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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.

Crematogaster modiglianii Crematogaster modiglianii has a widespread distribution in South-East Asia, but does not occur in Java, while another widespread species, C. seaeardi, occurs also in Java. What could explain the absence of C. modiglianii on Java? Menzel & Blüthgen (2010) revealed parabiotic associations between C. modiglianii and Camponotus (Myrmotarsus) rufifemur Emery, 1900. Camponotus rufifemur is known from the Indochinese Peninsula, Malay Peninsula, Borneo, Sumatra and Philippines, but not distributed in Java (Antmaps, 2021; AntWiki, 2021). The close relationship might have affected the absence of C. modiglianii in Java. Alternately, Java has a different climate from the other islands on the Sunda Shelf (Wilting et al., 2012). The climate of Java is drier than in other parts of the Sunda Shelf (Heaney, 1991). Veron et al. (2007) suggested that open and drier habitat in Java at the time might have affected the species distribution of the mongoose Herpestes species. Land bridges connecting the Sundaic regions might have allowed for the migration of C. modiglianii, but the species did not successfully colonize, probably due to ecological competition with related taxa., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 916, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Menzel F, Bluthgen N. 2010. Parabiotic associations between tropical ants: equal partnership or parasitic exploitation? Journal of Animal Ecology 79: 71 - 81.","Antmaps. 2021. Antmaps. Available at: http: // www. antmaps. org / (accessed on 1 March 2021).","Wilting A, Sollmann R, Meijaard E, Helgen KM, Fickel J. 2012. Mentawai's endemic, relictual fauna: is it evidence for Pleistocene extinctions on Sumatra? Journal of Biogeography 39: 1608 - 1620.","Heaney LR. 1991. A synopsis of climatic and vegetational change in South-East-Asia. Climatic Change 19: 53 - 61.","Veron G, Patou M-L, Pothet G, Simberloff D, Jennings AP. 2007. Systematic status and biogeography of the Javan and small Indian mongooses (Herpestidae, Carnovora). Zoologica Scripta 36: 1 - 10."]}

Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus, Pham, Thai Hong (2023): Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia. Zoological Journal of the Linnean Society 198 (3): 901-922, DOI: 10.1093/zoolinnean/zlad005, URL: https://academic.oup.com/zoolinnean/article/198/3/901/7197147

The Crematogaster difformis-subgroup The relationships of the sister-species in the C. difformis -subgroup were variable with respect to their geographical distribution patterns. Crematogaster ampullaris and C. seaeardi display an allopatric distribution, whereas C. difformis and C. tanakai display a sympatric distribution. The closely related C. ampullaris and C. seaeardi illustrate an interesting distribution pattern with C. ampullaris in Sulawesi and C. seaeardi widespread elsewhere in South-East Asia. The estimated dating suggests that C. ampullaris and C. seaeardi diverged approximately 4 Mya (Fig. 4; Table 2). The divergence implies a recent split of these populations and landmass distributions in that geological time (Hall, 2002) suggesting that the ancestor of both species dispersed from the Sundaic region to Sulawesi west-to-east across Wallace’s Line. Sulawesi is considered to have taken its present form due to the collision of different islands c. 15–5 Mya (Lohman et al., 2011). As the last possible connections between Borneo and Sulawesi were during the Pliocene or the Miocene (Morales & Melnick, 1998; Mercer & Roth, 2003), the colonization of Sulawesi by the common ancestor may have occurred much later after the island collision event. The ancestor of C. seaeardi and C. ampullaris may have expanded its range to western Sulawesi (eastern Sunda Shelf) during low sea-level periods and evolved into C. ampullaris. As the Makassar Strait is a deep trench, Borneo and Sulawesi were not connected when sea levels dropped. Their diversification resulted from ‘an inter-island dispersal combined with climatic vicariance event’ (Condamine et al., 2013). Crematogaster difformis and C. tanakai are sympatric in Borneo, and their habitats are spatially close to each other. Crematogaster difformis has an obligate mutualistic relationship with Lecanopteris and Platycerium ferns in canopy trees of lowland rainforests, and C. tanakai inhabits the same nests with C. difformis (Tanaka et al., 2009, 2012; Tanaka & Itioka, 2011). This restricted distribution demonstrates distinct endemism and is, therefore, of conservation significance. Lecanopteris fern species are found in the canopies of lowland tropical rainforests. The preservation of such primary forests will lead to the conservation of the endemic species. Molecular dating indicates that C. difformis and C. tanakai diverged approximately 1 Mya (Fig. 4; Table 2). The low divergence implies a recent speciation. As in C. aurita, the dispersal event and competition can explain similarly the position of C. mucronata, which is endemic to Sumatra (Fig. 4). Klaus et al. (2013) suggested early divergence of freshwater crabs Parathelphusa and subsequent appearance on Sumatra, since much of Sumatra was still submerged during the Late Miocene. In contrast, the ancestor of C. mucronata already appeared on fragmented Sumatra during the Late Miocene. Freshwater crabs require more land area for their habitats, but ants presumably do not require large areas as Yamane (2013) listed 126 ant species having colonized Krakatau Island after eruption. Faunal and genetic similarities between the Malay Peninsula and Sumatran populations have been documented for several taxa (Gorog et al., 2004; Lohman et al., 2011; Leonard et al., 2015), as the two areas were connected at sea level 20–30 m below the present level (Voris, 2000)., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 917, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Hall R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences 20: 353 - 431.","Lohman DJ, de Bruyn M, Page T, von Rintelen K, Hall R, Ng PKL, Shin H-T, Carvalho GR, von Rintelen T. 2011. Biogeography of the Indo-Australian Archipelago. Annual ReVieae of Ecology, EVolution, and Systematics 42: 205 - 226.","Morales JC, Melnick DJ. 1998. Phylogenetic relationships of the macaques (Cercopithecidae: Macaca), as revealed by high resolution restriction site mapping of mitochondrial ribosomal genes. Journal of Human EVolution 34: 1 - 23.","Mercer JM, Roth VL. 2003. The effects of Cenozoic global change on squirrel phylogeny. Science 299: 1568 - 1572.","Condamine FL, Toussaint EFA, Cotton AM, Genson GS, Sperling FAH, Kergoat GJ. 2013. Fine-scale biogeographical and temporal diversification processes of peacock swallowtails (Papilio subgenus Achillides) in the Indo-Australian Archipelago. Cladistics 29: 88 - 111.","Tanaka HO, Inui Y, Itioka T. 2009. Anti-herbivore effects of an ant species, Crematogaster difformis, inhabiting myrmecophytic epiphytes in the canopy of a tropical lowland rainforest in Borneo. Ecological Research 24: 1393 - 1397.","Tanaka HO, Yamane S, Itioka T. 2012. Effects of a ferndwelling ant species, Crematogaster difformis, on the ant assemblages of emergent trees in a Bornean tropical rainforest. Annals of the Entomological Society of America 105: 592 - 598.","Tanaka HO, Itioka T. 2011. Ants inhabiting myrmecophytic ferns regulate the distribution of lianas on emergent trees in a Bornean tropical rainforest. Biology Letters 7: 706 - 709.","Klaus S, Selvandran S, Goh JW, Wowor D, Brandis D, Koller P, Schubart CD, Streit B, Meier R, Ng PKL, Yeo DCJ. 2013. Out of Borneo: Neogene diversification of Sundaic freshwater crabs (Crustacea: Brachyura: Gecarcinucidae: Parathelphusa). Journal of Biogeography 40: 63 - 74.","Yamane SK. 2013. A review of the ant fauna of the Krakatau Islands, Indonesia. Bulletin of the Kitakyushu Museum of Natural History and Human History 11: 1 - 66.","Gorog AJ, Sinaga MH, Engstrom MD. 2004. Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys aehiteheadi). Biological Journal of the Linnean Society 81: 91 - 109.","Leonard JA, Den Tex R-J, Hawkins MTR, Munoz-Fuentes V, Thorington R, Maldonado JE. 2015. Phylogeography of vertebrates on the Sunda Shelf: a multi-species comparison. Journal of Biogeography 42: 871 - 879.","Voris HK. 2000. Maps of Pleistocene sea levels in South-East Asia: shorelines, river systems and time durations. Journal of Biogeography 27: 1153 - 1167."]}

The Crematogaster inflata-subgroup Crematogaster aurita is distributed in Indochina, and does not extend southward beyond the Isthmus of Kra. The phylogenetic position of the species was sister to C. onusta within the C. inflata -subgroup. Biogeographic analyses based on the distribution pattern and phylogenetic position of the species suggests that C. aurita appeared as a result of a dispersal event (Fig. 4). During the Pliocene to Pleistocene, terrestrial Indochinese animals might have had many opportunities to disperse to the Sundaic regions and vice versa, since these land areas were connected in the Sunda Shelf during glacial periods. The absence of C. aurita to the south of the Isthmus of Kra presumably resulted from limited dispersal capability and could also be a consequence of competition between related species. The distribution in the Indochinese Peninsula is possibly related to the preference of C. aurita for open, dry habitats. This preference could be explained by savannah corridors that are thought to have spanned Sundaland during glacial periods (Bird et al., 2005) and drier and seasonal habitats are currently present in the Indochinese Peninsula and Java. However, Crematogaster inflata and C. physothorax are sympatric in the Malay Peninsula. Our molecular dating analyses revealed that C. inflata and C. physothorax diverged approximately 6 Mya (Fig. 4; Table 2). This high divergence implies an old split of these populations. It is unclear whether this divergence resulted from a vicariant event or dispersal of the ancestor of both species. The sympatric distribution of these two species in the peninsula might be due to secondary contact after their speciation (Fig. 4). The estimated divergence suggests that the present geographic overlap of the two species is the result of independent colonization events from different geographic centres of origin. Hosoishi & Ogata (2009: figs 25, 28) represented the distribution records of Crematogaster inflata and C. physothorax based on the original descriptions and literature, but the allocations in the figures seem controversial. A record of C. inflata from Myanmar was based on misidentification. Yet extensive field surveys in the Indochinese Peninsula were not able to find C. physothorax (e.g. Khachonpisitsak et al., 2020), whose type locality is Thagata, Tenasserim, Myanmar and presumably this species is not distributed in the Indochinese Peninsula., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 916, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular South-East Asia during the Last Glacial Period: a savanna corridor in Sundaland? Quaternary Science ReVieaes 24: 2228 - 2242.","Hosoishi S, Ogata K. 2009. A taxonomic revision of the Asian endemic subgenus Physocrema of the genus Crematogaster (Hymenoptera: Formicidae). Zootaxa 2062: 15 - 36.","Khachonpisitsak S, Yamane S, Sriwichai P, Jaitrong W. 2020. An updated checklist of the ants of Thailand (Hymenoptera, Formicidae). ZooKeys 998: 1 - 182."]}

The Crematogaster Ʋacca-subgroup The phylogenetic relationship between the C. Ʋaccasubgroup and their ancestral state reconstruction suggests that their common ancestor occurred in the Sundaic region and its founder dispersed to Sumatra (Fig. 4). During glacial periods, the populations (C. yamanei) retracted into glacial forest refugia. When environmental conditions became favourable for expansion, the populations recolonized, but in the Sundaic region they were restricted to higher elevation areas due to competition with the C. inflata -subgroup and the C. difformis -subgroup, which were already present in this area at lower elevations. Crematogaster yamanei is endemic to Sumatra, but closely related to C. Ʋacca, which is distributed in the Malay Peninsula and Borneo. Meijaard’s (2004) mammal studies suggested that Sumatra shares elements of its fauna with both the Bornean/Javan and Malayan regions as a result of secondary contact. Those patterns are found in the younger taxon C. yamanei, but not in the older taxon C. mucronata. The restricted distributions of C. mucronata and C. yamanei could have resulted from another speciation event and extirpation of the relatives, respectively., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 917, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Meijaard E. 2004. SolVing mammalian riddles: a reconstruction of the Tertiary and Quaternary distribution of mammals and their palaeoenVironments in island South-East Asia. PhD Thesis, Australian National University, 349."]}

IS CREMATOGASTER SUBCIRCULARIS A RELICT SPECIES? Crematogaster subcircularis was revealed to have an interesting distribution pattern. This species is found in mountainous areas (600–1250 m elevation) of the Sundaic region: the Malay Peninsula (Fraser’s Hill, 1000 m), Borneo (Poring, 600–700 m; Crocker Range, 1100 m) and Java (Cikaniki, 1000–1250 m). This distribution suggests that the ancestor retreated to high-altitude forests during the ice age, whereas the other related species (C. modiglianii) remained in lowaltitude forests (0–600 m elevation). Despite a postulated savannah corridor through central Sundaland (Bird et al., 2005) that might have restricted the west– east dispersal of forest species, gallery forests in the Sunda Shelf could have served as corridors for forest species and facilitated the dispersal of C. subcircularis (Fig. 4). Rising sea-levels in the Late Pleistocene may have isolated the three populations. Molecular dating estimates and restricted highland distribution imply that C. subcircularis is a relict species. Several studies have hypothesized that continuous mountain ranges, which span Myanmar, Thailand, Laos, south China and Vietnam, acted as glacial refugia (Brandon-Jones, 1996; Gorog et al., 2004; Iyengar et al., 2005). Assuming that C. subcircularis is a relict with a wide distribution range in the glacial periods, those populations were fragmented into several geographical ones when the sea level was higher, then closely related or sister-species might have retreated to the high-elevation refugia of mainland South-East Asia (Surridge et al., 1999)., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on pages 917-918, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular South-East Asia during the Last Glacial Period: a savanna corridor in Sundaland? Quaternary Science ReVieaes 24: 2228 - 2242.","Brandon-Jones D. 1996. The Asian Colobinae (Mammalia, Cercopithecidae) as indicators of Quaternary climatic changes. Biological Journal of the Linnean Society 59: 327 - 350.","Gorog AJ, Sinaga MH, Engstrom MD. 2004. Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys aehiteheadi). Biological Journal of the Linnean Society 81: 91 - 109.","Iyengar A, Babu N, Hedges S, Venkataraman A, Maclean N, Morin PA. 2005. Phylogeography, genetic structure and diversity in the dhole (Cuon alpinus). Molecular Ecology 14: 2281 - 2297.","Surridge AK, Timmins RJ, Hewitt GM, Bell DJ. 1999. Striped rabbits in South-East Asia. Nature 400: 726 - 726."]}

The new genus Pumatiracia gen. nov. is described to accommodate a new species, P. venosa gen. et sp. nov. from Pu Mat National Park in Vietnam. The new genus is placed in the subtribe Parahiraciina of the Parahiraciini. It is compared with the genera Laohiracia Constant, 2021, Macrodarumoides Che, Zhang et Wang, 2012, Pseudochoutagus Che, Zhang et Wang, 2011, and Rostrolatum Che, Zhang et Wang, 2020 with which it shares possessing an elongate head. Illustrations of habitus, details, and male genitalia are given as well as a distribution map and photographs of the habitat. Laohiracia acuta Constant, 2021 is recorded for the first time from Vietnam, Pu Luong National Park; living specimens and habitat are illustrated, and the distribution map updated. The Parahiraciini fauna of Vietnam now comprises 14 species belonging to 11 genera.