwas taken using a digital camera Canon 5D Mark III (CanonTokyo Japan) coupled to a Canon MP-E 65 mm macro lensequipped with a polarizing filter The resulting photographwas optimized using Adobe Photoshop CS6

We follow the serial insect wing venation ground pat-tern (Lameere 1922 1923) The corresponding wing vena-tion nomenclature is repeated for convenience ScP posteriorsubcosta R radius RA anterior radius RP posterior radiusM media MA anterior media MP posterior media Cucubitus CuA anterior cubitus (indicated by an asterisk inFig 1) CuP posterior cubitus CuPa anterior branch of CuPCuPaα anterior branch of CuPa CuPaβ posterior branch ofCuPa CuPb posterior branch of CuP AA anterior analisBased on this ground pattern we follow topographic homol-ogy conjectures proposed by Beacutethoux and Nel (2002 2001)for total Orthoptera (see Du et al 2017 for references onalternative proposals and the associated discussion)

3 Description of new taxa

Taxon Archaeorthoptera Bethoux and Nel 2002

Taxon Panorthoptera Crampton 1928 (sensu Beacutethouxand Nel 2002)

Sinogerarus Gu Beacutethoux and Ren gen nov

(urnlsidzoobankorgactD42326AA-4A4A-4222-A2B2-7591FA91CF78) (The registration date is 27September 2017)

Type species Sinogerarus pectinatus spec nov

Gender Masculine

Etymology A combination of ldquoSinordquo referring to thegeographic origin of the holotype and Gerarus a genusname used for species sharing similarities with the new one

Diagnosis By monotypy as for the type species

Sinogerarus pectinatus Gu Beacutethoux and Ren spec nov

(urnlsidzoobankorgact008B9E61-F8FC-429F-A319-FD5914220387) (The registration date is 27September 2017)

(Fig 1)

Material Holotype only CNU-NX1-343 deposited at theCNU

Etymology The species epithet derives from the Latinldquopectinatusrdquo referring to the regular branching patterns ofRP MP and CuA + CuPaα all posteriorly pectinate

Diagnosis Forewing RP regularly posteriorly pecti-nate lack of connection of RP with MA (or the anteriorbranch of M) M (emerging from M + CuA) simple for along distance (first fork at ca the second fifth of M length)MP regularly posteriorly pectinate CuA + CuPaα straightregularly posteriorly pectinate CuPaβ simple CuPb forkeddistally cross-venation reticulated in most areas withloosely organized pseudo-veins in the distal part of the areasbetween RP branches

Description Holotype specimen (CNU-NX1-343) Iso-lated forewing veins elevation not evident (as a consequenceit cannot be determined whether it is a right or a left wing)base most of anterior wing margin and apical area missingwing apparently disrupted just basal of the first RP fork(putative correspondences indicated by double-headedarrows in Fig 1) preserved length 236 mm estimatedlength about 32 mm width about 8 mm area between ScPand RRA narrow with straight and distinct cross-veins RAsimple in preserved parts RP posteriorly pectinate with fivebranches preserved all simple (as preserved) M simple for80 mm before it forks into MA and MP (just basal of thefirst RP fork) MA simple MP posteriorly pectinate withthree branches reaching posterior wing margin all simplefree CuA (emerging from M + CuA) 10 mm long fusingwith CuPaα CuA + CuPaα straight simple for 24 mmthen posteriorly pectinate with a total of six branchesreaching the posterior wing margin all simple CuPa forked(into CuPaα and CuPaβ) shortly before the origin of freeCuA area between M and CuA + CuPaα broad with up tofour rows of cells CuPb forked distally with two terminalbranches most areas of preserved forewing covered by anetwork of cross-veins loosely organized in pseudo-veins inthe distal part of the areas between RP branches

Locality and horizon Xiaheyan village Zhongwei CityNingxia China Tupo Formation Namurian BC UpperCarboniferous

Discussion The new specimen can be assigned to Ar-chaeorthoptera as it exhibits the defining character state ofthis taxon (formally defined in Beacutethoux 2007) namely afusion of CuA (diverging from M + CuA) with the anteriorbranch of CuP The occurrence of a fork of CuPa basalto the fusion with CuA (itself emerging from M + CuA)indicates that within Archaeorthoptera it belongs to theclade Panorthoptera (not formally defined see diagnosisin Beacutethoux and Nel 2002) This clade includes Orthoptera

Foss Rec 20 253ndash257 2017 wwwfoss-recnet202532017

J-J Gu et al A new Panorthoptera from Xiaheyan 255

Figure 1 Sinogerarus pectinatus gen and spec nov holotype (CNU-NX1-343) habitus (a) interpretative drawing and (b) photograph(flipped horizontally) See text for symbols and abbreviations

and their closest fossil relatives such as ldquogeraridaeansrdquoand ldquooedischioideansrdquo whose reciprocal relationships arenot evident (the supposed ldquohemipteroidrdquo affinities of theformer were challenged by Beacutethoux and Briggs 2008)and a number of distinctive species such as H duyiwuerand Bruaylogus magnificus Coty Haacuteva Prokop Roquesand Nel 2014 The Caloneurodea an extinct Palaeozoicorder are also considered close relatives of Orthoptera(Beacutethoux et al 2004) As argued below the new specimenshares similarities with many of these taxa but cannot beconclusively affiliated to any

Most ldquogeraridaeansrdquo and all ldquooedischioideansrdquo display afork of M located shortly after the point of divergence of Mfrom M + CuA (Beacutethoux and Nel 2003 Carpenter 1992Gorochov 1995 Kukalovaacute-Peck and Brauckmann 1992Sharov 1968 1971 Zessin 2009) unlike the new specimenin which M is simple for ca two-fifths of its length Amongldquogeraridaeansrdquo Gerarus fischeri (Brongniart 1885) (revisedin Beacutethoux and Nel 2003) and Osnogerarus trecwithiensisKukalovaacute-Peck and Brauckmann 1992 (revised in Brauck-mann and Herd 2006 ndash including a drawing by O Beacutethoux)both display a comparatively distal first fork of M Yet thenew specimen can be distinguished from both species (andmost ldquooedischioideansrdquo) owing to the lack of ldquoconnectionrdquo(actual fusion or connection via a short cross-vein) betweenRP and MA Moreover cross-venation is denser in G fish-

eri and O trecwithiensis and does not tend to form inter-calary pseudo-veins in the apical area Finally the forkedCuPb precludes an assignment of the new specimen to theldquooedischioideansrdquo in which this vein is always simple

The very regular branching pattern of RP and the apparentdistal fork of M are reminiscent of the Caloneurodea but (1)according to Beacutethoux et al (2004) the apparent distal forkof M in Caloneurodea is actually the first fork of MA thefirst fork of M being super-imposed with the MCuA fork(there is no evidence of such fusion in the new specimen) and(2) in Caloneurodea CuA + CuPaα (actually MP + CuA +

CuPaα) is simple (if with an anterior branch it then corre-sponds to MP CuA + CuPaα is richly branched in the newspecimen) Note that Spiculum mendicum Brauckmann andHerd 2006 is a candidate Caloneurodea Affinities with thisorder can be excluded

Among the ldquounplacedrdquo Panorthoptera species B magnifi-cus has a fusion of CuPaα located opposite the point of diver-gence of M and CuA In other words there is no free part ofCuA emerging from M + CuA unlike in the new specimen(asterisk in Fig 1) Several other features (cross-venationbranching pattern of RP etc) allow the new specimen to bedifferentiated from B magnificus After its late branching ofM the new specimen resembles Heterologus langfordorumCarpenter 1944 but this species does not display the regularbranching pattern of RP MP and CuA + CuPaα it also has a

wwwfoss-recnet202532017 Foss Rec 20 253ndash257 2017

256 J-J Gu et al A new Panorthoptera from Xiaheyan

simple CuPb (forked distally in the new specimen) The newspecimen maybe shares the most numerous similarities withH duyiwuer (from the same locality provisionally assignedto the genus Heterologus in the original description) includ-ing the distal first fork of M and the forked CuPb The mostconspicuous difference regards the cross-venation (reticula-tion occurs only in the distal-most areas in the forewing inH duyiwuer) Also branches of CuA + CuPaα are emittedregularly in the new specimen while in H duyiwuer there isa long distance between the point of divergence of the firstand second posterior branches Based on the available datathe new specimen has a forewing more elongate than that ofH duyiwuer A number of additional species which are pu-tative stem Orthoptera (see Brauckmann and Herd 2006) aredocumented based on very fragmentary material and there-fore will not be discussed We conclude that erecting a newgenus and a new species is justified

Sinogerarus pectinatus displays a combination of charac-ter states whose polarities are difficult to assess due to (1)the scarcity of non-ldquogeraridaeanrdquo and non-ldquooedischioideanrdquoPanorthoptera material and (2) the lack of identified immedi-ate sister group of Panorthoptera Even though it remains per-plexing in some respects the newly described species addsto the knowledge of a poorly documented set of Upper Car-boniferous insects

Data availability No data sets were used in this article

Competing interests The authors declare that they have no conflictof interest

Acknowledgements We thank the anonymous reviewers for theiruseful comments We thank the numerous students who overthe years contributed to fieldwork at Xiaheyan This research issupported by the National Natural Science Foundation of China(nos 31360525 31730087 41688103) Program for ChangjiangScholars and Innovative Research Team in University (IRT17R75)

Edited by Johannes MuumlllerReviewed by two anonymous referees

References

Beacutethoux O Cladotypic taxonomy applied titanopterans are or-thopterans Arth Syst Phyl 65 135ndash156 2007

Beacutethoux O Gaps and nodes between fossil and extant in-sects Syst Ent 34 599ndash609 httpsdoiorg101111j1365-3113200900484x 2009

Beacutethoux O and Briggs D E G How Gerarus lost its head stem-group Orthoptera and Paraneoptera revisited Syst Ent 33 529ndash547 httpsdoiorg101111j1365-3113200800419x 2008

Beacutethoux O and Nel A Venation pattern of OrthopteraJ Orthopt Res 10 195ndash198 httpsdoiorg1016651082-6467(2001)010[0195VPOO]20CO2 2001

Beacutethoux O and Nel A Venation pattern and revision of Or-thoptera sensu nov and sister groups Phylogeny of Palaeo-zoic and Mesozoic Orthoptera sensu nov Zootaxa 96 1ndash88httpsdoiorg1011646zootaxa9611 2002

Beacutethoux O and Nel A Wing venation morphology and vari-ability of Gerarus fischeri (Brongniart 1885) sensu Burnham(Panorthoptera Upper Carboniferous Commentry France) withinferences on flight performances Org Div Evol 3 173ndash183httpsdoiorg1010781439-6092-00070 2003

Beacutethoux O Nel A and Lapeyrie J The extinct order Caloneu-rodea (Insecta Pterygota Panorthoptera) wing venation sys-tematics and phylogenetic relationships Ann Zool 54 289ndash318 2004

Beacutethoux O Cui Y Kondratieff B Stark B and Ren D At lasta Pennsylvanian stem-stonefly (Plecoptera) discovered BMCEvol Biol 11 248 httpsdoiorg1011861471-2148-11-2482011

Beacutethoux O Gu J-j and Ren D A new Upper Carboniferousstem-orthopteran (Insecta) from Ningxia (China) Insect Sci19 153ndash158 httpsdoiorg101111j1744-7917201101468x2012a

Beacutethoux O Gu J Yue Y and Ren D Miamia maimai n spa new Pennsylvanian stem-orthopteran insect and a case studyon the application of cladotypic nomenclature Foss Rec 15103ndash113 httpsdoiorg101002mmng201200008 2012b

Brauckmann C and Herd K J Insekten-Funde aus dem Westfal-ium D (Ober-Karbon) des Piesberges bei Osnabruumlck (Deutsch-land) Teil 2 Neoptera Osnabruumlcker nat Mitt 3031 (for 2005)19ndash65 2006

Brongniart C Les Insectes fossiles des terrains primaires Coupdrsquooeil rapide sur la faune entomologique des terrains paleacuteo-zoiumlques Bull Soc Amis Sci nat Rouen 1885 50ndash68httpsdoiorg105962bhltitle36372 1885

Carpenter F M Carboniferous insects from the vicinity of MazonCreek Illinois Illinois State Mus Sci Pap 3 1ndash20 1944

Carpenter F M Superclass Hexapoda In Treatise on InvertebratePaleontology edited by Kaesler R L The Geological Societyof America and the University of Kansas Boulder 1992

Coty D Haacuteva J Prokop J Roques P and Nel A Newarchaeorthopteran insects from the Late Carboniferous of theNord and Pas-de-Calais basins in northern France (InsectaCnemidolestodea Panorthoptera) Zootaxa 3878 462ndash470httpsdoiorg1011646zootaxa387854 2014

Crampton G C The grouping of the insect orders and their linesof descent The Entomologist 61 82ndash85 1928

Cui Y Beacutethoux O and Ren D Intraindividual variabil-ity in Sinonamuropteridae forewing venation (GrylloblattidaLate Carboniferous) taxonomic and nomenclatural implica-tions Syst Ent 36 44ndash56 httpsdoiorg101111j1365-3113201000545x 2011

Du S Beacutethoux O Gu J and Ren D Protomiamia yangi gen etsp nov (Early Pennsylvanian Xiaheyan China) a sexually di-morphic Palaeozoic stem-Orthoptera J Syst Pal 15 193ndash204httpsdoiorg1010801477201920161154899 2017

Foss Rec 20 253ndash257 2017 wwwfoss-recnet202532017

J-J Gu et al A new Panorthoptera from Xiaheyan 255

Figure 1 Sinogerarus pectinatus gen and spec nov holotype (CNU-NX1-343) habitus (a) interpretative drawing and (b) photograph(flipped horizontally) See text for symbols and abbreviations

and their closest fossil relatives such as ldquogeraridaeansrdquoand ldquooedischioideansrdquo whose reciprocal relationships arenot evident (the supposed ldquohemipteroidrdquo affinities of theformer were challenged by Beacutethoux and Briggs 2008)and a number of distinctive species such as H duyiwuerand Bruaylogus magnificus Coty Haacuteva Prokop Roquesand Nel 2014 The Caloneurodea an extinct Palaeozoicorder are also considered close relatives of Orthoptera(Beacutethoux et al 2004) As argued below the new specimenshares similarities with many of these taxa but cannot beconclusively affiliated to any

Most ldquogeraridaeansrdquo and all ldquooedischioideansrdquo display afork of M located shortly after the point of divergence of Mfrom M + CuA (Beacutethoux and Nel 2003 Carpenter 1992Gorochov 1995 Kukalovaacute-Peck and Brauckmann 1992Sharov 1968 1971 Zessin 2009) unlike the new specimenin which M is simple for ca two-fifths of its length Amongldquogeraridaeansrdquo Gerarus fischeri (Brongniart 1885) (revisedin Beacutethoux and Nel 2003) and Osnogerarus trecwithiensisKukalovaacute-Peck and Brauckmann 1992 (revised in Brauck-mann and Herd 2006 ndash including a drawing by O Beacutethoux)both display a comparatively distal first fork of M Yet thenew specimen can be distinguished from both species (andmost ldquooedischioideansrdquo) owing to the lack of ldquoconnectionrdquo(actual fusion or connection via a short cross-vein) betweenRP and MA Moreover cross-venation is denser in G fish-

eri and O trecwithiensis and does not tend to form inter-calary pseudo-veins in the apical area Finally the forkedCuPb precludes an assignment of the new specimen to theldquooedischioideansrdquo in which this vein is always simple

The very regular branching pattern of RP and the apparentdistal fork of M are reminiscent of the Caloneurodea but (1)according to Beacutethoux et al (2004) the apparent distal forkof M in Caloneurodea is actually the first fork of MA thefirst fork of M being super-imposed with the MCuA fork(there is no evidence of such fusion in the new specimen) and(2) in Caloneurodea CuA + CuPaα (actually MP + CuA +

CuPaα) is simple (if with an anterior branch it then corre-sponds to MP CuA + CuPaα is richly branched in the newspecimen) Note that Spiculum mendicum Brauckmann andHerd 2006 is a candidate Caloneurodea Affinities with thisorder can be excluded

Among the ldquounplacedrdquo Panorthoptera species B magnifi-cus has a fusion of CuPaα located opposite the point of diver-gence of M and CuA In other words there is no free part ofCuA emerging from M + CuA unlike in the new specimen(asterisk in Fig 1) Several other features (cross-venationbranching pattern of RP etc) allow the new specimen to bedifferentiated from B magnificus After its late branching ofM the new specimen resembles Heterologus langfordorumCarpenter 1944 but this species does not display the regularbranching pattern of RP MP and CuA + CuPaα it also has a

wwwfoss-recnet202532017 Foss Rec 20 253ndash257 2017

256 J-J Gu et al A new Panorthoptera from Xiaheyan

simple CuPb (forked distally in the new specimen) The newspecimen maybe shares the most numerous similarities withH duyiwuer (from the same locality provisionally assignedto the genus Heterologus in the original description) includ-ing the distal first fork of M and the forked CuPb The mostconspicuous difference regards the cross-venation (reticula-tion occurs only in the distal-most areas in the forewing inH duyiwuer) Also branches of CuA + CuPaα are emittedregularly in the new specimen while in H duyiwuer there isa long distance between the point of divergence of the firstand second posterior branches Based on the available datathe new specimen has a forewing more elongate than that ofH duyiwuer A number of additional species which are pu-tative stem Orthoptera (see Brauckmann and Herd 2006) aredocumented based on very fragmentary material and there-fore will not be discussed We conclude that erecting a newgenus and a new species is justified

Sinogerarus pectinatus displays a combination of charac-ter states whose polarities are difficult to assess due to (1)the scarcity of non-ldquogeraridaeanrdquo and non-ldquooedischioideanrdquoPanorthoptera material and (2) the lack of identified immedi-ate sister group of Panorthoptera Even though it remains per-plexing in some respects the newly described species addsto the knowledge of a poorly documented set of Upper Car-boniferous insects

Data availability No data sets were used in this article

Competing interests The authors declare that they have no conflictof interest

Acknowledgements We thank the anonymous reviewers for theiruseful comments We thank the numerous students who overthe years contributed to fieldwork at Xiaheyan This research issupported by the National Natural Science Foundation of China(nos 31360525 31730087 41688103) Program for ChangjiangScholars and Innovative Research Team in University (IRT17R75)

Edited by Johannes MuumlllerReviewed by two anonymous referees

References

Beacutethoux O Cladotypic taxonomy applied titanopterans are or-thopterans Arth Syst Phyl 65 135ndash156 2007

Beacutethoux O Gaps and nodes between fossil and extant in-sects Syst Ent 34 599ndash609 httpsdoiorg101111j1365-3113200900484x 2009

Beacutethoux O and Briggs D E G How Gerarus lost its head stem-group Orthoptera and Paraneoptera revisited Syst Ent 33 529ndash547 httpsdoiorg101111j1365-3113200800419x 2008

Beacutethoux O and Nel A Venation pattern of OrthopteraJ Orthopt Res 10 195ndash198 httpsdoiorg1016651082-6467(2001)010[0195VPOO]20CO2 2001

Beacutethoux O and Nel A Venation pattern and revision of Or-thoptera sensu nov and sister groups Phylogeny of Palaeo-zoic and Mesozoic Orthoptera sensu nov Zootaxa 96 1ndash88httpsdoiorg1011646zootaxa9611 2002

Beacutethoux O and Nel A Wing venation morphology and vari-ability of Gerarus fischeri (Brongniart 1885) sensu Burnham(Panorthoptera Upper Carboniferous Commentry France) withinferences on flight performances Org Div Evol 3 173ndash183httpsdoiorg1010781439-6092-00070 2003

Beacutethoux O Nel A and Lapeyrie J The extinct order Caloneu-rodea (Insecta Pterygota Panorthoptera) wing venation sys-tematics and phylogenetic relationships Ann Zool 54 289ndash318 2004

Beacutethoux O Cui Y Kondratieff B Stark B and Ren D At lasta Pennsylvanian stem-stonefly (Plecoptera) discovered BMCEvol Biol 11 248 httpsdoiorg1011861471-2148-11-2482011

Beacutethoux O Gu J-j and Ren D A new Upper Carboniferousstem-orthopteran (Insecta) from Ningxia (China) Insect Sci19 153ndash158 httpsdoiorg101111j1744-7917201101468x2012a

Beacutethoux O Gu J Yue Y and Ren D Miamia maimai n spa new Pennsylvanian stem-orthopteran insect and a case studyon the application of cladotypic nomenclature Foss Rec 15103ndash113 httpsdoiorg101002mmng201200008 2012b

Brauckmann C and Herd K J Insekten-Funde aus dem Westfal-ium D (Ober-Karbon) des Piesberges bei Osnabruumlck (Deutsch-land) Teil 2 Neoptera Osnabruumlcker nat Mitt 3031 (for 2005)19ndash65 2006

Brongniart C Les Insectes fossiles des terrains primaires Coupdrsquooeil rapide sur la faune entomologique des terrains paleacuteo-zoiumlques Bull Soc Amis Sci nat Rouen 1885 50ndash68httpsdoiorg105962bhltitle36372 1885

Carpenter F M Carboniferous insects from the vicinity of MazonCreek Illinois Illinois State Mus Sci Pap 3 1ndash20 1944

Carpenter F M Superclass Hexapoda In Treatise on InvertebratePaleontology edited by Kaesler R L The Geological Societyof America and the University of Kansas Boulder 1992

Coty D Haacuteva J Prokop J Roques P and Nel A Newarchaeorthopteran insects from the Late Carboniferous of theNord and Pas-de-Calais basins in northern France (InsectaCnemidolestodea Panorthoptera) Zootaxa 3878 462ndash470httpsdoiorg1011646zootaxa387854 2014

Crampton G C The grouping of the insect orders and their linesof descent The Entomologist 61 82ndash85 1928

Cui Y Beacutethoux O and Ren D Intraindividual variabil-ity in Sinonamuropteridae forewing venation (GrylloblattidaLate Carboniferous) taxonomic and nomenclatural implica-tions Syst Ent 36 44ndash56 httpsdoiorg101111j1365-3113201000545x 2011

Du S Beacutethoux O Gu J and Ren D Protomiamia yangi gen etsp nov (Early Pennsylvanian Xiaheyan China) a sexually di-morphic Palaeozoic stem-Orthoptera J Syst Pal 15 193ndash204httpsdoiorg1010801477201920161154899 2017

Foss Rec 20 253ndash257 2017 wwwfoss-recnet202532017

256 J-J Gu et al A new Panorthoptera from Xiaheyan

simple CuPb (forked distally in the new specimen) The newspecimen maybe shares the most numerous similarities withH duyiwuer (from the same locality provisionally assignedto the genus Heterologus in the original description) includ-ing the distal first fork of M and the forked CuPb The mostconspicuous difference regards the cross-venation (reticula-tion occurs only in the distal-most areas in the forewing inH duyiwuer) Also branches of CuA + CuPaα are emittedregularly in the new specimen while in H duyiwuer there isa long distance between the point of divergence of the firstand second posterior branches Based on the available datathe new specimen has a forewing more elongate than that ofH duyiwuer A number of additional species which are pu-tative stem Orthoptera (see Brauckmann and Herd 2006) aredocumented based on very fragmentary material and there-fore will not be discussed We conclude that erecting a newgenus and a new species is justified

Sinogerarus pectinatus displays a combination of charac-ter states whose polarities are difficult to assess due to (1)the scarcity of non-ldquogeraridaeanrdquo and non-ldquooedischioideanrdquoPanorthoptera material and (2) the lack of identified immedi-ate sister group of Panorthoptera Even though it remains per-plexing in some respects the newly described species addsto the knowledge of a poorly documented set of Upper Car-boniferous insects

Data availability No data sets were used in this article

Competing interests The authors declare that they have no conflictof interest

Acknowledgements We thank the anonymous reviewers for theiruseful comments We thank the numerous students who overthe years contributed to fieldwork at Xiaheyan This research issupported by the National Natural Science Foundation of China(nos 31360525 31730087 41688103) Program for ChangjiangScholars and Innovative Research Team in University (IRT17R75)

Edited by Johannes MuumlllerReviewed by two anonymous referees

References

Beacutethoux O Cladotypic taxonomy applied titanopterans are or-thopterans Arth Syst Phyl 65 135ndash156 2007

Beacutethoux O Gaps and nodes between fossil and extant in-sects Syst Ent 34 599ndash609 httpsdoiorg101111j1365-3113200900484x 2009

Beacutethoux O and Briggs D E G How Gerarus lost its head stem-group Orthoptera and Paraneoptera revisited Syst Ent 33 529ndash547 httpsdoiorg101111j1365-3113200800419x 2008

Beacutethoux O and Nel A Venation pattern of OrthopteraJ Orthopt Res 10 195ndash198 httpsdoiorg1016651082-6467(2001)010[0195VPOO]20CO2 2001

Beacutethoux O and Nel A Venation pattern and revision of Or-thoptera sensu nov and sister groups Phylogeny of Palaeo-zoic and Mesozoic Orthoptera sensu nov Zootaxa 96 1ndash88httpsdoiorg1011646zootaxa9611 2002

Beacutethoux O and Nel A Wing venation morphology and vari-ability of Gerarus fischeri (Brongniart 1885) sensu Burnham(Panorthoptera Upper Carboniferous Commentry France) withinferences on flight performances Org Div Evol 3 173ndash183httpsdoiorg1010781439-6092-00070 2003

Beacutethoux O Nel A and Lapeyrie J The extinct order Caloneu-rodea (Insecta Pterygota Panorthoptera) wing venation sys-tematics and phylogenetic relationships Ann Zool 54 289ndash318 2004

Beacutethoux O Cui Y Kondratieff B Stark B and Ren D At lasta Pennsylvanian stem-stonefly (Plecoptera) discovered BMCEvol Biol 11 248 httpsdoiorg1011861471-2148-11-2482011

Beacutethoux O Gu J-j and Ren D A new Upper Carboniferousstem-orthopteran (Insecta) from Ningxia (China) Insect Sci19 153ndash158 httpsdoiorg101111j1744-7917201101468x2012a

Beacutethoux O Gu J Yue Y and Ren D Miamia maimai n spa new Pennsylvanian stem-orthopteran insect and a case studyon the application of cladotypic nomenclature Foss Rec 15103ndash113 httpsdoiorg101002mmng201200008 2012b

Brauckmann C and Herd K J Insekten-Funde aus dem Westfal-ium D (Ober-Karbon) des Piesberges bei Osnabruumlck (Deutsch-land) Teil 2 Neoptera Osnabruumlcker nat Mitt 3031 (for 2005)19ndash65 2006

Brongniart C Les Insectes fossiles des terrains primaires Coupdrsquooeil rapide sur la faune entomologique des terrains paleacuteo-zoiumlques Bull Soc Amis Sci nat Rouen 1885 50ndash68httpsdoiorg105962bhltitle36372 1885

Carpenter F M Carboniferous insects from the vicinity of MazonCreek Illinois Illinois State Mus Sci Pap 3 1ndash20 1944

Carpenter F M Superclass Hexapoda In Treatise on InvertebratePaleontology edited by Kaesler R L The Geological Societyof America and the University of Kansas Boulder 1992

Coty D Haacuteva J Prokop J Roques P and Nel A Newarchaeorthopteran insects from the Late Carboniferous of theNord and Pas-de-Calais basins in northern France (InsectaCnemidolestodea Panorthoptera) Zootaxa 3878 462ndash470httpsdoiorg1011646zootaxa387854 2014

Crampton G C The grouping of the insect orders and their linesof descent The Entomologist 61 82ndash85 1928

Cui Y Beacutethoux O and Ren D Intraindividual variabil-ity in Sinonamuropteridae forewing venation (GrylloblattidaLate Carboniferous) taxonomic and nomenclatural implica-tions Syst Ent 36 44ndash56 httpsdoiorg101111j1365-3113201000545x 2011

Du S Beacutethoux O Gu J and Ren D Protomiamia yangi gen etsp nov (Early Pennsylvanian Xiaheyan China) a sexually di-morphic Palaeozoic stem-Orthoptera J Syst Pal 15 193ndash204httpsdoiorg1010801477201920161154899 2017

Foss Rec 20 253ndash257 2017 wwwfoss-recnet202532017

J-J Gu et al A new Panorthoptera from Xiaheyan 257

Gorochov A V System and evolution of the suborder Ensifera(Orthoptera) Part I Proc Zool Inst Russ Acad Sci 260 1ndash224 1995

Gu J Beacutethoux O and Ren D Longzhua loculata n gen andn sp one of the most completely documented PennsylvanianArchaeorthoptera (Insecta Ningxia China) J Pal 85 303ndash314httpsdoiorg10166610-0851 2011

Kukalovaacute-Peck J and Brauckmann C Most Paleozoic Pro-torthoptera are ancestral hemipteroids major wing braces asclues to a new phylogeny of Neoptera (Insecta) Can J Zool70 2452ndash2473 httpsdoiorg101139z92-330 1992

Lameere A Sur la nervation alaire des insectes Bull Cl SciAcad Roy Belg 8 138ndash149 1922

Lameere A On the wing-venation of insects Psyche 30 123ndash132httpsdoiorg101155192316920 1923

Li Y Beacutethoux O Pang H and Ren D Early Pennsylva-nian Odonatoptera from the Xiaheyan locality (Ningxia China)new material taxa and perspectives Foss Rec 16 117ndash139httpsdoiorg101002mmng201300006 2013

Lu L Fang X Ji S and Pang Q A contribution to the knowl-edge of the Namurian in Ningxia Acta Geosci Sin 23 165ndash168 2002

Pecharovaacute M Ren D and Prokop J A new palaeodicty-opteroid (Megasecoptera Brodiopteridae) from the Early Penn-sylvanian of northern China reveals unique morphologicaltraits and intra-specific variability Alcheringa 39 236ndash249httpsdoiorg101080031155182015993299 2015

Sharov A G Filogeniya orthopteroidnykh nasekomykh Trudy Pa-leontol inst Akad Nauk SSSR 118 1ndash216 1968

Sharov A G Phylogeny of the Orthopteroidea Israel Program forScientific Translations Jerusalem 1971

Truumlmper S Schneider J W Beacutethoux O Ren D and Linne-mann U G Paper in preparation 2017

Zessin W Ploetzgerarus krempieni n gen et sp ndash eine neue Ger-aride (Insecta Panorthoptera Geraridae) aus dem Oberkarbon(Stephanium C) von Ploumltz bei Halle (Deutschland) Virgo MittEntomol Ver Mecklenburg 13 22ndash29 2009

Zhang Z Schneider J W and Hong Y The most ancient roach(Blattodea) a new genus and species from the earliest Late Car-boniferous (Namurian) of China with a discussion of the phy-lomorphogeny of early blattids J Syst Pal 11 27ndash40 2013httpsdoiorg101080147720192011634443

wwwfoss-recnet202532017 Foss Rec 20 253ndash257 2017