Molecular phylogeny of Chinese raspy crickets (Orthoptera: Gryllacrididae) reveals incongruences in current classification Author Li, Shi-Yu School of Life Sciences, East China Normal University, Shanghai 200241, China Author Liu, Yi-Jiao School of Life Sciences, East China Normal University, Shanghai 200241, China Author Xu, Jing-Yi School of Life Sciences, East China Normal University, Shanghai 200241, China Author Yin, Zi-Xu College of Fisheries, Ocean University of China, 5 Yushan Road, Qingdao 266003, China zqhe@bio.ecnu.edu.cn Author He, Zhu-Qing Museum of Biology, School of Life Sciences, East China Normal University, Shanghai 200241, China zqhe@bio.ecnu.edu.cn text Zoological Journal of the Linnean Society 2024 Zool. J. Linn. Soc. 2024-08-01 201 4 1 31 http://dx.doi.org/10.1093/zoolinnean/zlae051 journal article 10.1093/zoolinnean/zlae051 0024-4082 13220291 C695228-ACB5-42FA-A29F-BC2D6B822196 Eugryllacris Karny 1937 Ḋaeā ( Figs 7H , 8D , 9F ) Eugryllacris Karny 1937: 161 . Eugryllacris Jin and Xia 1994: 17 . Eugryllacris Gorochov 1995: 212 . Eugryllacris Otte 2000: 13 . Eugryllacris Taewoo et al. 2014: 96 . Eugryllacris Storozhenko et al. : 90. Eugryllacris Bian and Shi 2016a: 439 . Eugryllacris Ingrisch 2018: 27 . Eugryllacris Cadena-Castañeda 2019: 67 . Eugryllacris Bian et al. 2021: 209 . Type species: Gryllacris ruficeps Serville 1831 = Eugryllacris ruficeps ruficeps (Serville 1831) . Diagnosis: Eugryllacris can be recognized by the combination of the following characters: veins of forewings black ( Fig. 9F ), hindwings almost transparent ( Fig. 8D ); posterior margin of male ninth abdominal tergite with spines extending inwards from the middle of each lobe at base, length of spines shorter than half the width of ninth abdominal tergite ( Fig. 7H ). Eugryllacris is similar to Magnigryllacris , Radigryllacris , and Prosopogryllacris , but can be distinguished by the overall transparent hindwings. Magnigryllacris and Radigryllacris also differ from Eugryllacris by green veins and yellowishbrown cells of forewings when alive. Prosopogryllacris also differs from Eugryllacris by the black face (see Table 5 for details). Included species: Eugryllacris ruficeps ruficeps (Serville, 1831) , E. comotti (Griffini, 1908) , E. gandaki Ingrisch , 2018 , E. inversa Ingrisch, 2018 , E. loriae (Griffini, 1908) , E. maculipennis bakeri (Griffini, 1915) , E. maculipennis laticauda (Karny, 1925) , E. maculipennis maculipennis (Stal, 1877) , E. maculipennis specularis (Karny, 1925) , E. moesta laurentii (Griffini, 1918) , E. moesta moesta ( Brunner von Wattenwyl, 1888 ) , E. moestissima ( Brunner von Wattenwyl, 1888 ) , E. panteli (Bolivar, 1900) , E. poultoniana (Griffini, 1909) , E. princeps (Stal, 1877) , E. ruficeps stigmaticeps (Karny, 1929) , E. ruficeps viridiceps (Karny, 1926) , E. sarawaccensis (Karny, 1928) , E. sordida (Fritze, 1908) , E. viridescens (Walker, 1870) , E. vittipes lineosa (Walker, 1869) , E. vittipes nasalis (Walker, 1869) , E. vittipes vittipes (Walker, 1869) . Remarks: The genera Eugryllacris and Prosopogryllacris were established by Karny (1937) . Gorochov (2004) proposed that the species of Prosopogryllacris from Japan [ P . japonica (Matsumura and Shiraki, 1908) , P . simulans Ichikawa 2001 , P . rotundimacula Ichikawa 2001 , P . okadai Ichikawa 2001 , and P . iriomote Gorochov 2002 ] should be transferred to the genus Eugryllacris , because the apex of their ovipositor is sharp. Taewoo et al. (2014) moved P . japonica to the genus Eugryllacris based on their genital characters. Bian et al. (2016) moved P . cylindrigera to genus Eugryllacris , and described six species of Eugryllacris from China ( E . bifoliata Bian & Shi, 2016 , E . elongata Bian & Shi, 2016 , E . fanjingshanensis Bian & Shi, 2016 , E . lobulis Bian & Shi, 2016 , E . longifissa Bian & Shi, 2016 , and E . xiei Bian & Shi, 2016 ). Ingrisch (2018) described eight new species and one new subspecies of Eugryllacris from South-East Asia ( E . crassicauda Ingrisch, 2018 , E . gandaki Ingrisch, 2018 , E . guomashan Ingrisch, 2018 , E . inversa Ingrisch, 2018 , E . serricauda Ingrisch, 2018 , E . sulcata Ingrisch, 2018 , E . trabicauda Ingrisch, 2018 , and E . vermicauda Ingrisch, 2018 , and one subspecies E . crassicauda cambodiana Ingrisch, 2018 ), and he thought that the genus was characterized by a rather wide and short head, and a pronotum combined with the following coloration of the hindwings: cross veins in light colour, green or light brown bordered by light, semitransparent bands, while the middle of cells is dark or blackish brown. The molecular results show that the genus Eugryllacris is not a monophyletic group. Therefore, two new genera Magnigryllacris gen. nov. and Radigryllacris gen. nov. are erected. The differences among these genera include: colouration of foreand hindwings, and size of ocelli ( Table 5 ). After our new classification, species of Euryllacris are mainly distributed in the Malay Archipelago, species of Magnigryllacris and Prosopogryllacris are mainly distributed in South-East Asia, species of Radigryllacris are mainly distributed in East Asia . Magnigryllacris Li, Yin & He gen. nov. ḪHaeā http://zoobank.org/ urn:lsid:zoobank.org:act: 26DC1ECCFAD6-4A47-B81F-E9C1414EE9C5 ( Figs 7F , 8A, B , 9D ) Type species: Eugryllacris guomashan Ingrisch 2018 = Magnigryllacris guomashan ( Ingrisch 2018 ) . Diagnosis: Magnigryllacris can be recognized by the combination of the following characters: ocelli small or invisible; veins of forewings green and cells yellowish brown when alive; veins of hindwings light brown, cells between cross veins brown ( Fig. 8A, B ); posterior margin of male ninth abdominal tergite with spines extending inwards from each lobe at base, length of spines shorter than half the width of ninth abdominal tergite ( Fig. 7F ). Magnigryllacris is similar to Eugryllacris , Radigryllacris , and Prosopogryllacris , but can be distinguished by small or invisible ocelli and centre area of cells on hindwings brown. Eugryllacris also differs from Magnigryllacris by overall transparent hindwings. Prosopogryllacris also differs from Magnigryllacris by a black face (see Table 5 for details). Description: Body large-sized ( 25–50 mm . approx.) and stout ( Fig. 9D ); fastigium verticis about twice as wide as scapus; ocelli small or invisible; anterior margin of pronotum slightly roundly projecting, posterior margin straight; macropterous, veins of forewings green and cells yellowish brown when alive; veins of hindwings light brown, cells between cross veins brown ( Fig. 8A, B ); posterior margin of male ninth abdominal tergite with spines extending inwards from each lobe at base, length of spines shorter than half the width of ninth abdominal tergite ( Fig. 7F ); male subgenital plate with styli, posterior margin wide and slightly excised; female ovipositor long and curved upwards, dorsal margin with an expansion before tip. Included species: Magnigryllacris guomashan ( Ingrisch, 2018 ) comb. nov. , Mag. crassicauda cambodiana ( Ingrisch, 2018 ) comb. nov., Mag. crassicauda crassicauda ( Ingrisch, 2018 ) comb. nov. , Mag.hainanensis ( Bin & Bian, 2021 ) comb.nov. , Mag. Malaccensis (Griffini, 1908) comb. nov. , Mag. serricauda ( Ingrisch, 2018 ) comb. nov. , Mag. sulcata ( Ingrisch, 2018 ) comb. nov. , Mag . tiga (Yin & Shen, 2021) comb. nov., Mag. vaginalis (Pictet & Saussure, 1893) comb. nov .. Etymology: The word ‘magni-’ is derived from the Latin word ‘ magnus ’, meaning huge, and refers to the large size of species in this family. Remarks: See remarks for Eugryllacris and Table 5 . Key to species of male Magnigryllacris Li, Yin & He gen. nov. from China 1. Male ninth abdominal tergite with a pair of rod-like projections; hind margin of female seventh abdominal sternite shallowly concave .............................................................................................................................................................................................................2 - Male ninth abdominal tergite with a pair of small discs with granular margin, a styliform rounded projection below disc; hind margin of female seventh abdominal sternite with a projection...................................................................... Mag . guomashan 2. Projections of male ninth abdominal tergite close to each other..........................................................................................................3 - Projections of male ninth abdominal tergite widely separate .................................................................................... Mag . hainanensis 3. Face, legs, and pronotum without distinct black dots...................................................................................................... Mag . vaginalis - Face, legs, and pronotum with distinct black dots ..................................................................................................................... Mag . tiga Table 5. Diagnosis of the genera Eugryllacris , Magnigryllacris , Radigryllacris , and Prosopogryllacris .

Eugryllacris Karny	Magnigryllacris Li, Yin & He	Radigryllacris Li, Yin &	Prosopogryllacris Karny
1937	gen. nov.	He gen. nov.	1937
Forewings	Veins deep darkening,	Veins green, cells yellowish	Veins green, cells	Veins yellowish brown,
cells yellowish	brown when alive	yellowish brown	cells yellowish
transparent	when alive	transparent
Hindwings	Veins and cells overall	Longitudinal veins light	Longitudinal veins deep	Cells of bilateral cross veins
transparent	brown, centre of cells brown,	darkening, other	darkening, other areas
other areas transparent	areas transparent	yellowish transparent
Ocelli	Small or invisible	Small or invisible	Prominent	Prominent
Face	Typical color	Typical color	Typical color	Black

Radigryllacris Li, Yin & He gen. nov. Ñaeā http://zoobank.org/urn:lsid:zoobank.org:act:7B19B683- EA81-4CC5-AA29-D8B56E94715E ( Figs 7I , 8E, F , 9G ) Type species: Eugryllacris xiei Bian and Shi 2016a = Radigryllacris xiei (Bian and Shi 2016) . Diagnosis: Radigryllacris can be recognized by the combination of the following characters: ocelli distinct when alive, median ocellus subcircular; veins of forewings green and cells of forewings yellowish brown when alive, longitudinal veins of hindwings black and radial, and cells of wings colourless and transparent ( Fig. 8E, F ); posterior margin of male ninth abdominal tergite with spines extending inwards from the middle of each lobe at base; length of spines less than half the width of abdominal tergite ( Fig. 7I ). Radigryllacris is similar to Eugryllacris , Magnigryllacris , and Prosopogryllacris , but can be distinguished by: prominent ocellus, transparent cross veins and cells, and black longitudinal veins on hindwings. Eugryllacris also differs from Radigryllacris by overall transparent hindwings. Prosopogryllacris also differs from Radigryllacris by a black face (see Table 5 for details). Description: Body medium- to large-sized ( 25–40 mm . approx.) and stout ( Fig. 9G ); fastigium verticis about twice as wide as scapus; ocelli distinct when alive, median ocellus subcircular; anterior margin of pronotum slightly roundly projecting in the middle, posterior margin straight; macropterous, veins of forewings green and cells of forewings yellowish brown when alive, longitudinal veins of hindwings black and radial, and the cells of wings colourless and transparent ( Fig. 8E, F ); male ninth abdominal tergite divided in the middle, forming two lobes, posterior margin with spines extending inwards from the middle of each lobe at base; length of spine extending from male ninth abdominal tergite less than half the width of abdominal tergite ( Fig. 7I ); male subgenital plate with styli; posterior margin of female seventh abdominal sternite with a process in the middle, directing backwards; female ovipositor long and curved upwards, dorsal margin with an expansion before tip. Included species: Radigryllacris xiei (Bian & Shi, 2016) comb. nov. , R . bifoliata (Bian & Shi, 2016) comb. nov. , R . cylindrigera (Karny, 1926) comb. nov. , R . elongata (Bian & Shi, 2016) comb. nov. , R. fanjingshanensis (Bian & Shi, 2016) comb. nov. , R . forficata (Pang, Zhang & Bian, 2023 ) comb. nov., R . iriomote (Gorochov, 2002) comb. nov. , R . japonica (Matsumura and Shiraki, 1908) comb. nov. , R . lobulis (Bian & Shi, 2016) comb. nov. , R . longifissa (Bian & Shi, 2016) comb. nov. , R . longiproceris (Pang, Zhang & Bian, 2023 ) comb. nov., R . nigriabdominis ( Bin & Bian 2021 ) comb. nov., R . okadai (Ichikawa, 2001) comb. nov. , R . rotundimacula (Ichikawa, 2001) comb. nov. , R . simulans (Ichikawa, 2001) comb. nov. , R . trabicauda ( Ingrisch, 2018 ) comb. nov. , R . vermicauda ( Ingrisch, 2018 ) comb. nov .. Etymology: The word ‘radi-’ is derived from the Latin ‘ radius ’, meaning radiate object, and refers to the radial longitudinal vein of the hindwings in this genus. Remarks: See remarks for Eugryllacris and Table 5 . Key to species of male Radigryllacris Li, Yin & He gen. nov. from China 1. Abdominal tergite not black.........................................................................................................................................................................2 - Abdominal tergite black ................................................................................................................................................... R . nigriabdominis 2. Female seventh abdominal sternite longer than wide.............................................................................................................................3 - Female seventh abdominal sternite wider than long ........................................................................................................ R . cylindrigera 3. Process of female seventh abdominal sternite long, reaching the middle area of subgenital plate ................................................4 - Process of female seventh abdominal sternite short, only reaching the basal area of subgenital plate..........................................6 4. Apical half of process of female abdominal sternite slightly curved backwards, its apex faintly concave in the middle............5 - Apical half of process of female abdominal sternite nearly straight, its apex with a shallow concavity in the middle ................. ........................................................................................................................................................................................................... R . elongata 5. Process of female abdominal sternite slender.................................................................................................................. R . longiproceris - Process of female abdominal sternite stout ...................................................................................................................................... R . xiei 6. Lateral lobes of male subgenital plate as long as styli ..............................................................................................................................7 - Lateral lobes of male subgenital plate shorter than styli ........................................................................................................... R . lobulis 7. Lateral lobes of male subgenital plate on posterior margin obviously curved ventral wards ..........................................................8 - Lateral lobes of male subgenital plate not curved ventral wards...........................................................................................................9 8. Tips of lateral lobes of male subgenital plate near to each other, the distance about half the length of a stylus ........................... ........................................................................................................................................................................................................... R . forficata - Tips of lateral lobes of male subgenital plate far from each other, the distance longer than the length of a stylus....................... ............................................................................................................................................................................................. R . fanjingshanensis 9. Posterior margin of male subgenital plate deeply concave, lateral lobes nearly narrowly rounded ............................ R . longifissa - Posterior margin of male subgenital plate with a triangular concavity, lateral lobes nearly triangular......................... R . bifoliata