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 Capnogryllacris Karny 1937 Oiaeā ( Figs 6E , 7D , 9C ) Capnogryllacris Karny 1937: 123 . Capnogryllacris Otte 2000: 8 . Capnogryllacris Gorochov 2003: 629 . Capnogryllacris Gorochov et al. 2015: 567 . Capnogryllacris Ingrisch 2018: 135 . Capnogryllacris Cadena-Castañeda 2019: 93 . Capnogryllacris Bian et al. 2021: 231 . Capnogryllacris Peng et al. 2021: 391 . Type species: Locusta ( Gryllacris ) fumigata Haan, 1843 = Capnogryllacris fumigata fumigata (Haan 1843) . Diagnosis: Capnogryllacris can be recognized by the combination of the following characters: hindwings almost fuscous ( Fig. 6E ); posterior margin of male ninth abdominal tergite with robust, straight, and conical spines extending inwards from the middle of each lobe at base, length of spines equal to or longer than half the width of ninth abdominal tergite ( Fig. 7D ). Capnogryllacris is similar to Borneogryllacris , Marthogryllacris , Dracogryllacris ,and Ultragryllacris , but can be distinguished by fuscous cells on forewings Fig. 13 , and cells overall fuscous or only centre of cells transparent on hindwings ( Fig. 6E ). Borneogryllacris also differs from Capnogryllacris by the large, black and yellow spot on basal area of forewings. Marthogryllacris and Dracogryllacris also differ from Capnogryllacris by the curved spines on male ninth abdominal tergite. Ultragryllacris also differs from Capnogryllacris by the large dorsal projection on male ninth abdominal tergite (see Table 4 for details). Included species: Capnogryllacris fumigata fumigata (Haan, 1843) , C . alivittata (Griffini,1911) , C . annulicornis (Hebard,1922) , C . buttikoferi (Karny, 1931) , C . elongata (Fritze, 1908) , C . fasciculata fasciculata (Pictet & Saussure, 1893) , C . fasciculata gonymelaena (Karny, 1928) , C . nigripennis trimaculata (Karny, 1925) , C . fumigate miniata (Karny, 1931) , C . fumigata sumatrae Gorochov,2003 , C . funebris (Brunner von Wattenwyl, 1898) , C . gigantea Karny, 1937 , C . multifracta (Griffini, 1914) , C . nigripennis nigripennis (Gerstaecker, 1860) , C . nigripennis trimaculata (Griffini, 1913) , C . obscurata Karny 1937 , C . pictipes (Karny, 1925) , C . primigenii (Griffini, 1918) , C . soror ( Brunner von Wattenwyl, 1888 ) , C . superba ( Brunner von Wattenwyl, 1888 ) . Figure 14. Tenuigryllacris huanglianensis sp. nov .. A, head in frontal view; B, pronotum in dorsal view; C, ovipositor in lateral view; D, living female individual. Remarks: The genera Borneogryllacris , Marthogryllacris , Dictyogryllacris , Erythrogryllacris , Cyanogryllacris , and Capnogryllacris were established by Karny (1937) . Gorochov (2003) suggested that Borneogryllacris and Dictyogryllacris might be synonyms of Capnogryllacris . Liu et al. (2010) considered that Dictyogryllacris should not be regarded as a subgenus under Capnogryllacris based on its enlarged median ocellus. Liu et al. (2010) classified Borneogryllacris and Capnogryllacris as subgenera under the genus Marthogryllacris , distinguished by Marthogryllacris possessing carina on each side of head, while Borneogryllacris and Capnogryllacris do not possess carina; hindwings of Borneogryllacris are dark near cross veins, while those of Capnogryllacris are transparent. Gorochov et al. (2015) regarded the genus Dictyogryllacris as a subgenus under Capnogryllacris , and considered the genera Borneogryllacris , Marthogryllacris , and Erythrogryllacris as synonyms of Capnogryllacris . Ingrisch (2018) regarded the genus Cyanogryllacris as a synonym of Capnogryllacris based on the characters of the type species ( Gryllacris grassii Griffini 1912 ) possessing black spots on the forewings. This character was consistent with Capnogryllacris borneoensis , the type species of Capnogryllacris . Cadena-Castañeda (2019) considered that the genus Dictyogryllacris should not be treated as a subgenus under Capnogryllacris , and again restored it to a separate genus with three species groups: species group borneoensis , fumigata , and rubrocellata . Liu et al. (2022a, b) studied seven species of Capnogryllacris by complete mtDNA and found that they can be divided into two clades: C . rufonotata and C . erythrocephala maculatis in one clade, and C . melanocrania , C . nigromarginata nigromarginata , C . nigromarginata hainanensis , C . nigromarginata rectispina , and C . spinosa in the other clade. Our molecular results show that the genus Capnogryllacris is not monophyletic, with U. pulchra rubricapitis nested with Capnogryllacris species. Ultragryllacris species possess a strongly modified and projecting male ninth abdominal tergite, which makes it different from Capnogryllacris . Thus, we treat Capnogryllacris species in our study as four monophyletic groups: (i) C . fumigata and Capnogryllacris sp. , (ii) C . sequestris and C . rufonotata , (iii) C . spinosa and C . melanocraina , and (iv) C . xujuni . The differences among these include: colour of pronotum, colour of fore- and hindwings, and shape of process on male ninth abdominal tergite (see Table 4 for details). The species of C . fumigata and Capnogryllacris sp. should be remained in the genus Capnogryllacris , because C . fumigata is the type species and its hindwings are overall fuscous. C . sequestris and C . rufonotata are moved to Marthogryllacris stat. resurr., because the bilateral cross veins of hindwings are deep dark. C . spinosa and C . melanocraina are moved to Dracogryllacris gen. nov. , because cells of hindwings are transparent, with only the margins of pronotum black. Capnogryllacris xujuni is moved to Borneogryllacris stat. resurr. based on large, black and yellow spots on forewings and separated spines on male ninth abdominal tergite. According to our new classification, species of Capnogryllacris are mainly distributed in Malay Archipelago, species of Marthogryllacris and Ultragryllacris are mainly distributed on the Indo-Chinese Peninsula, species of Dracogryllacris are mainly distributed in southern China . Figure 15. Male T . yingjiangensis sp. nov .. A, face; B, abdominal terminal; C, subgenital plate; D, holotype in dorsal view; E, living male individual in lateral view. Table 4. Diagnosis of the genera Capnogryllacris , Borneogryllacris , Marthogryllacris , Dracogryllacris , and Ultragryllacris .

Capnogryllacris	Borneogryllacris Karny 1937	Marthogryllacris Karny	Dracogryllacris	Ultragryllacris Gorochov
Karny 1937	1937	Li, Yin & He	and Dawwrueng, 2015
gen. nov.
Pronotum	Only margins	Anterior and posterior margin	Centre with a large	Only margins	With a pair of black spots
black, or	with a black band	V-shaped black spot	black	at anterior margin and a
without margin	black band at posterior
margin
Forewings	Veins and cells	Basal areas of forewing with a	Veins deep darkening,	Veins deep	Veins darkening, cells
overall fuscous	large, black and yellow spot,	cells yellowish trans-	darkening,	yellowish transparent
veins deep darkening, other	parent	cells yellowish
area yellowish transparent	transparent
Hind-	Veins fuscous, cells	Veins deep darkening, cells	Veins deep darkening,	Veins deep	Veins darkening, cells
wings	overall fuscous	of bilateral cross veins	cells of bilateral cross	darkening,	transparent
or only centre of	darkening, other areas	veins darkening, other	cells
transcells transparent	transparent	areas transparent	parent
Process on	A pair of robust,	A pair of slender spines basal	A pair of short and	A pair of long	A very large dorsal
promale ninth	straight, and	separated	curved spines basal	and curved	jection occupied most
abdominal	conical spines	crossed	spines basal	part of the tergite,
tergite	basal crossed	crossed	trapeziform or triangular

In Liu et al. (2022a), 11 specimens of Capnogryllacris were divided into two clades. Capnogryllacris spinosa and C . nigromarginata were found to be close, and then gathered with C . melanocrania . In the other clade, C . erythrocephala maculatis was paraphyletic with C . rufonotata . Two specimens of C . melanocrania did not clustere together. We considered that C . melanocrania ( KX 057731) was misidentified. It should belong to the genus Marthogryllacris . Thus, the study byLiu et al. (2022a) includes two clades:one clade ( C . melanocranis KX 057731, C . erythrocephala , C . rufonotata ) and another clade ( C . melanocrania , C . nigromarginata , C . spinosa ), which are Marthogryllacris and Dracogryllacris gen. nov. in our new classification.