Phryganogryllacris Karny 1937

Li, Shi-Yu, Liu, Yi-Jiao, Xu, Jing-Yi, Yin, Zi-Xu & He, Zhu-Qing, 2024, Molecular phylogeny of Chinese raspy crickets (Orthoptera: Gryllacrididae) reveals incongruences in current classification, Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 201 (4), pp. 1-31 : 1-31

Phryganogryllacris Karny 1937 IJaeā

( Figs 10A, B , 11C, D , 12H ) Phryganogryllacris Karny 1937: 118 .

Phryganogryllacris Jin and Xia 1994: 17 .

Phryganogryllacris Liu 1999: 178 .

Phryganogryllacris Otte 2000: 28 .

Phryganogryllacris Gorochov 2005: 806 .

Phryganogryllacris Tan 2012: 39 .

Phryganogryllacris Li et al. 2014 b: 508 .

Phryganogryllacris Li et al. 2016: 376 .

Phryganogryllacris Ingrisch 2018: 231 .

Phryganogryllacris Cadena-Castañeda 2019: 89 . Phryganogryllacris Bian et al. 2021: 237 .

Phryganogryllacris Zhao et al. 2022: 157 .

Type species: Gryllacris phryganoides Haan, 1843 = Phryganogryllacris phryganoides phryganoides (Haan 1843) .

Diagnosis: Phryganogryllacris can be recognized by a combination of the following characters: forewings almost yellowish brown ( Fig. 11C, D ); male ninth abdominal tergite undivided, with the middle part of triangular shape in lateral view; apical margin of the middle of male subgenital plate obviously prominent and slightly divided into two lobes with cylindrical styli ( Fig. 10A, B ). Phryganogryllacris is similar to Sericgryllacris and Bicornisgryllacris . Sericgryllacris differs from Phryganogryllacris by darkened veins and transparent cells on forewings. Bicornisgryllacris differs from Phryganogryllacris by a pair of horns projecting from female seventh abdominal sternite (see Table 6 for details).

Included species: Phryganogryllacris phryganoides phryganoides (Haan, 1843) , Ph. arctata Walker 1869 , Ph. arctatiformis (Karny, 1925) , Ph. aruana (Karny, 1931) , Ph. bengalensis (Griffini, 1913) , Ph. brevixipha (Brunner von Wattenwyl, 1893) , Ph. cambodjana (Karny, 1929) , Ph. differens (Griffini, 1908) , Ph. discus Gorochov, 2008 , Ph. extensa Ingrisch, 2018 , Ph. gialaiensis fovealis Ingrisch, 2018 , Ph. gialaiensis gialaiensis Gorochov, 2005 , Ph. griseola (Karny, 1930) , Ph. grobbeni (Karny, 1925) , Ph. lobulata Gorochov, 2005 , Ph. mascata Karny, 1937 , Ph. mioccana (Karny, 1935) , Ph. nivea ( Brunner von Wattenwyl, 1888) , Ph. nonangulata Ingrisch, 2018 , Ph. parva Li, Liu & Li, 2014 , Ph. phryganoides elefanensis Gorochov, 2005 , Ph. phryganoides thailandensis Gorochov, 2005 , Ph. pusilla (Karny, 1926) , Ph. sphegidipraeda (Karny, 1924) , Ph. subangulata Gorochov, 2005 , Ph. superangulata Gorochov, 2005 , Ph. truncilamina Bian, 2023 , Ph. trusmadi Gorochov, 2008 .

Remarks: Phryganogryllacris was erected by Karny in 1937, with Gryllacris phryganoides De Haan 1842 as its type species.The genus can be distinguished from other genera by a pair of hooks with apex directed sidewards on male ninth abdominal tergite. Zhang and Bian (2023) suggested that Phryganogryllacris could be divided into two groups based on the morphology of male ninth tergite. However, our results do not support their classification. The molecular results show that the genus Phryganogryllacris is not a monophyleticgroup, andincludesthreegroups:(i) Ph. sichuanensis and Ph. decempunctata , (ii) Ph. brevixipha and Phryganogryllacris sp. , and (iii) Ph. mellii and Ph. subrectis . The differences among them include colour of forewings and posterior margin of female seventh abdominal sternite with or without a pair of horns projecting (see Table 6 for details). The type specimen of the genus Phryganogryllacris is a female with a flat subgenital plate and forewings in overall light yellow. Ph. brevixipha and Phryganogryllacris sp. are also consistent with these characters. Thus, we consider this group as the genus Phryganogryllacris . Females of Ph. mellii and Ph. subrectis possess a pair of horns projecting on the subgenital plate, and we erect Bicornisgryllacris gen nov. for this group. Colour of forewing veins of Ph. sichuanensis and Ph. decempunctata are darkened, and we erect Sericgryllacris gen nov. for this group. In our new system, species of Phryganogryllacris are distributed in East Asia and South-East Asia, species of Sericgryllacris are distributed in China, and species of Bicornisgryllacris are distributed in southern China.

Sericgryllacris Li, Yin & He gen. nov. Ṃaeā

http://zoobank.org/urn:lsid:zoobank.org:act:D399B719-

E4AF-475D-AC5E-2A2F6EF73F08

( Figs 10D–G , 11A, B )

Type species: Phryganogryllacris unicolor Liu and Wang 1998 = Sericgryllacris unicolor (Liu and Wang 1998) .

Diagnosis: Sericgryllacris can be recognized by the combination of the following characters: mostly veins of forewings black and cells transparent ( Fig. 11A, B ); ventral area of male ninth abdominal tergite on both sides with a rather large sturdy projection incurved basally and excurved apically; posterior margin of male subgenital plate with an incision in the middle; styli stout and slightly flattened. Sericgryllacris is similar to Phryganogryllacris and Bicornisgryllacris . Bicornisgryllacris differs from Sericgryllacris by a pair of horns projecting on female seventh abdominal sternite. Phryganogryllacris differs from Sericgryllacris by overall yellowish brown forewings (see Table 6 for details).

Description: Body small-sized (20–30 mm. approx.) and slender ( Fig. 11A, B ); fastigium of vertex about 1.5 times as broad as scape; anterior margin of pronotum slightly projecting in the middle, posterior margin nearly straight; lateral lobes longer than deep; most macropterous; veins of forewings black and cells transparent ( Fig. 11A, B ); second and third abdominal tergites each with two rows of large serried stridulatory pegs on both sides; male ninth abdominal tergite neither divided nor curved in the middle, ventral area of baso-lateral surface on both sides with a rather large sturdy projection incurved basally and excurved apically; posterior margin of male subgenital plate with an incision in the middle, styli stout and slightly flattened ( Fig. 10D–F ); posterior margin of female seventh abdominal sternite straight ( Fig. 10G ); female ovipositor long and nearly straight ( Fig. 11B ).

Included species: Sericgryllacris unicolor (Liu & Wang, 1998) comb. nov., Se. brevipennis (Li, Liu & Li, 2016) comb. nov., Se. decempunctata (Liu, Bi & Zhang, 2010) comb. nov., Se. fanjingshanensis (Li, Liu & Li, 2016) comb. nov., Se. flavistria (Pang, Zhang & Bian, 2023) comb. nov., Se. hubeiensis (Li, Liu & Li, 2016) comb. nov., Se. laoshanica (Yin, Wang & Shen, 2021) comb. nov., Se. longicerca ( Li, Liu & Li, 2014) comb. nov., Se. problematica ( Gorochov, 2005) comb. nov., Se. sheni (Niu & Shi, 1999) comb. nov., Se. sichuanensis ( Li, Liu & Li, 2014) comb. nov., Se. sigillata (Li, Liu & Li, 2016) comb. nov., Se. truncata ( Li, Liu & Li, 2014) comb. nov., Se. xiai (Liu & Zhang, 2001) comb. nov..

Etymology: The word ‘seric-’ is taken from the Latin ‘ serica ’, which was used by ancient Romans to describe land and products like silk from China.

Remarks: See remarks for Phryganogryllacris and Table 6 .

Key to species of male Sericgryllacris Li, Yin & He gen. nov. from China

1. Frons unicolorous, without black spots .....................................................................................................................................................2

- Frons with 10 black spots................................................................................................................................................ Se. decempunctata

2. Pronotal disc with black spots......................................................................................................................................................................3

- Pronotal disc without black spots ...............................................................................................................................................................8

3. Occiput with yellowish brown spots ..........................................................................................................................................................4

- Occiput entirely black............................................................................................................................................................... Se. longicerca

4. Two black lateral stripes of pronotal disc closely connected in the middle........................................................................................5

- Two black lateral stripes of pronotal disc disconnected in the middle ..................................................................................... Se. xiai

5. Middle area of occiput with black spots.....................................................................................................................................................6

- Middle area of occiput without black spots ..............................................................................................................................................7

6. Male eighth and ninth abdominal tergite with shallow spots................................................................................ Se. fanjingshanensis

- Male eighth and ninth abdominal tergite entirely black...................................................................................................... Se. flavistria

7. Disc of pronotum with interrupted longitudinal stripes ............................................................................................. Se. problematica

- Disc of pronotum with continuous longitudinal stripes ....................................................................................................... Se. sigillata

8. Wings macropterous, surpassing hind margin of eighth abdominal tergite .......................................................................................9

- Wings mesopterous, not surpassing hind margin of eighth abdominal tergite............................................................ Se. laoshanica

9. Lateral margins and posterior margin of pronotum yellow brown ................................................................................................... 10

- Lateral margins and posterior margin of pronotum black................................................................................................... Se. truncata

10. Internal and external margins of lateral lobes of male subgenital plate equal in length................................................................. 11

- Internal margins of lateral lobes of male subgenital plate longer than external margins......................................... Se. sichuanensis

11. Tegmina obviously surpassing apex of hind femora............................................................................................................................. 12

- Tegmina not surpassing apex of hind femora..................................................................................................................... Se. brevipennis

12. Male cerci longer than 5.0 mm.................................................................................................................................................................. 13

- Male cerci shorter than 5.0 mm...................................................................................................................................................... Se. sheni

13. Posterior margin of male subgenital plate with a U-shaped concavity, apex of lateral lobes obtusely rounded............................ ....................................................................................................................................................................................................... Se. hubeiensis

- Posterior margin of male subgenital plate with a V-shaped concavity, apex of lateral lobes obliquely truncated..... Se. unicolor

Bicornisgryllacris Li, Yin & He gen. nov. NJmaeā http://zoobank.org/ urn:lsid:zoobank.org:act:BEC7AF5FB85C-455F-AA54-232402CC5C5B

( Figs 10C, H, I , 11E, F , 12G )

Type species: Gryllacris mellii Karny 1926 = Bicornisgryllacris mellii (Karny 1926) .

Diagnosis: Bicornisgryllacris can be recognized by the combination of the following characters: forewings almost yellowish brown ( Fig. 11E, F ); ventral area of male ninth abdominal tergite on both sides with a rather large sturdy projection incurved basally and excurved apically ( Fig.10C ); posterior margin of male subgenital plate slightly concave, styli long and cylindrical ( Fig. 10C ); female seventh abdominal sternite with a pair of horns projecting ( Fig. 10H, I ). Bicornisgryllacris is similar to Sericgryllacris and Phryganogryllacris , but can be distinguished by a pair of horns projecting from female seventh abdominal sternite. Sericgryllacris also differs from Bicornisgryllacris by darkened veins and transparent cells on forewings (see Table 6 for details).

Description: Body small-sized (20–25 mm. approx.) and slender ( Fig. 11E, F ); fastigium of verticis about 1.5 times as broad as scape ( Fig. 12E, G ); anterior margin of pronotum slightly projecting in the middle, posterior margin nearly straight; lateral lobes as long as deep ( Fig. 11E, F ); macropterous, forewings almost yellowish brown ( Fig. 11E, F ); male ninth abdominal tergite neither divided nor curved in the middle, ventral area of baso-lateral surface on both sides with a rather large sturdy projection incurved basally and excurved apically ( Fig. 10C ); posterior margin of male subgenital plate slightly concave, styli long and cylindrical ( Fig. 10C ); female seventh abdominal sternite with a pair of horns projecting ( Fig. 10H, I ); female ovipositor long and nearly straight ( Fig. 11E ).

Included species: Bicornisgryllacris mellii (Karny, 1926) comb. nov., Bi. subrectis (Matsumura & Shiraki, 1908) comb. nov..

Etymology: ‘bicornis-’ is taken from ‘bi-’ and ‘cornu’, ‘bi’ meaning double and ‘cornu’ meaning horns.

Remarks: See remarks for Phryganogryllacris and Table 6 .

Key to species of Bicornisgryllacris Li, Yin & He gen. nov. from China

1. Pronotum without blackish markings ...................................................................................................................................... Bi. subrectis

- Pronotum with blackish markings ................................................................................................................................................. Bi. mellii

Figures

Figure 10. Male abdominal terminal.A, B, Ph. brevixipha; C, Bi. mellii; D, E, Se. decempunctata; F, Se. laoshanica; G, Se. sichuanensis; H, Bi. subrectis; I, Bi. mellii.(Arrow in A indicates the middle of tergite that bulges sharply caudally, arrow in B indicates the bulge extending caudally from the root on each side of the posterior margin of male abdominal tergite.).

Figure 11. Living Gryllacrididae.A, Se. decempunctata; B, Se. sichuanensis; C, Ph. brevixipha; D, Ph. sp.; E, Bi. subrectis; F, Bi. mellii.

Figure 12. Head in frontal view.A, Ho. gladiate; B, Ho. obtusitubera; C, F. chirurga; D, W. procera; E, Se. decempunctata; F, Met. permodesta; G, Bi. subrectis; H, Ph. brevixipha; I, Ni. testaceus.