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A new species of Meconematini is described from Tabin Wildlife Reserve in eastern Sabah: Cercoteratura repens sp. nov. This is the first report of the genus Cercoteratura in Sabah. Two species of Phisidini, Carliphisis acutipennis (Carl, 1908) and Neophisis (Indophisis) longipennis Jin, 1992, are reported in eastern Sabah for the first time. The calling song of Neophisis (Indophisis) longipennis with a near-complete ultrasonic spectrum is also described.

Systolederus cinereus Brunner von Wattenwyl, 1893 (Figs. 18–20) Systolederus cinereus Brunner von Wattenwyl, 1893: 105 (holotype —female, Myanmar, Carin Cheba [= Karen Hills]; in the Natural History Museum of Genova). Systolederus cinereus — Kirby, 1910: 20; Kirby, 1914: 44; Günther, 1939: 170; Blackith, 1992: 171; Mahmood et al., 2007: 1282; Storozhenko & Dawwrueng, 2015: 536. Systolederus ridleyi Hancock, 1909: 401 (holotype —female, Singapore, Singapore Botanical Garden; in the Oxford University Museum of Natural History, UK), syn. nov. Teredorus ridleyi — Hancock, 1915: 110. Systolederus ridleyi — Günther, 1939: 169; Blackith, 1992: 174; Devriese & Husemann, 2023: 340. Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: •5Ô and 3♀; open stream near Bạch Mã National Park, N16.22036 E107.88702, 72.9± 6.3 m.a.s.l.; 8 March 2023, 11h01; on a boulder or rock; coll. M.K. Tan & V. T. Trung; VIET. 23.31–38 (ZRC) •1Ô; along stream near Bạch Mã National Park, N16.22419 E107.88233, 57.4± 4.5 m.a.s.l.; 8 March 2023, 19h37; on a tree trunk; coll. M.K. Tan & V. T. Trung; VIET.23.50 (VNMN). Remarks. Images of the holotypes of C. cinereus and C. ridleyi are available in the internet catalogue Orthoptera Species File (Cigliano et al., 2023). The comparison of original descriptions and photos of both species, as well as all materials examined by Sergey Yu. Storozhenko from Southeast Asia, allows proposing a new synonymy: Systolederus cinereus Brunner von Wattenwyl, 1893 = Systolederus ridleyi Hancock, 1909, syn. nov. The specimens from Bạch Mã National Park represent the first record of this species in Vietnam. Ecology. This species was found along rivers, on rocks and substrate near the fast-flowing water. Nevertheless, it has also been found along forest edges near a river and dwelling among tree trunks. Distribution. Myanmar, Thailand, Malaysia, Singapore; Vietnam: Thừa Thiên Huễ Province, Bạch Mã National Park [new]. Type locality. Myanmar, Carin Cheba.

Tan, Ming Kai, Vu, Trung Thanh, Le, Cuong Viet Canh, Pham, Thai Hong (2023): Pygmy grasshoppers (Orthoptera: Tetrigidae) from Bạch Mã National Park, central Vietnam. Zootaxa 5319 (2): 199-223, DOI: 10.11646/zootaxa.5319.2.3, URL: http://dx.doi.org/10.11646/zootaxa.5319.2.3

Zhengitettix albitarsus Storozhenko, 2013 (Fig. 11) Zhengitettix albitarsus Storozhenko, 2013: 206. Zhengitettix albitarsus — Storozhenko & Dawwrueng, 2014: 325; Dawwrueng & Dooddeum, 2014: 5; Storozhenko & Dawwrueng, 2015: 548; Chen & Deng, 2022: 73 (key to species of Zhengitettix). Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: •1Ô; open stream near Bạch Mã National Park, N16.22036 E107.88702, 72.9± 6.3 m.a.s.l.; 8 March 2023, 11h01; on a boulder or rock; coll. M.K. Tan & V. T. Trung; VIET.23.30 (ZRC) (Fig. 11). Remarks. Previously described and known only from Thailand, the male specimen from Bạch Mã National Park represents the first record of this species in Vietnam (Fig. 11). Ecology. This species was found along rivers, on rocks and substrate near the fast-flowing water (Fig. 3). Upon disturbance, this species would fly off. It is likely to be limno-terrestrial, being associated with open stream. Distribution. Thailand: Nakhon Ratchasima Province, Mae Hong Son Province; Vietnam: Thừa Thiên Huễ Province [new]. Type locality. Thailand, Nakhon Ratchasima Province, Khao Yai.

Epitettix parallelus (Podgornaya, 1986) (Fig. 12) Vaotettix parallelus Podgornaya, 1986: 17. Vaotettix parallelus — Blackith, 1992: 196; Kim & Pham, 2014: 59. Epitettix parallelus — Storozhenko, 2021: 182. Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: •1Ô; Rhododendron Trail, heading to Four Lakes, N16.18950 E107.85360, 1160.0± 7.7 m.a.s.l.; 9 March 2023, 11h02; among leaf litter; coll. M.K. Tan & V. T. Trung; VIET.23.63 (ZRC) (Fig. 12). Remarks. This species is widely distributed in Vietnam. The specimen from Bạch Mã National Park, nevertheless, differs by the emarginated apex of posterior process of the pronotum (in typical form apex of process rounded) (Fig. 12). This is likely intraspecific variability. Ecology. The leaf litter area at which the specimen was collected was not too far from a forest stream, but the area is clearly elevated and probably not part of the water table. Distribution. So far, known only from central Vietnam in the provinces of Gia Lai and Thừa Thiên Huễ [new]. Type locality. Vietnam, Gia Lai Province, Buon Luoi.

Bolivaritettix sculptus (Bolívar, 1887) (Fig. 14) Mazarredia sculpta Bolívar, 1887: 237. Mazarredia sculpta — Hancock, 1907b: 32; Kirby, 1914: 51; Hancock, 1915: 100; Bolívar, 1917: 285. Bolivaritettix sculptus — Günther, 1939: 59; Shishodia, 1991: 82; Blackith, 1992: 12; Paris, 1994: 249; Deng et al., 2009: 282; Deng et al., 2010: 48; Storozhenko & Dawwrueng, 2015: 533; Ding et al., 2018: 311; Storozhenko, 2018a: 18; Bhaskar et al., 2022: 422. Mazarredia indotata Bolívar, 1909: 398; synonymized by Günther, 1939: 59. Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: • 1♀; Summit Trail (forest trail), N16.19735 E107.86131, 1401.0±6.0 m.a.s.l.; 7 March 2023, 11h42; on clayey rock; coll. M.K. Tan & V. T. Trung; VIET.23.16 (ZRC) (Fig. 14). Ecology. The colouration of this species allows it to camouflage among orange clayey rock where it was found. Distribution. India, China, Myanmar, Cambodia, Thailand, Vietnam: provinces Gia Lai, Vĩnh Phúc, Bình Phýớc, Đắk Lắk, and Thừa Thiên Huễ [new]. Type locality. Vietnam, Gia Lai Province, Buon Luoi.

Hebarditettix magnus Storozhenko, 2017 (Figs. 6, 7) Hebarditettix magnus Storozhenko, 2017: 121. Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: • 1♀; Summit Trail (forest trail), N16.19802 E107.85991, 1457.0± 5.9 m.a.s.l.; 7 March 2023, 21h12; on a vertical surface; coll. M.K. Tan & V. T. Trung; VIET.23.26 (ZRC) •1Ô; Summit Trail (forest trail), N16.19829 E07.85970, 1447.0±6.0 m.a.s.l.; 7 March 2023, 21h20; on a tree trunk; coll. M.K. Tan & V. T. Trung; VIET.23.27 (VNMN) (Fig. 7). Remarks. This species was first described from Bạch Mã National Park. This represents the second record of this species. We provide a live image of this species (Fig. 6). Ecology. This species was found among shaded forested habitat (Fig. 2), often dwelling among tree trunks or rocky surfaces (covered with some mosses) (Fig. 6). It seems that this species tends to be found at higher altitudes (see Storozhenko, 2017). Distribution. So far, known only from central Vietnam in the provinces of Thừa Thiên Huễ and Gia Lai. Type locality. Vietnam, Th ừa Thiên Hu ễ Province, B ạch Mã National Park.

Criotettix bispinosus (Dalman, 1818) (Figs. 4, 5) Acrydium bispinosum Dalman, 1818: 77. Acridium (Tetrix) bispinosum — Haan, 1843: 169. Criotettix bispinosus [refer to OSF for complete literature]— Bolívar, 1887: 226; Günther, 1937a: 179; Tinkham, 1937: 230; Günther, 1937b: 121–122; Günther, 1939: 319; Shishodia, 1991: 36; Blackith, 1992: 34; Mahmood et al., 2007: 1281; Benediktov. 2013: 257 (vibrational signalling); Tumbrinck, 2015: 276; Deng et al., 2023: 208. Acanthalobus bispinosus — Hancock, 1907b: 29; Kirby 1910: 16; Kirby, 1914: 37; Karny, 1915: 78; Hancock, 1915: 93; Willemse, 1930: 18. Criotettix brachynotus Zheng & Jiang, 1994: 34; synonymized by Deng, 2021: 455. Formosatettix hainanensis Zheng, 2012: 2; synonymized by Deng et al., 2023: 2008. Tettix latispina Walker, 1871: 837; synonymized by Kirby, 1910: 18. Acanthalobus longinotus Hancock, 1907a: 221; synonymized by Blackith, 1992: 35. Tettix simplex Walker, 1871: 836; synonymized by Kirby, 1910: 18. Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: •1Ô, 1♀; road-side vegetation along road to summit, N16.24626 E107.86881, 85.9± 6.1 m.a.sl., 7 March 2023, 9h16; on a leaf of a shrub; coll. M.K. Tan & V. T. Trung; VIET. 23.12, 13 (ZRC) • 1♀; road-side vegetation along road to summit, N16.24198 E107.86661, 144.7± 6.9 m.a.s.l.; 7 March 2023, 10h09; on a rock with flowing water; coll. M.K. Tan & V. T. Trung; VIET.23.15 (VNMN) • 3Ô; open stream near Bạch Mã National Park, N16.22036 E107.88702, 72.9± 6.3 m.a.s.l.; 8 March 2023, 11h01; on boulders and rocks; coll. M.K. Tan & V. T. Trung; VIET.23.39–41 (ZRC) (Fig. 5). Ecology. This species was found along rivers, residing on rocks and substrate near the fast-flowing water. However, it has also been found on more vegetated area along road. Distribution. India, Myanmar, South China (including Hainan), Thailand, Vietnam, Peninsular Malaysia, and Indonesia (the Greater Sunda Islands). Type locality. Oriental region (without definite locality).

Bolivaritettix chinensis (Hancock, 1912) (Fig. 13) Mazarredia chinensis Hancock, 1912: 140. Bolivaritettix chinensis — Günther, 1939: 59; Günther, 1941: 149; Kim & Pham, 2014: 59; Storozhenko, 2018a: 504. Material examined. VIETNAM, Thừa Thiên Huễ province, Bạch Mã National Park: • 1♀; 22–23 September 2008, coll. V. G. Bezborodov (Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia). Remarks. This species was reported in Bạch Mã National Park (Storozhenko, 2018a). Distribution. Vietnam: provinces Cao Bằng, Hòa Bình, Thừa Thiên Huễ, Lang Son, Thái Nguyên, Vĩnh Phúc. Type locality. Tonkin, Than Moi (northern Vietnam).

Tegotettix bufocrocodil (Storozhenko & Dawwrueng, 2015) (Figs. 8–10) Gavialidium bufocrocodil Storozhenko & Dawwrueng, 2015: 540. Gavialidium bufocrocodil — Storozhenko, 2018b: 19. Tegotettix bufocrocodil — Muhammad et al., 2018: 31. Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: •1Ô, 4♀, 3 nymphs; along stream near Bạch Mã National Park, N16.22419 E107.88233, 57.4± 4.5 m.a.s.l.; 8 March 2023, 19h37; on a tree trunk; coll. M.K. Tan & V. T. Trung; VIET. 23.42–49 (ZRC) •1Ô; forest near Bạch Mã National Park, N16.22069 E107.88253, 74.9± 6.5 m.a.s.l.; 9 March 2023, 21h11; on a tree trunk; coll. M.K. Tan & V. T. Trung; VIET.23.73 (VNMN). Remarks. This represents the first record of this species in Vietnam. Ecology. This species was found along forest edge not too far from a river and dwelling among tree trunks, often staying still and camouflaging among the crevices (Figs. 8–10). Adults and nymphs can be found aggregating together on a few trees. Distribution. Thailand: Nakhon Ratchasima and Trat Provinces; Cambodia: Sen Monorom; Vietnam: Thừa Thiên Huễ Province [new]. Type locality. Thailand, Nakhon Ratchasima Province, Khao Yai.

Rhopalina bachma Tan & Storozhenko, sp. nov. (Fig. 17) Material examined. VIETNAM, Th ừa Thiên Hu ễ Province, B ạch Mã National Park: • ♀ holotype; Rhododendron Trail, heading to Four Lakes, N16.18950 E107.85360, 1160.0± 7.7 m.a.s.l.; 9 March 2023, 11h02; among leaf litter; coll. M.K. Tan & V. T. Trung; VIET.23.62 (VNMN) (Fig. 17). Remarks. This represents the second species from this little-known genus. The previously monotypic genus consisted of Rhopalina javana Tinkham, 1939 from Java. Here, a second species from this genus was discovered in Indochina (quite far away from Java) and is described. The fastigium of the holotype has its right margin rounded, but its left margin emarginated in the middle. It is unclear if this asymmetry is due to defects (e.g., damaged during individual development), injuries or genuine character. Diagnosis. The new species is similar to Rhopalina javana in the general habitus, colouration and the shape of fastigium but differs from latter by the antennal groove inserted at the lower margins of the eyes, the tegmen stouter, 2.4–2.5 times as long as wide, and the pronotum well-surpassing apices of ovipositor (in R. javana, the antennal groove instead of above lower margins of the eyes, the tegmen 2.6–2.7 times as long as wide, and the pronotum slightly surpassing apices of ovipositor). Etymology. The species is named after the type locality, Bạch Mã National Park; noun in apposition. Description. Habitus as shown in Figs. 17A, 17B. Different shades of brown, ranging from pale brown, dark brown to red brown, well camouflaged against leaf litter on the ground. Head: In frontal view (Fig. 17C): antennal groove inserted at lower margin of eyes. Fastigium produced into obtuse apex. Face with numerous small nodules and large oval nodules above clypeal triangle; with robust lateral carinae running from ventrad of eyes to labrum, slightly bent in middle. Frontal costa elongated, bifurcation of frontal costa in line with slightly dorsal of middle of eye. Facial carinae straight and slightly diverging ventrally. Compound eyes hemispheric, not exerted above vertex, 1.41 times taller than wide. Lateral ocelli slightly ventral of bifurcation of frontal costa and slightly above antennal groove in line with middle of eye. In dorsal view (Fig. 17A): apex of fastigium surpasses frontal margin of eyes; right margin of fastigium rounded, left margin emarginated in the middle. Vertex 1.47 times wider than eye width. Lateral carinae of fastigium not distinct; median carina more distinct, reaching apex of fastigium. In lateral view (Fig. 17B): frontal costa well protruding in front of eyes into obtuse apex. Mouthparts with labrum pale coloured. Maxillary palps pale coloured with tint of grey on anterior (or dorsal parts), elongated; with apical (fifth) segment more oval, longest following by third segment and then subapical (fourth) segment. Pronotum: Pronotum surpassing posterior femur, nearly reaching middle of posterior tibia, 3.26 times longer than wide (pronotal lateral lobe width). In lateral view (Fig. 17B): with median, humero-apical and interhumeral carina weakly produced. Infrascapular area broad. Angle of ventral sinus between lateral shoulder and lateral lobe nearly right-angled. In dorsal view (Fig. 17A): median carina distinct throughout length of pronotum (except very close to the posterior end). Lateral carinae distinct. Interhumeral carina distinct, diverging posteriorly. Lateral lobe of pronotum slightly produced, and broadly triangular; posterior margin of lateral lobe acute. Apical end of pronotal disc narrow and truncated. Legs: Fore and middle legs (Fig. 17A): femora generally marmoration of pale and dark spots; tibiae with dark and light bands. Hind legs (Fig. 17B): hind femur about 4.29 times longer than wide, about 1.09 times longer than hind tibia; with clear carinae along dorsal margin; marmoration of pale and dark spots and ventrad of ventroexternal carina dark coloured. Knee of posterior femur with antegenicular and genicular teeth acute. Hind tibia dark coloured with shades of pale colouration; with six spines on each lateral margin, with numerous spinules between these large dorsal spines. Basal article of hind tarsus generally pale coloured, 1.38 times longer than apical article; middle article dark coloured very short; apical article white, except apex being slightly darker in colouration. Abdomen: Tergites and sternites typically dark coloured with pale-coloured nodules. Ovipositor (Fig. 17D) with pale-coloured spinules with apex red brown; dorsal and ventral valves with sparse hairs along dorsal and ventral margin. Dorsal valve with eight dorsal spines, apex acute; ventral valve with six ventral spines typically smaller than dorsal spines; apical ventral spine of ventral valve more hooked at apex than apical dorsal spine of dorsal valve; spines on valves increasing larger and robust apically. Apices of ovipositor valves red brown. Subgenital plate short in lateral view (Fig. 17D), truncated with medial lobule in ventral view (Fig. 17E). Measurements (in mm). PL = 10.34; PW = 3.18; PH = 1.73; HWL = 8.13; HFL = 5.10; HFW = 1.19; TL = 1.20; VW = 0.66; EW = 0.45; SW = 0.19; IAH = 0.22; BL = 8.16; PAW = 1.45; PPW = 1.99; MAL = 0.36; MAW = 0.05; 1 stAW = 0.20; TW = 0.55; FFL = 1.92; FFW = 0.53; MFL = 2.09; MFW = 0.54; HTL = 4.67; 1 st TaL = 0.87; 3 rd TaL = 0.63; SGPL = 0.67; SGPW = 0.54; OVDL = 1.02; OVDW = 0.31; OVVL = 1.01; OVVW = 0.25 Ecology. The leaf litter area at which the specimen was collected was not too far from a forest stream, but the area is clearly elevated and probably not part of the water table. Distribution. So far known only from type locality. Type locality. Vietnam, Th ừa Thiên Hu ễ Province, B ạch Mã National Park.

Miriatroides luna Tan & Storozhenko, sp. nov. (Figs. 15, 16) Material examined. VIETNAM, Th ừa Thiên Hu ễ province, Bạch Mã National Park: •Ô holotype; Summit Trail (forest trail), N16.19667 E107.86169, 1428.0± 5.8 m.a.s.l.; 7 March 2023, 20h15; hiding among rotten tree trunk on the ground; coll. M.K. Tan & V. T. Trung; VIET.23.17 (VNMN) (Figs. 15, 16). Diagnosis. This new species is most similar to Miriatroides kannackiensis Storozhenko, 2016 from Gia Lai Province, but differs from latter (as well as from all other congeners) by the rostrum more projected in lateral view and the fore femur with margins (especially ventral margin) undulated. Etymology. This species is named after the moon (in Latin, luna = moon), as the holotype was collected near the summit on a clear night with a full moon. Description. Habitus as shown in Figs. 15, 16A, 16B. Different shades of brown, ranging from pale brown, and dark brown to red brown, well camouflaged against leaf litter and tree trunk on the ground. Head: In frontal view (Fig. 16C): antennal groove inserted below lower margin of eyes. Fastigium convex; sinuosoidal on anterior border. Face with numerous nodules. Frontal costa elongated. Bifurcation of frontal costa in line with middle of eye. Facial carinae faintly sinusoidal. Compound eyes hemispheric, not exerted above vertex, 1.55 times taller than wide. Lateral ocelli slightly ventral of bifurcation of frontal costa and slightly above antennal groove. In dorsal view (Fig. 16A): apex of fastigium clearly surpasses frontal margin of eyes, emarginated in the middle. Vertex 1.53 times wider than eye width. Lateral carinae of fastigium not distinct; median carina well surpassing apex of fastigium. In lateral view: frontal costa arched and well protruding in front of eyes. Mouthparts mostly brown to grey with small pale-coloured nodules. Maxillary palps pale coloured with tint of grey; with segments stout and apical segment somewhat oval. Pronotum: Pronotum reaching apex of posterior knee, 2.03 times longer than wide (pronotal lateral lobe width). In lateral view (Fig. 16B): with median, humero-apical, and interhumeral carinae irregularly undulated. Infrascapular area broad. Angle of ventral sinus between lateral shoulder and lateral lobe acute. In dorsal view (Fig. 16A): median carina distinct throughout length of pronotum, flanked by a network of irregular carinae. Lateral carinae distinct. Interhumeral carina distinct, diverging posteriorly. In dorsal view (Fig. 16A), lower margin of lateral lobe of pronotum leaf-shaped. Apical end of pronotal disc acute. Legs (Figs. 16A, 16B): Fore and middle legs: generally brown with either dark- or light-coloured bands. Fore femur with distinct dorsal carina straight, with ventral carina undulated, forming three nodules. Middle femur with less distinct dorsal and ventral carinae, clearly not undulating; lateral external carina undulated, forming two or three small nodules. Hind legs: posterior femur about 3.36 times longer than wide, about 1.34 times longer than hind tibia; with clear carinae along dorsal margin. Knee of hind femur with antegenicular tooth small but acute, genicular tooth rounded. Posterior tibia dark coloured anteriorly, becoming slightly lighter in colouration; with five spines on each lateral margin, with numerous spinules between these large dorsal spines. Basal article of hind tarsus generally pale coloured, 0.78 times as long as apical article; middle article dark coloured and very short; apical article white, except apex being black. Abdomen: Tergites and sternites typically dark coloured with some sparse patches of pale spots. Subgenital plate triangular (Figs. 16D, 16E). Measurements (in mm). PL = 7.17; PW = 3.54; PH = 1.20; HWL = NA; HFL = 4.53; HFW = 1.35; TL = NA; VW = 0.55; EW = 0.36; SW = 0.14; IAH = 0.52; BL = 7.34; PAW = 1.47; PPW = 2.16; MAL =?; MAW =?; 1 stAW = 0.18; TW = NA; FFL = 1.70; FFW = 0.55; MFL = 1.93; MFW = 0.44; HTL = 3.37; 1 st TaL = 0.49; 3 rd TaL = 0.63; SGPL = 0.56; SGPW = 0.59. Ecology. Only one specimen was encountered staying still and camouflaging itself in a decaying wood on the ground of a sheltered montane forest habitat at high altitude of Bạch Mã National Park (Fig. 3). Upon disturbance, it remained still. Distribution. So far known only from type locality. Type locality. Vietnam, Th ừa Thiên Hu ễ Province, B ạch Mã National Park.

Nisitrus danum Robillard & Tan, 2021 (Figs 1–8) Nisitrus danum Robillard & Tan, 2021, in Tan et al. 2021a: 31 Material examined MALAYSIA — Sabah • 1J; Kawag Forest Reserve, near rest house; N5.0502 E117.98286, 134.7 m.a.s.l. (GPS SAB22_6); 14.v.2022; afternoon; on plant; call recording MNHN-SO-2023-1478; M.K. Tan & T. Robillard; MNHN-EO-ENSIF11344 • 1♀; Kawag Forest Reserve, near rest house; N5.0502 E117.98286, 134.7 m.a.s.l. (GPS SAB22_ 6); 14.v.2022; afternoon; on plant; molecular sample N67; M.K. Tan & T. Robillard; MNHN-EO-ENSIF11345 • 1♀ (photographed, not collected);Tabin Wildlife Reserve,Lipad mud volcano; N5.20967 E118.50732, 168.9m.a.s.l. (GPS SAB22_11); 16.v.2022; day; T. Robillard observation • 2 juveniles; Tabin Wildlife Reserve; N5.19534 E118.50365, 119.7 m.a.s.l. (GPS SAB22_9); 15.v.2022, night; molecular samples N65–N66; MNHN-EO-ENSIF10978– ENSIF10979 • 1 juvenile (not collected); Sepilok, Rainforest Discovery Centre; N5.87428 E117.93970, 1.x.2019 (Fig. 3B) • 1 juvenile; Lahad Datu, N4.962598 E117.801956; 19.viii.2022; twan3253 observation; https://www. inaturalist.org/observations/133043313 • 1♀; Lahad Datu, N4.964485 E117.803471; twan3253 observation; https:// www.inaturalist.org/observations/133303271 • 1♀; Lahad Datu, N4.965797 E117.800246; twan3253 observation; https://www.inaturalist.org/observations/133431471 Additional description Habitus as shown in Fig. 1A. Face, lateral lobes of pronotum and wide lateral band on FWs vivid yellow in living specimens (Figs 1A, 1B). Antennae with 2 main white rings, including 3–6 antennomeres and 2–3 thinner rings. Male. Metanotal gland well developed, as shown in Fig. 1C. FWs (Figs 1D, 1E) dorsal field anterior part black (cells and veins); harp and mirror area infuriated, with black veins; apical field with black cells and yellow veins. Harp with two straight oblique veins. Mirror large, not rounded, separated by one transverse vein. Vein c1 wide. Lateral field with a wide vivid yellow band including veins M, R and Sc, and bases of veins bifurcating from Sc; ventral part of lateral field shiny black, as in females. Male genitalia (Fig. 2) typical of genus, including ectophallic fold, endophallic sclerite, ectophallic arc and rami, but with pseudepiphallic lophi triangular and relatively stout in dorsal and profile views, narrowly spaced between pseudepiphallic lophi; pseudepiphallic parameres strongly bent at interno-anterior end. Juvenile (Fig. 3). Colouration pale yellow or white, characterised by numerous dark stripes all around body in early instars; antennae with 2 wide whitish rings as in adults. Measurements (in mm, means in brackets, N = 5 males). BL = 13.9–15.3 (14.6), PronL = 1.8–2.1 (1.9), PronW = 2.7–3.1 (2.9), FWL = 10.1–11.0 (10.4), FWW = 3.8–4.1 (3.9), HWT = 4.5–7.6 (6.6), FIIIL = 13.4–15.5 (14.5), FIIIW = 2.5–2.9 (2.6), TIIIL = 13.5–15.5 (14.4), Stridulatory teeth = 94–95. Comparison with N. vittatus Larger, with dark shiny lateral field of FWs in both sexes, below a vivid yellow band; face yellow with dark brown to black mouthparts (entirely yellow with a few dark spots in N. vittatus); adults and juveniles (Fig. 3) are recognisable by the larger white rings on antennae (2 main rings, with 3–6 white antennomeres, while antennae are almost entirely black with in N. vittatus, with at best thin rings). Comparison with N. musicus Similar in size, but differing by dark shiny lateral field of FWs in both sexes, below a vivid yellow band; face vivid yellow (whitish in N. musicus). Ecology Found in syntopy with Nisitrus vittatus, the latter species being much more abundant, and has been observed to be outnumbered by at least 10 to 1. The males and females were found inter-mixing with N. vittatus on large foliage of Leea indica along roads (forest edge) (Fig. 4). Similar to N. vittatus, this species tends to occur in open areas (Fig. 5). Distribution (Fig. 6) Borneo (Sabah: Danum Valley, Tawau Hill Park, Kawag Forest Reserve [new locality record], Sepilok Kabili Forest Reserve [new locality record], Tabin Wildlife Reserve [new locality record]) Calling song (9♁, 23°C, in captivity) The calling song consists of long homogeneous bouts (= trill) which can last between 3 s to more than 16 min (Fig. 7). The trill is made up of a sequence of short echemes. Each echeme is made up of two syllables and has an average duration of 41.3±2.1 ms (36.6–45.9 ms). The average interval between consecutive echemes is 50.1±5.2 ms (42.6–62.1 ms). On average, the first syllable has a slightly shorter duration of 15.2±0.8 ms (13.2–17.3 ms) than the second syllable with a duration of 16.0±1.2 ms (12.6–18.7 ms). The average interval between the two syllables is 10.0±1.0 ms (7.9–12.3 ms). The call spectrum consists of a series of harmonics with three distinct peaks in the first three harmonics: first harmonic (fundamental frequency) at 6.13±0.14 kHz (5.72–6.47 kHz), which is typically also the dominant frequency; second harmonic at 12.21±0.31 kHz (11.34–12.84 kHz); third harmonic at 18.32±0.43 kHz (16.97–19.22 kHz). While second and third harmonics are not dominant, their amplitude is high and sometimes almost co-dominate with the fundamental frequency. The calling song of N. danum is very distinct from that of the syntopic species, Nisitrus vittatus (see Tan et al., 2021a), by the echeme structure being made up of two syllables, rather than three syllables. The echeme duration is also distinctly longer in N. danum than in N. vittatus (average duration = 41.3 ms vs. 32.5 ms). Consequently, the syllable durations are also longer in N. danum than in N. vittatus. The interval between consecutive echemes is also distinctly longer in N. danum than in N. vittatus (average duration = 50.1 ms vs. 22.7 ms). Although the frequencies of N. danum is lower than that of N. vittatus, it could be an artefact of the lower ambient temperature when N. danum was recorded (23°C for N. danum vs. 27°C for N. vittatus). Despite morphological similarities between N. danum and N. musicus (also from Sabah, but specifically from Mount Kinabalu), there are also differences in their calling song. Both species have similar echeme structure made up of two syllables, but the echeme duration is distinctly longer in N. danum than in N. muscius (average duration = 41.3 ms vs. 29.7 ms). While the durations of the first and second syllables in each echeme are somewhat similar in N. danum (15.2 ms and 16.0 ms), those in N. musicus are drastically different (7.5 ms and 18.7 ms). We found that N. danum is also a diurnal species, similar to other Nisitrus (Fig. 8). The circadian rhythm of the calling activity was as follows: calling began at 6:00h in the morning and its activity increases to a maximum at 8h in the morning (amount of sound produced per hour = ca. 1683 s). This follows by a drop in calling activity. A second smaller peak in calling activity (amount of sound produced per hour = ca. 915 s) was detected in the afternoon around 16:00h. Most calling activity ceases at 19:00h in the evening. The two diurnal peaks were also observed in N. vittatus (see Tan & Robillard, 2021b) but they differ in that the highest peak for N. danum was in the morning whereas that for N. vittatus was in the afternoon., Published as part of Robillard, Tony, Tan, Ming Kai, Japir, Razy & Chung, Arthur Y. C., 2023, Notes on the Eneopterinae (Orthoptera, Grylloidea, Gryllidae) from eastern Sabah, pp. 231-250 in Zootaxa 5315 (3) on pages 233-240, DOI: 10.11646/zootaxa.5315.3.2, http://zenodo.org/record/8130509, {"references":["Tan, M. K. & Robillard, T. (2021 b) Highly diversified circadian rhythms in the calling activity of eneopterine crickets (Orthoptera: Grylloidea: Gryllidae) from Southeast Asia. Bioacoustics, 31 (4), 470 - 489. https: // doi. org / 10.1080 / 09524622.2021.1973562"]}

Genus Nisitrus Saussure, 1878 Type species: Nisitrus marginatus Walker, 1869 – According to the ICZN, and contrary to what was mentioned in the recent revision of the genus (Tan et al., 2021a), the type species should remain N. marginatus even though this name is a junior synonym of N. vittatus Haan., Published as part of Robillard, Tony, Tan, Ming Kai, Japir, Razy & Chung, Arthur Y. C., 2023, Notes on the Eneopterinae (Orthoptera, Grylloidea, Gryllidae) from eastern Sabah, pp. 231-250 in Zootaxa 5315 (3) on page 233, DOI: 10.11646/zootaxa.5315.3.2, http://zenodo.org/record/8130509, {"references":["Saussure, H. (1878) Melanges orthopterologiques. VI. Fascicule Gryllides. Memoires de la Societe de Physique et d'Histoire naturelle de Geneve, 25 (2), 369 - 704 (505 - 834)."]}

Robillard, Tony, Tan, Ming Kai, Japir, Razy, Chung, Arthur Y.C. (2023): Notes on the Eneopterinae (Orthoptera, Grylloidea, Gryllidae) from eastern Sabah. Zootaxa 5315 (3): 231-250, DOI: 10.11646/zootaxa.5315.3.2, URL: http://dx.doi.org/10.11646/zootaxa.5315.3.2

Pendleburyella testacea Chopard, 1969 (Figs. 3, 7A, 8A) Pendleburyella testacea Chopard 1969: 218 Diagnosis (new). This species is characterised by a medium-sized habitus, the maxillary palps with the apical segment triangular, the mirror longer than broad, the dividing vein of the mirror roundly curved at the basal end, the apical part of FW longer. Material examined. Holotype: MALAYSIA • 1♁; Selangor, Bukit Kutu, 3300–3500 ft; 24 September 1932; coll. H.M. Pendlebury; 0019 (MP). Allotype: EAST MALAYSIA • 1♀; Sabah, Sandakan [N. Borneo, Bettotan, NR. Sandakan]; 7 August 1927; 0020 (MP). Remarks. The male genitalia of the holotype was not dissected and examined to minimise the risk of further damage to the old specimen. The female allotype from Borneo probably belongs a different species. Distribution. Malay Peninsula (Selangor). Type locality. MALAYSIA, Selangor, Bukit Kutu., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on page 138, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034, {"references":["Chopard, L. (1969) Grylloidea. In: Sewell, R. B. S. (Ed.), The fauna of India and the adjacent countries. Orthoptera. Vol. 2. Zoological Survey of India and Baptist Mission Press, Calcutta, pp. 1 - 421."]}

Pendleburyella vicina Chopard, 1969 (Figs. 7B, 8B, 10) Pendleburyella vicina Chopard 1969: 219 Diagnosis (new). This species is characterised by a small-sized habitus, the maxillary palps with the apical segment somewhat cylindrical to tear-shaped, the mirror about as long as broad, the dividing vein of the mirror broadly and roundly curved at the basal end, the apical part of FW shorter. Material examined. Holotype: MALAYSIA • 1♁; Kuala Lumpur, at light; 13 September 1932; coll. H.M. Pendlebury; 0018 (MP). Remarks. The type specimen is mouldy and badly damaged in the abdomen area. Distribution. Malay Peninsula (Selangor). Type locality. MALAYSIA, Selangor, Kuala Lumpur., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on page 138, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034, {"references":["Chopard, L. (1969) Grylloidea. In: Sewell, R. B. S. (Ed.), The fauna of India and the adjacent countries. Orthoptera. Vol. 2. Zoological Survey of India and Baptist Mission Press, Calcutta, pp. 1 - 421."]}

Pendleburyella sp. 2 (Figs. 7F, 8G, 14) Material examined. MALAYSIA • 1♁ (type); Selangor –Pahang, The Gap, 2700 ft, at light; 21 January 1939; coll. H.M. Pendlebury; (MP). Remarks. This old specimen from Selangor – Pahang shares the greatest resemblance with Pendleburyella eirmosa sp. nov., including the shapes of the apical and subapical segments of the maxillary palps, FWL/FWW, the shape of the dividing vein of mirror at the basal end and the extent of apical area of the FW. But there are also some morphological differences: smaller size, mirror length as long as broad (instead of longer than broad). As the two specimens were collected from two different parts of Southeast Asia and at different altitudes (and hence different habitats), we speculate that they belong to two different species. However, we also avoid describing and naming this species as the male genitalia is not available for the verification of the species status., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on pages 145-146, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034

Pendleburyella pimela sp. nov. (Figs. 4, 5A, 6A, 6C, 6E, 7C, 8C, 9A–C, 11) Diagnosis. This species is characterised by a large-sized habitus, the maxillary palps with the apical segment also triangular (but more stout and broadly widened than Pendleburyella testacea); the FWs relatively broad (FWL/ FWW 2.5 in other species specimens); the mirror about as long as broad, the dividing vein of the mirror broadly and roundly curved at the basal end, the apical part of the FW shorter relative to the FWL (all of these similar to Pendleburyella vicina but this species is distinctly larger in size). The genitalia differs from that of Pendleburyella eirmosa sp. nov. by the pseudepiphallic lophi, in profile, cylindrical with roundly truncated apices (instead of dorso-ventrally flattened), the pseudepiphallic paramere ventral processes more elongated and in ventral view having acute apices (instead of obtuse apices); and the emargination between the dorsal and ventral processes pseudepiphallic paramere, in profile, narrowly angular (instead of broadly rounded). Etymology. The species name refers to the large habitus and relatively broad FWs (low FWL/FWW); pimele = fat in Greek. Material examined. Holotype: SINGAPORE • 1♁; Nee Soon freshwater swamp forest, along Woodcutter Trail; on leaf litter; 21 December 2022; coll. M.K. Tan & I. Neo; call recorded; (ZRC). Description. Very large species for this genus. Maxillary palpi with apical segment longest and expanded (triangular); with subapical segment cylindrical and expanded slightly distally, somewhat similar length as apical and third segments (Fig. 7C). Pronotal disc brown about 2.0 times as wide as long, widening posteriorly (posterior margin about 1.6 times as wide as anterior margin); anterior margin of disc broadly concave; posterior margin of disc slightly convex (Fig. 6A). Pronotal lateral lobe 2.3 times as long as high (Fig, 7C). Male. FW 2.4 times as long as broad (Fig. 8C), covering abdomen and slightly surpassing apex of FIII. Venation (Fig. 8C): 1A vein transverse, faintly curved; diagonal substraight, with 3 distinct oblique veins in harp area; posterior two oblique veins substraight,join at base near1A,anterior most vein distinctly shorter and running nearly perpendicular to 1A. Mirror about 1.1 times as long as wide, dividing vein broadly curved at basal end. Lateral field around 10 branches on Sc (Fig. 6E). Apical field 0.09 times as long as FWL (Fig. 8C). Hind wings clearly surpassing FWs. ♁ genitalia (Figs. 9A–C). Pseudepiphallus [epiphallus] typical of genus. Posterior end of pseudepiphallus [epiphallus] produced into two tongue-shaped lobular pseudepiphallic lophi [posterolateral epiphallic lobe] with obtuse apices. These lophi, in profile, cylindrical with roundly truncated apices. Pseudepiphallic paramere [ectoparamere] very elongated with ventral process well surpassing pseudepiphallic lophi; all processes with apex acute. Dorsal process of pseudepiphallic paramere distinctly shorter than ventral process barely surpassing pseudepiphallic lophi. Ventral process of pseudepiphallic paramere distinctly more sclerotized along inner margin. In profile, emargination between dorsal and ventral processes narrowly angular. Ectophallic fold [rachis] typical of genus; in profile tapering into acute apex, with anterior margin concave and posterior margin straight. Measurements (♂, in mm). BL = 9.5; BWL = 13.3; HL = 1.1; PronL = 1.4; PronW = 2.8; FWL = 7.9; FWW = 3.3; HWT = 3.3; FIIIL = 5.4; TIIIL = 3.9; TaIIIL = 1.5. Ecology. The species was found dwelling among thick leaf litter in freshwater swamp forest. Distribution. Malay Peninsula (SINGAPORE). Type locality. SINGAPORE Calling song (1♂, in captivity, 29.0°C) (Fig. 11). The male was observed to call between 22h00 and 23h30, each night having 2–4 bouts of continuous trill. The calling song is a loud continuous trill, typically lasting around 2–5 min when left undisturbed. The trill is made up of closely-packed syllables with an average duration of 9.7±1.5 ms (5.9–14.2 ms). The average syllable period is 14.5±0.8 ms (10.8–17.1 ms). There is no distinct downtime between syllables. The frequency spectrum is pure-tonal and forms a harmonic, with the energy peaking at the fundamental frequency. The dominant frequency is 8.14±0.08 kHz (7.78–8.34 kHz)., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on pages 138-142, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034

Pendleburyella eirmosa sp. nov. (Figs. 1, 5B, 6B, 6D, 6F, 7D, 8D, 9D–F) Diagnosis. This species is characterised by a medium-sized habitus (similar to Pendleburyella testacea), the maxillary palps with the apical segment also somewhat rectangular (similar to that of Pendleburyella vicina but stouter); the mirror longer than broad, the dividing vein of the mirror broadly and roundly curved at the basal end; the apical part of FW longer relative to the FWL. The genitalia also differs from that of Pendleburyella pimela sp. nov. (see the diagnosis for the latter species). Etymology. The species name refers to the broadly rounded emargination between the dorsal and ventral processes of the pseudepiphallic paramere (distinguished from the narrowly angular emargination in Pendleburyella pimela sp. nov.); eirmós = continuity in Ancient Greek. Material examined. Holotype: BRUNEI DARUSSALAM • 1♁; Belait District, Wasai Wong Kadir Recreational Park; N4.34186, E114.44611, 35.8± 5.2 m.a.s.l.; 1 March 2019, 18h54; calling on a leaf; coll. M.K. Tan & H. Yeo; BRU.19.47 (UBDM) Photograph examined. BRUNEI DARUSSALAM • 1♁; Temburong District, Kuala Belalong Field Studies Centre, along Ashton Trail; N4.54619, E115.15690, 110.8± 6.9 m.a.s.l.; 26 July2017, dusk; on leaf litter; photographed M.K. Tan (Fig. 1). Description. Medium-sized species for this genus. Maxillary palpi with apical segment stout and cylindrical to somewhat rectangular; with subapical segment very stout, distinctly shorter than apical and third segments (Fig. 7D). Pronotal disc brown, about 1.7 times as wide as long, widening posteriorly (posterior margin about 1.6 times as wide as anterior margin); anterior margin of disc somewhat straight; posterior margin of disc slightly convex (Fig. 6B). Pronotal lateral lobe 2.0 times as long as high (Fig. 7D). Male. FW 2.6 times as long as broad (Fig. 8D), covering abdomen and slightly surpassing apex of FIII. Venation (Fig. 8D): 1A vein transverse, faintly sinuous; diagonal faintly bent, with 3 distinct oblique veins in harp area; posterior two oblique veins parallel to each other, anterior most vein distinctly shorter and running nearly along length of FW. Mirror about 1.2 times as long as wide, dividing vein broadly curved at basal end. Lateral field around 7 branches on Sc (Fig. 6F). Apical field 0.11 times as long as FWL (Fig. 8D). Hind wings clearly surpassing FWs. ♁ genitalia (Figs. 9D–F). Pseudepiphallus [epiphallus] typical of genus. Posterior end of pseudepiphallus [epiphallus] produced into two lobular pseudepiphallic lophi [posterolateral epiphallic lobe], in dorsal view tongue-shaped with apex obtuse; more broadly emarginated between latero-apical lobules; in profile distinctly flattened. Pseudepiphallic paramere [ectoparamere] not surpassing pseudepiphallic lophi, sclerotized, also slender and tapering into subacute apex. Dorsal process of pseudepiphallic paramere not surpassing ventral process. Ventral process of pseudepiphallic paramere obtuse at apex in ventral view. In profile, emargination between dorsal and ventral processes of pseudepiphallic paramere broadly rounded. Ectophallic fold [rachis] typical of genus; in profile slenderer. Measurements (♂, in mm). BL = 6.5; BWL = 10.1; HL = 0.9; PronL = 1.2; PronW = 2.0; FWL = 6.2; FWW = 2.4; HWT = 2.1; FIIIL = 4.0; TIIIL = 3.1; TaIIIL = 1.6. Ecology. The species was found dwelling among the leaf litter as well as on the leaves of shrub/small trees. Distribution. Borneo (BRUNEI DARUSSALAM, Belait). Type locality. BRUNEI DARUSSALAM, Belait.

Pendleburyella pimela sp. nov. (Figs. 4, 5A, 6A, 6C, 6E, 7C, 8C, 9A–C, 11) Diagnosis. This species is characterised by a large-sized habitus, the maxillary palps with the apical segment also triangular (but more stout and broadly widened than Pendleburyella testacea); the FWs relatively broad (FWL/ FWW 2.5 in other species specimens); the mirror about as long as broad, the dividing vein of the mirror broadly and roundly curved at the basal end, the apical part of the FW shorter relative to the FWL (all of these similar to Pendleburyella vicina but this species is distinctly larger in size). The genitalia differs from that of Pendleburyella eirmosa sp. nov. by the pseudepiphallic lophi, in profile, cylindrical with roundly truncated apices (instead of dorso-ventrally flattened), the pseudepiphallic paramere ventral processes more elongated and in ventral view having acute apices (instead of obtuse apices); and the emargination between the dorsal and ventral processes pseudepiphallic paramere, in profile, narrowly angular (instead of broadly rounded). Etymology. The species name refers to the large habitus and relatively broad FWs (low FWL/FWW); pimele = fat in Greek. Material examined. Holotype: SINGAPORE • 1♁; Nee Soon freshwater swamp forest, along Woodcutter Trail; on leaf litter; 21 December 2022; coll. M.K. Tan & I. Neo; call recorded; (ZRC). Description. Very large species for this genus. Maxillary palpi with apical segment longest and expanded (triangular); with subapical segment cylindrical and expanded slightly distally, somewhat similar length as apical and third segments (Fig. 7C). Pronotal disc brown about 2.0 times as wide as long, widening posteriorly (posterior margin about 1.6 times as wide as anterior margin); anterior margin of disc broadly concave; posterior margin of disc slightly convex (Fig. 6A). Pronotal lateral lobe 2.3 times as long as high (Fig, 7C). Male. FW 2.4 times as long as broad (Fig. 8C), covering abdomen and slightly surpassing apex of FIII. Venation (Fig. 8C): 1A vein transverse, faintly curved; diagonal substraight, with 3 distinct oblique veins in harp area; posterior two oblique veins substraight,join at base near1A,anterior most vein distinctly shorter and running nearly perpendicular to 1A. Mirror about 1.1 times as long as wide, dividing vein broadly curved at basal end. Lateral field around 10 branches on Sc (Fig. 6E). Apical field 0.09 times as long as FWL (Fig. 8C). Hind wings clearly surpassing FWs. ♁ genitalia (Figs. 9A–C). Pseudepiphallus [epiphallus] typical of genus. Posterior end of pseudepiphallus [epiphallus] produced into two tongue-shaped lobular pseudepiphallic lophi [posterolateral epiphallic lobe] with obtuse apices. These lophi, in profile, cylindrical with roundly truncated apices. Pseudepiphallic paramere [ectoparamere] very elongated with ventral process well surpassing pseudepiphallic lophi; all processes with apex acute. Dorsal process of pseudepiphallic paramere distinctly shorter than ventral process barely surpassing pseudepiphallic lophi. Ventral process of pseudepiphallic paramere distinctly more sclerotized along inner margin. In profile, emargination between dorsal and ventral processes narrowly angular. Ectophallic fold [rachis] typical of genus; in profile tapering into acute apex, with anterior margin concave and posterior margin straight. Measurements (♂, in mm). BL = 9.5; BWL = 13.3; HL = 1.1; PronL = 1.4; PronW = 2.8; FWL = 7.9; FWW = 3.3; HWT = 3.3; FIIIL = 5.4; TIIIL = 3.9; TaIIIL = 1.5. Ecology. The species was found dwelling among thick leaf litter in freshwater swamp forest. Distribution. Malay Peninsula (SINGAPORE). Type locality. SINGAPORE Calling song (1♂, in captivity, 29.0°C) (Fig. 11). The male was observed to call between 22h00 and 23h30, each night having 2–4 bouts of continuous trill. The calling song is a loud continuous trill, typically lasting around 2–5 min when left undisturbed. The trill is made up of closely-packed syllables with an average duration of 9.7±1.5 ms (5.9–14.2 ms). The average syllable period is 14.5±0.8 ms (10.8–17.1 ms). There is no distinct downtime between syllables. The frequency spectrum is pure-tonal and forms a harmonic, with the energy peaking at the fundamental frequency. The dominant frequency is 8.14±0.08 kHz (7.78–8.34 kHz).

Pendleburyella sp. 1 (Figs. 7E, 8F, 13) Material examined. MALAYSIA • 1♁ (type); Kuala Lumpur; 5 March 1939; 0021 (MP). Remarks. This specimen is small among the other specimens, and is most easily characterised by the very slender FWs and the distinctly longer than broad mirror. The dividing vein of the mirror at the basal end is strongly and distinctly angularly curved, unlike the other specimens examined. It is also clearly different from the holotype of Pendleburyella vicina from Kuala Lumpur by the apical segment of the maxillary palps and the slenderer FWs(Table1)., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on page 145, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034

Pendleburyella eirmosa sp. nov. (Figs. 1, 5B, 6B, 6D, 6F, 7D, 8D, 9D–F) Diagnosis. This species is characterised by a medium-sized habitus (similar to Pendleburyella testacea), the maxillary palps with the apical segment also somewhat rectangular (similar to that of Pendleburyella vicina but stouter); the mirror longer than broad, the dividing vein of the mirror broadly and roundly curved at the basal end; the apical part of FW longer relative to the FWL. The genitalia also differs from that of Pendleburyella pimela sp. nov. (see the diagnosis for the latter species). Etymology. The species name refers to the broadly rounded emargination between the dorsal and ventral processes of the pseudepiphallic paramere (distinguished from the narrowly angular emargination in Pendleburyella pimela sp. nov.); eirmós = continuity in Ancient Greek. Material examined. Holotype: BRUNEI DARUSSALAM • 1♁; Belait District, Wasai Wong Kadir Recreational Park; N4.34186, E114.44611, 35.8± 5.2 m.a.s.l.; 1 March 2019, 18h54; calling on a leaf; coll. M.K. Tan & H. Yeo; BRU.19.47 (UBDM) Photograph examined. BRUNEI DARUSSALAM • 1♁; Temburong District, Kuala Belalong Field Studies Centre, along Ashton Trail; N4.54619, E115.15690, 110.8± 6.9 m.a.s.l.; 26 July2017, dusk; on leaf litter; photographed M.K. Tan (Fig. 1). Description. Medium-sized species for this genus. Maxillary palpi with apical segment stout and cylindrical to somewhat rectangular; with subapical segment very stout, distinctly shorter than apical and third segments (Fig. 7D). Pronotal disc brown, about 1.7 times as wide as long, widening posteriorly (posterior margin about 1.6 times as wide as anterior margin); anterior margin of disc somewhat straight; posterior margin of disc slightly convex (Fig. 6B). Pronotal lateral lobe 2.0 times as long as high (Fig. 7D). Male. FW 2.6 times as long as broad (Fig. 8D), covering abdomen and slightly surpassing apex of FIII. Venation (Fig. 8D): 1A vein transverse, faintly sinuous; diagonal faintly bent, with 3 distinct oblique veins in harp area; posterior two oblique veins parallel to each other, anterior most vein distinctly shorter and running nearly along length of FW. Mirror about 1.2 times as long as wide, dividing vein broadly curved at basal end. Lateral field around 7 branches on Sc (Fig. 6F). Apical field 0.11 times as long as FWL (Fig. 8D). Hind wings clearly surpassing FWs. ♁ genitalia (Figs. 9D–F). Pseudepiphallus [epiphallus] typical of genus. Posterior end of pseudepiphallus [epiphallus] produced into two lobular pseudepiphallic lophi [posterolateral epiphallic lobe], in dorsal view tongue-shaped with apex obtuse; more broadly emarginated between latero-apical lobules; in profile distinctly flattened. Pseudepiphallic paramere [ectoparamere] not surpassing pseudepiphallic lophi, sclerotized, also slender and tapering into subacute apex. Dorsal process of pseudepiphallic paramere not surpassing ventral process. Ventral process of pseudepiphallic paramere obtuse at apex in ventral view. In profile, emargination between dorsal and ventral processes of pseudepiphallic paramere broadly rounded. Ectophallic fold [rachis] typical of genus; in profile slenderer. Measurements (♂, in mm). BL = 6.5; BWL = 10.1; HL = 0.9; PronL = 1.2; PronW = 2.0; FWL = 6.2; FWW = 2.4; HWT = 2.1; FIIIL = 4.0; TIIIL = 3.1; TaIIIL = 1.6. Ecology. The species was found dwelling among the leaf litter as well as on the leaves of shrub/small trees. Distribution. Borneo (BRUNEI DARUSSALAM, Belait). Type locality. BRUNEI DARUSSALAM, Belait., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on pages 142-144, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034

Genus Pendleburyella Chopard, 1969 Chopard, 1969: 217 Type species. Pendleburyella testacea Chopard, by original designation (Fig. 3) Diagnosis (new). Small brown crickets. Among Pentacentrinae, this genus differs most prominently by the male FWs with a fully developed stridulatory apparatus (the diagonal vein substraight, three oblique veins in the harp area, a large mirror with a distinct dividing vein). The male genitalia: the pseudepiphallus [epiphallus] is elongated and gently narrows posteriorly into two tongue-shaped, obtuse lobular pseudepiphallic lophi [posterolateral epiphallic lobe]. The pseudepiphallic parameres [ectoparameres] are strongly sclerotized, elongated and forming dorsal and ventral subacute processes. The ectophallic fold [rachis] is weakly sclerotized, forming a process with an acute apex pointing perpendicular to the pseudepiphallus (in profile view). The endophallic sclerite [formula (= mold of spermatophore attachment plate)] is small and elongated, with two lateral arms. The ectophallic apodeme [endoparameral apodeme] is very long. Redescription. Small cricket, generally brown or yellow brown (Figs. 1, 4, 5). Head rounded, a little wider than anterior margin of pronotum, with dorsum slightly flattened, vertex sloping, very finely pubescent (Figs. 6A, 6B). Frontal rostrum dark-coloured, about two times as wide as scapes, with apex truncated (in dorsal view) (Figs. 6A, 6B). Eyes globular, feebly projected anteriorly in dorsal view (Figs. 6A, 6B). Antennae inserted at level midpoint of eye in anterior view (Figs. 6C, 6D). Median ocellus round and small; lateral ocelli oval, located near eyes (Figs. 6C, 6D). Maxillary palpi with apical segment longest and expanded (triangular or slightly broadening apically); with subapical segment cylindrical and shorter than apical and third segments (Fig. 7). Face brown, in anterior view about 0.7–0.8 times as tall as wide (Figs. 6C, 6D). Pronotal disc brown about 1.6–2.0 times as wide as long, widening posteriorly (posterior margin about 1.3–1.6 times as wide as anterior margin); anterior margin of disc nearly straight to broadly concave; posterior margin of disc nearly straight to slightly convex (Figs. 6A, 6B). Pronotal lateral lobe about 1.8–2.3 times as long as high (Fig. 7). TI slightly swollen; with inner and outer tympana open and having oval tympanal membrane. TIII with 3 inner and 4 outer long subapical spurs; with 2–4 denticles between spines; 4 denticles before most proximal spines; and with 2 inner spurs (apical spines) longer than corresponding 3 outer ones. Legs generally pale brown. Thoracic and abdominal segments yellow brown to brown (Figs. 6E, 6F). Male. FWs covering abdomen and slightly surpassing apex of FIII. Colouration: yellow brown, with veins darker (Fig. 8). Venation: 1A vein transverse, faintly curved; diagonal substraight, with three distinct oblique veins in harp area; posterior two oblique vein more or less straight and long, anteriormost vein distinctly shorter and running nearly perpendicular to 1A. Mirror about 1.1–1.5 times as long as wide, dividing vein curved at basal end (Fig. 8). Lateral field around 7–10 branches on Sc (Figs. 6E, 6F). Hind wings clearly surpassing FWs. ♁ genitalia as shown in Fig. 9: Pseudepiphallus [epiphallus] elongated, gently narrowing posteriorly. Posterior end of pseudepiphallus produced into two lobular pseudepiphallic lophi [posterolateral epiphallic lobe], in dorsal view tongue-shaped; emarginated between latero-apical lobules in dorsal view; in profile variable (e.g., cylindrical or flattened), with a few setae. Pseudepiphallic paramere [ectoparamere] strongly sclerotized, elongated with dorsal and ventral processes (emarginated in profile); each process tapered into a slender and slightly hooked-like apical third with apex acute. Ectophallic fold [rachis] weakly sclerotized, rather long but weakly sclerotized, pointing perpendicular to pseudepiphallus; in profile tapering into acute apex. Endophallic sclerite [formula (= mold of spermatophore attachment plate)] small and elongated, with two lateral arms. Ectophallic apodeme [endoparameral apodeme] widened posteriorly, otherwise long, slender and substraight. Rachis weakly sclerotized. Distribution. Malay Peninsula, Borneo, Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on pages 134-137, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034, {"references":["Chopard, L. (1969) Grylloidea. In: Sewell, R. B. S. (Ed.), The fauna of India and the adjacent countries. Orthoptera. Vol. 2. Zoological Survey of India and Baptist Mission Press, Calcutta, pp. 1 - 421."]}

Pendleburyella sp. “ intermedia Chopard” (Figs. 8E, 12) Material examined. MALAYSIA • 1♁ (type); Pahang, Fraser’s Hill, at light; 12 July 1936; 0017 (MP). Remarks. The ‘type’ specimen was identified as “ Pendleburya intermedia ” by Chopard, but we did not find the original description of this species in Chopard (1969) where the genus and species were first described. We also did not find other publications where this species was mentioned, and consequently it was also not found in the Orthoptera Species File (Cigliano et al., 2023). This specimen is large and shows clear differences with Pendleburyella sp. 2 from Selangor – Pahang, The Gap (not too far from Fraser’s Hill) (Table 1). Unfortunately, the specimen also appeared damaged at the abdominal apex and the male genitalia may be lost. As the male genitalia was not available, we did not describe and name this species unless newer material from Fraser’s Hill can be collected., Published as part of Tan, Ming Kai, Muhammad, Amira Aqilah & Wahab, Rodzay Bin Haji Abdul, 2023, The taxonomy and bioacoustics of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae: Pentacentrinae), pp. 131-148 in Zootaxa 5277 (1) on page 144, DOI: 10.11646/zootaxa.5277.1.6, http://zenodo.org/record/7893034, {"references":["Chopard, L. (1969) Grylloidea. In: Sewell, R. B. S. (Ed.), The fauna of India and the adjacent countries. Orthoptera. Vol. 2. Zoological Survey of India and Baptist Mission Press, Calcutta, pp. 1 - 421.","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2023) Orthoptera Species File Online. Version 5 (5.0). Available from: http: // orthoptera. speciesfile. org / HomePage / Orthoptera / HomePage. aspx (accessed 14 March 2023)"]}

Genus Asiophlugis Gorochov, 1998 Asiophlugis Gorochov, 1998: 128 Phlugis — Hebard 1922: 263 (new species); Jin 1993: 1589 (taxonomic revision); Helfert & Sänger 1998: 61 (new species and calling song description) Asiophlugis — Gorochov & Tan 2011: 129 (new species and combinations); Tan 2011: 233 (species from Singapore); Gorochov 2012: 2 (literature review and new species); Gorochov 2019: 6 (new species and remarks); Tan et al. 2019: 510 (bioacoustics); Tan et al. 2020: 537 (redescription of Asiophlugis philippina) Type species. Asiophlugis rete Gorochov, by original designation, Published as part of Tan, Ming Kai, Salvador, Jewel Anne G. & Nuñeza, Olga Macas, 2023, A new species of Asiophlugis (Orthoptera: Tettigoniidae: Phlugidini) from Misamis Oriental (Philippines, Mindanao), pp. 475-484 in Zootaxa 5258 (4) on page 477, DOI: 10.11646/zootaxa.5258.4.8, http://zenodo.org/record/7784627, {"references":["Gorochov, A. V. (1998) New and little known Meconematinae of the tribes Meconematini and Phlugidini (Orthoptera: Tettigoniidae). Zoosystematica Rossica, 7 (1), 101 - 131.","Hebard, M. (1922) Studies in Malayan, Melanesian and Australian Tettigoniidae (Orthoptera). Proceedings of the Academy of Natural Sciences, Philadelphia, 74, 121 - 299, pls. 11 - 22.","Jin, X. B. (1993) Remarks on the tribe Phlugidini Eichler and recognition of new taxa from the Indo-Malayan region and East Africa (Grylloptera: Tettigonioidea: Meconematidae). Invertebrate Taxonomy, 7 (6), 1589 - 1610. https: // doi. org / 10.1071 / IT 9931589","Helfert, B. & Sanger, K. (1998) Phlugis thai sp. n., a new Phlugidini (Insecta: Ensifera, Tettigoniidae, Meconematinae) from Thailand. Annalen des Naturhistorischen Museums in Wien, 100 (B), 61 - 68.","Gorochov, A. V. & Tan, M. K. (2011) New katydids of the genus Asiophlugis Gorochov (Orthoptera: Tettigoniidae: Meconematinae) from Singapore and Malaysia. Russian Entomological Journal, 20 (2), 129 - 133. https: // doi. org / 10.15298 / rusentj. 20.2.03","Tan, M. K. (2011) The species of Asiophlugis Gorochov, 1998 in Singapore (Orthoptera: Tettigoniidae: Meconematinae). Nature in Singapore, 4, 233 - 239.","Gorochov, A. V. (2012) Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 5. Far Eastern Entomologist, 252, 1 - 26. https: // doi. org / 10.25221 / fee. 459.1","Gorochov, A. V. (2019) Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 12. Far Eastern Entomologist, 379, 1 - 24. https: // doi. org / 10.25221 / fee. 379.1","Tan, M. K., Montealegre-Z., F., Wahab, R. A., Lee, C. - Y., Belabut, D. M., Japir, R. & Chung, A. Y. C. (2019) Ultrasonic songs and stridulum anatomy of Asiophlugis crystal predatory katydids (Tettigonioidea: Meconematinae: Phlugidini). Bioacoustics, 18 (5), 510 - 524. https: // doi. org / 10.1080 / 14772000.2020.1769223","Tan, M. K., Jin, X. - B., Baroga-Barbecho, J. B. & Yap, S. A. (2020) Taxonomy and bioacoustics of Meconematinae (Orthoptera: Tettigoniidae) from Laguna (Philippines: Luzon). Zootaxa, 4732 (4), 527 - 544. https: // doi. org / 10.11646 / zootaxa. 4732.4.2"]}

Asiophlugis spatulata sp. nov. (Figs 2–5) Specimens examined. Holotype: PHILIPPINES • 1J; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, near Mt Sumagaya; N8.69952, E125.01376, 1243.0± 6.5 m.a.s.l.; 13 February 2023, 11h15; at the edge of a mossy forest, under the foliage of a shrub; coll. J.A.G. Salvador, M.K. Tan, A.M. Sabang, A. Gono, D. Bahoy & K. Grumo; PNM (MIN.23.06) Paratype: PHILIPPINES • 1J; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, near Mt Sumagaya; N8.70034, E125.00941, 1217.0± 9.9 m.a.s.l.; 17 February 2023, 10h11; vegetation along a river, under the leaf of a shrub; coll. M.K. Tan, J.A.G. Salvador, A.M. Sabang, A. Gono, D. Bahoy & K. Grumo; ZRC (MIN.23.77) Other material: PHILIPPINES • 1♀ subadult; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, foot of Mt Balatukan; N8.70531, E125.00652, 1209.0± 6.1 m.a.s.l.; 14 February 2023, 10h55; cleared forest patch, under the leaf of a small tree; coll. M.K. Tan, J.A.G. Salvador, A.M. Sabang, A. Gono, D. Bahoy & K. Grumo; MSU-IIT (MIN.23.35) • 1♀ subadult; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, foot of Mt Balatukan; N8.70389, E125.01028, 1209.0± 6.3 m.a.s.l.; 17 February 2023, 20h10; open, disturbed forest, on the leaf of a shrub; coll. J.A.G. Salvador, M.K. Tan, A.M. Sabang, D. Bahoy & K. Grumo; ZRC (MIN.23.91) Diagnosis. This new species is most similar to Asiophlugis philippina (Jin, 1993) from Luzon and shares similar unique characters among Asiophlugis (see Tan et al., 2020): brachypterous, and with the male tegmen being tear-shaped; the male stridulatory anatomy obscure; the presence of two long, cylindrical latero-apical processes at the posterior end of the male tenth abdominal tergite; and dark patterns on the dorsal and medial parts of some tergites. However, the new species differs from A. philippina by: the lateral processes on the male tenth abdominal tergite being more cylindrical (instead of being dorso-ventrally flattened) and pointing more dorsad and externally (instead of pointing posteriorly); the male cercus distinctly longer, slenderer and flattened, with a medial flattened tooth-like process produced along inner margin, but without any apical process; the basal process of the male cercus with spatulated apex. The female of this new species differs from that of A. philippina by the shape of subgenital plate being small and transverse (rather than long, narrowed distally with a rectangular lobe at the apex). The new species is also similar to the brachypterous Asiophlugis cercalis Gorochov, 2012 and Asiophlugis paracercalis Gorochov, 2019 from Sarawak (Borneo) in the presence of a slender male cercus with very short stout setae along its inner margin; but the cercus differs from those of the two species by the shape of the basal process and the presence of a medial flattened tooth-like process produced along the inner margin. The new species also differs from these two species by the presence of cylindrical latero-apical processes produced along the posterior margin of the male tenth abdominal tergite and the weakly visible venation on the male tegmen. The new species is also similar to Asiophlugis borneoensis (Jin, 1993) from Sabah (Borneo) in the greatly reduced male tegmen with weakly visible venation and the presence of latero-apical processes along the posterior margin of the male tenth abdominal tergite; but it differs distinctly by the slenderer latero-apical processes of the male tenth abdominal tergite and the shape of cercus. Etymology. The species name refers to the spatulated apex of the basal process of the male cercus. Description. Habitus typical of Asiophlugis, generally green when alive (and yellow when preserved), but with some brown and dark patterns on pronotum and tergites (Figs 2, 3). Eye very large with a bright red band (not always present), almost globular, somewhat longitudinally elongated (Fig. 4A). Scapus brown apically; antennal segments generally black, some with brown apices (Fig. 4A). Face high, oblique and slightly concave in profile (Fig. 4A). Pronotum dorsal disc slender and elongated, generally green; with posterior half of metazona brown (Fig. 2); having fairly straight posterior margin (Fig. 4B). Pronotum lateral lobe distinctly longer than tall; with metazona barely elevated (Figs 3, 4A). Thoracic auditory foramen small, thoracic spiracle slit (Fig. 4A). Outer and inner tympana on fore tibia open, oval and large. Fore coxa with long and thin spine which is slightly curved. Fore and middle legs generally green with tint of brown (particular around the tympana, basal and apical parts of middle tibiae) (Fig. 2). Fore femur with 4 and 1 ventral spines on the inner and outer margins respectively (outer spine at distal third distinctly shorter than inner ones). Fore tibia with 5 and 4 long and thin ventral spines on the inner and outer margins respectively. Middle leg with one tibial spine slightly after middle. Hind femur mostly green, posterior third slightly brown, black ring around knee; hind knee lobe blunt and black (Figs 2C, 3). Hind tibia brown, at the posterior end with a broad white ring, posterior of the white ring black; with only moderately numerous small spinules on both dorsal edges of tibia (Figs 2C, 3). Hind tarsus mostly black (Figs 2C, 3). Abdominal tergites generally green; some segments (especially the basal and posterior segments) with brown bands (either longitudinal along tergite length in middle, or transverse along posterior margin) and triangular black patch along posterior margin (Fig. 2). Male. Tegmina tear-shaped; transparent with anal and apical margins brown; barely surpassing beyond posterior margin of first tergite (Figs 3A, 3B, 4B). Venation and stridulatory anatomy obscure (Fig. 4B). Tenth abdominal tergite with posterior margin slightly concave in middle, with a pair of brown long latero-apical processes (Fig. 4C). Latero-apical process of tenth abdominal tergite cylindrical, long and surpassing epiproct, pointing dorsad and outward; apex obtuse and black (Figs 4C–F). Epiproct stoutly tongue-shaped; basal half slightly swollen laterally, posterior half rounded (Fig. 4D). Cercus complex: long, slender and flattened after basal third (Figs 4C, 4F). Inner basal process of cercus green and swollen basally, then black, flattened and strongly curved into spatulated apex; apex flattened, obtuse and with fine setae (Figs 4D, 4E). In middle, cercus dorso-ventrally flattened; with sparse setae very short and stout (spinule-like) along inner margin and inner-dorsal surface; with tooth-like and flattened medial process produced along inner margin. Medial process with apex brown, obtuse and with a few stout setae (Figs 4C, 4E). Before apex, cercus rotates inwards and slightly widens, with apical part more laterally flattened; with very short stout setae along inner margin; apex brown and rounded truncated (Figs 4C, 4E, 4F). Subgenital plate about as long as broad, with lateral margins rounded into posterior margin; posterior margin somewhat straight, with lateral lobe barely produced. Stylus green, fairly long and cylindrical; with obtuse apex brown to black (Figs 4E, 4F). Female (subadult) (Fig. 3C). Tegminal bud squarish (Fig. 5A). Cercus long and slender, typical of genus; apical half progressively darkened (Fig. 5B). Ovipositor also typical of genus (Fig. 5B). Subgenital plate small, transverse, broader than long; posterior margin broadly rounded and faintly emarginated (Fig. 5C). Measurements (in mm). Table 1. Ecology. This species was found on and under the leaves of small trees and shrubs (Fig. 2A), which is typically the case for Asiophlugis. The specimens were collected from forest edges (Fig. 1). Distribution. Philippines, Mindanao, Misamis Oriental (currently known only from type locality) Type locality. Philippines, Mindanao, Misamis Oriental Calling song. Unknown.

Asiophlugis spatulata sp. nov. (Figs 2–5) Specimens examined. Holotype: PHILIPPINES • 1J; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, near Mt Sumagaya; N8.69952, E125.01376, 1243.0± 6.5 m.a.s.l.; 13 February 2023, 11h15; at the edge of a mossy forest, under the foliage of a shrub; coll. J.A.G. Salvador, M.K. Tan, A.M. Sabang, A. Gono, D. Bahoy & K. Grumo; PNM (MIN.23.06) Paratype: PHILIPPINES • 1J; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, near Mt Sumagaya; N8.70034, E125.00941, 1217.0± 9.9 m.a.s.l.; 17 February 2023, 10h11; vegetation along a river, under the leaf of a shrub; coll. M.K. Tan, J.A.G. Salvador, A.M. Sabang, A. Gono, D. Bahoy & K. Grumo; ZRC (MIN.23.77) Other material: PHILIPPINES • 1♀ subadult; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, foot of Mt Balatukan; N8.70531, E125.00652, 1209.0± 6.1 m.a.s.l.; 14 February 2023, 10h55; cleared forest patch, under the leaf of a small tree; coll. M.K. Tan, J.A.G. Salvador, A.M. Sabang, A. Gono, D. Bahoy & K. Grumo; MSU-IIT (MIN.23.35) • 1♀ subadult; Mindanao, Misamis Oriental, Gingoog City, Barangay Lunotan, foot of Mt Balatukan; N8.70389, E125.01028, 1209.0± 6.3 m.a.s.l.; 17 February 2023, 20h10; open, disturbed forest, on the leaf of a shrub; coll. J.A.G. Salvador, M.K. Tan, A.M. Sabang, D. Bahoy & K. Grumo; ZRC (MIN.23.91) Diagnosis. This new species is most similar to Asiophlugis philippina (Jin, 1993) from Luzon and shares similar unique characters among Asiophlugis (see Tan et al., 2020): brachypterous, and with the male tegmen being tear-shaped; the male stridulatory anatomy obscure; the presence of two long, cylindrical latero-apical processes at the posterior end of the male tenth abdominal tergite; and dark patterns on the dorsal and medial parts of some tergites. However, the new species differs from A. philippina by: the lateral processes on the male tenth abdominal tergite being more cylindrical (instead of being dorso-ventrally flattened) and pointing more dorsad and externally (instead of pointing posteriorly); the male cercus distinctly longer, slenderer and flattened, with a medial flattened tooth-like process produced along inner margin, but without any apical process; the basal process of the male cercus with spatulated apex. The female of this new species differs from that of A. philippina by the shape of subgenital plate being small and transverse (rather than long, narrowed distally with a rectangular lobe at the apex). The new species is also similar to the brachypterous Asiophlugis cercalis Gorochov, 2012 and Asiophlugis paracercalis Gorochov, 2019 from Sarawak (Borneo) in the presence of a slender male cercus with very short stout setae along its inner margin; but the cercus differs from those of the two species by the shape of the basal process and the presence of a medial flattened tooth-like process produced along the inner margin. The new species also differs from these two species by the presence of cylindrical latero-apical processes produced along the posterior margin of the male tenth abdominal tergite and the weakly visible venation on the male tegmen. The new species is also similar to Asiophlugis borneoensis (Jin, 1993) from Sabah (Borneo) in the greatly reduced male tegmen with weakly visible venation and the presence of latero-apical processes along the posterior margin of the male tenth abdominal tergite; but it differs distinctly by the slenderer latero-apical processes of the male tenth abdominal tergite and the shape of cercus. Etymology. The species name refers to the spatulated apex of the basal process of the male cercus. Description. Habitus typical of Asiophlugis, generally green when alive (and yellow when preserved), but with some brown and dark patterns on pronotum and tergites (Figs 2, 3). Eye very large with a bright red band (not always present), almost globular, somewhat longitudinally elongated (Fig. 4A). Scapus brown apically; antennal segments generally black, some with brown apices (Fig. 4A). Face high, oblique and slightly concave in profile (Fig. 4A). Pronotum dorsal disc slender and elongated, generally green; with posterior half of metazona brown (Fig. 2); having fairly straight posterior margin (Fig. 4B). Pronotum lateral lobe distinctly longer than tall; with metazona barely elevated (Figs 3, 4A). Thoracic auditory foramen small, thoracic spiracle slit (Fig. 4A). Outer and inner tympana on fore tibia open, oval and large. Fore coxa with long and thin spine which is slightly curved. Fore and middle legs generally green with tint of brown (particular around the tympana, basal and apical parts of middle tibiae) (Fig. 2). Fore femur with 4 and 1 ventral spines on the inner and outer margins respectively (outer spine at distal third distinctly shorter than inner ones). Fore tibia with 5 and 4 long and thin ventral spines on the inner and outer margins respectively. Middle leg with one tibial spine slightly after middle. Hind femur mostly green, posterior third slightly brown, black ring around knee; hind knee lobe blunt and black (Figs 2C, 3). Hind tibia brown, at the posterior end with a broad white ring, posterior of the white ring black; with only moderately numerous small spinules on both dorsal edges of tibia (Figs 2C, 3). Hind tarsus mostly black (Figs 2C, 3). Abdominal tergites generally green; some segments (especially the basal and posterior segments) with brown bands (either longitudinal along tergite length in middle, or transverse along posterior margin) and triangular black patch along posterior margin (Fig. 2). Male. Tegmina tear-shaped; transparent with anal and apical margins brown; barely surpassing beyond posterior margin of first tergite (Figs 3A, 3B, 4B). Venation and stridulatory anatomy obscure (Fig. 4B). Tenth abdominal tergite with posterior margin slightly concave in middle, with a pair of brown long latero-apical processes (Fig. 4C). Latero-apical process of tenth abdominal tergite cylindrical, long and surpassing epiproct, pointing dorsad and outward; apex obtuse and black (Figs 4C–F). Epiproct stoutly tongue-shaped; basal half slightly swollen laterally, posterior half rounded (Fig. 4D). Cercus complex: long, slender and flattened after basal third (Figs 4C, 4F). Inner basal process of cercus green and swollen basally, then black, flattened and strongly curved into spatulated apex; apex flattened, obtuse and with fine setae (Figs 4D, 4E). In middle, cercus dorso-ventrally flattened; with sparse setae very short and stout (spinule-like) along inner margin and inner-dorsal surface; with tooth-like and flattened medial process produced along inner margin. Medial process with apex brown, obtuse and with a few stout setae (Figs 4C, 4E). Before apex, cercus rotates inwards and slightly widens, with apical part more laterally flattened; with very short stout setae along inner margin; apex brown and rounded truncated (Figs 4C, 4E, 4F). Subgenital plate about as long as broad, with lateral margins rounded into posterior margin; posterior margin somewhat straight, with lateral lobe barely produced. Stylus green, fairly long and cylindrical; with obtuse apex brown to black (Figs 4E, 4F). Female (subadult) (Fig. 3C). Tegminal bud squarish (Fig. 5A). Cercus long and slender, typical of genus; apical half progressively darkened (Fig. 5B). Ovipositor also typical of genus (Fig. 5B). Subgenital plate small, transverse, broader than long; posterior margin broadly rounded and faintly emarginated (Fig. 5C). Measurements (in mm). Table 1. Ecology. This species was found on and under the leaves of small trees and shrubs (Fig. 2A), which is typically the case for Asiophlugis. The specimens were collected from forest edges (Fig. 1). Distribution. Philippines, Mindanao, Misamis Oriental (currently known only from type locality) Type locality. Philippines, Mindanao, Misamis Oriental Calling song. Unknown., Published as part of Tan, Ming Kai, Salvador, Jewel Anne G. & Nuñeza, Olga Macas, 2023, A new species of Asiophlugis (Orthoptera: Tettigoniidae: Phlugidini) from Misamis Oriental (Philippines, Mindanao), pp. 475-484 in Zootaxa 5258 (4) on pages 477-483, DOI: 10.11646/zootaxa.5258.4.8, http://zenodo.org/record/7784627, {"references":["Jin, X. B. (1993) Remarks on the tribe Phlugidini Eichler and recognition of new taxa from the Indo-Malayan region and East Africa (Grylloptera: Tettigonioidea: Meconematidae). Invertebrate Taxonomy, 7 (6), 1589 - 1610. https: // doi. org / 10.1071 / IT 9931589","Tan, M. K., Jin, X. - B., Baroga-Barbecho, J. B. & Yap, S. A. (2020) Taxonomy and bioacoustics of Meconematinae (Orthoptera: Tettigoniidae) from Laguna (Philippines: Luzon). Zootaxa, 4732 (4), 527 - 544. https: // doi. org / 10.11646 / zootaxa. 4732.4.2","Gorochov, A. V. (2012) Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 5. Far Eastern Entomologist, 252, 1 - 26. https: // doi. org / 10.25221 / fee. 459.1","Gorochov, A. V. (2019) Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 12. Far Eastern Entomologist, 379, 1 - 24. https: // doi. org / 10.25221 / fee. 379.1"]}

Platybinthus striolatus (Brunner von Wattenwyl, 1898) gen. et comb. nov. Figs 2–3, 4A–D, 5B, E, 6B, 7A, C, 8A–C, H Paraeneopterus striolatus Brunner von Wattenwyl, 1898: 279. Lebinthus striolatus – Kirby 1906: 88. — Karny 1915: 68. — Shiraki 1930: 233. — Chopard 1968: 354. — Hsiung 1993: 26. — Ichikawa et al. 2000: 275. Diagnosis Platybinthus striolatus gen. et comb. nov. differs from P. punctatus gen. et comb. nov. by smaller size and by female copulatory papilla with apex narrowed and obtusely rounded. It also differs from Platybinthus sandyi gen. et sp. nov. by male genitalia with pseudepiphallus dorsal disk smaller, margins at the apex tapering (instead of straight) and by pseudepiphallic parameres with ventral lobe cylindrical (instead of appearing triangular and flattened). Material examined Holotype INDONESIA – Maluka • ♂ (juvenile); Halmaheira [Halmahera Island]; W. Kükenthal leg.; Brunner v. W. det.; DORSA FG; #Orth.57; SMFD. Additional specimens INDONESIA – Maluka • 1 ♀; Halmaheira [= Halmahera Island]; 1894; W. Kükenthal leg.; 21.852; MNHN-ENSIF1449 • 1 ♂; prov. Maluka Utara [northern Maluka], Halmahera Island, env. of vill. Subaim to S from vill. Lolobata (not far of it) near coast of Wasile Bay; 27 Jan.–1 Feb. 2011; A. Gorochov leg.; molecular sample L81-Lpc1; GenBank accession: OP684788 and OP686542; ZIN. Remarks The type (holotype) specimen in SMFD is a male juvenile. It is the only known specimen of the species, and probably the only specimen examined by Brunner von Wattenwyl (1898), according to the measurements that he provided. Given the general shape and colouration pattern on the legs, we hypothesise here that this juvenile belongs to the same species as the adult specimens from Halmahera found in MNHN and ZIN, and differing from the type of the species P. punctatus gen. et comb. nov. Although the evidence is weak, and not being able to sequence the old type juvenile, we refrained from describing a new species from these adult specimens; instead we propose to stabilise the name P. striolatus gen. et comb. nov. by redescribing the species based on them. As a consequence of the very brief original description and limited characters available on the type specimen, the name Lebinthus striolatus has been mentioned multiple times in databases, books and publications based on erroneous identifications of specimens of other species of Lebinthus from Taiwan (e.g., Shiraki 1930; Ichikawa et al. 2000). Those specimens and records probably correspond to one of the two species of Lebinthina documented in Taiwan, Lebinthus lanyuensis Oshiro, 1996 or Falcerminthus yaeyamensis (Oshiro, 1996), which were wrongly identified as Lebinthus striolatus based on their colouration matching the evocative ‘striolated’ name. Redescription Size small for the genus (Fig. 4A–D). Vertex and fastigium very dark brown; dorsum of head at posterior end with three longitudinal pale bands (Fig. 5B); back of eyes with a cream-coloured band. Scapes white, with some dark patterns (more in the ventral). Antennae orange brown. Fastigium verticis and frons dark grey-brown (Fig. 5E). Mouthparts whitish, mandibles grey-brown mottled with white; clypeus and labrum white; maxillary palpi with segments white with some dark stripes (more prominent on external surface) (Fig. 5E). Gena grey with a triangular white spot beneath eyes and two small white spots nearer to mouthparts. Lateral parts of head dark brown behind eyes (Fig. 6B). Pronotum dorsal disk yellow mottled with dark brown, anterior and posterior margins straight; lateral lobes very dark brown (Fig. 6B). Legs: FI–II pale with numerous dark brown spots and incomplete ring near apices, knee area dark brown; TI–II pale with three broad dark brown rings; tarsomeres 1 and 3 of each leg yellow basally then brown; FIII brown finely mottled with yellow and dark brown, knees dark brown; TIII homogeneously dark brown, spurs lighter brown with dark tips. Abdominal tergites dark brown with faint dark and light spots. Cerci brown with faint apical rings. Male FW reaching apex of third tergite. FW colouration (Fig. 7A): dorsal field cells mostly brown, plectrum area and basal area darker, with two faint pale spots at the basal and apical end of Cu and M; veins whitish or cream-coloured; M yellow brown, R orange brown. Lateral field cells dark brown in dorsal half and whitish in ventral half; longitudinal veins red brown with cross-veins cream-coloured. FW venation: harp vein bordering false mirror strongly sinuous. Anal area of dorsal field extending beyond apex lateral field, apex of dorsal field obliquely rounded. GENITALIA (Fig. 8A–C). Pseudepiphallus elongate and slender, its basal margin distinctly indented in the middle, posterior apex rounded, slightly swollen at the apex. Rami short, shorter than half the pseudepiphallus length. Pseudepiphallic parameres with ventral lobe cylindrical. Ectophallic apodemes parallel and long, slightly reaching beyond anterior margin of pseudepiphallic sclerite. Endophallic sclerite with long anterior region not reaching anterior margin of pseudepiphallic sclerite and a short median expansion and small lateral arms posteriorly. Endophallic apodeme with lateral lamellae curved and pointing anteriorly. Female FWs reaching middle of fourth tergite, faintly overlapping. Cells of dorsal field dark brown, not translucent, with strong orange longitudinal veins and weak yellow transverse ones; with 9 longitudinal veins on dorsal field, 4 on lateral field (Fig. 7C). Dorsal field apex truncated, lateral field posterior margin oblique. GENITALIA. Ovipositor as long as FIII, apex slightly denticulate dorsally. Copulatory papilla rounded; basal sclerite forming a semi-circular ring with a small basal plate; apex narrowed and obtusely rounded (Fig. 8H). Measurements See Table 1. Ecology Unknown. Distribution INDONESIA: Maluka: Halmahera Island. Type locality INDONESIA: Maluka: Halmahera Island. Calling song Unknown., Published as part of Robillard, Tony & Tan, Ming Kai, 2023, Platybinthus, a new genus of Lebinthina crickets (Orthoptera, Gryllidae, Eneopterinae) from Maluka, Indonesia, pp. 46-66 in European Journal of Taxonomy 856 (1) on pages 58-61, DOI: 10.5852/ejt.2023.856.2027, http://zenodo.org/record/7544313, {"references":["Brunner von Wattenwyl H. H. 1898. Orthopteren des Malayischen Archipels gesammelt von Prof. Dr. W. Kukenthal in den Jahren 1893 and 1894. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 24 (2): 193 - 288.","Kirby W. F. 1906. A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). Vol. 2. Orthoptera Saltatoria. Part I. (Achetidae et Phasgonuridae). British Museum (Natural History), London. https: // doi. org / 10.5962 / bhl. title. 6745","Karny H. H. 1915. Orthoptera et Oothecaria (H. Sauter's Formosa-Ausbeute). Supplementa Entomologica 4: 56 - 108.","Shiraki T. 1930. Orthoptera of the Japanese Empire. Part I. (Gryllotalpidae and Gryllidae). Insecta Matsumurana 4 (4): 181 - 252.","Chopard L. 1968. Fam. Gryllidae: Subfam. Mogoplistinae, Myrmecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Fam. Oecanthidae, Gryllotalpidae. In: Beier M. (ed.) Orthopterorum Catalogus 12: 213 - 500. Uitgeverij Dr. W. Junk N. V. ' s, Gravenhage.","Hsiung C. - C. 1993. Catalogue of species and type localities of Taiwanese crickets (Gryllodea, Grlloptera). Yushania 10: 19 - 29.","Ichikawa A., Murai T. & Honda E. 2000. Monograph of Japanese crickets (Orthoptera; Grylloidea). Bulletin of the Hoshizaki Green Foundation 4: 257 - 332."]}

Genus Platybinthus gen. nov. urn:lsid:zoobank.org:act: 31113AC2-ED96-4949-95A5-E8F0879B351B Type species Piestodactylus punctatus Brunner von Wattenwyl, 1898. Diagnosis Among the genera of Lebinthina, species of Platybinthus are average to large sized with a dark brown colouration. Platybinthus is mainly characterised by male genitalia with pseudepiphallus middle part forming a dorsal ovular disk. General shape close to that of Macrobinthus and Rugabinthus, both mostly from New Guinea. Eyes prominent and large as in Macrobinthus and Rugabinthus. Microptery in both sexes, FWs not reaching mid-length of abdomen. Female FWs shorter than in males, very slightly overlapping, rounded posteriorly as in Macrobinthus and Rugabinthus. Ovipositor nearly as long as FIII; apex faintly denticulate dorsally. Copulatory papilla conical, with a wide basal sclerite and a well-developed apex slightly sclerotized and indented. Male FWs more resemble that of Macrobinthus, from which it slightly differs by main oblique vein partly differentiating a false mirror in harp posterior angle. Likewise, mirror only slightly differentiated, as in Macrobinthus. CuA clearly curved inwards posteriorly, as in Macrobinthus (almost straight in Rugabinthus). Some characters of male genitalia more resemble some species of Rugabinthus in that it is elongated and slender, its basal margin distinctly indented in the middle and the posterior apex spoon-like, without paired lophi. In contrast, Platybinthus differs from Rugabinthus by the male FWs with a shield-shaped harp (triangleshaped in Rugabinthus) and absence of strong diagonal fold carrying diagonal vein and cell c1, and separating FWs in two distinct areas (characteristic feature of Rugabinthus). The new genus also differs by hind legs distinctly longer (twice as long as body size). Etymology Genus named after the old genus name ‘ Platydactylus ’ to which several early described eneopterine species belonged, including the type species, before the name was recognised as an homonym of a genus of Squamata Oppel, 1811. Therefore we use the prefix ‘ Platy ’ (referring to Platydactylus) with the suffix ‘ binthus ’ referring to Lebinthina. Description Size medium for the subtribe (Figs 1–2, 4). Dorsum of head rounded, prolonged by rectangular fastigium distinctly wider than long (Fig. 4A–C). Head dorsum uniformly black (Fig. 5A–C). Eyes large and prominent. Head roundly triangular in facial view, almost as wide as high (Fig. 5D–F). Face finely rugulose, colouration usually cream-coloured and with lateral parts dark (Fig. 5D–F). Maxillary palpi with apical and subapical segments cream-coloured (slightly darkened apically), other segments black. Ocelli large, pale-coloured, forming a rather wide triangle; median ocellus rounded, lateral ocelli similar sized and rounded. Scapes small. Antenna with basal segments cream-coloured with tint of black, otherwise dark. Lateral part of head black with a narrow yellow band behind eyes (Fig. 6). Pronotum dorsal disk trapezoidal, on average 1.6 times as wide as long, its anterior margin distinctly emarginated in the middle, posterior margin straight, mostly black. Lateral lobes longer than wide, uniformly black (Fig. 6). Legs (Figs 1–2, 4): TI with two tympana; inner tympanum covered by a flat sclerotized expansion, its membrane visible along a small longitudinal slit only; outer tympanum ellipsoidal, its membrane transversally plicate in dorsal half. TI with two inner and two outer apical spurs. TII with two inner and two outer spurs. Hind legs very long (twice as long as body size); FIII muscular; TIII serrulated on their whole length, with four pairs of subapical spurs and three pairs of apical spurs; inner spurs long and curved, outer spurs shorter and straight. Legs I and II light brown to yellow brown, femora with brown spots and longitudinal patterns, tibiae with rings. TaIII-1 with 4–5 spines on dorsal outer edge and without any on dorsal inner edge. TIIIs and tarsomeres brown with spines and spurs with dark apices. FWs short, barely reaching abdomen mid-length; hind wings absent. Cerci well developed. Abdomen: tergites dark brown, with faint pale spots along posterior margins. Male Metanotal glands absent. Dorsal field of FWs slightly shorter than lateral field. FWs 1.4–1.6 times as long as wide, on average 1.7 times as long as pronotum dorsal disk. FW venation (Fig. 7A–B): 1A vein (file) transverse part straight, bisinuate anteriorly to angle, forming a notch in most species. Diagonal vein strong. Harp shield-shaped, with a characteristic fold, longer than wide, occupying approximately half of dorsal field surface, with a strong transverse, bisinuate harp vein, and a secondary one in anterior corner of harp. Cell c1 widened posteriorly. CuA anterior part strong throughout, slightly curved inward near apex. Mirror (d1) little differentiated from other cells of D alignment. Apical field very short, including one cell alignment posterior to mirror. Lateral field with four strong longitudinal veins ventral to Sc; Sc without projection. Subgenital plate elongated, clog-shaped. Epiproct and subgenital plate brown. GENITALIA (Fig. 8A–F). Pseudepiphallus elongate and slender, its basal margin distinctly indented in the middle; median part of pseudepiphallus forming a dorsal ovular disk dorsally; posterior apex elongated, without paired lophi, forming a median spoon-like expansion shorter than rest of pseudepiphallus. Rami short, shorter than half of pseudepiphallus length. Pseudepiphallic parameres average sized, ventral lobe strongly sclerotized. Ectophallic apodemes parallel and long, usually reaching beyond anterior margin of pseudepiphallic sclerite. Ectophallic arc well sclerotized, curved. Ectophallic fold with two ventral sclerites of variable shapes, extending posteriorly but not reaching posterior apex of pseudepiphallus. Endophallic sclerite comprising a long and straight anterior region, a short median expansion and short lateral arms posteriorly. Endophallic apodeme with well-developed lateral lamellae and a triangular dorsal median crest. Female FWs very short, not reaching base of first tergite, very slightly overlapping. Cells of dorsal field brown, dorsal field with strong orange longitudinal veins and very weak yellow transverse ones; lateral field with four longitudinal veins (Fig. 7C–D). Posterior corner between dorsal and lateral fields forming an acute angle. Dorsal field apex oblique. GENITALIA (Fig. 8G–H). Ovipositor rather short, apex faintly denticulate dorsally. Copulatory papilla conical, with a wide basal sclerite and a well-developed apex slightly sclerotized and indented. Distribution INDONESIA: Halmahera Island of Maluka and nearby island of Morotai. Included species Platybinthus punctatus (Brunner von Wattenwyl, 1898) gen. et comb. nov. from Halmahera Island, Indonesia. Platybinthus striolatus (Brunner von Wattenwyl, 1898) gen. et comb. nov. from Halmahera Island, Indonesia. Platybinthus sandyi gen. et sp. nov. from Morotai Island, Indonesia., Published as part of Robillard, Tony & Tan, Ming Kai, 2023, Platybinthus, a new genus of Lebinthina crickets (Orthoptera, Gryllidae, Eneopterinae) from Maluka, Indonesia, pp. 46-66 in European Journal of Taxonomy 856 (1) on pages 51-56, DOI: 10.5852/ejt.2023.856.2027, http://zenodo.org/record/7544313, {"references":["Brunner von Wattenwyl H. H. 1898. Orthopteren des Malayischen Archipels gesammelt von Prof. Dr. W. Kukenthal in den Jahren 1893 and 1894. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 24 (2): 193 - 288."]}

Platybinthus punctatus (Brunner von Wattenwyl, 1898) gen. et comb. nov. Figs 1, 3, 5A, D, 6A, 8G Piestodactylus punctatus Brunner von Wattenwyl, 1898: 278. Eurepa punctata – Kirby 1906: 89. — Chopard 1968: 355. Lebinthus punctatus – Robillard 2011: 58. Diagnosis Platybinthus punctatus gen. et comb. nov. differs from Platybinthus striolatus gen. et comb. nov. by larger size, and female copulatory papilla with apex narrowed and bilobed (instead of narrowed and obtusely rounded). Only faint differences of female FW venation and colouration with Platybinthus sandyi gen. et sp. nov. Material examined Syntype INDONESIA – Maluka • 1 ♀; Halmaheira [= Halmahera Island]; 1894; W. Kükenthal leg.; molecular sample L176; GenBank accession: OP684789 and OP686541; MNHN-EO-ENSIF1448. Additional specimens INDONESIA – Maluka • 1 ♀; Halmahera [Island], Gamkonora; 4 May 1931; G. Heinrich leg.; MNHN- EO-ENSIF1430. Remark The only type specimen examined from the original type series is a female from MNHN, but male syntypes exist according to the male measurements and the drawing of male FW venation presented in Brunner von Wattenwyl (1898). Due to the lack of male characters, the comparison with other species of the genus remains limited. Molecular data obtained for the three species of the genus support the fact that they should be considered as different taxa, despite the lack of detailed information on P. punctatus gen. et comb. nov. Further information will have to be provided by future studies. Redescription Female FWs reaching middle of fourth tergite, faintly overlapping. Cells of dorsal field dark brown, not translucent, with orange strong longitudinal veins and white weak transverse ones; with 9 veins on dorsal field, 4 on lateral field. Dorsal field apex truncated, lateral field posterior margin oblique. GENITALIA. Ovipositor as long as FIII, apex not denticulate dorsally. Copulatory papilla rounded; basal sclerite forming a semi-circular ring with a small basal plate; bent at apical third in lateral view; apex narrowed and bilobed (Fig. 8G). Measurements See Table 1. Ecology Unknown. Distribution INDONESIA: Maluka: Halmahera Island. Type locality INDONESIA: Maluka: Halmahera Island. Calling song Unknown., Published as part of Robillard, Tony & Tan, Ming Kai, 2023, Platybinthus, a new genus of Lebinthina crickets (Orthoptera, Gryllidae, Eneopterinae) from Maluka, Indonesia, pp. 46-66 in European Journal of Taxonomy 856 (1) on pages 56-58, DOI: 10.5852/ejt.2023.856.2027, http://zenodo.org/record/7544313, {"references":["Brunner von Wattenwyl H. H. 1898. Orthopteren des Malayischen Archipels gesammelt von Prof. Dr. W. Kukenthal in den Jahren 1893 and 1894. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 24 (2): 193 - 288.","Kirby W. F. 1906. A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). Vol. 2. Orthoptera Saltatoria. Part I. (Achetidae et Phasgonuridae). British Museum (Natural History), London. https: // doi. org / 10.5962 / bhl. title. 6745","Chopard L. 1968. Fam. Gryllidae: Subfam. Mogoplistinae, Myrmecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Fam. Oecanthidae, Gryllotalpidae. In: Beier M. (ed.) Orthopterorum Catalogus 12: 213 - 500. Uitgeverij Dr. W. Junk N. V. ' s, Gravenhage.","Robillard T. 2011. Centuriarus n. gen., a new genus of Eneopterinae crickets from Papua (Insecta, Orthoptera, Grylloidea). Zoosystema 33: 49 - 60. https: // doi. org / 10.5252 / z 2011 n 1 a 2"]}

Discotettix (Discotettix) belzebuth (Serville, 1838) (Figs 5–14) Vernacular name: Bornean Spiky Pygmy Devil Tetrix belzebuth Serville, 1838: 759 [original description, type locality: originally Java but actually probably Borneo]. Tettix belzebuth: Stål 1873: 152 [listed in the catalog]. Discotettix belzebuth: Bolívar 1887: 306 [included in the revision]; Rehn, 1904: 670 [new records]; Hancock 1907a: 6 [included in the revision]; Hancock 1907b: 213 [new records]; Kirby 1910: 2 [included in the catalog]; Hancock 1913: 39 [new records]; Willemse 1930: 8 [new records]; Günther 1938: 301 [included in the revision]; Steinmann 1970: 216 [listed in the catalog]; Blackith 1992: 46 [listed in the catalog]; Yin et al. 1996: 866 [listed in the catalog], Otte 1997: 32 [listed in the catalog], Kočárek et al. 2015: 289 [new records, data on variability]; Kuřavova et al. 2017: 120–128 [data on the antennae morphology]; Tan & Wahab 2018: 123 [new records in Brunei Darussalam]. Discotettix armatus Costa, 1864: 59 [original description, type locality: Borneo]; synonymized by Bolívar (1887). Discotettix adenanii Mahmood et al., 2007: 1276 [original description, type locality: Borneo: Kuching]; synonymized by Kočárek et al. (2015). Type locality. According to the original description the type locality is Java, but since no locality label is present under the holotype (the only found specimen originating from Serville's collection in MNHN), we believe that the specimen originates from Borneo, from where the majority of the records of this species are. In MNCN, there are four specimens labeled ‘Java’ so it is not fully clear whether this species is/was present on Java. Borneo Island is the type locality of D. armatus, which is the type species of the genus Discotettix (a junior synonym of D. belzebuth). The type specimen of D. belzebuth was considered lost, but was found in March 2016 by JS in MNHN. For D. armatus, we believe that the specimen still exists in the Naples collection (Italy), but we were not able to contact the Museum or to get information on Costa's collection. Material examined. Type material. HOLOTYPE of D. belzebuth 1♀ (locality, date and collector labels missing), red label 'TYPE' and Günther's label ' Discotettix belzebuth Serv. K. Günther det.' present (MNHN); PARATYPE of D. adenanii 1♂ Malaysia: Sarawak: “Adanan Bukra”, Gunung Serapi 3.VI.1988. (UKM) (according to Kočárek et al. (2015) the holotype and the rest of the type series has probably been destroyed). Additional museum material. 2♀♀, 1♂ Borneo: Malaysia: Sarawak: Mattang Collector Frivaldsky, det. J. Skejo (MNCN); 1♀, 2♂♂ Borneo: Malaysia: Sarawak Collector Mjöberg, det. J. Tumbrinck (NHRS); 1♀ Borneo: Malaysia: Sarawak: Bidi 1908. Collector. C. J. Brooks, det. J. Tumbrinck (NCM); 1♂ Borneo: Malaysia: [Sarawak, Bidi] Collector: C. J. Brooks, det. J. Tumbrinck (NCM); 1 ♀ Borneo: Malaysia: Sarawak: 75 km S of Miri Town, Niah Nat. Park [100 m a.s.l., forest around Niah Great Cave] 30.III.2012. Collectors A.V. Gorochov and M. Berezin, det. J. Skejo et T. Pushkar (ZISP); 2♀♀ Borneo: Malaysia: Sabah: Kina-Balu-Gebirge Collector: Waterstradt, det. Hancock (AMS); 1♀ Borneo: Malaysia: Sabah: Kinabalu NP: Poring [forest clearing] 8.VIII.1984. Collector S. Ingrisch, det. S. Ingrisch (CJT); 2♀♀, 2♂ ♂ Malaysia: Sabah [labeled Nord-Borneo Collector Waterstradt det. J. Skejo (MNCN); 3♂♂ Borneo: Malaysia: Sabah: Pajau River, Collector Mjöberg, det. J. Tumbrinck (NHRS); 1♂ Borneo: Malaysia: Sabah: Kajan River Collector Mjöberg, det. J. Tumbrinck (NHRS); 1♀, 2♂♂ Malaysia: Sabah [on original label written North Borneo] Collector: Watterstradt, det. K. Günther (SMTD); 1♀ Borneo: Malaysia: Sabah: Batu Niah 4.VIII.1984. Collector S. Ingrisch, det. S. Ingrisch (ZFMK); 1♀ Borneo: Malaysia: Sabah: Kinabalu NP: Poring: Bergil 10.IV.1997. [6º5'N, 116º33'E] [collected by fogging] Collector A. Floren, det. J. Tumbrinck (ZFMK); 1 ♀ Malaysia: Sabah: Crocket Range, 80 km S of Kota Kinabalu City [environment of the village Ula Kumanis at 800 m a.s.l.] 5 – 10.V.2006. Collector A. Sochivko, det. J. Skejo et T. Pushkar (ZISP); 1♀, 3♂ Borneo: Malaysia: Sabah: [original labels „ Nord Borneo, Kina-Balu-Gebirge [at 1500 m a.s.l.] Collector Waterstradt, det. Brunner von Wattenwyl (ZISP); 1♀ Malaysia: Sabah: Mt. Trus Madi: Tambunan distr [975 m a.s.l.] 25.IV–10.V.2006, Collector P. Udivichenko, det. J. Skejo et T. Pushkar (ZISP); 4 ♂♂ Malaysia: Sabah: Mt. Trus Madi: Tambunan district [975 m a.s.l.] 25.IV–10.V.2006. Collector P. Udivichenko, det. J. Skejo et T. Pushkar (ZISP); 2♀, 2♂ Borneo: Malaysia: Sabah: Mt. Trus Madi [1000 m a.s.l.] 13.–25.V.2007. Collector A.V. Gorochov, det. J. Skejo et T. Pushkar (ZISP); 1 ♀ Borneo: Malaysia: Sabah: Mt. Trus Madi [1200 m a.s.l.] 13–24.I.2007. Collector A. Sochivko, det. J. Skejo et T. Pushkar (ZISP); 1♂ label '99. 10064.' [without specified data] det. J. Skejo (MNCN); 1♂ [without specified data] det. J. Skejo (MNCN); 1♀, 1♂ Malaysia: Borneo [without specified other data] (NMNH NASU); 2♀♀ Indonesia: Kalimatan: Kalimantan Timur [= East]: Marah 12.XI.1925. Collector: H. C. Siebers, det. K. Günther (SMTD); 1 nymph (sex unteterminable) Indonesia: Kalimatan: West Kalimatan: Pontianak det. J. Skejo (MNCN); 2♀♀, 2♂ ♂ Indonesia: Java [without other data specified] det. J. Skejo (MNCN); Additional material from online social media. All the records of D. belzebuth from iNaturalist may be found on the following link: https://www.inaturalist.org/observations?taxon_id=637240 (all records submitted by July 2022 are shown in Table 1); while for Flickr, ProjectNoah, and SpinelessWonders all the observations (Figs 6–14) are listed in the Table 1. ......Continued on the next page TABLE 1. (Continued) ......Continued on the next page TABLE 1. (Continued) Distribution. The species is widespread in Borneo (many data from Malaysian and Brunei part, but few from the Indonesian part, probably due to the lack of research) and adjacent small islands (such as Labuan). This species was previously considered widely distributed on all the Greater Sunda Islands of the Malay Archipelago, namely on Borneo, Java, and Sumatra (Serville 1838; De Haan 1843; Rehn 1904; Hancock 1907a; 1907b; Günther 1938; Mahmood et al. 2007; Kočárek et al. 2015; Tan & Wahab 2018). Distribution of the species in Java was/is possible, as in the Bolívar's collection in MNCN Madrid there are several specimens labeled ‘Java’, but its presence nowadays should be confirmed as there have been no records for more than a century. The species is not present in Sumatra (De Haan 1843), with De Haan’s (1843) records likely belonging to D. selysi, which was not described at the time. Taxonomic notes. Discotettix adenanii is considered a junior synonym of D. belzebuth. Seven well-developed pronotal projections (Mahmood et al. 2007) are the diagnostic character of D. belzebuth as well. Günther (1938) pointed out the variability of this polymorphic species with certain specimens that are not easy to identify as D. belzebuth, but as a form with certain characteristics somewhat similar to Kraengia. The description of D. adenanii, on the other hand, completely fits D. belzebuth description and diagnosis. Variability of the species can be observed solely by comparison of Bolívar's drawing (Hancock, 1907a: 6) with Günther's D. belzebuth figure (1938: p. 301, Fig. 2). Kočárek et al. (2015) examined type specimen of D. adenanii and the variability of the pronotal morphology in D. belzebuth populations collected in different parts of Borneo, showing that all the studied specimens fit the variability of D. belzebuth. We agree with the conclusion of Kočárek et al. (2015) that D. adenanii is a pure synonym of D. belzebuth. Diagnosis. This is the species with the highest (in lateral view) and spikiest/sharpest pronotal projections in the entire genus and by its general appearance it can readily be distinguished from all the other species of the genus. It is somewhat similar to D. kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n. from NE Borneo, and to D. sumatrensis Skejo, Pushkar et Tumbrinck sp. n. from Sumatra. From D. scabridus the new species can be separated by the following characters: (I) bifurcation of the frontal costa between the eyes (on the lower margin of the compound eyes in D. scabridus), (II) FM high and developed (present as a small tubercle in D. scabridus), (III) MM elevated into spines (lower and saw-like in D. scabridus), (IV) MML elevated into spines (present as low and triangular, compressed elevations in D. scabridus), (V) ML strong and long, tooth-like (absent in D. scabridus), (VI) interscapular area with parallel margins (wide and triangular in D. scabridus). Discotettix belzebuth can be separated from D. selysi, D. doriae, and D. aruanus Skejo, Pushkar et Tumbrinck sp. n. by the following characters: (I) more than one well-developed protuberance on the pronotal disc, (II) long frontomedial projection of the anterior margin of the pronotum, (III) fore and mid femora are more slender, (IV) antennal segments are more specialized in morphology than in D. selysi, the widest and the most flattened is the 10 th segment (unlike the 9 th segment in D. selysi and D. doriae). From D. sumatrensis Skejo, Pushkar et Tumbrinck sp. n. the species can be easily separated by the following characteristics: (I) larger body size, (II) all antennae segments are of black/dark color, (III) the 8 th antennal segment is the widest one (unlike the 7 th segment in D. sumatrensis Skejo, Pushkar et Tumbrinck sp. n.), (IV) the hind femur bears small lappets. D. kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n. can be easily separated from D. belzebuth by the following set of characters: (I) in D. kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n. antennae with more robust segments gradually become wider distally, not as specialized as in D. belzebuth, (II) D. kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n. has short FM, not covering the whole head as in D. bellzebuth, (III) projections on the dorsal surface of the pronotum are smaller in D. kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n., (IV) D. kirscheyi sp. n. has weak ML, (V) D. kirscheyi sp. n. has robust and serrated fore and mid femora, not as slender as in D. belzebuth and (VI) D. kirscheyi sp. n. has smaller body size. Redescription. General characters. Medium to large sized, robust, species (13.01–17.02 mm), texture granulated, rugose; pronotum wrinkled, with numerous small tubercles, medium-sized and large protuberances on dorsal and lateral sides. Epizoic symbiotic bryophytes and algae are often present on the pronotal surfaces, so usually the specimens are characterized by cryptic colors (Figs 8, 10). Macropronotal. Coloration. Coloration variable: from black and dark brown to brighter tints of brown: grayish, greenish, yellowish, reddish, somewhat purple; pronotal projections usually darker. The entire body, including antennae, may be of the same color, except brightly colored frontal carina on the head and the median carina of pronotum, while fore and mid tibiae and tarsi usually bear 1–3 lighter rings. Maxillary palpi black with pale colored joints between the segments (Figs 5–14). Head. In dorsal and frontal view, vertex 2.21–2.76 times as wide as an eye. Fossula elliptic and deep. The lower margin of the lateral ocelli a bit below the level of the lower margin of a compound eye. In frontal view, frontal costa narrow, bifurcated above lateral ocelli into subparallel, finely granulated facial carinae forming a very narrow scutellum. Scutellum slightly narrower than the antennal groove. Antennal groove below the lower margin of the compound eye (Fig. 5C). Antenna with 13 antennomeres. A detailed description of the antennal morphometrics and morphology was published recently by Kuřavová et al. (2017). Antennal segments as follows: scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 6 th) elongated and circular in cross-section; central or subapical segments (7 th and 8 th) strongly widened, pennate, 8 th being the widest antennal segment (in holotype 2.22 times as long as wide); apical segment 9 th much smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th reduced, very small, and borders between them barely visible. Antennomeres 3 rd to 9 th bearing saw-like margins, because of the presence of large basiconic sensilla (Fig. 10). When the body and the antennae are covered with algae and moss, the 8 th segment is always free of epizoic organisms (Fig. 10). Pronotum. Pronotum wrinkled and granulated, covered in numerous small tubercles and larger projections. The posterior process of the pronotum slender, surpassing the hind knee for about half of the hind femur length. Disc of the pronotum at almost the same level along all length, and unlike other species of the genus, without a distinct depression behind the shoulder. Pronotum not descending backward. Morphology of the pronotal disc variable, usually with 4–7 unpaired projections of variable size on the median carina (FM, PM, and 2–5 MMs); 2–3 pairs of FL; 4–7 pairs of more or less distinct PMLs and MMLs; 3 pairs of PLs and MLs; and a pair of VL. Prozona short, subsquare. The anterior margin of the pronotum projected as a large digitate FM directed upwards and forwards above the head, covering the whole area of the fastigium of the vertex. FM often decurved with excised apex. Prozonal carinae slightly elevated, surpassing the anterior margin of the pronotum as dentiform FL1. Extralateral carinae elevated, surpassing anterior margin of pronotum as dentiform FL2. FL3 dentiform, small and weak, more distinct than FL1. Median carina extended along the whole length of the pronotum, tuberculated, but with smooth areas. Median carina bearing FM and 3–6 of large digitate medial projections well seen in profile. PM small, distinct in about a third of the examined specimens. MM1, MM2, MM3, and MM4 large and digitate (MM1MM3>MM4). MM4 well visible in most of the specimens, but in some completely reduced. MM5 present as a spike in about every tenth specimen, but usually very small and almost invisible. Among mediolateral projections, PML1 is a small tubercle, almost indistinct; PML2 small; MML1 small; MML2 large; MML3 and MML4 distinct in a few examined specimens. PL1 and PL2 elongated, small, and almost indistinct. In the metazona humero-apical carinae forms a sharp humeral angle, projected outwards as strong spine-like or digitate ML. Interhumeral carinae hardly observable because of numerous net-like elevations and tubercles present in the whole disc. The apex of pronotum blunt, and shallowly excised. The lower part of the lateral lobe with finely serrate anterior and coarsely serrate posterior margin, elongated as VL of variable shapes from sharp spine-like to saw-like form, directed strongly outwards or in rare cases somewhat backward (Fig. 5A, B). Wings. The visible part of tegmen elongated and oval. Hind wing long, often not reaching the pronotal apex. Legs. Femora more or less robust, and compressed laterally, but elongated in comparison to other species. The rough surface of the legs usually bears outgrowths and tubercles of variable size and sharpness. The dorsal and ventral margins serrate. Genicular teeth visible on all the knees, and additionally 1–3 weak teeth present on the dorsal and ventral margins of the fore and mid femora. Hind femur with small lappets on both dorsal and ventral margins. Lateral area of the hind femur with net-like elevations and weak carinae, ventro-external carina with teethlike outgrowths. Genicular teeth of the hind femora larger than the antegenicular. Both sides of the dorsal margin of the hind tibia finely serrated, additionally with 4–5 outer and 3–4 inner larger teeth. Abdominal apex. Female subgenital plate with a triangular protrusion in the middle of the posterior margin. Ovipositor of variable shapes, usually elongated, but can be more robust, probably due to the ecological factors. Measurements. BL ♂♂ 13.1–15.01 mm, ♀♀ 14.3–17.02 mm; PnL ♂♂ 16.04–19.14 mm, ♀♀ 18.5–22.34mm; PnW ♂♂ 7.99–8.24 mm, ♀♀ 9.04–9.66 mm; AnL ♂♂ 6.88–7.8 mm, ♀♀ 7.11–7.99 mm; TL ♂♂ 1.9–2.54 mm, ♀♀ 2.1–3.01 mm; TW ♂♂ 0.78–1.14 mm, ♀♀ 0.99–1.23 mm; fFL ♂♂ 3.29–4.46 mm, ♀♀ 4.1–5.16 mm; fFW ♂♂ 0.77–1.01 mm, ♀♀ 0.89–1.02 mm; mFL ♂♂ 3.49–4.72 mm, ♀♀ 4.2–5.09 mm; mFW ♂♂ 0.98–1.02 mm, ♀♀ 0.99–1.11 mm; hFL ♂♂ 6.99–10.03 mm, ♀♀ 8.1–10.28 mm; hFW ♂♂ 2.28–3.45 mm, ♀♀ 2.26–3.24 mm; OvL ♀♀ 1.38–2.11 mm; AnL/fFL ♂♂ 1.58–2.09, ♀♀ 1.55–1.71; VW ♂♂ 1.02–1.48 mm, ♀♀ 1.35–1.91 mm; EW ♂♂ 0.38–0.49 mm, ♀♀ 0.46–0.82 mm; VW/EW ♂♂ 2.21–2.52, ♀♀ 2.31–2.76; SW ♂♂ 0.31–0.37 mm, ♀♀ 0.32– 0.42 mm; AgW ♂♂ 0.19–0.34 mm, ♀♀ 0.29–0.41 mm; ScW ♂♂ 0.21–0.29 mm, ♀♀ 0.23–0.28 mm; SW/AgW ♂♂ 1.46–1.51, ♀♀ 1.41–1.56; SW/ScW ♂♂ 1.20–1.47, ♀♀ 1.21–1.35; As—L/W ♂♂ 2.41–3.1, ♀♀ 2.39–3.12; PrzW ♂♂ 3.35–3.59 mm, ♀&fema, Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 15-26, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Serville, J. G. A. (1838) Histoire naturelle des insectes. Orthopteres. Roret, Paris, 776 pp. https: // doi. org / 10.5962 / bhl. title. 95609","Stal, C. (1873) Recensio Orthopterorum. P. A. Norstedt and Soner, Stockholm, 152 pp.","Bolivar, I. (1887) Essai sur les Acridiens de la tribu des Tettigidae. Annales de la Societe Entomologique de Belgique, 31, 175 - 313.","Rehn, J. A. G. (1904) Studies in the Orthopterous subfamilies Acrydiinae (Tettiginae), Eumastacinae and Proscopinae. Proceedings of the Academy of Natural Sciences, Philadelphia, 56, 658 - 683.","Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Hancock, J. L. (1907 b) Studies of the Tetriginae (Orthoptera) in the Oxford University Museum. Transactions of the Entomological Society of London, 1907, 213 - 244. https: // doi. org / 10.1111 / j. 1365 - 2311.1907. tb 01760. x","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Hancock, J. L. (1913) Studies of Tetriginae (Acrydinae) from Sarawak Museum, Borneo. The Sarawak Museum Journal, 1 (3), 39 - 54.","Willemse, C. J. M. (1930) Fauna Sumatrensis (Bijdrage Nr. 62). Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie, 73, 1 - 210.","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Steinmann, H. (1970) Check-list of the Tetricidae (Orthoptera) of the Oriental faunal region. Acta Zoologica Academiae Scientiarum Hungaricae, 16, 215 - 240.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261.","Kocarek, P., Kuravova, K., Musiolek, D., Wahab, R. A. & Kahar, S. R. A. (2015) Synonymy of Discotettix adenanii Mahmood, Idris & Salmah, 2007 with D. belzebuth (Serville, 1838) (Orthoptera: Tetrigidae). Zootaxa, 4057 (2), 288 - 294. https: // doi. org / 10.11646 / zootaxa. 4057.2.10","Kuravova, K., Wahab, R. A. & Kocarek, P. (2017) External morphology of the antennae and sense organs of the groundhopper Discotettix belzebuth (Orthoptera, Tetrigidae). Zoologisher Anzeiger, 266, 120 - 128. https: // doi. org / 10.1016 / j. jcz. 2016.11.003","Tan, M. K. & Wahab, R. A. (2018) Preliminary study on the diversity of Orthoptera from Kuala Belalong Field Studies Centre, Brunei Darussalam, Borneo. Journal of Orthoptera Research, 27 (2), 119 - 142. https: // doi. org / 10.3897 / jor. 27.24152","Costa, A. (1864) Acquiste fatti durante l-anno 1862. Annuario del Museo zoologico della Universita di Napoli, 2, 8 - 94.","Mahmood, K., Idris, A. B. & Salmah, Y. (2007) Tetrigidae (Orthoptera: Tetrigoidea) from Malaysia with the description of six new species. Acta Entomologica Sinica, 50 (12), 1272 - 1284.","De Haan, W. (1843) Bijgragen tot de kennis der Orthoptera. In: Temminck, D. (Ed.), Verhangelingen over de Natuurlijke Geschiedenis der Nederlansche Overzeesche Bezittingen, de Leden der Natuurkundige Commissie in Indie en andere Schrijvers. s. n., Leiden, 165 - 228."]}

Discotettix (Discotettix) doriae Bolívar, 1898 stat. resurr. (Fig. 18) Vernacular name: Mentawai Unicorn Pygmy Devil Discotettix doriae Bolívar, 1898: 80 [original description, type locality: Mentawai: Sipura]; Hancock 1907a: 6 [listed in the catalog]; Kirby 1910: 2 [listed in the catalog]; Willemse 1930: 207 [listed in the catalog]; París 1994: 236 [data on type series]; Muhammad et al. 2018: 20. Discotettix selysi (partim): Günther 1938: 302 [synonymized D. doriae with D. selysi; it is not accepted here]; Blackith 1992: 46 [listed in the catalog]; Yin et al. 1996: 866 [listed in the catalog]; Otte 1997: 32 [listed in the catalog]. Type locality. Indonesia: Sumatra: Mentawai: Sipura Island Material examined. Type material. LECTOTYPE (Fig. 18) 1♀ Mentawei: Sipora, Sereinu [= Indonesia: Mentawai: Sipura island] V.–VI.[18]84. Leg. E. Modigliani (MCSN); PARALECTOTYPE 1♀ Mentawei: Sipora, Sereinu [= Indonesia: Mentawai: Sipura island] V.–VI.[18]84. Leg. E. Modigliani (MNCN). Distribution. This species inhabits the rainforest of the small island of Sipura (845 km 2), the smallest of the four large islands of the Mentawai Archipelago, west of Sumatra. Only two females of this species have been known hitherto. Diagnosis. This species is morphologically similar to D. selysi from Sumatra and the Malayan peninsula and to D. aruanus sp. n. from the Aru Islands because of the presence of low projections, or absence of the high ones, flat dorsum of the pronotum, and coloration. It seems to be closely related to D. selysi, but can be easily recognized from the latter by the following set of characters: (1) FM small and narrow, covering vertex only partially (in D. selysi it is large, long, and covering entire vertex), (2) smaller body size (females have a slightly shorter body and pronotum than the ones of D. selysi), (3) shorter and stouter antennae with swollen 6 th, 7 th and 8 th antennal segments (no swollen segments in D. selysi), and (4) stronger teeth of the fore and mid femora. From D. aruanus sp. n. from Aru Island, which also seems related to D. selysi and D. doriae, D. doriae can be easily told apart by the much smaller FM (large and reaching in front of the head in D. aruanus sp. n.), and much stouter and more armed fore and mid femora (less armed in D. aruanus sp. n.). From D. sumatrensis sp. n. inhabiting neighboring Sumatra, the species can be separated by (1) dark antennae with stouter antennomeres, (2) lower projections, especially FM and MMs, (3) larger body size (approximately 2 mm longer), and (4) stouter femora. For comparison with other species (D. belzebuth and D. kirscheyi from Borneo and D. scabridus from the Philippines), please consult the diagnosis of D. selysi, where a comparison to all the species of the genus Discotettix has been given, as well as the detailed diagnoses of D. scabridus and D. belzebuth containing comprehensive comparisons. Redescription. (Fig. 18) General features. Medium-sized (about 15 mm), robust species. Body finely granulated; pronotum rugose, with numerous small tubercles and net-like elevations, in parts smooth and without tubercles, anterior part of the pronotum bearing a few larger protuberances (Fig. 18). Macropronotal. Coloration. Body dark brown. Carinae and projections darker than the rest of the body. Median pronotal carina with orange patches. Antenna dark brown, without pale colored parts. Maxillary palpi dark brown. The visible part of the tegmen dark brown and without any pale spots. Legs brown except for the pale rings on tibiae and tarsi. The body probably covered with epizoic symbionts, just as in other species, giving an animal greenish camouflage when alive, after preservation in alcohol and/or drying the green color disappears. Head. In dorsal and frontal view, vertex 2.21–2.36 times as wide as the eye. Lateral carinae raised and granulated. Fossula deep. Lateral ocelli situated just below the level of the lower margin of the compound eye. Antennal groove significantly below the lower margin of the compound eye. In frontal view, frontal costa bifurcated into facial carinae, forming a scutellum with narrow and parallel borders, bifurcation just above the level of lateral ocelli. Antennal groove slightly wider than frontal costa. Antenna 13-segmented, all segments robust in comparison to other species of the genus: scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 5 th) robust, less elongated than in other species of the subgenus Discotettix, and circular in cross-section; basal segment 6 th widened and swollen; central or subapical segments (7 th and 8 th) strongly widened, pennate and swollen, 8 th being the widest antennal segment (about 2.6 times as long as wide); apical segment 9 th elongated and pennate, but smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th reduced, very small, and borders between them barely visible. Antennomeres 3 rd to 9 th bearing saw-like margins, but weaker than in the other species of the subgenus (Fig. 18C). Pronotum. Pronotum rugose, granulated, with numerous tubercles and net-like elevations, somewhere smooth and without tubercles (some parts of the pronotal disc and parts of the median carina of the pronotum). The anterior margin of the pronotum bearing a medium-sized FM, smaller than in the other species of the subgenus Discotettix. The posterior process of the pronotum surpassing the hind knees for at most a half-length of a hind femur. Disc of the pronotum with a small depression between the tegminal bases, then slightly elevated again. Caudad, pronotum gradually descending backward. Net-like elevations of omnipresent on the pronotal disc, all along the median carina of the pronotum and connecting it to all other carinae and the projections of the dorsal surface of the pronotum. Netlike elevations very distinct in the interhumeral region of the disc. The median carina of the pronotum continuous from the anterior margin to the pronotal apex, undulated irregularly because of the projections. The median carina of the pronotum bearing four unpaired projections of variable size: medium-sized digitate FM, directed upwards and forwards and covering the fastigium of the vertex (but not completely); PM small and triangular; MM1 large and triangular; MM2 and MM3 evident, but decreasing in size caudad, MM4 reduced. Prozona subsquare. Prozonal and extralateral carinae in prozona distinct, surpassing anterior margin of pronotum as dentiform FL1 and FL2, with FL2 more distinct. FL3 dentiform, small, and weak. Among the mediolateral projections PML1 more or less distinct; PML2 very small; MML1 small; MML2 large; MML3 v ery small; MML4 and MML5 wanting. PL1 and PL2 present as small triangular tubercles forming posterior elongation of the extralateral carinae in the line joining the humero-apical carinae. Humeral angle obtuse with pointed apex. ML present, but reduced. Pronotal apex narrow, shallowly excised. The lower part of the lateral pronotal lobe with serrate anterior and posterior margins, elongated as spine-like VL, directed strongly outwards and backward (Fig. 18A, B). Wings. Tegmina and alae present and visible. The visible part of tegmen elliptical, granulated, and tuberculated. Hind wing not reaching apex of pronotum, a few millimeters of length are lacking to touch the tip. Legs. Fore and mid femora robust, compressed laterally, dorsal and ventral margins serrated; with genicular tooth on the knees and additionally with two to three sharp teeth on the dorsal and ventral margin. Mid femora with stronger teeth than the fore femora. Hind femur with lappets, and a small protuberance situated on the ventral external carina. The genicular tooth large, while the antegenicular one small. Both sides of the dorsal margins of the hind tibia finely serrated, with 3–4 outer and 1–3 inner large teeth. Abdominal apex. Female subgenital plate in ventral view with a triangular protrusion in the middle of the posterior margin. Ovipositor robust, dorsal valvae robust, ventral slender, and serrate. Cerci robust, hairy, with a long tip. Measurements (♀♀ only). BL 14.51–14.79 mm; PnL 16.06–17.01 mm; PnW 8.45–8.61 mm; AnL 6.88– 7.01 mm; TL 1.93–2.11 mm; TW 0.75–0.91 mm; fFL 3.86–4.01 mm; fFW 0.78–0.82 mm; mFL 3.47–3.61 mm; mFW 0.86–0.93 mm; hFL 7.59–8.01 mm; hFW 2.68–2.73 mm; OvL 1.62–1.71 mm; AnL/fFL 1.75–1.81; VW 1.18–1.23 mm; EW 0.51–0.55 mm; VW/EW 2.21–2.36; SW 0.26–0.31 mm; AgW 0.31–0.34 mm; ScW 0.21–0.27 mm; SW/AgW 0.67–0.81; SW/ScW 1.19–1.24; As–L/ W 2.59–2.63; PrzW 3.44–3.51 mm; PrzL 2.22–2.38 mm; Prz–W/L 1.47–1.60; TL/TW 2.57–2.90; mFW/TW 1.08–1.09; fFL/fFW 4.03–4.12; mFL/mFW 4.01–4.51; hFL/ hFW 2.91–3.22; T1L/T3L 1.02–1.04., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 30-33, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Bolivar, I. (1898) Contributions a l'etude des Acridiens especes de la Faune indo et austro-malaisienne du Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 39, 66 - 101. https: // doi. org / 10.5962 / bhl. part. 9541","Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Willemse, C. J. M. (1930) Fauna Sumatrensis (Bijdrage Nr. 62). Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie, 73, 1 - 210.","Paris, M. (1994) Catalogo de tipos de ortopteroides (Insecta) de Ignacio Bolivar, I: Blattaria, Mantodea, Phasmoptera y Orthoptera (Stenopelmatoidea, Rhaphidophoroidea, Tettigonioidea, Grylloidea, Tetrigoidea). Eos, Revista espanola de Entomologia, 69, 143 - 264.","Muhammad, A. A., Tan, M. K., Abdullah, N. A., Azirun, M. S., Bhaskar, D. & Skejo, J. (2018) An annotated catalogue of the pygmy grasshoppers of the tribe Scelimenini Bolivar, 1887 (Orthoptera: Tetrigidae) with two new Scelimena species from the Malay Peninsula and Sumatra. Zootaxa, 4485 (1), 1 - 70. https: // doi. org / 10.11646 / zootaxa. 4485.1.1","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261."]}

Tribe Discotettigini Hancock, 1907 stat. resurr. Discotettigiae: Hancock 1907a: 5; 1907b: 213; Willemse 1930: 4, 7; Günther 1938: 301. Discotettigini: Kevan 1966: 380. Discotettiginae: Steinmann 1970: 216; Storozhenko 2013: 158; Tumbrinck 2014: 349. Discotettinae: Otte 1997: 32; Mahmood et al. 2007: 1275. Discotettigidae: Liang & Zheng 1998: 23; Zheng 2005: 15; Deng et al. 2007: 400. Type genus: Discotettix Costa, 1864. Differential diagnosis. The tribe Discotettigini is a sister tribe to Scelimenini; their representatives share many apomorphies inherited from the common ancestor. The main difference is the shape of the pronotum (elongated, flat, and hydrodynamic in Scelimenini; and robust, wrinkled, and cryptic in Discotettigini; Rebrina et al. in preparation), as well as the shape of the fore and mid femora (elongated in Scelimenini, short and toothed in Discotettigini), and the shape of the hind tibia and tarsi (widened into a paddle in Scelimenini, while of regular shape in Discotettigini). Scelimenini members have shorter projections of the pronotal disc in comparison to Discotettigini members. Discotettigini are mostly corticolous, and Scelimenini members are mostly amphibious grasshoppers (e.g., Muhammad et al. 2018). Description. Head. Antenna with 11–15 antennomeres, filiform or with widened preapical segments (Fig. 1). Frontal costa visible above the bifurcation; the bifurcation and the lateral ocelli placed low, between the compound eyes, in line with the lower margin or visibly below the compound eye; the antennal groove located below the lower margins of the compound eyes; lateral carinae of the vertex more or less elevated; anterior margin of the vertex truncated, slightly indrawn from the level of the outer margin of the compound eyes; fossula present; medial carina of the vertex visible in the upper third cephalad (Fig. 5C). Pronotum. Prozonal, extralateral, median, humeral and lateral carinae present; interhumeral carinae present, but sometimes not visible because of the pronotal projections in their place. Pronotum strongly granulated and wrinkled, armed with numerous strong projections (FM, FL, PM, MM1, MM2, MML1, MML2, and ML) that can be of different shapes and sizes: low, high, and wart-like or high triangular, saw-like, or spine-like protuberances. Paranota triangular, laterally projected, usually bearing strong VL projection, sometimes without a spine (Figs 2, 5A, B). Legs. Dorsal and ventral margins of all the legs with small, medium-sized, or large teeth. Tibiae rectangular in cross-section, not widened into the paddles. The dorsal margin of the fore and mid femora carinated. Hind tarsi not widened into a paddle (Fig. 29B). Composition and distribution. The tribe gathers corticolous Scelimeninae genera with an undulated pronotum, and without widened hind tibiae. Altogether, 10 genera and 68 species are herewith assigned to this tribe, and these are Austrohancockia (19 species in PR China, China, and Taiwan), Bidentatettix (2 species in PR China), Disconius gen. n. (1 species in Borneo) Discotettix (7 species in Peninsular Malaysia, Sumatra, Borneo, Mindanao, Aru), Eufalconius (1 species in Peninsular Malaysia), Gavialidium (2 species in Sri Lanka and Southern India), Gibbotettix (13 species in PR China), Kraengia (1 species in Sulawesi), Paragavialidium (14 species in PR China), and Tegotettix (8 species in Indochina, Borneo, Philippines, Sulawesi, New Guinea) (Muhammad et al. 2018, this study)., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on page 10, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Hancock, J. L. (1907 b) Studies of the Tetriginae (Orthoptera) in the Oxford University Museum. Transactions of the Entomological Society of London, 1907, 213 - 244. https: // doi. org / 10.1111 / j. 1365 - 2311.1907. tb 01760. x","Willemse, C. J. M. (1930) Fauna Sumatrensis (Bijdrage Nr. 62). Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie, 73, 1 - 210.","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Kevan, D. K. (1966) Some Orthoptera-Caelifera from the Philippine, Bismarck and Solomon Islands, with a few interesting records from New Guinea and the Moluccas. Entomologiske Meddelelser, 34, 375 - 420.","Steinmann, H. (1970) Check-list of the Tetricidae (Orthoptera) of the Oriental faunal region. Acta Zoologica Academiae Scientiarum Hungaricae, 16, 215 - 240.","Storozhenko, S. Y. (2013) Review of the subfamily Tripetalocerinae Bolivar, 1887 (Orthoptera: Tetrigidae). Zootaxa, 3718 (2), 158 - 170. https: // doi. org / 10.11646 / zootaxa. 3718.2.4","Tumbrinck, J. (2014) Taxonomic revision of the Cladonotinae (Orthoptera: Tetrigidae) from the islands of South-East Asia and from Australia, with general remarks to the classification and morphology of the Tetrigidae and descriptions of new genera and species from New Guinea and New Caledonia. In: Telnov, D., Barclay, M. V. L. & Pauwels, O. S. G. (Eds.), Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea. Vol. 2. Entomological Society of Latvia, Riga, pp. 345 - 396.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261.","Mahmood, K., Idris, A. B. & Salmah, Y. (2007) Tetrigidae (Orthoptera: Tetrigoidea) from Malaysia with the description of six new species. Acta Entomologica Sinica, 50 (12), 1272 - 1284.","Liang, G. & Zheng, Z. (1998) Fauna Sinica, Insecta, Orthoptera: Tetrigoidea. Vol. 12. Science Press, Beijing, 278 pp.","Zheng, Z. - M. (2005) Fauna of the Tetrigoidea from Western China. Science Press, Beijing, 501 pp.","Deng, W. - A., Zheng, Z. & Wei, S. - Z. (2007) Fauna of Tetrigoidea from Yunnan and Guangxi. Guangxi Science and Technology Press, Nanning, 458 pp.","Costa, A. (1864) Acquiste fatti durante l-anno 1862. Annuario del Museo zoologico della Universita di Napoli, 2, 8 - 94.","Muhammad, A. A., Tan, M. K., Abdullah, N. A., Azirun, M. S., Bhaskar, D. & Skejo, J. (2018) An annotated catalogue of the pygmy grasshoppers of the tribe Scelimenini Bolivar, 1887 (Orthoptera: Tetrigidae) with two new Scelimena species from the Malay Peninsula and Sumatra. Zootaxa, 4485 (1), 1 - 70. https: // doi. org / 10.11646 / zootaxa. 4485.1.1"]}

Discotettix (Mnesarchus) scabridus (Stål, 1877) (Fig. 33) Vernacular name: Filipino Spiky Pygmy Devil Mnesarchus scabridus Stål, 1877: 55 [original description, type locality: the Philippines]; Casto de Elera 1895: 207. Discotettix scabrides: Hancock 1907a [lapsus calami]. Discotettix scabridus: Bolivar 1877: 307 [in revision]; Kirby 1910: 2 [listed in the catalog]; Günther 1938: 302 [in revision]; Yin et al. 1996: 866 [listed in the catalog]; Blackith 1992: 46 [listed in the catalog]; Otte 1997: 32 [listed in the catalog]. Discotettix (Mnesarchus) scabridus: Kevan 1966: 380 [new records]. This is the type species of the subgenus Mnesarchus and the only species presently assigned to this subgenus. The species diagnosis and the description may hence be used as subgeneric diagnosis and description of Mnesarchus. Type locality. The Philippines (without exact locality). Lectotype designation. The lectotype for this species is designated herewith, that being the female deposited in NHRS, under the inventory number NRM-ORTH 00112879, designated by J. Tumbrinck. The lectotype bears the red label ‘lectotypus’. Lectotype designation is important because of the future analysis of intraspecific differences among D. scabridus populations in Mindanao and other islands that could result in the discovery of new species or subspecies. Since Tetrigidae species are usually morphologically well-defined, the lectotype may provide a basis for the identification of the nominotypical and/or already described taxon. Material examined. Type material. LECTOTYPE (Fig. 33): ♀ Ins. Philipp. Collector: Semper, det. C. Stål, inventory number NRM-ORTH 00112879 (NHRS); labeled Mnesarchus scabridus; here designated. Additional museum material. 2♀♀ the Philippines, Mindanao, Zambaonga, date uknown, Collector: W. Schultze, det. K. Günther (SMTD); 1♂ Ins. Philipp. Collector: Semper, det. K. Günther (MFN); 1♀, 1♂ the Philippines, Mindanao, Pt. Bango, det. J. Tumbrinck (NHRS); 1♀ labeled ‘100.4115’ (MNCN) det. J. Skejo; 1♀ nymph, 2♂♂ the Philippines, Mindanao, Zambonga Del Norte, VIII.2020. Collector: Sandayong, det. J. Tumbrinck (ZMHU); 1♀, 1♂ the Philippines, Luzon, N-Luzon Island, Hueva, Belance, III.2014, det. J. Tumbrinck (ZMHU); Additional material from eBay. 3♀♀ the Philippines, Mindanao, Zambaonga del Sur, IX.2013. Collector: unknown (not in museum) (found on eBay on 18.VI.2016) det. J. Skejo; 2♂♂, 2♀♀ the Philippines: Samar Collector: unknown (not in museum) (found on eBay on 21.VII.2016) det. J. Skejo; 4♀♀ the Philippines: Mindanao: Bukidnon Collector: unknown (not in museum) (found on eBay on 04.VIII.2016) det. J. Skejo; 1♂ the Philippines: Mindanao: Davao Collector: unknown (not in museum) (found on eBay on 15.VIII.2016) det. J. Skejo; 3♀♀ the Philippines: Mindanao: Surigao unknown (not in museum) (found on eBay on 01.IX.2016) det. J. Skejo. Distribution. This species inhabits the Philippines, where it has been reported from the islands of Mindanao, Samar, and Luzon. Numerous records from Mindanao, while few from Luzon might indicate that the species has higher abundance in Mindanao. Subgeneric diagnosis of Mnesarchus and specific diagnosis of Discotettix scabridus . The species is morphologically very different from other Discotettix members and is thus assigned to its own subgenus, Mnesarchus. It can be easily separated from other species of the genus by the set of the following characters: (I) frontal costa bifurcates on the lower margin of the compound eyes (bifurcation is in the lower third of the compound eyes in D. aruanus sp. n., D. belzebuth, D. doriae, D. kirscheyi sp. n., D. selysi, and D. sumatrensis sp. n.), (II) FM present as a low tubercle (developed and elevated in D. aruanus sp. n., D. belzebuth, D. doriae, D. kirscheyi sp. n., D. selysi, and D. sumatrensis sp. n.), (III) MM laterally compressed and elevated (spine-like in D. belzebuth, triangular protrusions in D. sumatrensis sp. n., D. kirscheyi sp. n., while very low in D. aruanus sp. n., D. doriae and D. selysi), (IV) MML low and triangular, compressed elevations (spine-like in D. belzebuth, triangular protrusions in D. sumatrensis sp. n., similarly formed in D. selysi), (V) interscapular area triangular, with large concavity (similarly to D. selysi, and D. sumatrensis sp. n., almost parallel in D. belzebuth), (VI) lateral and humeral carinae strongly toothed (similar to D. belzebuth, granulated in D. selysi, and D. sumatrensis sp. n.), (VII) weak ML (almost absent in D. scabridus, while well visible in other species), (VIII) VL complex, with a few spines (in other species usually with one main spine, and a saw-like margin) and (IX) tegmen more elongated (TL/TW > 5) than in other species of the genus (TL/TW Redescription (Fig. 33). General features. Medium-sized flattened species (body 12.3–12.75 mm) (15–17 mm cited in the original description, but in this measurement, pronotum was probably included as well), relatively robust in dorsal view. The entire body finely granulated, rugose; pronotum wrinkled and with numerous small tubercles and medium-sized projections. Macropronotal form. Coloration. Body color from almost black, dark brown, and ferruginous brown to yellowish-brown. Pronotal protuberances usually colored in different colors (black, reddish, yellow, whitish) and often different in color from the rest of the pronotal surface. Antenna with pale colored joints between the antennomeres; scapus, pedicel, basal and medial segments dark brown, while subapical and apical even darker. Maxillary palpi dark brown, sometimes with even darker distal margins of the last segments. The visible part of the tegmen dark brown, without spots. Tibiae and tarsi with unclear pale colored rings. Head. In dorsal and frontal view, vertex 2.4–2.65 times as wide as the eye. Lateral carinae of the vertex granulated. Lateral ocelli situated at the level of the lower margin of the compound eye. In frontal view, frontal costa narrow, bifurcating a bit above the lateral ocelli into slightly divergent facial carinae forming a very narrow scutellum.Antennal groove slightly wider than the scutellum, situated just below the lower margin of the compound eye (Fig. 33C). Antenna 13-segmented (in male seems 12-segmented, because 13 th segment very small and not visible under an optical microscope, only under SEM): scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 6 th) elongated and circular in cross-section; central or subapical segments (7 th and 8 th) widened, weakly pennate, 7 th and 8 th antennal segments almost equal in width (2–2.4 times as long as wide); apical segment 9 th elongated, but much smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th reduced, very small, and borders between them barely visible. Antennomeres 3 rd to 9 th bear weak saw-like margins (weaker than in the other species of the genus), because of the presence of large basiconic sensilla. Antennomeres of D. scabridus are less significantly widened than in other species of the genus and less saw-like in appearance, supposedly because of fewer or smaller basiconic sensilla. Pronotum. Pronotum wrinkled and granulated, covered in numerous small tubercles, medium-sized and large triangular protuberances. The posterior process of the pronotum slender, surpassing the hind knee for about half the length of the hind femur (macropronotal form). Disc of the pronotum a bit depressed behind the well-developed shoulder, gradually descending backward. Pronotum with 5 unpaired projections of variable size on the medial carina (FM, PM and 3 MMs), three pairs of FLs, two pairs of more or less distinguished PMLs, and two of MMLs; three pairs of lateral and a pair of VL (well visible in profile). Anterior margin of the pronotum truncated, with small FM directed upwards. FM considerably lower than in the other species of the genus. Prozonal carina elevated and decurved caudad in the direction of the median carina, in females projected anteriorly as a small dentiform FL1. Extralateral carinae distinct in males, saw- or fan-like and elevated along the entire length much more than the prozonal carina, surpassing the anterior margin of the pronotum as a dentiform FL2. FL3 weak, but distinct. Prozona of variable length, from subsquare to wider than long, but still very short when compared to other species of the genus. Behind the FM median carina extended along the whole length of pronotum. PM larger than FM, and MM1 larger than PM1. MM2 medium-sized in males or small in females. MM3 lies in the middle of pronotum length, in females as large as PM, while in males smaller. MM4 small and indistinct. PML1 and PML2 very small. MML1 small, situated between the shoulders. MML2 large. MML3 very small, better visible in females than in males. MML4 almost indistinct in both sexes. PL1 and PL2 small and triangular tubercles between the sulci on the line joining the extralateral carinae and the humero-apical carinae. ML reduced, almost absent. The apex of the posterior pronotal process very narrow, and rounded; in females weakly excised. The posterior part of the pronotal process (about a fifth of the pronotum length) directed slightly upwards or in the level of the rest of the pronotum. The lower part of the lateral lobe with serrate margins (2–3 larger teeth on the posterior margin, and numerous smaller teeth on the anterior margin), elongated as strong spine-like VL directed outwards and slightly forwards (Fig. 33A, B, D, E). Wings. The visible part of the tegmen shaped quite elongated oval, lower side slightly curved. The tegmen is not always visible. Based on the photographs, it is hard to say whether tegmina are visible or not, in both sexes they can be noticed only after careful examination under the stereomicroscope. Hind wing present, visible under pronotum, not reaching the apex of the pronotum. Legs. Femora robust and compressed laterally. Dorsal and ventral margins of the fore and the mid femora roughly serrate, with genicular tooth on knees, and additionally with 1–3 strongly projected teeth on each margin. Teeth of ventral margin projected outwards and downwards. Hind femur with small teeth on the dorsal and ventral margins; one large tooth projected outwards situated on the ventro-external carina. Genicular and antegenicular teeth small, but recognizable. Both sides of the dorsal margins of the hind tibia finely serrated, additionally with 5–8 outer and 2–4 inner bit larger teeth. The first tarsal segment of the hind leg slightly longer than the third segment. Abdominal apex. Ovipositor elongated. Measurements. BL ♂ 10.56 mm, ♀♀ 12.31–12.74 mm; PnL ♂ 14.99 mm, ♀♀ 15.48–16.52 mm; PnW ♂ 8.16 mm, ♀♀ 7.99–8.95 mm; AnL ♂ 5.08 mm, ♀♀ 6.89–7.01 mm; TL ♂ 2.49 mm, ♀♀ 2.51–2.7 mm; TW ♂ 0.49 mm, ♀♀ 0.47–0.52 mm; fFL ♂ 3.45 mm, ♀♀ 3.86–4.01 mm; fFW ♂ 0.84 mm, ♀♀ 0.68–0.77 mm; mFL ♂ 3.52 mm, ♀♀ 3.42–3.93 mm; mFW ♂ 0.84 mm, ♀♀ 0.76–1.01 mm; hFL ♂ 6.69 mm, ♀♀ 7.39–7.91 mm; hFW ♂ 2.05 mm, ♀♀ 2.16–2.22 mm; OvL ♀♀ 1.39–1.55 mm; AnL/fFL ♂ 1.48, ♀♀ 1.68–1.72; VW ♂ 0.93 mm, ♀♀ 1.11–1.18 mm; EW ♂ 0.49 mm, ♀♀ 0.42–0.49 mm; VW/EW ♂ 1.9, ♀♀ 2.51–2.65 mm; SW ♂ 0.22 mm, ♀♀ 0.19–0.23 mm; AgW ♂ 0.39 mm, ♀♀ 0.31–0.36 mm; ScW ♂ 0.25 mm, ♀♀ 0.18–0.21 mm; SW/AgW ♂ 0.64, ♀♀ 0.51–0.7; SW/ScW ♂ 1.48, ♀♀ 0.96–1.02; As—L/W ♂ 2.41, ♀♀ 2.06–2.11; PrzW ♂ 3.25 mm, ♀♀ 3.18–3.24 mm; PrzL ♂ 2.41 mm, ♀♀ 1.34–1.74 mm; Prz—W/L ♂ 1.35, ♀♀ 1.84–2.41; TL/TW ♂ 5.08, ♀♀ 4.68–5.71; mFW/TW ♂ 1.75, ♀♀ 0.47– 0.51; fFL/fFW ♂ 4.01, ♀♀ 5.18–5.34; mFL/mFW ♂ 4.09, ♀♀ 3.88–3.91; hFL/hFW ♂ 3.26, ♀♀ 3.49–3.66; T1L/ T3L ♂ 1.14, ♀♀ 1.22–1.31., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 49-52, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Stal, C. (1877) Orthoptera nova ex Insulis Philippinis descripsit. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlinger, 34 (10), 33 - 58.","Casto de Elera, R. P. F. (1895) Ortopteros. Catalogo sistematico de toda la fauna de Filipinas. Monograph, 2, 189 - 223. https: // doi. org / 10.5962 / bhl. title. 58548","Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261.","Kevan, D. K. (1966) Some Orthoptera-Caelifera from the Philippine, Bismarck and Solomon Islands, with a few interesting records from New Guinea and the Moluccas. Entomologiske Meddelelser, 34, 375 - 420."]}

Genus Discotettix Costa, 1864 Discotettix: Costa 1864: 59; Bolívar 1887: 306; Rehn 1904: 670; Hancock 1907a: 6; Hancock 1907b: 213; Kirby 1910: 2; Willemse 1930: 7; Steinmann 1970: 216; Blackith 1992: 46; Yin et al. 1996: 866; Otte 1997: 32; Mahmood et al. 2007: 1275; Ko&ccaron;árek et al. 2015: 288–294. Mnesarchus Stål, 1877: 55; synonymized with Discotettix by Bolívar (1887). Type species: Discotettix armatus Costa, 1864, by original monotypy, a junior synonym of Discotettix belzebuth (Serville, 1838). Nomenclatural note. Many authors recently treat tettix as a noun originally of feminine gender. It is incorrect, as in all the Ancient Greek dictionaries the noun “ tettix, tettigos or tettikos, ho ” is of masculine gender. The word “ tetrix, tetrigos, he ” is however of feminine gender in Ancient Greek. Latreille (1802) introduced the name Tetrix (vernacular tétrix), but did not explain why he used the Ancient Greek name of a bird (tetrix is the Ancient Greek name of the Pipit, still present in the name of the Black Grouse Tetrao tetrix). Since then, the name Grouse Locust has been coined in the US, while the vernacular name pygmy grasshoppers has become more widely used. Even if Latreille did not want to relate pygmy grasshoppers to the bird and randomly invented the word “ tetrix ”, he used the word as feminine gender, which has to be followed (ICZN 1999, Art. 30.1.4.2). Tettix is a masculine Ancient Greek word for grasshopper, introduced by Berthold (1827) as an (unjustified) emendation of Latreille’s name. The epitheta of all the Tetrigidae species whose genus is coined out from the word tettix should be in the grammatic masculine gender. Diagnosis. The genus can be distinguished from all the other genera by the following set of characters: (I) frontal costa bifurcates between the lower third of the compound eye height (bifurcates below the lower third in other Discotettigini), (II) scutellum narrower than scapus (of the same width or wider in Gavialidium, Paragavialidium, and Tegotettix), (III) antenna 13-segmented (15-segmented in Gavialidium, Paragavialidium, and Tegotettix), (IV) subapical antennal segments widened (filiform in most of Discotettigini), (V) margins of the antenna saw-like (smooth in most of other Discotettigini). Comparison to former Discotettiginae genera. Among the former Discotettiginae genera (see Skejo 2017) the genus is similar to Kraengia and certain members of the genus Hirrius, i.e., H. montanus Günther, 1937 and H. sarasinorum Günther, 1937 from Sulawesi. Discotettix is similar to Kraengia in the general arrangement of pronotal protuberances (FM, FLs, MM, ML). However, in Discotettix the lower part of the lateral pronotal lobe is directed outwards forming a spine-like VL projection, while in Kraengia the lower part of the lateral pronotal lobe has a truncated margin. ML is more or less distinct in Discotettix species, while fully reduced in Kraengia, the humeral angle being obtuse. Additionally, Discotettix can be distinguished from Kraengia by the following set of characters: (1) 13 antennal segments (11 in Kraengia), (2) large body size (more than 11 mm in Discotettix, less than 9 mm in Kraengia), (3) presence of tegmen and wing in all Discotettix species (Kraengia is wingless), and (4) distinct prozona with carinae (in Kraengia prozona is very short and carinae are usually not distinct). Discotettix can be distinguished from Hirrius montanus and H. sarasinorum by the following characteristics: (1) dorsal surface of the pronotum with protuberances and projections (in Hirrius the pronotum is almost flat, medial, mediolateral, and lateral projections are considerably reduced in size, hump-like or fully absent); (2) the lower part of the lateral lobe of the pronotum forms a sharp spine-like or saw-like VL projection (VL spine wanting or weak in Hirrius); (3) tegmen and wing visible (not visible in Hirrius). Comparison to similar Scelimeninae: Discotettigini genera. The genus is morphologically similar to other Discotettigini genera, especially winged Bidentatettix, Disconius gen. n., Gavialidium, Eufalconius, Paragavialidium, and Tegotettix. Of all the mentioned genera, Discotettix is most similar to Disconius. From all the genera except for the Disconius, Discotettix can be easily distinguished by the widened antennomeres, while from Disconius it can be distinguished by the visible FM (reduced in Disconius), by tuberculated median carina (continuous in Disconius) and by strong FLs (almost absent in Disconius). Redescription of the genus Discotettix General features. Medium and large sized species, robust in appearance. All the surfaces rough and granulated, rugose; pronotal disc wrinkled with numerous small tubercles and protuberances of different sizes and shapes. Macropronotal. Coloration. Body color dark brown, ferruginous brown, or with brighter tints of brown; pronotal projections darker or differently colored than the rest (e.g., reddish or yellowish). Antenna black or dark brown, sometimes with pale-colored joints between the segments or with yellowish apical segments. Maxillary palpi dark brown, sometimes with darker distal margins of the last segment, or black with pale-colored joints between the segments. The visible part of the tegmen dark brown without spots. Legs dark brown except more or less distinct pale rings on tibia and tarsi and whitish 1 st tarsal pads. Head. Head not elevated above the pronotum in lateral view. In dorsal view, the fastigium of the vertex considerably broader than a compound eye; the anterior margin of the fastigium truncated, widely excised, with protruded medial carina of the vertex, reaching not far from the anterior margin of a compound eye. In frontal view, the vertex slightly concave, indrawn from the considerably raised lateral carinae on the level of the upper margin of a compound eye; the medial carina of the vertex distinct in the anterior part of the vertex. Fossula present. Supraocular lobe absent. Lateral ocelli at the level of lower margin or between the compound eyes. Median ocellus far below the level of the lower margin of a compound eye, just between the facial carinae in the place where they end. Antennal groove just above the median ocellus, below or on the level of the lower margin of a compound eye. Frontal costa narrow, with the bifurcation a bit above or between the lateral ocelli. Frontal costa bifurcates into slightly divergent facial carinae forming a narrow scutellum, in lateral view with two concavities: the first large between the lateral ocelli and the second smaller below the antennal grooves. Maxillary palpi flattened. Compound eye in frontal view subglobular, in lateral and dorsal view drop-like, not protruding above the pronotum in lateral view. The occipital area between the eye and the anterior margin of the pronotum narrow, partly visible (more often not) from above (Fig. 5C). Antenna 13-segmented (but in male looks like 12-segmented, because 13 th segment very small and not visible under an optical microscope, only under SEM). Antennal segments as follows: 1 st massive scapus; 2 nd large pedicel; 3 th to 6 th basal elongated antennomeres; 7 th and 8 th central or subapical antennomeres, widened; apical 9 th small; 10 th to 13 th apical segments small, very reduced in comparison to others (Fig. 1). Pronotum. Pronotum wrinkled and granulated, covered by numerous small tubercles and larger projections. Posterior process of the pronotum slender, surpassing the hind knee for about a half of the hind femur length or more (macropronotal); covering the whole abdomen. Disc of pronotum: 1) more or less depressed behind the well-developed shoulder and gradually descending backward, or 2) almost at the same level along all length, without distinct depression behind the shoulder, and not descending backward. General arrangement of pronotal disc projections: pronotum with 4–7 unpaired projections of variable size on the medial carinae (FM and 3–6 medial projections); 1–3 pairs of FL projections; 1–7 pairs of more or less distinguished mediolateral projections; 1–3 pairs of lateral and a pair of more or less distinct VL (better seen in profile). In some species, some of the projections lacking or reduced. Prozona subsquare or wider than long (not taking into account FM). Anterior margin of pronotum truncated or projected, with a small or a large FM directed mainly upwards or forwards, sometimes covering a part of or the whole vertex. Prozonal and extralateral carinae in the prozona distinct, more or less elevated, surpassing the anterior margin of the pronotum as dentiform FL1 and FL2, where FL2 more distinct. FL3 dentiform, small and weak, sometimes indistinct. Median carina behind FM extended along the whole length of the pronotum, with 3–6 unpaired medial projections 4 of variable size, more or less distinct (seen very well in profile). PM small and triangular. MM1 large and triangular. MM2, MM3, and MM4 decreasing in size towards the apex of the pronotum (sometimes MM3 and MM4 reduced). MM5 present only in a few specimens of D. belzebuth. Usually, 1–7 pairs of the mediolateral projections increase in size towards MML1 (largest) and then decrease towards the tip of the pronotum (PML1 < PML2 < MML1 > MML2 > MML2 > MML4 > MML5). PML1 more or less distinct; PML2 distinct; MML1 small; MML2 large; MML3, MML4, and MML5 small, decreasing caudad (sometimes 1–3 of these posterior projections reduced). PL1 and PL2 small and triangular. ML more or less sharp, usually projected outwards. Interhumeral carinae indistinct, weak. External lateral carinae raised upwards above the base of the tegmen, in the posterior half smooth, not reaching the apex of the pronotum. Internal lateral carinae smooth, weak, usually indistinct. The infrascapular area triangular, as wide as the mid femur, fused to the lateral area. Lateral area narrower than the infrascapular and running towards the apex of the pronotum. The apex of the posterior pronotal process in the dorsal view shallowly excised or rounded. Hind margin of the pronotal lateral lobe bisinuate, ventral sinus deep, tegminal sinus small. The lower part of the lateral lobe with serrate anterior and posterior margins. VL elongated as spine-like, directed strongly outwards, sometimes forward or even slightly backward, but never downward (Figs 2, 5A, B). 4 The description of medial, mediolateral and lateral projections is given in the order from the anterior to the posterior part of the body. Wings. The visible part of the tegmen oval and elongated. Hind wing with scalloped inner margin, usually shorter than the pronotal process, not reaching its apex. Legs. Femora robust, compressed laterally; with smooth or rough surface; dorsal and ventral margins finely or roughly serrate (Fig. 29B); genicular teeth visible on the knees; additional one to three teeth present on each margin. Fore and mid tarsi with distal segments longer than the proximal ones. Both sides of the upper margin of the hind femur finely serrated with distinct or indistinct lappets. Lateral area of the hind femur bears weak carinae with net-like elevations and outgrowths, especially on the ventro-external carina. Genicular teeth equal to or larger than the antegenicular. Hind tibia in dorsal view very slightly widened in basal and apical part. Both sides of the dorsal margin of the hind tibia finely serrated, usually with a few outer and large inner teeth. 1 st tarsal segment of the hind leg longer than 3 rd (without claws); 1 st and 2 nd basal pads of 1 st tarsal segment short and triangular, 3 rd (apical) elongated (Fig. 1). Abdominal apex. Male subgenital plate in ventral view triangular, longer than wide (Fig. 31A, B). Female subgenital plate in ventral view subsquare. Ovipositor elongated or robust. Valves of the ovipositor narrow, serrate (Fig. 31C, D). Epiproct in females as long as wide near the base, apex pointed. Cerci conical with narrowly rounded apex. Composition and classification. The genus Discotettix is divided into two subgenera: (1) nominotypical Discotettix (type species D. armatus = D. belzebuth) characterized by a long FM projected over the vertex; and (2) Mnesarchus Stål, 1877 stat. resurr. (type species Mnesarchus scabridus = Discotettix scabridus) characterized by a minute FM, not projected over the vertex. The subgenus Discotettix includes six species. One species formerly assigned to Discotettix, that is D. shelfordi, has been transferred to a new genus, Disconius Skejo, Pushkar et Tumbrinck gen. n. The distribution of all the species is presented in Fig. 3. The annotated checklist of Discotettix species 1) Discotettix (Discotettix) aruanus Skejo, Pushkar et Tumbrinck sp. n. [Aru: Tanahbesar], 2) Discotettix (Discotettix) belzebuth (Serville, 1838) [Borneo, Java (?)], 3) Discotettix (Discotettix) doriae Bolívar, 1898 stat. resurr. [Mentawai: Sipora], 4) Discotettix (Discotettix) kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n. [Northeastern Borneo], 5) Discotettix (Discotettix) selysi Bolívar, 1887 [Peninsular Malaysia, Sumatra], 6) Discotettix (Discotettix) sumatrensis Skejo, Pushkar et Tumbrinck sp. n. [Southern Sumatra], 7) Discotettix (Mnesarchus) scabridus (Stål, 1877) [Philippines: Mindanao, Samar]. A key for the identification of Discotettix subgenera and species (Fig. 4) 1A) FM not projected above the vertex in lateral view (red arrow in Fig. 4). Shoulders unarmed. (Subgenus Mnesarchus). The Philippines.............................................................................. D. (M.) scabridus 1B) FM projected above the vertex in lateral view (red arrow in Fig. 4). Shoulders armed with ML. (Subgenus Discotettix)..... 2 2A) Dorsum of the pronotum with high projections, as high or almost as high as the FM (compare the grey line in Fig. 4)....... 3 2B) Dorsum of the pronotum flattened, usually no projection higher than the FM (gray line in Fig. 4, exception is D. doriae where FM is reduced)....................................................................................... 5 3A) FM small (red arrow in Fig. 4). Dorsum of the pronotum with triangular projections (blue arrows in Fig. 4). Widest antennomere 8 th. NE Borneo..................................................................... D. (D.) kirscheyi sp. n. 3B) FM large (red arrow in Fig. 4)........................................................................... 4 4A) Dorsum of the pronotum with high spikes (blue arrows in Fig. 4). Widest antennomere 8 th. Borneo........ D. (D.) belzebuth 4B) Dorsum of the pronotum with triangular projections (blue arrows in Fig. 4). Widest antennomere 7 th. Sumatra............................................................................................ D. (D.) sumatrensis sp. n. 5A) FM small, not exceeding the head (red arrow in Fig. 4). Mentawai Isl.................................. D. (D. ) doriae 5B) FM large, directed more upwards than forwards, usually not exceeding the head (red arrow in Fig. 4)................... 6 6A) Larger species, pronotum length more than 20 mm in females. PM and MM1 lower and more oblique. Sumatra, Peninsular Malaysia.................................................................................. D. (D.) selysi 6B) Smaller species, pronotum length less than 17 mm in females. PM and MM1 elevated and more triangular. Aru Isl............................................................................................ D. ( D.) aruanus sp. n., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 11-15, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Costa, A. (1864) Acquiste fatti durante l-anno 1862. Annuario del Museo zoologico della Universita di Napoli, 2, 8 - 94.","Bolivar, I. (1887) Essai sur les Acridiens de la tribu des Tettigidae. Annales de la Societe Entomologique de Belgique, 31, 175 - 313.","Rehn, J. A. G. (1904) Studies in the Orthopterous subfamilies Acrydiinae (Tettiginae), Eumastacinae and Proscopinae. Proceedings of the Academy of Natural Sciences, Philadelphia, 56, 658 - 683.","Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Hancock, J. L. (1907 b) Studies of the Tetriginae (Orthoptera) in the Oxford University Museum. Transactions of the Entomological Society of London, 1907, 213 - 244. https: // doi. org / 10.1111 / j. 1365 - 2311.1907. tb 01760. x","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Willemse, C. J. M. (1930) Fauna Sumatrensis (Bijdrage Nr. 62). Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie, 73, 1 - 210.","Steinmann, H. (1970) Check-list of the Tetricidae (Orthoptera) of the Oriental faunal region. Acta Zoologica Academiae Scientiarum Hungaricae, 16, 215 - 240.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261.","Mahmood, K., Idris, A. B. & Salmah, Y. (2007) Tetrigidae (Orthoptera: Tetrigoidea) from Malaysia with the description of six new species. Acta Entomologica Sinica, 50 (12), 1272 - 1284.","Kocarek, P., Kuravova, K., Musiolek, D., Wahab, R. A. & Kahar, S. R. A. (2015) Synonymy of Discotettix adenanii Mahmood, Idris & Salmah, 2007 with D. belzebuth (Serville, 1838) (Orthoptera: Tetrigidae). Zootaxa, 4057 (2), 288 - 294. https: // doi. org / 10.11646 / zootaxa. 4057.2.10","Stal, C. (1877) Orthoptera nova ex Insulis Philippinis descripsit. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlinger, 34 (10), 33 - 58.","Serville, J. G. A. (1838) Histoire naturelle des insectes. Orthopteres. Roret, Paris, 776 pp. https: // doi. org / 10.5962 / bhl. title. 95609","Latreille, P. A. (1802) Histoire Naturelle, generale et particuliere, des Crustaces et des Insectes. Vol. 3. De L'imprimerie de F.","ICZN [International Commision on Zoological Nomenclature] (1999) International Code of Zoological Nomenclature. 4 th Edition. The International Trust for Zoological Nomenclature, London, 306 pp. https: // doi. org / 10.5962 / bhl. title. 50608","Berthold, A. A. (1827) Latreille's Naturliche Familien des Thierreichs, aus dem Franzosischen mit Anmerkungen und Zusatzen. Verlage des Gr. H. S. priv. Landes-Industrie-Compositoirs, Weimar, VIII + 602 pp. https: // doi. org / 10.5962 / bhl. title. 11652","Gunther, K. (1937) Orthoptera celebica sarasiniana, Fam. Acrididae, Subfam. Acrydiinae. Treubia, 16, 165 - 195.","Bolivar, I. (1898) Contributions a l'etude des Acridiens especes de la Faune indo et austro-malaisienne du Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 39, 66 - 101. https: // doi. org / 10.5962 / bhl. part. 9541"]}

Discotettix (Discotettix) kirscheyi Skejo, Pushkar, Tumbrinck et Tan sp. n. (Figs 22–29) Vernacular name: Kirschey’s Spiky Pygmy Devil Discotettix belzebuth (Serville, 1838) [partim]: all older records from NE Borneo need revision Type locality. Borneo: Malaysia: East Sabah: Sepilok, lowland. Material examined. Type material. HOLOTYPE (Fig. 22) 1&female; Borneo: Malaysia: East Sabah: Sepilok XI.1994. (ZFMK); PARATYPES (Fig. 23) 2&male;, 1&female; Borneo: Malaysia: East Sabah: Kawag Forest Reserve, N5.04861, E117.07355, 118.6± 8.7 m.a.s.l., 14.V.2022, 19h27, on tree trunk, leg. M.K. Tan, T. Robillard & R. Japir, SBH.22.76–78 (ZRC); 1&male;, 2&female; Borneo: Malaysia: East Sabah: Kawag Forest Reserve, N5.05020, E117.98286, 134.7± 6.8 m.a.s.l., 14.V.2022, 13h16, on leaf litter on track, leg. M.K. Tan, SBH.22.69–71 (FRC); 1&male; Borneo: Malaysia: East Sabah: Sepilok: Rainforest Discovery Centre, N5.87414, E117.93782, 96.8± 6.8 m.a.s.l., 18.V.2022, 21h28, foliage of seedling near ground, leg. M.K. Tan & T. Robillard, SBH.22.155 (FRC); 1&male; Borneo: Malaysia: East Sabah: Tabin Wildlife Reserve, N5.19465, E118.50310, 92.7± 7.5 m.a.s.l., 15.V.2022, 20h01, on foliage near ground, leg. M.K. Tan, T. Robillard & R. Japir, SBH.22.86 (ZRC); 1&female; Borneo: Malaysia: Sabah state (North Borneo), env. of Kinabatangan, 29.II.2008, leg. V.G. Bezborodov (ZISP); 1&female; Borneo: Malaysia: Borneo: East Sabah: Sepilok, 1–6.II.2014, leg. M. Berezin (ZISP). Additional material from online social media. 1&female; Borneo: Malaysia: East Sabah: Sepilok (Fig. 27) photo: T. Kirschey; 2&male;&male;, 1&female; Borneo: Malaysia: East Sabah: Danum Valley, 04.II.2011. photo: A. Anker (Flickr) (Figs 25, 26); 1 adult Borneo: Malaysia: East Sabah: Danum Valley, 08.II.2014. photograph: P. Bertner (Flickr); 1 adult Borneo: Malaysia: East Sabah: Tawau district, 03.IV.2009. photo: author unknown (found in the album ‘ Fauna of Sabah’) (Flickr); 1 adult Borneo: Malaysia: East Sabah: Kinabatangan River Area 26–27.XI.2016. photograph: C. Odonnell (Facebook); 1 adult Borneo: Malaysia: East Sabah: near the Tawau Hills Park 21.IV.2018. photograph: A. Bouma (Facebook) (Fig, 29); 1 adult Borneo: Malaysia: East Sabah: Tawau Hills Park (4.34N, 117.89E) observed by Sustainable Strategies Network (@hobatahalmahera) on 22.I.2020. (inaturalist.org/observations/37844288) (Fig. 24); 1&male; Borneo: Malaysia: East Sabah: Kinabatangan (5.44N, 117.75E) observed by @simben on 21.VII.2016. (inaturalist.org/observations/102814348) (Fig. 28). Material that likely belongs to this species but was not checked by the authors. 1&female; Borneo: Malaysia: Sabah state (North Borneo), env. of Kinabatangan, 29.II.2008, leg. V.G. Bezborodov (ZISP). Type series depository. Museum Alexander Koenig in Bonn (ZFMK). Etymology. Named after the German biologist Tom Kirschey, our friend and a well-known researcher of the Oriental region, who currently serves as the Head of the International Peatlands and Southeast Asia Programme at NABU (Naturschutzbund Deutschland) headquarters. Distribution and habitat. The species is restricted to NE Borneo, East Sabah. It is found in Danum Valley (and adjacent Kawag Forest Reserve), Tabin Wildlife Reserve, Tawau Hills, Sepilok District, and around the Kinabatangan River Area. The species inhabits the rainforest, where it can be found on the bark of the roots and trunks of trees, among leaf litter, and sometimes on the foliage of seedlings near the ground. According to the photographer, biologist Tom Kirschey, who observed the species in its natural habitat in the lowlands and the mountains of Sepilok, the microhabitat of D. kirscheyi are the roots of the big (old) trees close to pools filled with rain water (Fig. 28). The distribution areas of this species and that of D. belzebuth do not appear to overlap. Diagnosis. The species is similar to D. belzebuth, but can be easily distinguished from the latter by the following set of characters: (1) FM shorter, narrower, and directed more upwards, not covering the entire vertex as in D. belzebuth, which has a large and long digitate FM covering the entire vertex, (2) pronotal projections are much shorter, and not as spiky as in D. belzebuth, but stouter and triangular, (3) femora are stouter and bear stronger teeth than observed in D. belzebuth, and (4) tegmina are smaller, partly covered and not as evident and wide as in D. belzebuth. Description (holotype). General features. Medium-sized, robust species (body length 14.82 mm); pronotum granulated and wrinkled, with one digitate FM on anterior margin and numerous triangular protuberances on dorsal and lateral sides. Almost the whole body (except the eye, labrum, fore and mid tarsi, second and third segments of hind tarsus, and inner side of the hind femur) covered with numerous small tubercles. Macropronotal. Coloration. Body dark brown, almost completely covered by numerous small tubercles. Pronotum, projections, and protuberances dark brown (Figs 23–26, 28, 29). Antenna dark brown, almost black. Proximal segments of the palpi light brown, and distal segments black. The visible part of the tegmen dark brown and without spots. Legs dark brown with lighter rings and patches. Fore and mid femora and tibiae dark brown with numerous small tubercles with lighter apices, fore and mid tarsi black, second segment with a light ring in the middle; 1 st tarsal pads whitish, claws brown. Hind femur dark brown with numerous small tubercles with lighter apices. Hind tibia dark brown with two weak light rings, one in the basal part, and another in the distal third. 1 st tarsus of hind leg pale brown, pads whitish; third segment pale brown with a dark ring in its distal part, claws brown. Tergites, sternites, epiproct, and cerci dark brown. Head. In dorsal and frontal view, vertex 1.95 times as wide as an eye. Fossula elliptic and deep. Lateral ocelli at the level of the lower margin of a compound eye. In frontal view, frontal costa narrow, bifurcated slightly above lateral ocelli into finely granulated facial carinae, forming a narrow scutellum. Scutellum slightly narrower than antennal groove (Fig. 22E). Antennal groove considerably below the lower margin of the compound eye. Antennae 13-segmented, short in appearance compared to D. belzebuth: scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 6 th) elongated, but shorter than in D. belzebuth, and circular in cross-section; central or subapical segments (7 th and 8 th) strongly widened, pennate, 8 th being the widest antennal segment (about 3–4 times as long as wide), but only slightly wider than the 7 th; subapical segments less widened than in the other species of the subgenus Discotettix; apical segment 9 th small and pennate, smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th reduced, very small, and borders between them barely visible. Antennomeres 3 rd to 9 th bearing saw-like margins, because of the presence of large basiconic sensilla. When the body and the antennae are covered with algae and moss, the 8 th segment is free of epizoic organisms. Pronotum. Pronotum wrinkled and granulated, covered by numerous small and medium-size tubercles. The posterior process of the pronotum extended beyond the hind knees for less than half of the hind femur length. Disc of the pronotum slightly depressed behind the well-developed shoulder. Pronotum with 5(–7) unpaired projections of variable size on median carina (digitate FM and 4 to 6 medial projections), 3 pairs of FL, up to 7 pairs of more or less distinct mediolateral projections, 3 pairs of lateral projections, and a pair of VL (well seen in profile). Prozona robust, wider than long (because of the short FM). Anterior margin of pronotum projected into a medium-sized digitate FM protuberance directed mainly upwards than slightly forwards, and not covering the whole vertex above. Prozonal and extralateral carinae forming small FL1 and larger FL2, more distinct. FL3 dentiform. Behind the FM medial carina extended along the whole length of pronotum, with 5 unpaired large and small medial projections (well visible in profile). PM triangular and smaller than other medial projections. MM1 is the largest projection; MM2 and MM3 distinct, while MM4 and MM5 small. PML1 and PML2 distinct, tubercle-like. MML1 and MML2 large and triangular; MML3, MML4, and MML5 smaller, decreasing in size caudad. MML4 and MML5 sometimes absent. PL1 and PL2 triangular, tubercle-like. ML sharp and triangular. The apex of the posterior pronotal process in dorsal view shallowly excised. The lower part of the lateral lobe with serrate anterior and posterior margins, elongated as spine-like VL, with saw-like margins, directed outwards (Fig. 22A–C). Wings. The visible part of the tegmen small, oval with a tuberculated surface, visible part 2.57 times as long as wide; the ratio of the maximum width of the mid femur (without teeth)/visible part of tegmina width 1.21. Hind wing not reaching the apex of the posterior pronotal process, ending a few millimeters before the tip. Legs. Femora robust, compressed laterally, surface rough, dorsal, and ventral margins roughly serrate. Fore and mid femora bearing one genicular tooth on knees on each margin, and additionally 3 strong teeth on upper carina and 2–3 on lower margin, almost equal in size; teeth on fore femur equal or larger than on mid femur. Upper and lower margin of hind femur finely serrated, with 2 lappets on each margin, and with numerous small tubercles. Lateral area of hind femur with weak carinae that have up to 3 outgrowths, especially in ventro-external carina. Genicular teeth larger than antegenicular. teeth. Fore femur length/width ratio 4.69. Mid femur length/ width ratio 3.94. Hind femur length/width ratio 3.28. Both sides of the upper margin of the hind tibia finely serrated, with 2–3 outer and 2–3 inner larger teeth (spines). Abdominal apex. Male subgenital plate in ventral view with shallowly excised apex, slightly longer than wide. Female subgenital plate in ventral view with a keel in middle and with a triangular protrusion in the middle of posterior margin. Ovipositor elongated, upper valve 5.0 times as long as wide. The lower valve of ovipositor about 6.0 times as long as wide (maximal width). Cerci stout, with thin apex, hairy. Measurements. All the measurements of Discotettix kirscheyi sp. n. are given in the Table 2., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 37-40, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Serville, J. G. A. (1838) Histoire naturelle des insectes. Orthopteres. Roret, Paris, 776 pp. https: // doi. org / 10.5962 / bhl. title. 95609"]}

Discotettix (Discotettix) selysi Bolívar, 1887 (Figs 15–17) Vernacular name: Sumatran Unicorn Pygmy Devil Discotettix selysi Bolívar, 1887: 307 [original description, type locality: Sumatra]; Hancock 1907a: 6 [listed in catalog]; Kirby 1910: 2 [listed in catalog]; Willemse 1930: 207 [new records], Günther 1938: 301 [partim; new records] Blackith 1992: 46 [listed in catalog], París 1994: 236 [data on the type specimens]; Yin et al. 1996: 866 [listed in the catalog]; Otte 1997: 32 [listed in the catalog]. Discotettix selangori Mahmood et al., 2007: 1276 [original description, type locality: Selangor]; syn. n. Discotettix selangorei: Mahmood et al. 2007: 1275 [lapsus calami]. Acridium (Tetrix) belzebuth (nec Serville): De Haan 1843: 166 [reported for Sumatra, misidentification]. Type locality. Sumatra: Padang Panjang (= originally “Padang Pandjang” on the label) [approximate coordinates 0.45S, 100.416667E]. Note on Discotettix selysi identification history. The species was recorded by De Haan (1843) for the first time under the name D. belzebuth, since only D. belzebuth was described at the time (D. selysi being described 44 years later). De Haan’s (1843) drawing of D. belzebuth agrees in morphology with D. selysi. The author, in the description, noted the possession of only one well distinctive anterior pronotal projection. Examination of specimens confirmed that De Haan’s records of D. belzebuth from Sumatra belong to D. selysi and there is no evidence of the presence of D. belzebuth on the island. Material examined. Type material. SYNTYPES of D. selysi 1&male; Indonesia: Sumatra: Padang Panjang (= on the label “Padang Pandjang”) Collector H. Rolle [the type lacks antennae] (Fig. 16) (MNCN); 1&male; Indonesia: Sumatra 25.XII. [18]84. Soerian (MHNG). HOLOTYPE of D. selangori 1&male; Malaysia: Selangor: leg. Brokurtak (UKM), Sabah Forestry Department, East Malaysia). Misidentified museum material. Identified by Günther (1938) as D. selysi: 1&male; Indonesia: Sumatra: west coast, Anai Kloof [500 m a.s.l.] 1926. Collector E. Jacobson, det. K. Günther (SMTD); (5–6) 2&female;&female; Indonesia: Sumatra: Excell. v. Studf., collector and date unknown, det. K. Günther (MFN); Identified by De Haan (1843) as D. belzebuth: 1&male; + 2&female;&female; + 2 nymphs (sex indeterminable) Indonesia: Sumatra: Bat. Sing. [= W Sumatra Isl., Mt. Singgalang volcano] (collector and date not specified in labels), as D. selysi det. J. Skejo et J. Tumbrinck (Fig. 16) (NCB-RMNH); Additional museum material. 2&female;&female; Indonesia: Sumatra: Maninjau, Puncak, Lawang [600–950 m a.s.l. forest] 17.IV.1995. Collector Sigfrid Ingrisch, det. J. Tumbrinck (CJT, ZFMK); 3&male;&male; Indonesia: Sumatra: Mt. Tandikat [600–900 m a.s.l.] VII.2009. Collector Jakl, det. J. Tumbrinck (ZMUH); 1&male; Indonesia: Sumatra: North Sumatra Prov., Roburan Dolok, Panyabungan Selatan, Mandailing, Natal Regency (bamboo stand) 0°44’52.83’’N 99°31’30.68’’E [723 m a.s.l.] 7.X.2019. Collector Fajar Kaprawi, det. J. Tumbrinck (CJT); Additional material from online social media. 2 specimens, sex unidentifiable (lateral and dorsal habitus) Malaysia: Peninsular Malaysia: Kuala Lumpur XII.2014. Photographer Pang Way, det. J. Skejo et J. Tumbrinck (Figs 17, 18) (Facebook). Distribution. This species inhabits the rainforests of southern Peninsular Malaysia and of Sumatra, where it can be found on tree bark and roots (Bolívar 1887; Hancock 1907a; Günther 1938; Mahmood et al. 2007, our data). Taxonomic notes on Discotettix selangori and D. doriae . The description and the measurements of the recently described Discotettix selangori Mahmood, Idris et Salmah, 2007 (type locality Malaysia: Selangor) completely fit that of D. selysi. The new name for the D. selysi population on the Malaysian peninsula does not provide any new information thus we synonymize D. selangori syn. n. with D. selysi. The authors of the former were not aware of D. selysi morphological variability and distribution. In the description (Mahmood et al. 2007), the epitheton was “selangori”, while in the key and under the drawing it was written as “selangorei” (Mahmood et al. 2007). As the first reviewers, according to the ICZN, we pick selangori as the original spelling, while “ selangorei ” is considered a misspelling. We do not agree with Günther’s (1938) synonymy of D. doriae and D. selysi, since a few clear morphological differences can be found (small FM in D. doriae, and swollen antennal segment in D. doriae) thus we regard D. doriae as a separate species with its own traits and distribution (see a paragraph on D. doriae). Diagnosis. The species is morphologically similar to its congeners that do not have strong pronotal projections of the disc, i.e. to D. doriae from Mentawai and D. aruanus from Aru Isl. Discotettix selysi can be distinguished from D. doriae by the following set of characters: (1) D. doriae has FM small and narrow, covering vertex only partially, while in D. selysi FM is large, long, and covering entire vertex); (2) D. doriae is of smaller body size (pronotum length only 16 to 17 mm in females) than D. selysi (pronotum longer than 20 mm in females); (3) D. doriae has shorter and stouter antennae with swollen 6 th, 7 th and 8 th antennal segments, while D. selysi does not have swollen segments and (4) D. selysi has more elongated and slender fore and mid femora, while D. doriae has strong teeth on their ventral and dorsal margins. From D. aruanus sp. n., D. selysi can be distinguished by (I) different shape of the antennae (margins stronger, more saw-like in D. aruanus); (II) different shape of FM (not exceeding the head in D. selysi, exceeding the head in D. aruanus sp. n.); (III) less elevated and less triangular PM and MM1; (IV) legs more robust and toothed in D. selysi than in D. aruanus sp. n., and (V) by larger body size (pronotum length more than 20 mm in females of D. selysi, while less than 17 mm in females of D. aruanus sp. n.). The species is easily distinguished from D. sumatrensis sp. n. from Sumatra, D. belzebuth from Borneo, D. kirscheyi sp. n. from NE Borneo by the lack of strong spines on the pronotal disc and from the Filipino D. scabridus by numerous characters: (I) bifurcation of the frontal costa between the eyes (on the lower margin of the compound eyes in D. scabridus), (II) FM high and developed (present as a small tubercle in D. scabridus), (III) reduced MM, not highly protruded (higher, compressed and saw-like in D. scabridus), (IV) lateral and humeral carinae are granulated (toothed in D. scabridus) and (V) larger body size. From D. belzebuth, D. selysi can be easily separated by (I) less widened antennal segments, (II) FM being the only large medial projection (disc of pronotum with many spine-like MM, ML and MML in D. belzebuth), (III) smoother lateral and humeral carinae (not as equipped and toothed as in D. belzebuth), (IV) hind femora bearing large lappets, (V) fore and mid femora being more armed and stouter, FM shorter and not decurved as in D. belzebuth. From D. sumatrensis sp. n. the species can be distinguished by (I) completely black antennae, (II) FM being the only large medial projection, pronotum without elevated MM, ML and MML and (III) larger body size. Discotettix kirscheyi sp. n. has (1) smaller FM than D. selysi, (II) larger MM1 and MM2, (II) higher MML2s, (III) more robust fore and mid femora with stronger lobes, (IV) less specialized subapical antennal segments, (V) and is smaller in size. Finally, from D. doriae the species can be separated by (I) longer FM, (II) more slender appearance, (III) not swollen subapical antennal segments, and (IV) larger body size. Redescription (Fig. 15). General features. Large-sized and relatively robust species (16–20 mm). Body finely granulated; pronotum slightly rugose, with numerous small tubercles and net-like elevations. The chitinous surfaces are smooth and without tubercles in places, while the rest is strongly granulated. The anterior part of the pronotum bears several strong protuberances, while other projections are reduced (Fig. 15). Macropronotal. Coloration. The general color dark brown, but may be of brighter tints of brown or even dark greenish. Pronotal carinae and projections darker and of a different color than the rest of the body. Median pronotal carina from dark orange to bright red (Figs 16, 17). Antennae completely black or dark brown. Maxillary palpi dark brown. The visible part of the tegmen dark brown without any spots. Legs dark brown except for more or less distinct pale rings on tibiae and tarsi. The body usually covered with algae that give a greenish appearance to the specimens. After preservation in alcohol or drying the greenish color disappears. Head. In dorsal and frontal view, vertex 2.3 times as wide as an eye. Lateral carinae considerably raised and granulated. Fossula deep, but not easily observable because it is covered by large FM of the anterior pronotal margin. Lateral ocelli situated just below the level of the lower margin of a compound eye. Antennal groove significantly below the lower margin of a compound eye. In frontal view, frontal costa bifurcated at the level of lateral ocelli into facial carinae, forming narrow parallel scutellum (Fig. 15E). Antennal groove slightly wider than the frontal costa. Antenna 13-segmented (but in male looks like 12-segmented, because the 13 th segment is very small and not visible under an optical microscope, only under SEM): scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 6 th) elongated and circular in cross-section; central or subapical segments (7 th and 8 th) strongly widened, pennate, 8 th being the widest antennal segment (about 2.7–2.8 times as long as wide); apical segment 9 th elongated and pennate, smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th reduced, very small, and borders between them barely visible. Antennomeres 3 rd to 9 th bearing saw-like margins, because of the presence of the large basiconic sensilla. When the body and the antennae are covered with algae and moss, the 8 th segment is always free of epizoic organisms. Pronotum. Pronotum rugose and granulated with numerous small tubercles and net-like elevations, but in some parts smooth and without tubercles (parts of the pronotal disc and some intervals of the median carina of the pronotum). The anterior part of the pronotum bears large FM, and a few medium-sized and small protuberances. The posterior process of the pronotum slender, surpassing hind knees for more than a half-length of the hind femur or less. The disc of the pronotum almost completely flat. A small depression on the disc positioned between the near bases, then the disc becomes slightly elevated again. Caudad, pronotum gradually descending. Disc rich in net-like elevations, more distinct in the places of the interhumeral carinae. The median carina of the pronotum bears unpaired projections of variable size: high and digitate FM, directed upwards and forwards above the head and covering the whole area of the fastigium of the vertex; small triangular PM, a bit larger triangular MM1, small MM2; and completely reduced MM3 and MM4. Prozona subsquare. Prozonal and extralateral carinae in prozona distinct, surpassing anterior margin of pronotum as dentiform small FL1 and larger FL2, FL2. FL3 small, dentiform. Behind the FM medial carina extended along the whole length of pronotum from the anterior margin to the pronotal apex, slightly undulated by the low aforementioned projections. PML1 small, PML2 even smaller. MML1 almost indistinct; MML2 distinct; MML3 very small, while MML4 and MML5 completely indistinct. PL1 and PL2 small. Humeral angle obtuse with pointed apex. ML small, reduced. Pronotum apex narrow, shallowly excised. The lower part of the lateral pronotal lobe with serrate anterior and posterior margins, elongated as spine-like VL, directed strongly outwards and backward (Fig. 15A, B). Wings. The visible part of the tegmen elliptical. Hind wing with scalloped inner margin, a few millimeters shorter than the apex of the pronotal process. In living specimens, the tegmina are covered by photosynthetic microbes (most probably algae and cyanobacteria), so are greenish in appearance. Legs. Fore and mid femora relatively robust, compressed laterally; with dorsal and ventral margins serrated with genicular tooth on the knees and additional 2–3 strongly projected and sharp teeth on each margin. Hind femur with wrinkled margins, one small protuberance situated on the ventral external carina. Genicular tooth large, while antegenicular tooth almost indistinct. Both sides of the dorsal margin of the hind tibia finely serrated, with 3–4 outer and 1–3 inner larger teeth. Abdominal apex. Male subgenital plate in lateral view about two times as long as high. Female subgenital plate in ventral view with a longitudinal keel in the median region and with a triangular protrusion in the middle of the posterior margin. Ovipositor robust, dorsal valvae robust, ventral valvae slender; all valvae serrate. Measurements. BL &male;&male; 14.41–14.88 mm, &female;&female; 16.01–16.28 mm; PnL &male;&male; 17.85–18.35 mm, &female;&female; 20.46–21.11 mm; PnW &male;&male; 8.06–8.14 mm, &female;&female; 9.27–9.48 mm; AnL &male;&male; 7.68– 7.22 mm, &female;&female; 7.18–7.49 mm; TL &male;&male; 2.44–2.51 mm, &female;&female; 2.78–2.91 mm; TW &male;&male; 0.77–0.81 mm, &female;&female; 1.03–1.11 mm; fFL &male;&male; 4.49–4.55 mm, &female;&female; 5.07–5.12 mm; fFW &male;&male; 0.78–0.84 mm, &female;&female; 0.92–0.94 mm; mFL &male;&male; 4.52–4.71 mm, &female;&female; 4.77–5.01 mm; mFW &male;&male; 0.99–1.01 mm, &female;&female; 1.03–1.14 mm; hFL &male;&male; 8.51–8.68 mm, &female;&female; 9.97–10.12 mm; hFW &male;&male; 2.65–2.72 mm, &female;&female; 3.05–3.2 mm; OvL &female;&female; 1.39–1.43 mm; AnL/fFL &male;&male; 1.68–1.7, &female;&female; 1.59–1.69; VW &male;&male; 1.01–1.05 mm, &female;&female; 1.36–1.48 mm; EW &male;&male; 0.39– 0.47 mm, &female;&female; 0.51–0.71 mm; VW/EW &male;&male; 2.23–2.49, &female;&female; 2.43–2.67; SW &male;&male; 0.32–0.36 mm, &female;&female; 0.33–0.42 mm; AgW &male;&male; 0.21–0.33 mm, &female;&female; 0.34–0.39 mm; ScW &male;&male; 0.22–0.27 mm, &female;&female; 0.24–0.29 mm; SW / AgW &male;&male; 1.49–1.53, &female;&female; 1.44–1.58; SW/ScW &male;&male; 1.21–1.45, &female;&female; 1.2–1.34; As–L/W &male;&male; 2.69–2.78, &female;&female; 2.71–2.81; PrzW &male;&male; 3.44–3.52 mm, &female;&female; 4.68–4.71 mm; PrzL &male;&male; 3.93–4.01 mm, &female;&female; 4.74–4.79 mm; Prz–W/L &male;&male; 0.86–0.88, &female;&female; 0.98–1.02; TL/ TW &male;&male; 2.91–3.09, &female;&female; 2.7–2.88; mFW/TW &male;&male; 1.19–1.25, &female;&female; 0.98–1.09; fFL/fFW &male;&male; 5.42–5.61, &female;&female; 5.51–5.71; mFL/mFW &male;&male; 4.32–4.66, &female;&female; 4.46–4.63; hFL/hFW &male;&male; 3.21–3.28, &female;&female; 3.21–3.27; T1L/T3L &male;&male; 1.01–1.08, &female;&female; 1.01–1.02., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 26-28, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Bolivar, I. (1887) Essai sur les Acridiens de la tribu des Tettigidae. Annales de la Societe Entomologique de Belgique, 31, 175 - 313.","Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Willemse, C. J. M. (1930) Fauna Sumatrensis (Bijdrage Nr. 62). Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie, 73, 1 - 210.","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Paris, M. (1994) Catalogo de tipos de ortopteroides (Insecta) de Ignacio Bolivar, I: Blattaria, Mantodea, Phasmoptera y Orthoptera (Stenopelmatoidea, Rhaphidophoroidea, Tettigonioidea, Grylloidea, Tetrigoidea). Eos, Revista espanola de Entomologia, 69, 143 - 264.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261.","Mahmood, K., Idris, A. B. & Salmah, Y. (2007) Tetrigidae (Orthoptera: Tetrigoidea) from Malaysia with the description of six new species. Acta Entomologica Sinica, 50 (12), 1272 - 1284.","De Haan, W. (1843) Bijgragen tot de kennis der Orthoptera. In: Temminck, D. (Ed.), Verhangelingen over de Natuurlijke Geschiedenis der Nederlansche Overzeesche Bezittingen, de Leden der Natuurkundige Commissie in Indie en andere Schrijvers. s. n., Leiden, 165 - 228."]}

Disconius shelfordi (Hancock, 1907), comb. n. (Figs 34–36) Vernacular name: Bornean Fallen Pygmy Devil Discotettix shelfordi Hancock, 1907b: 214 [original description, type locality: Borneo]; Kirby 1910: 575 [listed in the catalog]; Günther 1938: 303 [in review]; Yin et al. 1996: 866 [listed in the catalog]; Blackith 1992: 47 [listed in the catalog]; Otte 1997: 32 [listed in the catalog]. Type locality. Malaysia: Borneo: Kuching Note on the syntypes. The species description was based on two syntypes (male and female) labeled as “NW Borneo: Kuching, 22.IX.1899, Collector Dyak (Bornean native), det. J.L. Hancock ” and kept in the Oxford University Museum. As both specimens have been collected at the same locality and date, it is unlikely that they belong to different species. Thus, there is no need for lectotype designation. Material examined. Type material. SYNTYPE 1&female; NW Borneo: Kuching, 22.IX.1899., Collector Dyak, det. J. L. Hancock (Fig. 34) (OUMNH); SYNTYPE 1&male; NW Borneo: Kuching, 22.IX.1899., Collector Dyak, det. J. L. Hancock (OUMNH); Additional museum material. 2&female;&female;, 1&male; Indonesia: NE Borneo: Pajau River, leg. Mjöberg, det. J. Tumbrinck (Fig. 35) (NHRS). Additional material from online social media (iNaturalist): 1 adult Borneo: Malaysia: N Sarawak: Marudi (N4.041847, E114.8144) observed 15.X.2019., submitted on 5.II.2020. Photograph by Kinmatsu Lin (@kinmatsu), available at link inaturalist.org/observations/38326084 (Fig. 36). Distribution: Known only from N Borneo; in Sabah from the surroundings of Kuching (type locality) and from the region of the Pajau River; and from Sarawak reported from Marudi (Hancock 1907b; our data). Redescription. (Figs 34, 35) General features. Medium-sized species (body length 14.0– 15.8 mm) (in the original description stated as 17.5–19 mm, but it meant from the tip of the head to the tip of the pronotum ), relatively slender. The entire body finely granulated, covered by numerous small tubercles and with a few larger ones on the margins of the pronotal disc and the lateral lobes; in lateral view the pronotum almost flat, except for the median carina undulated by a few small wart-like and medium-sized semicircular compressed protuberances (Figs 34, 35). Macropronotal form. Coloration. Body color from dark brown and ferruginous brown to brown with an inexpressive grayish tint. Some parts of the body pale colored, previously yellowish: tubercles on the margin of the disc and lateral lobe of pronotum, VL and ML, connections of dark antennal segments, patches on femora, more or less distinct ring in the middle of fore and mid tibiae, two rings (I) in basal and (II) distal third of the hind tibia, distal segments and claws of fore and mid tarsi, and usually yellowish hind tarsus (except for the darker connections of tarsal segments). Living specimens have much more vivid coloration than the museum material. It is visible that the dorsum of the pronotum has an alternation of dark and pale tones (see iNaturalist observation in Material examined). Head. In dorsal and frontal view, vertex about 2.4 times as wide as the eye. Fossula visible, but not deep. Lateral ocelli situated between the compound eyes. The antennal groove situated at the level of the lower margin of the compound eye (in males) or a bit below (in females). In frontal view frontal costa narrow, bifurcates a bit above the lateral ocelli into a slightly divergent facial carinae, slightly concave inside in medium length and forming a narrow hour-glass shaped scutellum (Figs 34C, 35C). Antennal groove considerably wider than scutellum. Antennae 15 segmented, long. Antennomeres are shaped as follows: scapus (1 st antennomere) massive; pedicel (2 nd antennomere) large, basal antennomeres (3 rd to 7 th) elongated, 7 th being extremely elongated; central or subapical segments 8 th to 10 th pennate, 8 th slightly widened, while 9 th, and 10 th significantly widened and flattened; apical segment 11 th small; while apical segments 12 th to 15 th reduced in size, filiform. Pronotum. Pronotum finely granulated, covered by numerous small and few larger tubercles on the margin of the disc and the lateral lobe of the pronotum; almost flat, except for the wrinkled and scalloped median carina with small wart-like and medium-sized, compressed laterally, semicircular protuberances (different in specimens from different geographical populations); posterior process of pronotum very long, surpassing the hind knee for more than a half hind femur length (macropronotal form). Disc of the pronotum almost of the same height in the anterior part, slightly depressed behind the level of the tegmen apex and gradually descending backwards. Morphology of the pronotal projections variable. Pronotum with 5 unpaired projections of variable size on medial carina (FM and 4 medial projections); 2–3 pairs of FL; a pair of VL (better seen in profile); while among the mediolateral and lateral projections only one projection distinct per group. Prozona very short. Anterior margin of pronotum truncated, bearing small and weak triangular FM directed more upwards, then forwards. Prozonal and extralateral carinae low, tuberculated, not forming sharp saw-like or fan-like ridge, with small FL1 and more distinct dentiform. FL3 indistinct. Behind FM medial carina extended along the whole length of the pronotum, low, bearing four more or less distinct medial projections of variable size (well visible in lateral view). PM triangular, equal to or larger than FM and joined with the latter as a continuous two-humped structure in specimens of some populations. MM1, large. MM2 is the largest and the most massive projection, usually compressed laterally as a semicircular triangular protuberance. MM3 relatively large, while MM4 almost indistinct, marked by spot and darker than rest of the pronotal disc surface. Unlike in Discotettix species, Disconius shelfordi comb. n. has only one distinct projection from the mediolateral group, MML2, situated in the place where most Tetrigidae (Tetriginae) species have a posthumeral spot. Similar to the previous projections’ group, only one projection exists from the lateral group of the projections. In the metazona the humero-apical carinae forms a moderately sharp humeral angle, projected outwards as a small ML tubercle of the humeral angle, much larger than other tubercles along the margin of the pronotal disc and the lateral lobe; and behind this point the humero-apical carinae is joining the external lateral carinae. The apex of the posterior pronotal process in the dorsal view shallowly excised. The lower part of the lateral lobe with smooth anterior and posterior margins, without smaller teeth. The lateral lobe elongated as spine-like VL, directed exactly outwards, blunt (Figs 34A, B, 35A, B). Wings. The visible part of the tegmen elongated and oval, distinctly acuminate towards the apex, about 2.75 times as long as wide. Hind wings reach the apex of the pronotal process. Legs. Femora relatively slender, compressed laterally, and finely granulated, with numerous small teeth-like tubercles on the whole surface. Fore and mid femora bearing a hardly noticeable genicular tooth on the knees, and additionally 1–3 small teeth on the dorsal and ventral margin. Hind femur significantly compressed laterally, finely granulated, without any lappets on dorsal and ventral margins, smooth, and not bearing any recognizable outgrowth on the external carinae. Genicular and antegenicular teeth small. Both sides of the dorsal margin of the hind tibia only finely serrated, without any large teeth. Abdominal apex. Male subgenital plate in ventral view about 1.5 times as long as wide, in lateral view about 2 times as long as tall. Ovipositor elongated. Observed variability and differences found among populations, with implications for taxonomy. In the different populations of D. shelfordi comb. n., we have observed that the projections of the pronotum differ in size and shape. For example, among the specimens from the banks of the Pajau river the pronotum projections are much more expressed than in the specimens from Kuching (the type locality of the species). This primarily applies to lower and weaker medial protuberances in the specimens from Kuching. Especially FM and PM are quite small and wart-like, MM is medium size and slightly compressed laterally. In specimens from the Pajau river FM and PM are well expressed, and form a continuous double-hump structure. Other MMs are much more expressed, and more compressed laterally, they have semicircular form, especially the largest MM2. Only the examination of a larger series will allow drawing conclusions about the specific value of the aforementioned traits. Measurements (female syntype and a non-type male). BL &male; 13.99 mm, &female; 15.87 mm; PnL &male; 18.74 mm, &female; 20.81 mm; PnW &male; 7.35 mm, &female; 7.42 mm; AnL &male; 9.39 mm, &female; 9.68 mm; TL &male; 2.43 mm, &female; 2.62 mm; TW &male; 0.85 mm, &female; 0.96 mm; fFL &male; 3.22 mm, &female; 3.33 mm; fFW &male; 0.57 mm, &female; 0.62 mm; mFL &male; 4.01 mm, &female; 4.15 mm; mFW &male; 0.55 mm, &female; 0.61 mm; hFL &male; 7.99 mm, &female; 8.76 mm; hFW &male; 2.34 mm, &female; 2.71 mm; OvL &female; 1.41 mm; AnL/fFL &male; 2.91, &female; 2.9; VW &male; 1.1 mm, &female; 1.29 mm; EW &male; 0.45 mm, &female; 0.53 mm; VW/EW &male; 2.44, &female; 2.43; SW &male; 0.14 mm, &female; 0.19 mm; AgW &male; 0.39 mm, &female; 0.47 mm; ScW &male; 0.26 mm, &female; 0.38 mm; SW/AgW &male; 0.36, &female; 0.4; SW/ScW &male; 0.54, &female; 0.5; As-L/W &male; 2.35, &female; 2.62; PrzW &male; 3.18 mm, &female; 3.47 mm; PrzL &male; 1.5 mm, &female; 1.71 mm; Prz-W/L &male; 2.12, &female; 2.03; TL/TW &male; 2.86, &female; 2.73; mFW/TW &male; 0.65, &female; 0.64; fFL/fFW &male; 5.65, &female; 5.37; mFL/mFW &male; 7.29, &female; 6.8; hFL/hFW &male; 3.41, &female; 3.23; T1L/T3L &male; 1.27, &female; 1.19.

Discotettix (Discotettix) aruanus Skejo, Pushkar et Tumbrinck sp. n. (Figs 19–20) Vernacular name: Aruan Unicorn Pygmy Devil Type locality: Indonesia: SE Moluccas: Aru Island: Wokam: 10–15 km NEE of Wakua. Material examined. Type material. HOLOTYPE (Fig. 19) 1&female; Indonesia: SE Moluccuas: Aru Island, S coast of Wokam I., 10–15 km NEE of Wakua vill., 0–50 m alt. 21–30.I.2015., St. Jakl leg. (ZMUH); PARATYPE (Fig. 20) 1&female; Indonesia: SE Moluccuas: Aru Island, S coast of Wokam I., 10–15 km NEE of Wakua vill., 0–50 m alt. 21–30.I.2015., St. Jakl leg. (FM broken in this specimen) (ZMUH). Type series depository. Zoologisches Museum Hamburg, Germany. Type series depository. Zoologisches Museum Hamburg, Germany (ZMUH). Etymology. Named after the island of Aru. The specific epitheton is a masculine gender adjective in the Nominative case made from the Latinisation of the name Aru (aruanus, aruana, aruanum). Distribution. This species is known to inhabit the Peninsula of Wokam (Wokam Palau), a part of the island of Aru in the southeastern Moluccas. This is the easternmost species of the genus Discotettix. Specific diagnosis. The new species is very similar to Discotettix selysi from the southern Malaysian peninsula and from Sumatra. It shares many traits with D. selysi, but a few subtle differences exist. Due to morphological differences among D. aruanus, D. selysi and D. doriae (Fig. 21) and because of high geographical isolation, D. aruanus sp. n. is proposed as a new species within the genus Discotettix, and not a subspecies of D. selysi. From D. doriae this species is easily separated by (I) the large FM covering the whole vertex and being produced even in front of the head (in D. doriae FM is much smaller) and (II) by much longer antennae without swollen segments. From D. selysi, the new species can separated by the following set of characteristics: (I) FM somewhat larger and in dorsal and lateral views projected before the head (in D. selysi, there are specimens without and with long projected FM); (III) MMLs more elevated than in D. selysi; (IV) PM and MM1 more elevated and more triangular (usually lower and more oblique in D. selysi); (V) VL spine more distinct in D. aruanus sp. n. because the posterior lobe of the lateral pronotal lobe has smaller spikes than in D. selysi (in D. selysi some specimens are similar to D. aruanus); (VI) femora more slender in D. aruanus sp. n. than in D. selysi; and (VII) humeral angles more projected in D. selysi (well visible in frontal view), and infrascapular area thus wider in D. selysi than in D. aruanus sp. n. For a comprehensive comparison with other species of the genus please consult the diagnoses of D. scabridus and D. belzebuth. Description (holotype). General features. Medium-sized, robust species (body length 11.5–13.5 mm in females, pronotum length about 16.5 mm); pronotum granulated and wrinkled, with numerous small tubercles and net-like elevations. The chitinous surfaces are smooth and without tubercles in some places of the dorsum, while the rest is strongly granulated. One large digitate FM on the anterior margin and numerous small triangular protuberances on the dorsal and lateral sides of the pronotum. Almost the entire body (except the eye, labrum, fore and mid tarsi, second and third segments of hind tarsus, and inner side of the hind femur) covered with numerous small tubercles (Figs 19, 20). Only the macropronotal form is known. Coloration. Body dark brown, only some parts of the pronotal net-like elevations orange-red in color. Antenna dark brown, almost black. Proximal segments of the palpi light brown, and distal black. The visible part of the tegmen dark brown and without spots. Legs dark brown with lighter rings and patches. Fore and mid femora and tibiae dark brown with numerous small tubercles with somewhat lighter apices; fore and mid tarsi black, second segment with a light ring in the middle; 1 st tarsal pads whitish, claws brown. Hind femur dark brown with numerous small tubercles with lighter apices. Hind tibia dark brown with two weak light rings, one in the basal part, and another in the distal third. 1 st tarsus of hind leg pale brown, pads whitish; third segment pale brown with a dark ring in its distal part, claws brown. Tergites, sternites, epiproct, and cerci dark brown. Head. In dorsal and frontal view, vertex 1.7 times as wide as an eye. In frontal view, lateral carinae of the vertex considerably raised and granulated. Fossula elliptic and deep, but not easily observable because it is covered by large FM of the anterior pronotal margin. Lateral ocelli situated just below the level of the lower margin of a compound eye. Antennal groove significantly below the lower margin of a compound eye. In frontal view frontal costa bifurcated at the level of lateral ocelli into facial carinae, forming narrow parallel scutellum. Scutellum narrower than the antennal grove and the scapus (Figs 19E, 20E). Antenna elongated 13-segmented: scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 6 th) elongated and circular in crosssection; central or subapical segments (7 th and 8 th) strongly widened, pennate, 8 th being the widest antennal segment (about 2.8–3 times as long as wide); apical segment 9 th elongated and pennate, smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th reduced, very small, and borders between them barely visible. Antennomeres 3 rd to 9 th bear strong saw-like margins, likely because of the presence of large basiconic sensilla (Figs 19D, 20D). Pronotum. Pronotum rugose and granulated with numerous small tubercles and net-like elevations, but in some parts smooth and without tubercles (parts of the pronotal disc and some intervals of the median carina of the pronotum). The anterior part of the pronotum bears one large FM, and a few medium-sized and small protuberances. The posterior process of the pronotum slender, surpassing the hind knees for more than a half-length of the hind femur (macropronotal form). The disc of the pronotum is visually flat, but upon a closer inspection, many triangular elevations (projections) may be observed. A depression is visible on the pronotal disc positioned between the bases of the tegmina, then the disc becomes slightly elevated again. Caudad, pronotum gradually descending. The pronotal disc is rich in chitinous net-like elevations, more distinct in the interhumeral region. Medial carina extended along the whole length of the pronotum from the anterior margin to the pronotal apex, slightly undulated by the unpaired projections of variable size: the largest is high and digitiform massive FM, directed upwards and forwards above and before the head and covering the whole area of the fastigium of the vertex; triangular PM, a large triangular MM1, small triangular MM2; small MM3, and almost completely reduced MM4. Prozona subsquare. Prozonal and extralateral carinae in the prozona distinct, surpassing anterior margin of the pronotum as a small dentiform FL1 and large FL2, which is also well visible in frontal view. FL3 small and dentiform. PML1 and PML2 small. MML1 and MML2 distinct; MML3 very small; MML4 and MML5 indistinct. PL1 and PL2 small. Humeral angle obtuse with a pointed apex. ML small, reduced. Pronotum apex narrow, shallowly excised in the middle. The lower part of the lateral pronotal lobe bearing small teeth on the anterior and the posterior margins, with the middle region elongated as a large spine-like VL, directed mostly outwards and slightly backward (Figs 19A–C, 20A–C). Wings. Tegmina partly covered, not as easily discernible as in D. doriae and D. selysi. The visible part of the tegmen elliptical. Hind wing not reaching the pronotal tip for a few millimeters. Legs. Fore and mid femora elongated, but with robust armature on the dorsal and ventral margins. Fore and mid femora compressed laterally; with dorsal and ventral margins serrated with genicular tooth on the knees and additional 2–3 strongly projected and sharp teeth on each margin. Hind femur bearing small lappets on dorsal and ventral margins and one small protuberance on the ventral external carina. Genicular tooth large, while antegenicular tooth very small. Both sides of the dorsal margin of the hind tibia finely serrated, with 3–4 outer and 1–3 inner larger teeth. Abdominal apex. Female subgenital plate in ventral view bearing a longitudinal keel in the middle and has a triangular protrusion in the middle of the posterior margin. Ovipositor, in ventral view, seems to be elongated, but dorsal valvae are not visible because of the decurvation of the pronotal tip, which is directed downward. Ventral valvae are elongated, but for example in D. selysi the dorsal are more robust, which can be expected in D. aruanus sp. n. as well. Measurements (&female;&female; only, HT— holotype, PT— paratype). BL HT 13.45 mm, PT 11.80 mm; PnL HT 16.29 mm, PT 14.94 mm (without FM); PnW HT 8.40 mm, PT 8.49 mm; AnL HT 6.74 mm, PT 7.10 mm; TL HT 1.4 mm, PT 1.0 mm; TW HT 0.4 mm, PT 0.3 mm; fFL HT 4.40 mm, PT 4.22 mm; fFW HT 0.90 mm, PT 0.72 mm; mFL HT 4.41 mm, PT 4.21 mm; mFW HT 0.88 mm, PT 0.75 mm; hFL HT 8.63 mm, PT 7.42 mm; hFW HT 2.50 mm, PT 1.98 mm; OvL HT (Not visible, more than 1 mm), PT (not visible, more than 1 mm); AnL/fFL HT 1.51, PT 1.68; VW HT 1.00 mm, PT 1.05 mm; EW HT 0.59 mm, PT 0.61 mm; VW/EW HT 1.7, PT 1.7; SW HT 0.23 mm, PT 0.26 mm; AgW HT 0.45 mm, PT 0.40 mm; ScW HT 0.42 mm, PT 0.38 mm; SW/AgW HT 0.5, PT 0.65; SW/ScW HT 0.55, PT 0.68; As—L/W HT 2.8, PT 3.0; PrzW HT 3.71 mm, PT 3.43 mm; PrzL HT 1.85 mm, PT 1.78 mm; Prz—W/L HT 2.00, PT 1.92; TL/TW HT 3.5, PT 3.3; mFW/TW HT 2.2, PT 2.5; fFL/fFW HT 4.9, PT 5.6; mFL/mFW HT 5.0, PT 5.8; hFL/hFW HT 3.4, PT 3.7; T1L/T3L HT 1.1, PT 1.1., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 33-36, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Bolivar, I. (1898) Contributions a l'etude des Acridiens especes de la Faune indo et austro-malaisienne du Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 39, 66 - 101. https: // doi. org / 10.5962 / bhl. part. 9541","Bolivar, I. (1887) Essai sur les Acridiens de la tribu des Tettigidae. Annales de la Societe Entomologique de Belgique, 31, 175 - 313."]}

Subgenus Mnesarchus Stål, 1877, nom. resurr. Mnesarchus Stål, 1877: 55; synonymized with Discotettix by Bolívar (1887). Mnesarchus (as subgenus of Discotettix): Kevan 1966: 380. Discotettix (partim): Bolívar 1887: 306; Kirby 1910: 2 [listed in the catalog]; Blackith 1992: 46 [listed in the catalog]; Yin et al. 1996: 866 [listed in the catalog], Otte 1997: 32 [listed in the catalog]. Type species: Mnesarchus scabridus Stål, 1877, by original monotypy. Taxonomic notes. The genus Mnesarchus have been synonymized with Discotettix by Bolívar (1887). Kevan (1966) recognized it as distinct subgenus but later Mnesarchus has been considered a synonym of Discotettix again (Blackith, 1992; Yin et al., 1996; Otte, 1997). The type species is morphologically very different from other Discotettix members and is thus assigned to its own subgenus, Mnesarchus. It can be easily separated from other species of the genus by the set of the following characters: (I) frontal costa bifurcates at the lower margin of the compound eyes (bifurcation is in the lower third of the compound eyes in D. aruanus sp. n., D. belzebuth, D. doriae, D. kirscheyi sp. n., D. selysi, and D. sumatrensis sp. n.), (II) FM present as a low tubercle (developed and elevated in D. aruanus sp. n., D. belzebuth, D. doriae, D. kirscheyi sp. n., D. selysi, and D. sumatrensis sp. n.), (III) MM laterally compressed and elevated (spine-like in D. belzebuth, triangular protrusions in D. sumatrensis sp. n., D. kirscheyi sp. n., while very low in D. aruanus sp. n., D. doriae and D. selysi), (IV) MML low and triangular, compressed elevations (spine-like in D. belzebuth, triangular protrusions in D. sumatrensis sp. n., similarly formed in D. selysi), (V) interscapular area triangular, with large concavity (similarly to D. selysi, and D. sumatrensis sp. n., almost parallel in D. belzebuth), (VI) lateral and humeral carinae strongly toothed (similar to D. belzebuth, granulated in D. selysi, and D. sumatrensis sp. n.), (VII) weak ML (almost absent in D. scabridus, while well visible in other species), (VIII) VL complex, with a few spines (in other species usually with one main spine, and a saw-like margin) and (IX) tegmen more elongated (TL/TW > 5) than in the other species of the genus (TL/TW Composition. The type species of Mnesarchus is the only species presently assigned to this subgenus., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on page 49, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Stal, C. (1877) Orthoptera nova ex Insulis Philippinis descripsit. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlinger, 34 (10), 33 - 58.","Bolivar, I. (1887) Essai sur les Acridiens de la tribu des Tettigidae. Annales de la Societe Entomologique de Belgique, 31, 175 - 313.","Kevan, D. K. (1966) Some Orthoptera-Caelifera from the Philippine, Bismarck and Solomon Islands, with a few interesting records from New Guinea and the Moluccas. Entomologiske Meddelelser, 34, 375 - 420.","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261."]}

Discotettix (Discotettix) sumatrensis Skejo, Pushkar et Tumbrinck sp. n. (Figs 30–32) Vernacular name: Sumatran Spiky Pymgy Devil Type locality. Indonesia: Sumatra: Jambi province, 35 km N of Sungai Penuh, NP Kerinci-Seblat, Mt. Kerinci, 1500–2000 m a.s.l. Material examined. Type material. HOLOTYPE: 1&female; Indonesia: Sumatra: Jambi province, 35 km N of Sungai Penuh, NP KerinciSeblat, Mt. Kerinci, 1500–2000 m a.s.l. 8–22.XI.1999. Leg. A.V. Gorochov (Figs 30, 31) (ZISP); PARATYPES: 1&female;, 1&male; Indonesia: Sumatra: Jambi province, 35 km N of Sungai Penuh, NP Kerinci-Seblat, Mt. Kerinci, 1500–2000 m a.s.l. 8–22.XI.1999. Leg. A.V. Gorochov (Figs 30, 31) (ZISP). Type series depository. ZISP: Orthoptera collection of the Zoological Institute of the Russian Academy of Sciences (St. Petersburg), Russia. Additional material. 1&male; Indonesia: Sumatra: Siolak Daras.: Korinchi Valley (= Kerinci) 3100 ft. (= 945 m a.s.l.) III.1914. identified as D. selysi by K. Günther, photographs (Fig. 32) barcode NHMUK012498526 available at https://data.nhm.ac.uk/dataset/56e711e6-c847-4f99-915a-6894bb5c5dea/resource/05ff2255-c38a-40c9-b657- 4ccb55ab2feb/record/8248606 (NHMUK). Etymology. The new species is named sumatrensis (adjective masculine, third, vocal declension) after the type locality, Sumatra Island. Distribution and habitat. The species is currently known only from Mt. Kerinci, Sumatra, the highest vulcano in Indonesia and the Sumatra’s highest peak (above 1500 m). The mountain is surrounded by lush forest. It is an isolated mountain, so there is a chance of D. sumatrensis being a local endemic. Distribution is shown in Fig. 3. Specific diagnosis. The species is morphologically similar to D. kirscheyi sp. n. from Borneo and can be easily distinguished from other species. From D. scabridus the species differs by: (I) frontal costa bifurcates between the eyes, (II) FM well developed, (III) MM triangular and elevated, (IV) MML triangular and elevated, (V) lateral and humeral carinae granulated. The species can be distinguished from D. belzebuth by the following characters: (I) smaller body size, (II) peculiar coloration of antennae with lighter colored segments, (III) antenna with 7 th segment being the widest (not the 8 th as in most of the other species), (IV), pronotum with smaller triangular projections, except the digitate FM, (V) hind femur with large lappets (smaller in D. belzebuth), and (VI) ovipositor generally slenderer than in D. belzebuth. The species differs from D. selysi by (I) peculiar coloration of antennae with lighter colored segments, (II) widened antennal segments narrower than in D. selysi, (III) pronotal disc with numerous triangular protuberances and (IV) large lappets of hind femur (medium sized in D. selysi). The species is similar to D. kirscheyi sp. n. but can be easily distinguished from the mentioned by (I) longer FM, (II) stronger pronotal projections, (III) specialized and colorful antennal segments and (VI) smaller body size. Holotype description. (Figs 30, 31) General features. Medium sized, robust species (body length 11.7–13.5 mm); pronotum granulated and wrinkled, with one digitate FM on anterior margin and numerous triangular protuberances on dorsal and lateral sides.Almost the entire body (except eye, labrum, fore and mid tarsi, second and third segments of hind tarsus, inner side of the hind femur) covered with numerous small tubercles (Figs 31, 32). Macropronotal. Coloration. Body dark brown, almost completely covered by numerous small tubercles with lighter apices. Pronotum dark brown with darker projections and protuberances, tubercle-shaped ML of humeral angles on shoulders lighter. Antenna darkish with yellowish colored apical segments (after widened segments 8 th black with yellowish apex, 9 th yellow, 10 th –13 th black. Maxillary palpi dark brown. The visible part of the tegmen dark brown and without spots. Legs generally dark brown with lighter rings and patches. Fore and mid femora and tibiae dark brown with numerous small tubercles with lighter apices, fore and mid tarsi darkish, second segment with light ring near middle; 1 st tarsal pads whitish, claws brown. Hind femur dark brown with numerous small tubercles with lighter apices. Hind tibia in &female; dark brown with two light rings, one in basal part, and another—in distal third; while in &male; blackish brown, with weak (not distinguishable) light rings. 1 st tarsus of hind leg darkish, with light colored ring near apex, pads whitish; third segment blackish brown with a light ring near the middle, claws brown. Tergites, sternites, epiproct, and cerci dark brown. Head. In dorsal and frontal views, vertex 2.6 times in &male;, 2.5 times in &female; as wide as an eye. Fossula deep. Lateral ocelli situated at the level of the lower margin of the compound eye. In frontal view, frontal costa narrow, bifurcated between the lateral ocelli into a distinctly divergent, finely granulated facial carinae, concave in about the middle of its length, forming sand-clock-shaped scutellum. Frontal costa in &male; 1.6 times, in &female; 1.5 times, wider than antennal groove, 1.25 times in &male;, 1.2 times in &female; as long as antennal groove width. Antennal groove considerably below the lower margin of a compound eye (Fig. 30B). Antenna 13-segmented: scapus (1 st antennomere) and pedicel (2 nd antennomere) massive; basal segments (3 rd to 6 th) dark, elongated and circular in cross-section; central segment 7 th strongly widened and the widest antennal segment (3–3.6 times as long as wide), 8 th segment reduced, weakly pennate, dark, and with yellow apex; apical segment 9 th small, yellow, elongated, and pennate, smaller than the subapical and much larger than the rest of the apical segments; apical segments 10 th to 13 th again dark reduced, small, elongated in comparison to other species of the subgenus, and the borders between them barely visible. Antennomeres 3 rd to 9 th bearing saw-like margins, because of the presence of large basiconic sensilla. Pronotum. Pronotum wrinkled and granulated, covered by numerous small and medium-size tubercles. Posterior process of pronotum extended beyond hind knees for less than half of hind femur length. Disc of the pronotum depressed behind the well-developed shoulder, slightly descending backwards. Pronotum with 6 unpaired projections of variable size on medial carina (large digitate FM and 5 medium-sized medial), 3 pairs of FL projections, 7 pairs of more or less distinct mediolateral, 3 pairs of lateral and 1 pair of VL projections (well seen in profile). Prozona subsquare: prozonal length/width ratio 1.0 in &male;, 1.15 in &female;. Anterior margin of pronotum projected into a large digitate FM protuberance directed mainly forwards than upwards, covering whole vertex above. Prozonal and extralateral carinae in the prozona surpassing the anterior margin of the pronotum as dentiform FL1 and FL2. FL2 more distinct. Downwards, at the anterior margin of the pronotum less developed dentiform FL3. Behind FM medial carina extended along the whole length of the pronotum, with 5 unpaired large and small medial projections (better seen in profile): behind digitate FM, in the prozona, first small triangular PM protuberance. Next large triangular protuberance, between spinae of lateral lobes— MM1, then MM2, MM3 and MM4) protuberances, generally decreasing in size towards apex of pronotal process (sometimes posterior projection almost wanting). Smaller and lower then medial: 7 distinct double triangular MML. Mediolateral projections in the prozona present as more or less distinct PML1 double tubercle as a posterior elongation of prozonal carinae, and on same line PML2 double tubercle near border between prozona and metazona. Small MML1 between the shoulders, and large MML2 projection on the place where most Tetrigidae (Tetriginae) species have a posthumeral spot. Next three— MML3, MML4 and MML5 smaller, decreasing caudad. In the prozona lateral projections present as small double triangular PL1 and PL2 tubercles situated between the sulci on the line joining extralateral and humero-apical carinae. &Iukcy;n metazona, humero-apical carinae forming moderately sharp humeral angle, projected outwards as a small ML tubercle of humeral angle, behind this point joining the external lateral carinae. Apex of posterior pronotal process in dorsal view shallowly excised. Lower part of lateral lobe with serrate anterior and posterior margins, elongated as spine-like VL, directed outwards (Fig. 30A, C, E, F). Wings. Visible part of tegmen slightly elongated, oval with champlevé surface, visible part 3.2 times as long as wide in &male; and 2.9 in &female;; ratio of maximum width of mid femur (without teeth) / visible part of tegmina width 1.6 times in &male; and 1.29 times in &female;. Hind wing almost reaching the apex of the posterior pronotal process (ending 1.2–1.3 mm before it). Legs. Femora robust, compressed laterally, surface from smooth to rough, dorsal and ventral margins roughly serrate. Fore and mid femora bearing one genicular tooth on the knees on each margin, and additionally 3 strong teeth on the upper carina and 2–3 on the lower margin, lower usually smaller than upper; teeth on fore femur equal or smaller than on mid femur. Upper and lower margin of hind femur finely serrated, with 2–3 lappets on each margin, and with numerous small tubercles. Lateral area of hind femur with weak carinae that have 2–3 outgrowths, especially in ventro-external carina. Genicular teeth larger than antegenicular. Fore femur length/width ratio 3.9 in &male; and 3.8 in &female;. Mid femur length/width ratio 4.1 in &male; and 4.2 in &female;. Hind femur length/width ratio 3.0 in &male; and 3.2 in &female;. Both sides of the upper margin of hind tibia finely serrated, additionally with 2–3 outer and 2–3 inner (bit larger) teeth. Abdominal apex. Male subgenital plate in ventral view with shallowly excised apex, 1.4 times as long as wide, in lateral view 2.4 times as long as tall (Fig. 31A, B). Female subgenital plate in ventral view with triangular protrusion in middle of posterior margin. Ovipositor elongated, upper valve 5.0 times as long as wide. Lower valve of ovipositor 6.0 times as long as wide (maximal width) (Fig. 31C, D). Cerci length/width ratio near base 1.8 in &female; and 1.9 in &male;. Measurements (male paratype and female holoype). BL &male; 11.7 mm, &female; 13.5 mm; PnL &male; 12.8 mm, &female; 15.6 mm; PnW &male; 6.63 mm, &female; 7.86 mm; AnL &male; 7.2 mm, &female; 7.2 mm; TL &male; 1.6 mm, &female; 2.0 mm; TW &male; 0.91 mm, &female; 1.1 mm; fFL &male; 3.5 mm, &female; 3.8 mm; fFW &male; 0.89 mm, &female; 1.01 mm; mFL &male; 3.3 mm, &female; 3.8 mm; mFW &male; 0.8 mm, &female; 0.91 mm; hFL &male; 6.4 mm, &female; 7.5 mm; hFW &male; 2.13 mm, &female; 2.49 mm; OvL &female; 2.4 mm; AnL/fFL &male; 2.05, &female; 1.89; VW &male; 1.91 mm, &female; 2.05 mm; EW &male; 0.73 mm, &female; 0.82 mm; VW/EW &male; 2.6, &female; 2.5; SW &male; 0.36 mm, &female; 0.41 mm; AgW &male; 0.23 mm, &female; 0.27 mm; ScW &male; 0.29 mm, &female; 0.23 mm; SW/AgW &male; 1.6, &female; 1.5; SW/ScW &male; 1.25, &female; 1.2; As—L/W &male; 3.6, &female; 3.3; PrzW &male; 4.11 mm, &female; 5.62 mm; PrzL &male; 4.09 mm, &female; 4.91 mm; Prz—W/L &male; 1, &female; 1.15; TL/TW &male; 3.2, &female; 2.9; mFW/TW &male; 1.6, &female; 1.29; fFL/fFW &male; 3.9, &female; 3.8; mFL/mFW &male; 4.1, &female; 4.2; hFL/hFW &male; 3, &female; 3.2; T1L/T3L &male; 1.15, &female; 1.07., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 44-47, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418

Disconius shelfordi (Hancock, 1907), comb. n. (Figs 34–36) Vernacular name: Bornean Fallen Pygmy Devil Discotettix shelfordi Hancock, 1907b: 214 [original description, type locality: Borneo]; Kirby 1910: 575 [listed in the catalog]; Günther 1938: 303 [in review]; Yin et al. 1996: 866 [listed in the catalog]; Blackith 1992: 47 [listed in the catalog]; Otte 1997: 32 [listed in the catalog]. Type locality. Malaysia: Borneo: Kuching Note on the syntypes. The species description was based on two syntypes (male and female) labeled as “NW Borneo: Kuching, 22.IX.1899, Collector Dyak (Bornean native), det. J.L. Hancock ” and kept in the Oxford University Museum. As both specimens have been collected at the same locality and date, it is unlikely that they belong to different species. Thus, there is no need for lectotype designation. Material examined. Type material. SYNTYPE 1&female; NW Borneo: Kuching, 22.IX.1899., Collector Dyak, det. J. L. Hancock (Fig. 34) (OUMNH); SYNTYPE 1&male; NW Borneo: Kuching, 22.IX.1899., Collector Dyak, det. J. L. Hancock (OUMNH); Additional museum material. 2&female;&female;, 1&male; Indonesia: NE Borneo: Pajau River, leg. Mjöberg, det. J. Tumbrinck (Fig. 35) (NHRS). Additional material from online social media (iNaturalist): 1 adult Borneo: Malaysia: N Sarawak: Marudi (N4.041847, E114.8144) observed 15.X.2019., submitted on 5.II.2020. Photograph by Kinmatsu Lin (@kinmatsu), available at link inaturalist.org/observations/38326084 (Fig. 36). Distribution: Known only from N Borneo; in Sabah from the surroundings of Kuching (type locality) and from the region of the Pajau River; and from Sarawak reported from Marudi (Hancock 1907b; our data). Redescription. (Figs 34, 35) General features. Medium-sized species (body length 14.0– 15.8 mm) (in the original description stated as 17.5–19 mm, but it meant from the tip of the head to the tip of the pronotum ), relatively slender. The entire body finely granulated, covered by numerous small tubercles and with a few larger ones on the margins of the pronotal disc and the lateral lobes; in lateral view the pronotum almost flat, except for the median carina undulated by a few small wart-like and medium-sized semicircular compressed protuberances (Figs 34, 35). Macropronotal form. Coloration. Body color from dark brown and ferruginous brown to brown with an inexpressive grayish tint. Some parts of the body pale colored, previously yellowish: tubercles on the margin of the disc and lateral lobe of pronotum, VL and ML, connections of dark antennal segments, patches on femora, more or less distinct ring in the middle of fore and mid tibiae, two rings (I) in basal and (II) distal third of the hind tibia, distal segments and claws of fore and mid tarsi, and usually yellowish hind tarsus (except for the darker connections of tarsal segments). Living specimens have much more vivid coloration than the museum material. It is visible that the dorsum of the pronotum has an alternation of dark and pale tones (see iNaturalist observation in Material examined). Head. In dorsal and frontal view, vertex about 2.4 times as wide as the eye. Fossula visible, but not deep. Lateral ocelli situated between the compound eyes. The antennal groove situated at the level of the lower margin of the compound eye (in males) or a bit below (in females). In frontal view frontal costa narrow, bifurcates a bit above the lateral ocelli into a slightly divergent facial carinae, slightly concave inside in medium length and forming a narrow hour-glass shaped scutellum (Figs 34C, 35C). Antennal groove considerably wider than scutellum. Antennae 15 segmented, long. Antennomeres are shaped as follows: scapus (1 st antennomere) massive; pedicel (2 nd antennomere) large, basal antennomeres (3 rd to 7 th) elongated, 7 th being extremely elongated; central or subapical segments 8 th to 10 th pennate, 8 th slightly widened, while 9 th, and 10 th significantly widened and flattened; apical segment 11 th small; while apical segments 12 th to 15 th reduced in size, filiform. Pronotum. Pronotum finely granulated, covered by numerous small and few larger tubercles on the margin of the disc and the lateral lobe of the pronotum; almost flat, except for the wrinkled and scalloped median carina with small wart-like and medium-sized, compressed laterally, semicircular protuberances (different in specimens from different geographical populations); posterior process of pronotum very long, surpassing the hind knee for more than a half hind femur length (macropronotal form). Disc of the pronotum almost of the same height in the anterior part, slightly depressed behind the level of the tegmen apex and gradually descending backwards. Morphology of the pronotal projections variable. Pronotum with 5 unpaired projections of variable size on medial carina (FM and 4 medial projections); 2–3 pairs of FL; a pair of VL (better seen in profile); while among the mediolateral and lateral projections only one projection distinct per group. Prozona very short. Anterior margin of pronotum truncated, bearing small and weak triangular FM directed more upwards, then forwards. Prozonal and extralateral carinae low, tuberculated, not forming sharp saw-like or fan-like ridge, with small FL1 and more distinct dentiform. FL3 indistinct. Behind FM medial carina extended along the whole length of the pronotum, low, bearing four more or less distinct medial projections of variable size (well visible in lateral view). PM triangular, equal to or larger than FM and joined with the latter as a continuous two-humped structure in specimens of some populations. MM1, large. MM2 is the largest and the most massive projection, usually compressed laterally as a semicircular triangular protuberance. MM3 relatively large, while MM4 almost indistinct, marked by spot and darker than rest of the pronotal disc surface. Unlike in Discotettix species, Disconius shelfordi comb. n. has only one distinct projection from the mediolateral group, MML2, situated in the place where most Tetrigidae (Tetriginae) species have a posthumeral spot. Similar to the previous projections’ group, only one projection exists from the lateral group of the projections. In the metazona the humero-apical carinae forms a moderately sharp humeral angle, projected outwards as a small ML tubercle of the humeral angle, much larger than other tubercles along the margin of the pronotal disc and the lateral lobe; and behind this point the humero-apical carinae is joining the external lateral carinae. The apex of the posterior pronotal process in the dorsal view shallowly excised. The lower part of the lateral lobe with smooth anterior and posterior margins, without smaller teeth. The lateral lobe elongated as spine-like VL, directed exactly outwards, blunt (Figs 34A, B, 35A, B). Wings. The visible part of the tegmen elongated and oval, distinctly acuminate towards the apex, about 2.75 times as long as wide. Hind wings reach the apex of the pronotal process. Legs. Femora relatively slender, compressed laterally, and finely granulated, with numerous small teeth-like tubercles on the whole surface. Fore and mid femora bearing a hardly noticeable genicular tooth on the knees, and additionally 1–3 small teeth on the dorsal and ventral margin. Hind femur significantly compressed laterally, finely granulated, without any lappets on dorsal and ventral margins, smooth, and not bearing any recognizable outgrowth on the external carinae. Genicular and antegenicular teeth small. Both sides of the dorsal margin of the hind tibia only finely serrated, without any large teeth. Abdominal apex. Male subgenital plate in ventral view about 1.5 times as long as wide, in lateral view about 2 times as long as tall. Ovipositor elongated. Observed variability and differences found among populations, with implications for taxonomy. In the different populations of D. shelfordi comb. n., we have observed that the projections of the pronotum differ in size and shape. For example, among the specimens from the banks of the Pajau river the pronotum projections are much more expressed than in the specimens from Kuching (the type locality of the species). This primarily applies to lower and weaker medial protuberances in the specimens from Kuching. Especially FM and PM are quite small and wart-like, MM is medium size and slightly compressed laterally. In specimens from the Pajau river FM and PM are well expressed, and form a continuous double-hump structure. Other MMs are much more expressed, and more compressed laterally, they have semicircular form, especially the largest MM2. Only the examination of a larger series will allow drawing conclusions about the specific value of the aforementioned traits. Measurements (female syntype and a non-type male). BL &male; 13.99 mm, &female; 15.87 mm; PnL &male; 18.74 mm, &female; 20.81 mm; PnW &male; 7.35 mm, &female; 7.42 mm; AnL &male; 9.39 mm, &female; 9.68 mm; TL &male; 2.43 mm, &female; 2.62 mm; TW &male; 0.85 mm, &female; 0.96 mm; fFL &male; 3.22 mm, &female; 3.33 mm; fFW &male; 0.57 mm, &female; 0.62 mm; mFL &male; 4.01 mm, &female; 4.15 mm; mFW &male; 0.55 mm, &female; 0.61 mm; hFL &male; 7.99 mm, &female; 8.76 mm; hFW &male; 2.34 mm, &female; 2.71 mm; OvL &female; 1.41 mm; AnL/fFL &male; 2.91, &female; 2.9; VW &male; 1.1 mm, &female; 1.29 mm; EW &male; 0.45 mm, &female; 0.53 mm; VW/EW &male; 2.44, &female; 2.43; SW &male; 0.14 mm, &female; 0.19 mm; AgW &male; 0.39 mm, &female; 0.47 mm; ScW &male; 0.26 mm, &female; 0.38 mm; SW/AgW &male; 0.36, &female; 0.4; SW/ScW &male; 0.54, &female; 0.5; As-L/W &male; 2.35, &female; 2.62; PrzW &male; 3.18 mm, &female; 3.47 mm; PrzL &male; 1.5 mm, &female; 1.71 mm; Prz-W/L &male; 2.12, &female; 2.03; TL/TW &male; 2.86, &female; 2.73; mFW/TW &male; 0.65, &female; 0.64; fFL/fFW &male; 5.65, &female; 5.37; mFL/mFW &male; 7.29, &female; 6.8; hFL/hFW &male; 3.41, &female; 3.23; T1L/T3L &male; 1.27, &female; 1.19., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 54-55, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Hancock, J. L. (1907 b) Studies of the Tetriginae (Orthoptera) in the Oxford University Museum. Transactions of the Entomological Society of London, 1907, 213 - 244. https: // doi. org / 10.1111 / j. 1365 - 2311.1907. tb 01760. x","Kirby, W. F. (1910) A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). 3 (2). British Museum (Natural History), London, 674 pp.","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Yin, X. - C., Shi, J. & Yin, Z. (1996) Synonymic Catalogue of Grasshoppers and their Allies of the World (Orthoptera: Caelifera). China Forestry Publishing House, Beijing, 1266 pp.","Blackith, R. E. (1992) Tetrigidae (Insecta; Orthoptera) of Southeast Asia: Annotated catalogue with partial translated keys and bibliography. Ashford, Wicklow, 248 pp.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261."]}

Subfamily Scelimeninae Bolívar, 1887 Discotettiginae: Steinmann 1970, Storozhenko 2013, Tumbrinck 2014, syn. n. Discotettigiae: Hancock 1907a, Hancock 1907b, Willemse 1930, Günther 1938, syn. n. Discotettinae: Otte 1997, Mahmood et al. 2007, syn. n. Discotettigidae: Liang et Zheng 1998, Zheng 2005, Deng et al. 2007, syn. n. Notes. Herewith proposed definition of Scelimeninae is different from recent literature. Bolívar (1887) created the section (now subfamily) to include all Tetrigidae species with outwardly directed spines. Günther (1938) defined two groups within Scelimeninae —1) “Scelimenae verae” to include what are now Scelimeninae, i.e., the genera related to the type genus Scelimena, and 2) “Scelimenae spuriae” to include genera with lateral spines that are not closely related to the genus Scelimena. These “Scelimenae spuriae” are what we know today as tribes Criotettigini and Thoradontini. Kevan (1966) was the one to create the tribes Scelimenini, Thoradontini, Discotettigini, and Criotettigini. Thoradontini and Criotettigini were already then (Kevan 1966) defined as tribes for the taxa from the heterogeneous genera of section “Scelimenae spuriae”. Ad&zcaron;i&cacute; et al. (2020) confirmed the existence of two related tribes, Thoradontini and Criotettigini, which do not belong to Scelimeninae. The current definition of Discotettigini is very different from the Kevan’s (1966) definition, which was unfortunately based on a very partial list of genera by Günther (1955), so it may be regarded as the start of a highly chaotic taxonomy in Otte (1997) and later in Orthoptera Species File (Cigliano et al. 2022). Criotettigini and Thoradontini require revision and might be synonymous. Liang & Zheng (1998) and consequent Chinese authors ignored the taxonomic treatment by Kevan (1966), which we regard as useful, as far as the genera related to the type genus are included in the tribe. Muhammad et al. (2018) provided the most recent treatment for Scelimeninae: Scelimenini, which we follow in this paper. Composition and distribution (sensu Günther 1938, with additions from Muhammad et al. 2018 ). The subfamily currently counts 180 species within 23 genera. The tribe Scelimenini includes amphibious Asian and Papuan taxa with flat pronotum (genera Amphibotettix Hancock, 1906, Euscelimena Günther, 1938, Indoscelimena Günther, 1938, Paramphibotettix Günther, 1938, Platygavialidium Günther, 1938, Scelimena Serville, 1838, Tagaloscelimena Günther, 1938, and Tefrinda Bolívar, 1906), while the tribe Discotettigini stat. resurr. gathers Asian and Papuan corticolous taxa with undulated pronotum (genera Austrohancockia Günther, 1938, Bidentatettix Zheng, 1992, Disconius Skejo, Pushkar et Tumbrinck, gen. n. Discotettix Costa, 1864, Eufalconius Günther, 1938, Gavialidium Saussure, 1862, Gibbotettix Zheng, 1992, Kraengia Bolívar, 1909, Paragavialidium Zheng, 1994, Tegotettix Hancock, 1913). Genera Zhengitettix Liang, 1994, Hebarditettix Günther, 1938, Falconius Bolívar, 1898, and Dengonius Ad&zcaron;i&cacute;, Deranja, Franjevi&cacute; et Skejo, 2020 are of uncertain position within Scelimeninae, while Arulenus Stål, 1877 and Hirrius Bolívar, 1887 do not belong to this subfamily and should be removed from it, thus remaining without subfamily assignment for now., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on page 9, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418, {"references":["Bolivar, I. (1887) Essai sur les Acridiens de la tribu des Tettigidae. Annales de la Societe Entomologique de Belgique, 31, 175 - 313.","Steinmann, H. (1970) Check-list of the Tetricidae (Orthoptera) of the Oriental faunal region. Acta Zoologica Academiae Scientiarum Hungaricae, 16, 215 - 240.","Storozhenko, S. Y. (2013) Review of the subfamily Tripetalocerinae Bolivar, 1887 (Orthoptera: Tetrigidae). Zootaxa, 3718 (2), 158 - 170. https: // doi. org / 10.11646 / zootaxa. 3718.2.4","Tumbrinck, J. (2014) Taxonomic revision of the Cladonotinae (Orthoptera: Tetrigidae) from the islands of South-East Asia and from Australia, with general remarks to the classification and morphology of the Tetrigidae and descriptions of new genera and species from New Guinea and New Caledonia. In: Telnov, D., Barclay, M. V. L. & Pauwels, O. S. G. (Eds.), Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea. Vol. 2. Entomological Society of Latvia, Riga, pp. 345 - 396.","Hancock, J. L. (1907 a) Orthoptera Fam. Acridiidae. Subfam. Tetriginae. Genera Insectorum, 48, 1 - 79.","Hancock, J. L. (1907 b) Studies of the Tetriginae (Orthoptera) in the Oxford University Museum. Transactions of the Entomological Society of London, 1907, 213 - 244. https: // doi. org / 10.1111 / j. 1365 - 2311.1907. tb 01760. x","Willemse, C. J. M. (1930) Fauna Sumatrensis (Bijdrage Nr. 62). Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie, 73, 1 - 210.","Gunther, K. (1938) Revision der Acrydiinae, I. Sectiones Tripetalocerae, Discotettigiae, Lophotettigiae, Cleostrateae, Bufonidae, Cladonotae, Scelimenae verae. Mitteilungen aus dem Zoologischen Museum in Berlin, 23, 299 - 437.","Otte, D. (1997) Tetrigoidea and Tridactyloidea (Orthoptera: Caelifera) and Addenda to OSF. Vols. 1 - 5. Orthoptera Species File, 6, 1 - 261.","Mahmood, K., Idris, A. B. & Salmah, Y. (2007) Tetrigidae (Orthoptera: Tetrigoidea) from Malaysia with the description of six new species. Acta Entomologica Sinica, 50 (12), 1272 - 1284.","Liang, G. & Zheng, Z. (1998) Fauna Sinica, Insecta, Orthoptera: Tetrigoidea. Vol. 12. Science Press, Beijing, 278 pp.","Zheng, Z. - M. (2005) Fauna of the Tetrigoidea from Western China. Science Press, Beijing, 501 pp.","Deng, W. - A., Zheng, Z. & Wei, S. - Z. (2007) Fauna of Tetrigoidea from Yunnan and Guangxi. Guangxi Science and Technology Press, Nanning, 458 pp.","Kevan, D. K. (1966) Some Orthoptera-Caelifera from the Philippine, Bismarck and Solomon Islands, with a few interesting records from New Guinea and the Moluccas. Entomologiske Meddelelser, 34, 375 - 420.","Adzic, K., Deranja, M., Franjevic, D. & Skejo, J. (2020) Are Scelimeninae (Orthoptera: Tetrigidae) monophyletic and why it remains a question. Entomological News, 129, 128 - 146. https: // doi. org / 10.3157 / 021.129.0202","Gunther, K. (1955) Uber die Dornschrecken (Orth. Acrid. Tetrigidae) von Sumba und Flores mitfaunenhistorischen Anmerkungen zur Verbreitung einiger Gattungsgruppen der Tetrigidae imsudostasiatischen Inselbereich. Verhandlungen der Naturforschenden Gesellschaft in Basel, 66, 147 - 175.","Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2022) Orthoptera Species File. Version 5.0 / 5.0. Available from: http: // Orthoptera. SpeciesFile. org (accessed 10 August 2022)","Muhammad, A. A., Tan, M. K., Abdullah, N. A., Azirun, M. S., Bhaskar, D. & Skejo, J. (2018) An annotated catalogue of the pygmy grasshoppers of the tribe Scelimenini Bolivar, 1887 (Orthoptera: Tetrigidae) with two new Scelimena species from the Malay Peninsula and Sumatra. Zootaxa, 4485 (1), 1 - 70. https: // doi. org / 10.11646 / zootaxa. 4485.1.1","Hancock, J. L. (1906) Description of new genera and species of the orthopterous tribe Tettigidae. Entomological News, 17, 86 - 91.","Serville, J. G. A. (1838) Histoire naturelle des insectes. Orthopteres. Roret, Paris, 776 pp. https: // doi. org / 10.5962 / bhl. title. 95609","Bolivar, I. (1906) Rectificaciones y observaciones ortopterologicas. Boletin de la Real Sociedad Espanola de Historia Natural, 6, 384 - 393.","Zheng, Z. - M. (1992) New genera and new species of Tetrigidae (Orthoptera) from Sichuan and Yunnan. Entomotaxonomia, 14 (1), 1 - 7.","Costa, A. (1864) Acquiste fatti durante l-anno 1862. Annuario del Museo zoologico della Universita di Napoli, 2, 8 - 94.","Saussure, H. de (1862) Etudes sur quelques orthopteres du Musee de Geneve. Annales de la Societe Entomologique de France, 1, 469 - 494.","Bolivar, I. (1909) Nouvelles especes d'Acridiens du Musee de Geneve. Boletin de la Real Sociedad Espanola de Historia Natural, 9, 393 - 400.","Zheng, Z. - M. (1994) A new genus and three new species of Scelimenidae from China (Orthoptera: Terigoidea [Tetrigoidea]). Journal of Hubei University (Natural Science), 16 (1), 1 - 5.","Hancock, J. L. (1913) Studies of Tetriginae (Acrydinae) from Sarawak Museum, Borneo. The Sarawak Museum Journal, 1 (3), 39 - 54.","Liang, G. (1994) A new genus and a new species of Scelimenidae from Hainan, China (Orthoptera: Tetrigoidea). Entomological Research, 1, 33 - 34.","Bolivar, I. (1898) Contributions a l'etude des Acridiens especes de la Faune indo et austro-malaisienne du Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 39, 66 - 101. https: // doi. org / 10.5962 / bhl. part. 9541","Stal, C. (1877) Orthoptera nova ex Insulis Philippinis descripsit. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlinger, 34 (10), 33 - 58."]}

Genus Disconius Skejo, Pushkar et Tumbrinck gen. n. Type species: Discotettix shelfordi Hancock, 1907 (= Disconius shelfordi comb. n.). Justification for the establishment of the new genus. Head and pronotum morphology of Disconius shelfordi comb. n. differs too widely from other Discotettix species for it to be regarded as a member of the latter genus, so a new monotypic genus is established for this species, Disconius Skejo, Pushkar et Tumbrinck gen. n. Fastigial horns are lower, while the bifurcation of the frontal costa, lateral ocelli and antennal grooves have a much higher position than in any Discotettix species. Unlike FM and high pronotal projections in Discotettix, Disconius gen. n. lacks elevated FM (present as a small tubercle on the anterior pronotal margin); FL1 and FL3 are not strongly projected forwards as in Discotettix, and the strongest projections are ML of the shoulders area (similar to Tegotettix). The new genus is assigned to the subfamily Scelimeninae and to the tribe Discotettigini stat. resurr. on the basis of the width of the vertex (wide) and the weak elevation of the lateral carinae of the vertex; the typical arrangement of the pronotal projections (FM, MM, MML, ML); armed femora; and not widened hind tibiae and tarsi. The new genus is similar and likely related to the genera Tegotettix and Discotettx, but also superficially resembles Falconius. From Tegotettix and Falconius it can be easily distinguished by widened antennal segments, while from Discotettix, to which it was formerly assigned, it can be distinguished by a number of characteristics stated above. Molecular and comprehensive morphometric comparison of Disconius shelfordi comb. n. with large series of Discotettix, Tegotettix, and Falconius specimens is needed in the future, to elucidate the evolution of this curious taxon. Composition and distribution. A monotypic genus, including only Disconius shelfordi (Hancock, 1907) comb. n., known only from northern Borneo. Etymology. Because of the former taxonomic placement within Discotettix, and because of the superficial similarity to certain members of the genus Falconius due to its slender appearance and the shape of the pronotal projections, the names of two genera were combined into Disconius, meaning that this is both Discotettix -like and Falconius -like genus. Differential diagnosis. For the comparison with Discotettix, from which Disconius gen. n. has been removed, see the justification above, as well as the diagnosis of the genus Discotettix. From the genus Tegotettix (including T. armatus and T. bufocrocodil) the genus can be separated by head morphology — lateral ocelli positioned higher, frontal costa short before the bifurcation, vertex not bearing high horns, antennae with widened subapical antennal segments, and tibiae not armed. From the genus Falconius, Disconius gen. n. can be separated by the arrangement of the pronotal projections, the lack of flattened hind tarsi, and by widened apical segments of the antennae. Description. Head. Frontal costa bifurcation between the compound eyes; scutellum narrower than scapus; upper margin of the antennal groove above the lower margins of the compound eye; lateral (paired) ocelli between the compound eyes; eyes protruded above the vertex Antennae 15-segmented (1 st scapus; 2 nd pedicel; basal 3 rd –7 th elongated; central or subapical 8 th weakly compressed, 9 th elongated and strongly compressed; 10 th compressed; apical segment 11 th small; apical 12 th –15 th reduced, smooth and cylindrical). Vertex wider than a compound eye. Lateral carinae of the vertex in frontal view weakly elevated, medial carina of the vertex visible, anterior margin of the vertex slightly indrawn. Pronotum. Body robust, the ratio of the humeral angles’ width to the prozonal width more than 3.5. Anterior margin of the pronotum truncated or slightly excised, without strongly elevated FM; prozonal carinae distinct and parallel; extralateral carinae strong, with FL2 as a small elevation; FL3 weak; medial carina continuous along all the pronotum, tuberculated; MM high compressed elevations; PMLs and MMLs distinct; MML2 well developed as a high tubercle; ML triangular protrusion with a tubercle on its tip; interhumeral carinae distinct; interscapular area distinct and with parallel margins; lateral area as wide as the interscapular area; humero-apical, humeral, and lateral carinae with triangular or spine-like projections; VL protruded as a weak spine; paranota triangular; dorsum of the pronotum without Discotettix — characteristic net-like elevations, but still rough. Legs. Fore and mid femora carinated above, armed with a few small teeth on the dorsal and ventral margins; the dorsal margin of hind femora strongly armed; ventral margin with undulated carinae; the external surface of the hind femora with recognizable transverse ridges; hind tibia finely, densely serrate with numerous small teeth, but without large teeth; distal part of the hind tibia slightly widened, proximal tarsal segment slightly widened; first and the third tarsal segments of the hind legs almost equal in length; pulvilli typical for Scelimenini —first two angular and the third obtuse., Published as part of Skejo, Josip, Pushkar, Taras I., Kasalo, Niko, Pavlović, Marko, Deranja, Maks, Adžić, Karmela, Tan, Ming Kai, Rebrina, Fran, Muhammad, Amira Aqilah, Abdullah, Nurul Ashikin, Japir, Razy, Chung, Arthur Y. C. & Tumbrinck, Josef, 2022, Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae, pp. 1-64 in Zootaxa 5217 (1) on pages 52-53, DOI: 10.11646/zootaxa.5217.1.1, http://zenodo.org/record/7403418

Ectatoderus nigrofasciatus Tan & Wahab, 2021 (Fig. 3) Ectatoderus nigrofasciatus Tan et al., 2021: 412 Ectatoderus nigrofasciatus — Tan et al. 2022b: 600 Ectatoderus sp. — Tan & Wahab 2018: 132 Specimen examined. Holotype: BRUNEI DARUSSALAM • &male;; Belait District, Jalan Labi near Andulau Forest Reserve, N4.63354, E114.51047, 76.6±5.0 m.a.s.l.; 7 July 2019, 20h42; on rattan foliage; coll. M.K. Tan; BRU.19.70 (UBDM) Additional specimens examined. BRUNEI DARUSSALAM • 1&male;; Temburong District, Kuala Belalong Field Studies Centre, along Sungei Mata Ikan, N4.54742, E115.15715, 73.8± 27.9 m.a.s.l.; 22 September 2016, 22h33; on a foliage of a tree after rain; coll. M.K. Tan; KB.16.4 (ZRC) • 1&male;; Temburong District, Kuala Belalong Field Studies Centre, along Sungei Mata Ikan, N4.54733, E115.15720, 113.4± 14.2 m.a.s.l.; 23 September 2016, 19h00; on a foliage of a tree after rain; coll. M.K. Tan; KB.16.15 (ZRC) Remarks. When Ectatoderus nigrofasciatus Tan et al., 2021 was first described, the basal part of the male genitalia of the holotype was not examined (Tan et al., 2021: Fig. 7). The apical part of the male genitalia, previously thought to be the complete genitalia, was considered to resemble that of Ectatoderus argentatus Ingrisch, 2006 from Thailand. However, after examining more specimens collected from Brunei Darussalam, it was found that the basal half of the male genitalia of E. nigrofasciatus has in fact the medial valve curved to a spiral. This character is more similar to that in Ectatoderus angusticollis Chopard, 1969 from Singapore (Fig. 3). Below, the diagnosis is revised accordingly. Revised diagnosis. The species is characterised by the scapus being dorsally black, but ventrally and laterally brown, and the antennae being entirely black basally; the head has a distinct black band behind eyes; and the male phallic complex is very large and forming a spiral. The new species is most similar and closest in distribution to E. angusticollis. In both species, the pronotum is more strongly widening toward apex, and the TIs and TIIs are provided with black dorsal stripes. The male phallic complex is very similar to that of E. angusticollis, including the spiral-shaped medial valve and the shape of the ventro-external sclerite. The species differs from E. angusticollis by the male phallic complex being exceptionally larger despite of having very similar body size (Fig. 3); the scapus and antennae being black (instead of testaceous); and the head being provided with distinct black and white bands behind eyes. The new species is also similar to E. argentatus having the pronotum strongly widening apically and the frons dark; it differs by the scapus and basal antennal segments being darkened instead of yellowish brown, and by the male phallic complex forming a spiral at base instead of being merely curved. Distribution. Borneo, BRUNEI DARUSSALAM: Belait and Temburong [new locality record] Districts Type locality. BRUNEI DARUSSALAM, Belait District, near Andulau Forest Reserve, Published as part of Tan, Ming Kai, Japir, Razy, Chung, Arthur Y. C., Wahab, Rodzay Bin Haji Abdul & Robillard, Tony, 2022, Taxonomy and bioacoustics of scaly crickets (Orthoptera, Mogoplistidae Mogoplistinae) from Borneo and Singapore, pp. 177-189 in Zootaxa 5213 (2) on pages 181-182, DOI: 10.11646/zootaxa.5213.2.6, http://zenodo.org/record/7354377, {"references":["Tan, M. K., Japir, R., Chung, A. Y. C. & Wahab, R. A. (2021) New species and taxonomic notes of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Borneo. Zootaxa, 5048 (3), 407 - 421. https: // doi. org / 10.11646 / zootaxa. 5048.3.6","Tan, M. K., Japir, R., Chung, A. Y. C. & Wahab, R. A. (2022 b) Erratum: MING KAI TAN, RAZY JAPIR, ARTHUR Y. C. CHUNG & RODZAY BIN HAJI ABDUL WAHAB (2021) New species and taxonomic notes of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Borneo. Zootaxa, 5048, 407 - 421. Zootaxa, 5100 (4), 600. https: // doi. org / 10.11646 / zootaxa. 5100.4.9","Tan, M. K. & Wahab, R. A. (2018) Preliminary study on the diversity of Orthoptera from Kuala Belalong Field Studies Centre, Brunei Darussalam, Borneo. Journal of Orthoptera Research, 27 (2), 119 - 142. https: // doi. org / 10.3897 / jor. 27.24152","Ingrisch, S. (2006) New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae). Revue Suisse de Zoologie, 113 (1), 133 - 227. https: // doi. org / 10.5962 / bhl. part. 80345","Chopard, L. (1969) Family Gryllidae: Subfamilies Mogoplistinae, Myrecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Family Oecanthidae, Gryllotalpidae. In: Beier, M. (Ed.), Orthopterorum Catalogus. Vol. 12. Uitgeverij Dr. W. Junk N. V. ' s, Gravenhage, pp. 215 - 500."]}

Cycloptiloides bimaculatus Tan, Japir & Chung, 2021 (Figs 1A, 2) Cycloptiloides bimaculata Tan et al., 2021: 410 Cycloptiloides bimaculatus — Tan et al. 2022b: 600 Specimen examined. Holotype: EAST MALAYSIA • &male;; Sabah State, Sandakan, Sepilok, Rainforest Discovery Centre; N5.87580, E117.94299, 39.1± 5.4 m.a.s.l.; 9 January 2019, 19h11; on the leaf litter; coll. M.K. Tan, R. Japir, M. Binti & J.L. Yukang; SDK.19.20 (FRC) Additional specimens examined. EAST MALAYSIA • 1&male;; Sabah State, Sandakan, Sepilok, Rainforest Discovery Centre; N5.87554, E117.94135, 54.5± 6.1 m.a.s.l.; 11 May 2022, 20h58; on the leaf litter; coll. M.K. Tan & T. Robillard; SBH.22.19 (FRC) • 1&male;; Sabah State, Sandakan, Sepilok, Rainforest Discovery Centre; N5.87416, E117.93786, 89.2± 6.8 m.a.s.l.; 18 May 2022, 20h07; on the leaf litter; coll. M.K. Tan & T. Robillard; SBH.22.152 (MNHN) • 1&male;; Sabah State, Sandakan, Sepilok, Rainforest Discovery Centre; N5.87328, E117.93749, 95.1± 5.6 m.a.s.l.; 18 May 2022, 20h57; on the leaf litter; coll. M.K. Tan & T. Robillard; SBH.22.153 (FRC) • 1&male;; Sabah State, Sepagaya Waterfall, near Lahad Datu; N4.98354, E118.14498, 182.4± 7.4 m.a.s.l.; 13 May 2022, 11h39; on the leaf litter; coll. M.K. Tan & T. Robillard; SBH.22.50 (ZRC) • 1&male;, 1&female;; Sabah State, Mount Silam, lowland forest; N4.97589, E118.19060, 341 m.a.s.l.; 13 May 2022, 20h; on the leaf litter; coll. M.K. Tan & T. Robillard; TR22-10 (MNHN) Distribution. Borneo, Sabah State: Sepilok, Mount Silam [new locality record], Sepagaya near Lahad Datu [new locality record] Type locality. EAST MALAYSIA, Sabah State, Sepilok Calling song (1 &male;, in captivity, 26.4°C) (Fig. 2). The calling song generally consists of a sequence of echemes which can be highly variable in duration (8–26 s). The echeme sequence typically begins with the echemes more erratic in terms of number of syllables, syllable durations and intervals. Each echeme is made up of 4–10 syllables (= pulses), and has an average duration of 140.9±65.3 ms (78.0–249.1 ms). Consecutive echemes have an average interval of 371.6±86.3 ms (285.6–474.3 ms). The average syllable duration is 14.1±1.5 ms (11.9–15.6 ms) and the average interval between consecutive syllables is 7.2±1.6 ms (5.4–9.7 ms). The dominant frequency is 8.34 kHz. Compared to the continuous trill of Cycloptiloides timah Ingrisch, 2006 from Singapore (see Tan et al., under review), the calling song of C. bimaculatus also differs by a distinctly shorter syllable duration (14.1 ms in C. bimaculatus vs. 25.7 ms in C. timah) and by the intervals between consecutive syllables (7.2 ms vs. 104.5 ms). The dominant frequency is also distinctly lower (8.3 kHz vs. 9.1 kHz). It should be noted that the average temperature of the recordings for the two species are different (26.5°C in C. bimaculatus vs. 30.5°C in C. timah), which may have contributed partly to the differences in the call parameters between the two species., Published as part of Tan, Ming Kai, Japir, Razy, Chung, Arthur Y. C., Wahab, Rodzay Bin Haji Abdul & Robillard, Tony, 2022, Taxonomy and bioacoustics of scaly crickets (Orthoptera, Mogoplistidae Mogoplistinae) from Borneo and Singapore, pp. 177-189 in Zootaxa 5213 (2) on pages 178-181, DOI: 10.11646/zootaxa.5213.2.6, http://zenodo.org/record/7354377, {"references":["Tan, M. K., Japir, R., Chung, A. Y. C. & Wahab, R. A. (2021) New species and taxonomic notes of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Borneo. Zootaxa, 5048 (3), 407 - 421. https: // doi. org / 10.11646 / zootaxa. 5048.3.6","Tan, M. K., Japir, R., Chung, A. Y. C. & Wahab, R. A. (2022 b) Erratum: MING KAI TAN, RAZY JAPIR, ARTHUR Y. C. CHUNG & RODZAY BIN HAJI ABDUL WAHAB (2021) New species and taxonomic notes of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Borneo. Zootaxa, 5048, 407 - 421. Zootaxa, 5100 (4), 600. https: // doi. org / 10.11646 / zootaxa. 5100.4.9","Ingrisch, S. (2006) New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae). Revue Suisse de Zoologie, 113 (1), 133 - 227. https: // doi. org / 10.5962 / bhl. part. 80345"]}

Ornebius lupus Tan, sp. nov. (Figs 1C, 1D, 6, 7) Ornebius sp. — Tan et al. under review [calling songs from Singapore] Specimens examined. Holotype: SINGAPORE • 1&male;; Pasir Ris mangrove boardwalk; N1.37805, E103.95147; 19 March 2022, night time; on foliage of mangrove tree; coll. M.K. Tan; ZRC Paratypes: SINGAPORE • 1&male;; Pasir Ris mangrove boardwalk; N1.37805, E103.95147; 2 May 2022, night time; on foliage of mangrove tree; coll. M.K. Tan; MNHN-EO-ENSIF11343 Diagnosis. The new species has genitalia which appear most similar to those in the species group Ornebius (rufonigrus) Ingrisch, 1987, but differs by the dark-grey colouration; the maxillary palps being not so elongated (more similar to Ornebius alvarezi Tan, Ingrisch, Baroga-Barbecho & Yap, 2019 from the Philippines; elongated oblong in Ornebius dumoga Ingrisch, 2006 from Sulawesi and in Ornebius consternus Ingrisch, 2006 from Sulawesi), the FW colouration darker (orange in O. dumoga and in Ornebius rufonigrus Ingrisch, 1987 from Indochina and Malay Peninsula) and less covered by the pronotum (compared to O. consternus). The male genitalia of this new species differ from congeners from the species group by the internal sclerites at base being smaller instead of expanded. We also compared the new species with other species currently not placed in any species group (i.e., with unknown male genitalia). Our species resembles species with wholly dark/ black or smoky FWs, such as Ornebius obscuripennis (Chopard, 1930) from Sarawak and Ornebius fuscipennis (Chopard, 1929) from Sumatra (including Padang, Pulau Sipura, Pulau Telo and Batu Islands). The new species differs from O. obscuripennis by the pronotal lateral lobe and lateral parts of the head being dark-grey (i.e., unicolorous with dorsal parts) instead of forming a yellow lateral band. It also differs from O. fuscipennis by much smaller body size (BL 8.4 vs. 10.5 mm; PL 2.8 vs. 4.5 mm; TL 2.5 vs. 2 mm; FIIIL 4.4 vs. 7 mm); by the FW more uniformly coloured instead of smoky with posterior margin blackish; by the apical segment of the maxillary palps longer than subapical and third; by the pronotal disc not covered with yellow-brown scales; and by the legs that are not unicolourous yellowish. Lastly, the new species also differs from a syntopic species, Ornebius tampines Tan & Robillard, 2012 by the overall body and FW colouration, the body shape is more typical of Ornebius, instead of slenderly built, the shape of the male genitalia and by the calling songs (see Tan et al., under review). It is also somewhat similar to a sympatric species, O. pullus by the wholly black FWs, but differs by the lack of white margins along the lateral lobe and the posterior margin of the dorsal disc of the pronotum, by the shape of male genitalia and by the calling songs (see Tan et al., under review) Etymology. The species name refers to the dark-grey colouration, resembling the grey-wolf, Canis lupus. Description. Small species; habitus of male as shown in Fig 1C, 1D. Dorsum of head flattened, covered with dark-grey scales (Figs. 6A). Frontal rostrum 2.0 times wider than scapus (Fig. 6A). Scapus dark-grey (Fig 6A); basal antennal segments black, thereafter brown with some segments black (Figs 6B, 6C). Frons and mouthparts of dark colour (Fig. 6B). Maxillary palps pale grey; with apical (= fifth) segment longer than subapical (= fourth) segment, third segment of subequal length as subapical segment; apical segment pyriform (Fig. 6C). Lateral parts of head, including genae, of dark colour, without bands (Fig. 6C). Pronotal disc brown and wholly covered with dark-grey scales, about 1.3 times longer than wide, with anterior margin narrow and straight; lateral margin widening slightly posteriorly; covering only base of mirror of FW; posterior margin convex (Fig. 6D). Lateral lobe of pronotum also wholly covered with dark-grey scales (Figs 1D, 6C). TI with internal tympanum small and oval; without external tympanum. FIII 1.4–1.5 times longer than TIII; TIII 2.6 times longer than MTaIII. Femora generally pale, covered with marmoration of dark-grey scales except in basal parts. Tibiae and tarsi generally grey. Abdominal tergites covered with dark-grey scales; sternites pale. Abdominal apex darkened. Cercus covered with grey scales. Male. FW mostly infumated black, dorsal field darkest at posterior margin (Fig. 6D); lateral field likewise in some areas with infumated white veins (Fig. 6E). Supra-anal plate with last abdominal tergite and epiproct distinctly separated by a transverse suture. Last abdominal tergite transverse, posterior margin abruptly indented in middle; with oval pale spot in middle (not obvious in dry-pinned specimen) (Fig. 6F vs. 6G); posterior margin on either side of the indentation with two patches of short and strong setae. Supra-anal plate less transverse than last abdominal tergite, black coloured, apex subtruncate (Figs 6F, 6G). Subgenital plate black (Fig. 6H). Paraproct process black, elongate, cylindrical and tapering into a subacute apex (Fig. 6H). Genitalia as in Fig. 7. Central lobe of phallus with lateral valves (lv) elongate, apical area curved slightly ventrad in lateral view; tapering into obtuse apex, in apical area with inner margin curved in dorsal view; both sides together almost forming a dorsally and ventrally open tube. External sclerite (es) well developed, with external part more strongly sclerotized than inner part (in dorsal view). Medial valve (mv) membranous with base widened (in dorsal view) and upcurved (in lateral view). Internal sclerites (is) at base small but widened and curled, afterwards straight and elongate. Female. Unknown. Measurements (2&male;, in mm). &male; Holotype: BL = 8.4; PronL = 2.8; PronW = 2.2; FWL = 2.5; FWW = 2.3; FIIIL = 4.4; TIIIL = 3.1; MaTIIIL = 1.2. &male; paratype: BL = 8.4; PronL = 2.8; PronW = 2.2; FWL = 2.5; FWW = 2.3; FIIIL = 4.4; TIIIL = 2.9; MaTIIIL = 1.1. Ecology. This species is found among foliage of mangrove trees, occurring in syntopy with another mangrovespecialist, Ornebius tampines. Distribution. SINGAPORE Type locality. SINGAPORE, Pasir Ris mangrove Calling song. Described in Tan et al. (under review). The calling song of this species is similar to that of Ornebius rufonigrus Ingrisch, 1987 from the same species group with regard to echeme duration and dominant frequency, although the new species produces echeme-sequences composed of triple-chirps instead of double-chrips as in O. rufonigrus., Published as part of Tan, Ming Kai, Japir, Razy, Chung, Arthur Y. C., Wahab, Rodzay Bin Haji Abdul & Robillard, Tony, 2022, Taxonomy and bioacoustics of scaly crickets (Orthoptera, Mogoplistidae Mogoplistinae) from Borneo and Singapore, pp. 177-189 in Zootaxa 5213 (2) on pages 185-187, DOI: 10.11646/zootaxa.5213.2.6, http://zenodo.org/record/7354377, {"references":["Ingrisch, S. (1987) Neue Grillen von Borneo und aus Thailand (Insecta: Saltatoria: Grylloidea). Senckenbergiana Biologica, 68, 163 - 185.","Tan, M. K., Ingrisch, S., Baroga-Barbecho, J. B. & Yap, S. A. (2019) New species of Ornebius (Orthoptera; Mogoplistidae; Mogoplistinae) from Siargao Island of the Philippines. Zootaxa, 4590 (1), 166 - 176. https: // doi. org / 10.11646 / zootaxa. 4590.1.7","Ingrisch, S. (2006) New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae). Revue Suisse de Zoologie, 113 (1), 133 - 227. https: // doi. org / 10.5962 / bhl. part. 80345","Chopard, L. (1930) The Gryllidae of Sarawak. The Sarawak Museum Journal, 4 (12), 1 - 42.","Chopard, L. (1929) Spolia Mentawiensia: Gryllidae. Bulletin of the Raffles Museum, 2, 98 - 118.","Tan, M. K. & Robillard, T. (2012) Two new cricket species (Orthoptera: Gryllidae and Mogoplistidae) from the mangrove areas of Singapore. The Raffles Bulletin of Zoology, 60 (2), 411 - 420."]}

Ornebius lupus Tan, sp. nov. (Figs 1C, 1D, 6, 7) Ornebius sp. — Tan et al. under review [calling songs from Singapore] Specimens examined. Holotype: SINGAPORE • 1&male;; Pasir Ris mangrove boardwalk; N1.37805, E103.95147; 19 March 2022, night time; on foliage of mangrove tree; coll. M.K. Tan; ZRC Paratypes: SINGAPORE • 1&male;; Pasir Ris mangrove boardwalk; N1.37805, E103.95147; 2 May 2022, night time; on foliage of mangrove tree; coll. M.K. Tan; MNHN-EO-ENSIF11343 Diagnosis. The new species has genitalia which appear most similar to those in the species group Ornebius (rufonigrus) Ingrisch, 1987, but differs by the dark-grey colouration; the maxillary palps being not so elongated (more similar to Ornebius alvarezi Tan, Ingrisch, Baroga-Barbecho & Yap, 2019 from the Philippines; elongated oblong in Ornebius dumoga Ingrisch, 2006 from Sulawesi and in Ornebius consternus Ingrisch, 2006 from Sulawesi), the FW colouration darker (orange in O. dumoga and in Ornebius rufonigrus Ingrisch, 1987 from Indochina and Malay Peninsula) and less covered by the pronotum (compared to O. consternus). The male genitalia of this new species differ from congeners from the species group by the internal sclerites at base being smaller instead of expanded. We also compared the new species with other species currently not placed in any species group (i.e., with unknown male genitalia). Our species resembles species with wholly dark/ black or smoky FWs, such as Ornebius obscuripennis (Chopard, 1930) from Sarawak and Ornebius fuscipennis (Chopard, 1929) from Sumatra (including Padang, Pulau Sipura, Pulau Telo and Batu Islands). The new species differs from O. obscuripennis by the pronotal lateral lobe and lateral parts of the head being dark-grey (i.e., unicolorous with dorsal parts) instead of forming a yellow lateral band. It also differs from O. fuscipennis by much smaller body size (BL 8.4 vs. 10.5 mm; PL 2.8 vs. 4.5 mm; TL 2.5 vs. 2 mm; FIIIL 4.4 vs. 7 mm); by the FW more uniformly coloured instead of smoky with posterior margin blackish; by the apical segment of the maxillary palps longer than subapical and third; by the pronotal disc not covered with yellow-brown scales; and by the legs that are not unicolourous yellowish. Lastly, the new species also differs from a syntopic species, Ornebius tampines Tan & Robillard, 2012 by the overall body and FW colouration, the body shape is more typical of Ornebius, instead of slenderly built, the shape of the male genitalia and by the calling songs (see Tan et al., under review). It is also somewhat similar to a sympatric species, O. pullus by the wholly black FWs, but differs by the lack of white margins along the lateral lobe and the posterior margin of the dorsal disc of the pronotum, by the shape of male genitalia and by the calling songs (see Tan et al., under review) Etymology. The species name refers to the dark-grey colouration, resembling the grey-wolf, Canis lupus. Description. Small species; habitus of male as shown in Fig 1C, 1D. Dorsum of head flattened, covered with dark-grey scales (Figs. 6A). Frontal rostrum 2.0 times wider than scapus (Fig. 6A). Scapus dark-grey (Fig 6A); basal antennal segments black, thereafter brown with some segments black (Figs 6B, 6C). Frons and mouthparts of dark colour (Fig. 6B). Maxillary palps pale grey; with apical (= fifth) segment longer than subapical (= fourth) segment, third segment of subequal length as subapical segment; apical segment pyriform (Fig. 6C). Lateral parts of head, including genae, of dark colour, without bands (Fig. 6C). Pronotal disc brown and wholly covered with dark-grey scales, about 1.3 times longer than wide, with anterior margin narrow and straight; lateral margin widening slightly posteriorly; covering only base of mirror of FW; posterior margin convex (Fig. 6D). Lateral lobe of pronotum also wholly covered with dark-grey scales (Figs 1D, 6C). TI with internal tympanum small and oval; without external tympanum. FIII 1.4–1.5 times longer than TIII; TIII 2.6 times longer than MTaIII. Femora generally pale, covered with marmoration of dark-grey scales except in basal parts. Tibiae and tarsi generally grey. Abdominal tergites covered with dark-grey scales; sternites pale. Abdominal apex darkened. Cercus covered with grey scales. Male. FW mostly infumated black, dorsal field darkest at posterior margin (Fig. 6D); lateral field likewise in some areas with infumated white veins (Fig. 6E). Supra-anal plate with last abdominal tergite and epiproct distinctly separated by a transverse suture. Last abdominal tergite transverse, posterior margin abruptly indented in middle; with oval pale spot in middle (not obvious in dry-pinned specimen) (Fig. 6F vs. 6G); posterior margin on either side of the indentation with two patches of short and strong setae. Supra-anal plate less transverse than last abdominal tergite, black coloured, apex subtruncate (Figs 6F, 6G). Subgenital plate black (Fig. 6H). Paraproct process black, elongate, cylindrical and tapering into a subacute apex (Fig. 6H). Genitalia as in Fig. 7. Central lobe of phallus with lateral valves (lv) elongate, apical area curved slightly ventrad in lateral view; tapering into obtuse apex, in apical area with inner margin curved in dorsal view; both sides together almost forming a dorsally and ventrally open tube. External sclerite (es) well developed, with external part more strongly sclerotized than inner part (in dorsal view). Medial valve (mv) membranous with base widened (in dorsal view) and upcurved (in lateral view). Internal sclerites (is) at base small but widened and curled, afterwards straight and elongate. Female. Unknown. Measurements (2&male;, in mm). &male; Holotype: BL = 8.4; PronL = 2.8; PronW = 2.2; FWL = 2.5; FWW = 2.3; FIIIL = 4.4; TIIIL = 3.1; MaTIIIL = 1.2. &male; paratype: BL = 8.4; PronL = 2.8; PronW = 2.2; FWL = 2.5; FWW = 2.3; FIIIL = 4.4; TIIIL = 2.9; MaTIIIL = 1.1. Ecology. This species is found among foliage of mangrove trees, occurring in syntopy with another mangrovespecialist, Ornebius tampines. Distribution. SINGAPORE Type locality. SINGAPORE, Pasir Ris mangrove Calling song. Described in Tan et al. (under review). The calling song of this species is similar to that of Ornebius rufonigrus Ingrisch, 1987 from the same species group with regard to echeme duration and dominant frequency, although the new species produces echeme-sequences composed of triple-chirps instead of double-chrips as in O. rufonigrus.

Ornebius pullus Ingrisch, 2006 (Figs 1B, 4, 5) Ornebius pullus Ingrisch, 2006: 157 Ornebius pullus — He et al. 2021: 88 [assigning to species group Ornebius (flori) Ingrisch, 1998]; Tan et al. 2021: 416 [notes on new locality and variations with Singapore specimens]; Tan et al. under review [calling songs from Singapore] Ornebius cf. pullus — Tan 2012: 34; Tan & Ingrisch 2013: 25; Tan 2017: 55; Tan 2019: 335 Specimens examined. EAST MALAYSIA • 1&male;; Sabah State, Tabin Wildlife Reserve; N5.19496, E118.50326, 117.1± 10.3 m.a.s.l.; 15 May 2022, 22h17; on the branch of a small tree; coll. M.K. Tan, T. Robillard & R. Japir; SBH.22.95 (FRC) Remarks. The specimen collected from Sabah exhibits some differences with O. pullus collected from Belait (Brunei Darussalam) and Singapore: legs with more whitish marmoration, supra-anal plate whitish when alive. However, we could not identify clear-cut differences in the genitalia, including both the sclerotized median valve and the membranous lateral valves between the specimen from Sabah and those from Brunei Darussalam (Fig. 5). As Tan et al. (2021) have also suggested that there might be a polymorphic species and that more evidence was needed (e.g., DNA or acoustics) to better understand the species boundaries. At the moment, the external morphology and genitalia morphology are not congruent between specimens from Brunei Darussalam, Singapore and Sabah; we also do not have acoustic data for the Brunei population; and no molecular data are available. Furthermore, the specimen from Sabah also shares some similarities (but also with clear differences) in the external morphology of Ornebius minusculus (Chopard, 1929) from Sipura Island (Sumatra) (http://coldb.mnhn.fr/ catalognumber/mnhn/eo/ensif4531), although the genitalia for this species has not been examined. As such, without more specimens from Sabah to examine variations, we refrained from describing it as a new species, tentatively identified it as O. pullus and preferred to revise its taxonomy when the chance arises. Distribution. Brunei Darussalam (Belait and Brunei-Muara Districts), East Malaysia (Sabah State) [new locality record], Singapore. Type locality. BRUNEI DARUSSALAM, Brunei-Muara District, near bridge over river Sungai Lubang Barus on road coming from Tutong. Calling song (1 &male;, in captivity, 31.5°C) (Fig. 4). The calling song generally consists of a sequence of syllables (= pulses), which can be highly variable in duration (2–14 s or 4–25 syllables). The average syllable duration is 37.5±5.0 ms (31.0–46.0 ms) and the average interval between consecutive syllables is 501.2±20.4 ms (469.3–538.4 ms). The dominant frequency is 7.97±0.13 kHz (7.78–8.16 kHz). Compared to the calling song of Ornebius pullus from Singapore (see Tan et al., under review), the calling song of this specimen from Sabah has a similar call structure (i.e., the calls of both species are made up of a sequence of syllables). The biggest difference is in the interval between consecutive syllables being distinctly longer (501.2 ms vs. 400.0 ms) than that from Singapore. The syllable is only marginally longer in duration on average (37.5 ms vs. 32.4 ms) and the dominant frequency is not different (8.0 kHz vs. 8.1 kHz)., Published as part of Tan, Ming Kai, Japir, Razy, Chung, Arthur Y. C., Wahab, Rodzay Bin Haji Abdul & Robillard, Tony, 2022, Taxonomy and bioacoustics of scaly crickets (Orthoptera, Mogoplistidae Mogoplistinae) from Borneo and Singapore, pp. 177-189 in Zootaxa 5213 (2) on pages 182-185, DOI: 10.11646/zootaxa.5213.2.6, http://zenodo.org/record/7354377, {"references":["Ingrisch, S. (2006) New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae). Revue Suisse de Zoologie, 113 (1), 133 - 227. https: // doi. org / 10.5962 / bhl. part. 80345","He, Z., Ma, G. E., Long, J., Wang, Y., Zhang, T. & Ma, L. (2021) Taxonomy of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from China: five new species groups and three new species of the genus Ornebius Guerin-Meneville, 1844. Zootaxa, 4942 (1), 72 - 94. https: // doi. org / 10.11646 / zootaxa. 4942.1.3","Ingrisch, S. (1998) New Mogoplistinae from the Kinabalu region in Sabah, North Borneo (Insecta: Ensifera: Grylloidea: Mogoplistidae). Senckenbergiana Biologica, 77, 225 - 234.","Tan, M. K., Japir, R., Chung, A. Y. C. & Wahab, R. A. (2021) New species and taxonomic notes of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Borneo. Zootaxa, 5048 (3), 407 - 421. https: // doi. org / 10.11646 / zootaxa. 5048.3.6","Tan, M. K. & Ingrisch, S. (2013) New taxa and notes of some described species of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Singapore. Zootaxa, 3637 (1), 17 - 28. https: // doi. org / 10.11646 / zootaxa. 3637.1.2","Tan, M. K. (2017) Orthoptera in the Bukit Timah and Central Catchment Nature Reserves (Part 2): Suborder Ensifera. 2 nd Edition. Lee Kong Chian Natural History Museum, National University of Singapore, Singapore, 101 pp. [uploaded 16 June 2017]","Chopard, L. (1929) Spolia Mentawiensia: Gryllidae. Bulletin of the Raffles Museum, 2, 98 - 118."]}