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8) Phortica (Ashima) sagittaristula Chen & Wen, 2005 Phortica sagittaristula Chen & Wen in Chen et al., 2005a: 3954. Phortica (Ashima) sagittaristula: Chen & M��ca, 2012: 507. Diagnosis. Setulae just before prescutellar acrostichal setae as short as other acrostichal setulae (ch.15-0); anterior, sclerotized postgonite on right lateral lobe of aedeagal sheath bifurcated into subequal, long projections (��� Fig. 2C ��� in Chen et al. 2005a). Supplementary description (not repeating characters common to P. foliisetoides). Supracervical setae approximately 9. Cibarial, medial sensilla 7���8 per side; posterior sensilla approximately 6 per side. Specimens examined. Malaysia: 1♂, Ulu Gombak, Selangor, 8.xii.2013, M.J. Toda leg. (SEHU). Indonesia: 1♂, Bogor, West Java, 13���18.xi.2009, M.J. Toda leg. (SEHU). Distribution. China (Guangxi, Yunnan), Peninsular Malaysia *, Java*. Remarks. This species is closely related to the foregoing species, P. foliisetoides, in the molecular phylogeny reconstructed by Cao et al. (2011), but can be distinguished from it by the diagnostic characters., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on page 16, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Chen, H. W., Gao, J. J. & Wen, S. Y. (2005 a) Species diversity of the genus Phortica Schiner in Yunnan, China, with descriptions of nine new species (Diptera, Drosophildae [sic]). Journal of Natural History, 39, 3951 - 3978. https: // doi. org / 10.1080 / 00222930500533534","Chen, H. W. & Maca, J. (2012) Ten new species of the genus Phortica from the Afrotropical and Oriental regions (Diptera: Drosophilidae). Zootaxa, 3478 (1), 493 - 509. https: // doi. org / 10.11646 / zootaxa. 3478.1.42","Cao, H. L., Wang, X. L., Gao, J. J., Prigent, S. R., Watabe, H., Zhang, Y. P. & Chen, H. W. (2011) Phylogeny of the African and Asian Phortica (Drosophilidae) deduced from nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 61, 677 - 685. https: // doi. org / 10.1016 / j. ympev. 2011.08.002"]}

6) Phortica (Ashima) phyllochaeta (Tsacas & Okada, 1983) Amiota (Phortica) phyllochaeta Tsacas & Okada, 1983: 231. Phortica (Ashima) phyllochaeta: Chen & M��ca, 2012: 507. Diagnosis. Aedeagal sheath with two sclerotized, marginally serrated postgonites on anterior portion of left lateral lobe (��� Fig. 9 ��� in Tsacas & Okada 1983). Supplementary description (not repeating characters common to P. biclavata). Supracervical setae 12���14. Dorsomedial, tentorial apodeme 2/5 as long as basal, parallel portion of dorsolateral, tentorial apodeme. Cibarial, medial sensilla 9���11 per side; posterior sensilla approximately 6 per side. Specimen examined. Papua New Guinea: 1♂ paratype, Wau, Morobe district, 1,200 m a.s.l., 18.iii.1977, T. Okada leg. (NSMT). Distribution. Papua New Guinea. Remarks. This species closely resembles the foregoing species, P. biclavata, in having the aedeagal sheath with two marginally serrated postgonites on left lateral lobe and one anterior, large, marginally serrated postgonite and posterior, small, not serrated one on right lobe. However, these two species can be distinguished from each other by their diagnoses., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on pages 15-16, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Tsacas, L. & Okada, T. (1983) On the Oriental and New Guinean species of the genus Amiota Loew originally described by Duda as Phortica foliiseta, with descriptions of three new species (Diptera, Drosophilidae). Kontyu, 51, 228 - 237.","Chen, H. W. & Maca, J. (2012) Ten new species of the genus Phortica from the Afrotropical and Oriental regions (Diptera: Drosophilidae). Zootaxa, 3478 (1), 493 - 509. https: // doi. org / 10.11646 / zootaxa. 3478.1.42"]}

15) Phortica (Ashima) nudiarista Cheng & Chen, 2008 Phortica (Phortica) nudiarista Cheng & Chen in Cheng et al., 2008: 619. Phortica brachychaeta Chen & Toda in Chen et al., 2005b: 422 (part, misidentification). Phortica (Ashima) nudiarista: Chen & Máca, 2012: 507. Diagnosis. Arista apically with distinct, leaf-like expansion narrower than long and apically tapering and pointed but neither dorsal nor ventral branches (“ Fig. 8 ” in Cheng et al. 2008); one lateral lobe of aedeagal sheath with 2 anterior postgonites widely separated; the other lateral lobe with 1 posterior and 1 anterior postgonites (“Fig. 11” in Cheng et al. 2008). Supplementary description (not repeating characters common to P. pavriarista). Supracervical setae 9–11. Cibarial, medial sensilla approximately 9 per side; posterior sensilla approximately 5 per side. The antisymmetry is observed in the postgonites: in A-type, the left lateral lobe bears 2 anterior postgonites, and the right lobe 1 posterior and 1 anterior postgonites (“Fig. 11” in Cheng et al. 2008); but in B-type, vice versa. Specimens examined. China: 1♂ (B-type), Zhengxing, Jingdong, Yunnan, 24.vii.2009 (SEHU). Thailand: 1♂ (B-type), above Sangwal, Doi Suthep, Chiang Mai, 1,250 m a.s.l., 24.vii.2008, H. Bänziger leg. (SEHU); 1♂ (Atype), above Sangwal, Doi Suthep, Chiang Mai, 1,250 m a.s.l., 9.i.2008, H. Bänziger leg. (SEHU); 1♂ (B-type), path to Pha Lad, Doi Suthep, Chiang Mai, 5.x.2008, H. Bänziger leg. (SEHU); 1♂ (B-type), path to Bung La U O, Ban Thi Pho Ji, Umphang Distr., Tak, ~ 800 m, 22.i.2008, H. Bänziger leg. (SEHU). Malaysia: 1♂ (B-type), Ulu Gombak, Selangor, 9.xii.2013, M.J. Toda leg. (SEHU). India: 1♂ (B-type), Uttarakhand, Chamoli Distr., Simli (Narainbagar), 5.ix.2010, P.C. Sati leg. (SEHU). Distribution. China (Yunnan), Thailand *, Peninsular Malaysia *, India (Uttarakhand)*. Remarks. This species resembles P. pavriarista in the morphology of the aedeagal sheath and postgonites, but can be distinguished from it by the diagnostic characters.

1. Phortica (Ashima) afoliolata species group, new Diagnosis. Aedeagal, median rod distally diffentiated into two strongly sclerotized, asymmetric pieces (ch.58-4; Fig. 3F); arista with 4 or more ventral branches (ch.6-0; Fig. 2A); hypandrial phragma broader than lateral gonocoxites (ch.41-0; Fig. 3F). Shared characters. Some interfrontal setulae as long as lowest setulae on ocellar triangle (ch.3-0). Arista apically without leaf-like expansion (ch.7-0); longest, dorsal branch longer than longest seta on pedicel (Fig. 2A). Cibarial, medial sensilla all trichoid, apically pointed (ch.11-0; Fig. 2C). Prementum without pubescent patches (ch.12-0). Pseudotracheae in labellum 6 per side (ch.14-2; Fig. 2D). Setulae just before prescutellar acrostichal setae somewhat longer than other acrostichal setulae (ch.15-1). Mid and hind femora with gray patch submedially or largely dark brown (ch.16-1; Fig. 2G); all tibia with dark gray ring(s) or largely dark brown (ch.17,18,19-1; Fig. 2G). Apical, downward curved setula on fore tarsomere V neither thicker than surrounding setulae nor longer than tarsomere V (ch.20-0; Fig. 2E). Caudal bands on tergites III and IV medially interrupted/constricted (ch.21-2; Fig. 2F). Abdominal sternites III–V longer than wide; III not wider than IV; setulae on medial surface of III–V not so dense (ch.22–24,26–28-0; Fig. 2H). Epandrium widely round on ventral margin (ch.31-1; Fig. 3D); setae on and/or near ventral margin of epandrium not denser than those on other portions (ch.32-0; Fig. 3D), longer than others (ch.33- 1; Fig. 3D). Surstylus without expansion at caudoventral corner (ch.35-0; Fig. 3E), with peg-like teeth (ch.37-0; Fig. 3E). Hypandrium symmetric (ch.40-0; Fig. 3F). Pregonite (Fig. 3G,H) partly strongly sclerotized (ch.43-0), rod-like, with apical knob (ch.44-1), apically narrowing basal extension nearly rectangular to main body (ch.49-1) and sensilla on submedial and basal portions (ch.52,53-1) but no serration on ventrodistal portion (ch.50-0); apical knob without fine serrations (ch.45-0), ventrosubapically protruded, forming apically pointed barbule (ch.46,47-0). Aedeagal, outer membrane bearing neither spiny ornamentation nor apical, long, narrow tube (ch.55-1, ch.57-0; Fig. 3 F–H). Posterior, rod-like process of phallapodeme not longer than aedeagal median rod (ch.62-0; Fig. 3G,H). Aedeagal sheath and/or postgonites asymmetric (ch.63-1; Fig. 3F). Aedeagal sheath without lateral lobe(s) bearing strongly sclerotized postgonites (ch.65-0; Fig. 3G,H). Included species. Phortica (Ashima) afoliolata; P. (A.) eseta Chen; P. (A.) qingsongi An & Chen*; P. (A.) xishuangbanna Cheng & Chen; and P. (A.) efragmentata sp. nov. * The assignment was judged from the original description by An et al. (2015). Remarks. This species group first branched off in the subgenus Ashima (Fig. 1), retaining a number of plesiomorphic characters such as the plumose (with 5 and 4 or more dorsal and ventral branches, respectively; ch.5,6-0) arista but lacking the apical, leaf-like expansion (ch.7-0), cibarial, medial sensilla all trichoid, apically pointed (ch.11-0), the fore tarsomere V apically without specially developed, downward curved setula (ch.20-0), and the abdominal sternites III–V small, not so conspicuously differentiated in shape and setation from each other (ch.22– 24,26–28-0).

14) Phortica (Ashima) pavriarista Cheng & Chen, 2008 Phortica (Phortica) pavriarista Cheng & Chen in Cheng et al., 2008: 620. Phortica speculum (Maca & Lin, 1993): Chen et al., 2005b: 420 (part, misidentification). Phortica (Ashima) pavriarista: Chen & Máca, 2012: 507. Diagnosis. Arista only very slightly expanded apically (“Fig. 14” in Cheng et al. 2008); all postgonites strongly sclerotized, apically more or less pointed; posterior postgonite on only one lateral lobe of aedeagal sheath; lateral lobes submedially separated from each other; one lateral lobe of aedeagal sheath with 2 relatively close anterior postgonites (“Fig. 17” in Cheng et al. 2008). Supplementary description (not repeating characters common to P. foliiseta). Supracervical setae 12–14. Dorsomedial, tentorial apodeme 1/2 as long as basal, parallel portion of dorsolateral, tentorial apodeme. Longest, dorsal branch of arista shorter than longest seta on pedicel. Cibarial, medial sensilla 9–10 per side; posterior sensilla 4–6 per side. All tarsi with gray tarsomere V. The antisymmetry is observed in the postgonites: in A-type, the left lateral lobe bears 2 anterior postgonites, and the right lobe 1 posterior and 1 anterior postgonites (“Fig. 17” in Cheng et al. 2008); but in B-type, vice versa. Specimen examined. Thailand: 1♂ (B-type), above Sangwal, Doi Suthep, Chiang Mai, 1,250 m a.s.l., 6.i.2008, H. Bänziger leg. (SEHU). Distribution. China (Yunnan), Thailand *. Remarks. This species resembles P. andreagigoni in having the less expanded apex of arista, but can be distinguished from it by the diagnostic characters.

11) Phortica (Ashima) nakanoi Chen, 2012 Phortica (Ashima) nakanoi Chen in Chen & Máca, 2012: 497. Diagnosis. Aedeagal sheath with the following sclerotized postgonites: anterior one longest and strongly curved ventrad, and posterior one minute and apically pointed on left lateral lobe; anterior one broadest, somewhat triangular and gently curved ventrad, and posterior one long and apically pointed on right lateral lobe (“Figs 13, 14” in Chen & Máca 2012). Supplementary description (not repeating characters common to P. foliisetoides). Supracervical setae approximately 14. Dorsomedial, tentorial apodeme 1/2 as long as basal, parallel portion of dorsolateral, tentorial apodeme. Cibarial, medial sensilla 7–8 per side; posterior sensilla 6–7 per side. Setulae just before prescutellar acrostichal setae as short as other acrostichal setulae (ch.15-0). Specimen examined. Indonesia: 3♂ paratypes, Padang, West Sumatra, 11.xii.1989, S. Nakano leg. (SEHU). Distribution. Sumatra. Remarks. This species resembles P. saltiaristula in having both posterior postgonites apically pointed and neither anterior one bifurcated, but can be distinguished from it by the diagnostic characters.

3. Phortica (Ashima) nigrifoliiseta species group, new Diagnosis. Pseudotracheae in labellum 7 per side (ch.14-1; Fig. 9C); aedeagal, outer membrane apically with long, narrow tube (ch.57-1; Figs 6H, 8I); aedeagal, median rod moderately sclerotized (ch.58-1; Fig. 8K). Shared characters. Some interfrontal setulae as long as lowest setulae on ocellar triangle (ch.3-0). Arista with 1–3 dorsal branches (ch.5-1; Figs 8A, 9B) and apical, leaf-like expansion (ch.7-1; Figs 8A, 9B). Some anterior, cibarial, medial sensilla short, apically blunt, peg-like (ch.11-1; Fig. 8C). Prementum without pubescent patches (ch.12-0; Fig. 9D). Setulae just before prescutellar acrostichal setae somewhat longer than other acrostichal setulae (ch.15-1; Fig. 8B). Mid and hind femora and all tibiae entirely yellow (ch.16–19-0). Apical, downward curved setula on fore tarsomere V stout, as long as tarsomere V (ch.20-1; Fig. 8D). Caudal bands on tergites III and IV medially interrupted/constricted (ch.21-2). Abdominal sternite V wider than long (ch.25-1; Figs 8E, 9E). Epandrium widely round on ventral margin (ch.31-1; Figs 8G, 9F); setae on and/or near ventral margin of epandrium longer than those on other portions (ch.33-1; Figs 8G, 9F). Surstylus without expansion at caudoventral corner (ch.35-0; Fig. 9F). Hypandrial phragma as narrow as lateral gonocoxites (ch.41-1; Figs 8 I–K, 9G–I). Pregonite (Figs 8I, 9G) rod-like, with apical knob (ch.44-1); apical knob without fine serrations (ch.45-0). Aedeagal free sclerite absent (ch.59-0; Figs 8 I–K, 9G–I). Posterior, rod-like process of phallapodeme not longer than aedeagal median rod (ch.62-0). Aedeagal sheath and/or postgonites asymmetric (ch.63-1; Figs 8J, 9I), with lateral lobes bearing strongly sclerotized postgonites (ch.65-1; Figs 8 I–K, 9G–I). Included taxa. Phortica (Ashima) foliacea; P. (A.) longipenis Chen & Wen; P. (A.) montipagana An & Chen; P. (A.) semiannulata Gong, Zhu & Chen; P. (A.) spinosa Chen & Gao; P. (A.) glabra species subgroup; and P. (A.) nigrifoliiseta subgroup.

30) Phortica (Ashima) idiasta Gong, Zhu & Chen, 2018 Phortica (Ashima) idiasta Gong, Zhu & Chen, 2018: 440. Diagnosis. Pregonite partly less sclerotized (ch.43-0), not serrated on ventrodistal portion (ch.50-0); ventrosubapical portion of apical knob more or less protruded, forming barbule (ch.46-0); posterior postgonite dorsad curved, claw-like process, and anterior one basally divided into 3 long, outward oriented processes (“Figs 24, 25” in Gong et al. 2018). Supplementary description. Supracervical setae approximately 14. Dorsomedial, tentorial apodeme 1/2 as long as basal, parallel portion of dorsolateral, tentorial apodeme. Arista with arb = 2/0; longest, dorsal branch as long as longest seta on pedicel; apical, leaf-like expansion narrower than long, apically roundish and with minute projection. Cibarial, medial sensilla 6–7 per side; posterior sensilla 3–4 per side. The antisymmetry is observed in the phallic organs: in A-type, the posterior postgonite is present on the right side, the anterior one on the left side, and left gonocoxite of hypandrium more stretched out (“Fig. 25” in Gong et al. 2018); but in B-type, vice versa. Specimen examined. Thailand: 1♂ (B-type), Umphang, Tak, 30.v.2008, H. Bänziger leg. (SEHU). Distribution. China (Yunnan), Thailand *. Remarks. This species first branched off in the glabra subgroup (Fig. 1). The remaining three species form a cluster (BP = 96%) with three synapomorphies (Fig. 1): pregonite (Fig. 9G) nearly entirely sclerotized (ch.43-1), only angulated in ventrosubapical portion of apical knob (ch.46-1), and coarsely serrated on ventrodistal portion (ch.50-1).

25) Phortica (Ashima) longipenis Chen & Gao, 2005 (Fig. 6H) Phortica longipenis Chen & Gao in Chen et al., 2005a: 3957. Phortica (Ashima) longipenis: Chen & M��ca, 2012: 507. Diagnosis. Apical, leaf-like expansion of arista much narrower than long, apically with 1 long, medial and 2 minute, lateral, awn-like projections (��� Fig. 4A ��� in Chen et al. 2005a); strongly sclerotized arch on anterior margin of aedeagal sheath recurved at anterior ends as postgonites, posteriorly with one large and one small postgonites (projections); one anterior postgonite curved inward, longer, strongly sclerotized, apically with small claw, but the other curved dorsad, shorter, less sclerotized, without claw (Fig. 6H). Supplementary description (not repeating characters common to P. semiannulata). Supracervical setae approximately 16. Arista with arb = 1���3/0. Cibarial, posterior sensilla 6���7 per side. Pregonite without sensilla on submedial portion (ch.52-0). The antisymmetry is observed in the postgonites: in A-type, the large, posterior postgonite and strongly sclerotized, inward curved, anterior postgonite is present on the right side (Fig. 6H), while the small, posterior one and less sclerotized, dorsad curved, anterior one on the left side; but in B-type, vice versa. Specimens examined. China: 4♂ (A-type) 6♂ (B-type) paratypes, Mt. Jizu, Binchuan, Yunnan, 1,840 m a.s.l., 17.viii.2000, H.-W. Chen leg. (SEHU). Thailand: 1♂ (A-type), above Sangwal, Doi Suthep, Chiang Mai, 1,250 m a.s.l., 25.vii.2008, H. B��nziger leg. (SEHU). Distribution. China (Yunnan), Thailand *., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on page 28, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Chen, H. W., Gao, J. J. & Wen, S. Y. (2005 a) Species diversity of the genus Phortica Schiner in Yunnan, China, with descriptions of nine new species (Diptera, Drosophildae [sic]). Journal of Natural History, 39, 3951 - 3978. https: // doi. org / 10.1080 / 00222930500533534","Chen, H. W. & Maca, J. (2012) Ten new species of the genus Phortica from the Afrotropical and Oriental regions (Diptera: Drosophilidae). Zootaxa, 3478 (1), 493 - 509. https: // doi. org / 10.11646 / zootaxa. 3478.1.42"]}

Toda, Masanori J., Bänziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit, Katoh, Toru (2020): Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species. Zootaxa 4789 (1): 1-54, DOI: https://doi.org/10.11646/zootaxa.4789.1.1

29) Phortica (Ashima) kerinciensis Toda, sp. nov. (Fig. 8) Diagnosis. Surstylus without peg-like teeth (ch.37-1); aedeagal sheath with 1 posterior and 2 anterior postgonites all strongly sclerotized, recurved, long and thick (Fig. 8 I–K). Description (not repeating characters common to P. nigrifoliiseta). Supracervical setae 25–27. Dorsomedial, tentorial apodeme slightly longer than 1/2 basal, parallel portion of dorsolateral, tentorial apodeme. Arista with arb = 1–3/0. Cibarial, medial sensilla approximately 10 per side; posterior sensilla approximately 7 per side. All tarsomere Is slightly longer than rest tarsomeres together. One anterior postgonite deeply bifurcated, but the other with triangular flap submedially (Fig. 8 I–K). Measurements (holotype /range of 6♂ paratypes, in mm): BL = 4.13/3.94–4.32, ThL = 2.07/1.92–2.14, WL = 3.31/3.11–3.47, WW = 1.39/1.34–1.45. Indices (holotype /range of 6♂ paratypes, in ratio): FW/HW = 0.44/0.44–0.47, ch/o = 0.12/0.10–0.11, prorb = 1.24/1.05–1.28, rcorb = 0.52/0.48–0.60, vb = 0.55/0.36–0.67, dcl = 0.51/0.51–0.62, presctl = 0.66/0.60–0.76, sctl = 0.99/0.92–1.09, sterno = 1.00/0.92–0.99, orbito = 2.04/1.56–1.90, dcp = 0.26/0.25–0.31, sctlp = 1.12/1.03–1.29, C = 2.78/2.25–2.71, 4c = 1.21/1.31–1.48, 4v = 2.75/2.69–3.33, 5x = 0.91/0.90–1.13, ac = 3.00/3.00–3.45, M = 0.54/0.59–0.69, C3F = 0.71/0.66–0.76. The antisymmetry is observed in the postgonites: in A-type, the posterior postgonite and the bifurcated, anterior one are present on the right side, but only the other anterior postgonite on the left side; but in B-type, vice versa (Fig. 8J). Holotype. ♂ (B-type; MZB), “ Mt. Kerinci, Jambi, Indonesia, 1°45’5.9”S 101°15’34.6”E, 1800–2000 m a.s.l., 6.x.2004, M.J. Toda ”. Paratypes. Indonesia: 1♂ (A-type) 5♂ (B-type), same data as the holotype (3♂ MZB, 3♂ SEHU). Distribution. Sumatra (Jambi). Etymology. Referring to the type locality.

Genus Phortica Schiner, 1862 Phortica Schiner, 1862: 433 (as genus); M��ca, 2003: 251; B��chli et al., 2004: 70; Chen et al., 2005a: 3953. Type species: Drosophila variegata Fall��n, 1823. Phortica (as subgenus of Amiota Loew): Wheeler, 1952: 167; Okada, 1971: 96; M��ca, 1977: 116. Diagnosis. Ocellar tubercle with additional pair of setulae below ocellar setae; proclinate orbital seta nearer to inner vertical seta than to ptilinal fissure; palpus with hollow-shaped or depressed sense organ (modified from Chen et al. 2005a). Characters commonly seen across the genus were described by Chen et al. (2005a), and therefore are not repeated here., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on page 9, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Schiner, I. R. (1862) Vorlaufiger Commentar zum dipterologischen Theile der \" Fauna Austriaca \", mit einer naheren Begrundung der in derselben aufgenommenen neuen Dipteren-Gattungen. V. [part]. Wiener Entomologische Monatschrift, 6, 428 - 436.","Maca, J. (2003) Taxonomic notes on the genera previously classified in the genus Amiota Loew (Diptera: Drosophilidae, Steganinae). Acta Universitatis Carolinae Biologica, 47, 247 - 274.","Bachli, G., Vilela, C. R., Andersson Escher, S. & Saura, A. (2004) The Drosophilidae (Diptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica. Vol. 39. Brill, Leiden, 362 pp.","Chen, H. W., Gao, J. J. & Wen, S. Y. (2005 a) Species diversity of the genus Phortica Schiner in Yunnan, China, with descriptions of nine new species (Diptera, Drosophildae [sic]). Journal of Natural History, 39, 3951 - 3978. https: // doi. org / 10.1080 / 00222930500533534","Fallen, C. F. (1823) Geomyzides Sveciae. Berlingianis, Lundae (Lund), 8 pp.","Wheeler, M. R. (1952) XI. The Drosophilidae of the Nearctic region, exclusive of the genus Drosophila. The University of Texas Publication, 5204, 162 - 218.","Okada, T. (1971) A revision and taxometric analysis of the genus Amiota Loew of Japan and adjacent countries (Diptera, Drosophilidae). Kontyu, 39, 82 - 98.","Maca, J. (1977) Revision of Palaearctic species of Amiota subg. Phortica (Diptera, Drosophilidae). Acta Entomologica Bohemoslovaca, 74, 115 - 130."]}

21) Phortica (Ashima) halimunensis Toda, sp. nov. (Fig. 6) Diagnosis. Apical piece of aedeagal, median rod small and flat plate (Fig. 6E,F). Description (not repeating characters common to P. tanabei). Supracervical setae 8–11. Dorsomedial, tentorial apodeme slightly shorter than basal, parallel portion of dorsolateral, tentorial apodeme. Arista with arb = 1–2/1; apical, leaf-like expansion apically somewhat roundish and with short, awn-like projection (Fig. 6A). Cibarial, medial sensilla 5–6 per side; posterior sensilla approximately 5 per side. Fore and hind tarsomere Is as long as 3 succeeding tarsomeres together; mid one slightly longer than rest tarsomeres together. Measurements (holotype /range of 4♂ paratypes, in mm): BL = 2.90/2.46–2.74, ThL = 1.39/1.15–1.37, WL = 2.11/1.83–2.11, WW = 0.95/0.86–0.98. Indices (holotype /range of 4♂ paratypes, in ratio): FW/HW = 0.46/0.45–0.47, ch/o = 0.07/0.07–0.08, prorb = 1.13/1.11–1.29, rcorb = 0.66/0.47–0.69, vb = 0.31/0.33–0.46, dcl = 0.44/0.37–0.47, presctl = 0.41/0.45–0.51, sctl = 1.00/1.00–1.07, sterno = 0.88/0.85–0.99, orbito = 1.84/2.00–2.39, dcp = 0.22/0.21–0.26, sctlp = 1.08/0.93–1.12, C = 2.31/2.06–2.20, 4c = 1.44/1.43–1.70, 4v = 2.87/2.62–3.21, 5x = 1.30/1.04–1.28, ac = 3.72/3.34–4.03, M = 0.70/0.59–0.80, C3F = 0.54/0.61–0.72. Holotype. ♂ (MZB), “ Cikaniki, Mt. Halimun, West Java, Indonesia, 6°44’42.5” S 106°32’14.5” E, 1,051 m a.s.l., 4.xi.2009, ex around eyes, M.J. Toda ”. Paratypes. Indonesia: 1♂, same data as the holotype, except 5.xi.2009 (MZB); 1♂, ditto, except ex tree trunk (SEHU); 2♂, same data as the holotype, except 9.xi.2009 (SEHU). Distribution. Java (West Java). Remarks. This species is coupled with P. tanabei (BP = 72%; Fig. 1), sharing the aedeagal, median rod distally diffentiated into one asymmetric sclerite piece (ch.58-3; Fig. 6E,F), but can be distinguished from it by the diagnostic character. Etymology. Referring to the type locality.

17) Phortica (Ashima) brachychaeta Chen & Toda, 2005 Phortica brachychaeta Chen & Toda in Chen et al., 2005b: 422. Phortica (Ashima) brachychaeta: Chen & M��ca, 2012: 507. Diagnosis. Arista apically with somewhat triangular, leaf-like expansion but neither dorsal nor ventral branches (��� Fig. 5 ��� in Chen et al. 2005b); all postgonites nearly straight in ventral view; posterior postgonite apically pointed, as long as anterior ones; 2 anterior postgonites on one lateral lobe nearly parallel (��� Fig. 8 ��� in Chen et al. 2005b). Supplementary description (not repeating characters common to P. speculum). Supracervical setae 14���16. Cibarial, medial sensilla approximately 8 per side; posterior sensilla 6���7 per side. The antisymmetry is observed in the postgonites: in A-type, the left lateral lobe bears 2 anterior postgonites, and the right lobe 1 posterior and 1 anterior postgonites; but in B-type, vice versa (��� Fig. 8 ��� in Chen et al. 2005b). Specimen examined. Thailand: 1♂ (B-type), Chang Khian Stream gully, Doi Suthep, Chiang Mai, ~ 1,100 m a.s.l., 13.ii.2009, H. B��nziger leg. (SEHU). Distribution. China (Guangdong, Yunnan), Thailand *. Remarks. This species resembles P. speculum within the foliiseta complex, in having the abdominal sternite III wider than long (ch.22-1), widest among sternites (ch.23-1), and with dense setulae on medial surface (ch.26-1), but can be distinguished from it by the diagnostic characters., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on page 21, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Chen, H. W., Toda, M. J. & Gao, J. J. (2005 b) The Phortica (s. str.) foliiseta species-complex (Diptera, Drosophilidae) from China and its adjacent countries. Acta Zootaxonomica Sinica, 30, 419 - 429.","Chen, H. W. & Maca, J. (2012) Ten new species of the genus Phortica from the Afrotropical and Oriental regions (Diptera: Drosophilidae). Zootaxa, 3478 (1), 493 - 509. https: // doi. org / 10.11646 / zootaxa. 3478.1.42"]}

32) Phortica (Ashima) glabra Chen & Toda, 2005 Phortica glabra Chen & Toda in Chen et al., 2005b: 424. Phortica (Ashima) glabra: Chen & Máca, 2012: 507. Diagnosis. Arista with 1 or 2 ventral branches (ch.6-1); posterior postgonite present only on right side of aedeagal sheath, short, apically blunt, and left lateral lobe long, broad, somewhat sinuated, apically with very large, trifurcated, anterior postgonite (“Figs 17, 18” in Chen et al. 2005b). Supplementary description (not repeating characters common to P. zhangyuanae). Supracervical setae 20–22. Apical, leaf-like expansion of arista apically pointed or with minute, awn-like projection. Cibarial, medial sensilla 7–8 per side. All tarsi with gray tarsomeres IV and V. Specimens examined. China: 5♂ paratypes, Nanlling, Luyuan, Guangdong, 2.v.2004, H.-W. Chen leg. (SEHU). Distribution. China (Jiangxi, Guangdong, Guangxi).

4) Phortica (Ashima) efragmentata Toda, sp. nov. (Figs 2, 3 D–G) Diagnosis. Posterior piece of aedeagal median rod widely bifurcated (Fig. 3F); aedeagal free sclerite absent (ch.59- 0; Fig. 3F). Description. Supracervical setae 18–19. Dorsomedial, tentorial apodeme 3/4 as long as basal, parallel portion of dorsolateral, tentorial apodeme.Arista with arb = 8/4 (Fig. 2A). Palpus with hollow-shaped sense organ (Fig. 2B). Cibarial, medial sensilla 12–17 per side; posterior sensilla 6–7 per side (Fig. 2C). All tarsi entirely yellow (Fig. 2G); fore tarsomere I as long as 3 succeeding tarsomeres together; mid one slightly longer than rest tarsomeres together; hind one as long as rest together. Aedeagal sheath ventro-apically with a pair of strongly sclerotized postgonites much smaller than bifurcated pieces of aedeagal median rod (Fig. 3F). Measurements (holotype, in mm): BL (straight distance from anterior edge of pedicel to tip of abdomen) = 3.65, ThL (distance from anterior notal margin to apex of scutellum) = 1.56, WL (distance from humeral cross vein to wing apex) = 2.46, WW (maximum wing width) = 1.08. Indices (holotype, in ratio): FW/HW (frontal width/head width) = 0.44, ch/o (maximum width of gena/maximum diameter of eye) = 0.08, prorb (proclinate orbital seta length/posterior reclinate orbital seta length) = 1.78, rcorb (anterior reclinate orbital seta length /posterior reclinate orbital seta length) = 0.91, vb (subvibrissal seta length /vibrissa length) = 0.30, dcl (anterior dorsocentral seta length/posterior dorsocentral seta length) = (n/a), presctl (prescutellar seta length /posterior dorsocentral seta length) = 0.52, sctl (basal scutellar seta length /apical scutellar seta length) = (n/a), sterno (anterior katepisternal seta length/posterior katepisternal seta length) = (n/a), orbito (distance between proclinate and posterior reclinate orbital setae/distance between inner vertical and posterior reclinate orbital setae) = 2.08, dcp (distance between ipsilateral dorsocentral setae/distance between anterior dorsocentral setae) = 0.24, sctlp (distance between ipsilateral scutellar setae/distance between apical scutellar setae) = 1.20, C (2nd costal section between subcostal break and R 2+3 /3rd costal section between R 2+3 and R 4+5) = 2.35, 4c (3rd costal section between R 2+3 and R 4+5 /M 1 between r-m and dm-cu) = 1.50, 4v (M 1 between dm-cu and wing margin/M 1 between r-m and dm-cu) = 3.03, 5x (CuA 1 between dm-cu and wing margin/dm-cu between M 1 and CuA 1) = 1.00, ac (3rd costal section between R 2+3 and R 4+5 /distance between distal ends of R 4+5 and M 1) = 3.88, M (CuA 1 between dm-cu and wing margin/M 1 between r-m and dm-cu) = 0.66, C3F [length of heavy setation in 3rd costal section/(length of heavy setation in 3rd costal section+length of light setation in 3rd costal section)] = 0.71. Holotype. ♂ (SEHU), “ Ulu Gombak, Selangor, Malaysia, 8.xii.2013, M.J. Toda ”. Distribution. Peninsular Malaysia. Remarks. This species resembles P. afoliolata in having the small postgonites (Fig. 3 F–H), but can be distinguished from it by the diagnostic characters. Etymology. Referring to the lack of free sclerite within aedeagal outer membrane.

33) Phortica (Ashima) takehiroi Toda, sp. nov. (Fig. 9) Diagnosis. Arista with arb = 2/0 (ch.6-2) and apical, leaf-like expansion narrower than 1/2 length and apically roundish or slightly notched (Fig. 9B); anterior postgonite on left lateral lobe of aedeagal sheath apically trifurcated into one very large, sickle-like and two small, claw-like projections (Fig. 9I). Description (not repeating characters common to P. glabra). Supracervical setae 16–17. Dorsomedial, tentorial apodeme 1/2 as long as basal, parallel portion of dorsolateral, tentorial apodeme. Cibarial, medial sensilla approximately 6 per side; posterior sensilla 3–4 per side. All tarsi with gray tarsomere V; all tarsomere Is slightly longer than 3 succeeding tarsomeres together. Measurements (holotype / 1♂ paratype, in mm): BL = 3.34/3.43, ThL = 1.55/1.37, WL = 2.46/2.24, WW = 1.08/1.03. Indices (holotype / 1♂ paratype, in ratio): FW/HW = 0.43/0.48, ch/o = 0.11/0.09, prorb = 1.23/1.45, rcorb = 0.53/0.59, vb = 0.55/0.40, dcl = 0.49/0.50, presctl = 0.68/0.56, sctl = 1.01/1.03, sterno = 0.92/0.98, orbito = 1.91/1.95, dcp = 0.25/0.23, sctlp = 1.33/1.36, C = 2.48/2.32, 4c = 1.55/1.47, 4v = 3.37/(n/a), 5x = 1.17/1.08, ac = 3.40/(n/a), M = 0.76/0.68, C3F = 0.68/0.64. Holotype. ♂ (SEHU), “ Matsuyama, Ehime Pref., Japan, 33°51’19” N 132°46’21” E, 100 m a.s.l., 23.x.2010, T. Katoh”. Paratype. Japan: 1♂, Kakei, Aki-ota, Hiroshima Pref., 18.vii.2010, T. Katoh (SEHU). Distribution. Japan (Chugoku, Shikoku). Remarks. This species resembles P. glabra in having the apically blunt posterior postgonite on the right side of aedeagal sheath and the long, broad, sinuated lateral lobe on the left side, but can be distinguished from it by the diagnostic characters. Etymology. Patronym, dedicated to Dr. Takehiro Katoh, who collected the specimens.

2.1. Phortica (Ashima) foliiseta species subgroup, new Diagnosis. Surstylus without peg-like teeth (ch.37-1; Figs 4B, 5C); pregonite basally with recurved, apically truncate extension (ch.49-3; Figs 4C, 5D); posterior, rod-like process of phallapodeme longer than aedeagal median rod (ch.62-1; Figs 4C, 5D). Shared characters. Prementum without pubescent patches (ch.12-0). Setulae on medial surface of abdominal sternites IV and V not so dense (ch.27,28-0; Fig. 5B). Setae on and/or near ventral margin of epandrium longer than others (ch.33-1; Figs 4B, 5C). Aedeagal, outer membrane without spiny ornamentation (ch.55-1; Fig. 5D,E); aedeagal, median rod less sclerotized (ch.58-2; Fig. 5D). Aedeagal sheath with lateral lobes bearing strongly sclerotized postgonites (ch.65-1; Figs 4D, 5E). Included taxa. Phortica (Ashima) biclavata Chen; P. (A.) dianmianensis Gong, Zhu & Chen; P. (A.) foliisetoides Chen & Toda; P. (A.) libo Zhu, Qin & Chen*; P. (A.) menglian Gong, Zhu & Chen*; P. (A.) nakanoi Chen; P. (A.) phyllochaeta; P. (A.) sagittaristula Chen & Wen; P. (A.) saltiaristula Chen & Wen; P. (A.) symmetria *; P. (A.) xyleboriphaga *; P. (A.) yalianae Zhu, Qin & Chen*; P. (A.) yiqini Zhu, Qin & Chen*; and P. (A.) foliiseta species complex. * The assignment was judged from the original descriptions by Senior-White (1921), Chen et al. (2005b), Zhu et al. (2017) and Gong et al. (2018)., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on page 15, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Senior-White, R. A. (1921) New Ceylon Diptera. Spolia Zeylanica, 11, 381 - 396. https: // doi. org / 10.5962 / bhl. part. 12654","Chen, H. W., Toda, M. J. & Gao, J. J. (2005 b) The Phortica (s. str.) foliiseta species-complex (Diptera, Drosophilidae) from China and its adjacent countries. Acta Zootaxonomica Sinica, 30, 419 - 429.","Zhu, L., Qin, T. & Chen, H. W. (2017) Three new species of the subgenus Ashima (Diptera, Drosophilidae, Phortica) from southwestern China, with DNA barcoding information. Journal of Natural History, 51, 71 - 82. https: // doi. org / 10.1080 / 00222933.2016.1247480","Gong, L., Zhu, L. & Chen, H. W. (2018) Six new species of the subgenera Alloparadisa and Ashima (Diptera: Drosophilidae: Phortica) from Yunnan, China, with DNA barcoding information. Annales de la Societe entomologique de France, New Series, 54, 434 - 446. https: // doi. org / 10.1080 / 00379271.2018.1491327"]}

35) Phortica (Ashima) machoruka Prigent & Chen, 2008 (Fig. 3B,C) Phortica (Phortica) machoruka Prigent & Chen, 2008: 25. Phortica (Ashima) machoruka: Chen & Máca, 2012: 507. Diagnosis. Aedeagal, median rod deeply bifurcated from base: ventral process strongly sclerotized, apically curved ventrad, but dorsal one less sclerotized, apically curved dorsad (“Fig. 21” in Chen & Máca 2012). Supplementary description (not repeating characters common to P. angulata). Supracervical setae approximately 14. Arista with arb = 3–4/0; apical, leaf-like expansion narrower than 1/2 length, apically with long, awn-like projection. Cibarial, medial sensilla approximately 11 per side; posterior sensilla 5–6 per side. The antisymmetry is observed in the postgonites: in A-type, the left postgonite bears apically 1 long and 1 small, strongly sclerotized projections and 1 long, less sclerotized process, and subbasally 1 thick, apically blunt, strongly sclerotized projection, but the right one 2 long, strongly sclerotized projections and 1 long, less sclerotized process in the apical portion; but in B-type, vice versa (“Figs 21, 22” in Chen & Máca 2012). Specimens examined. Kenya: holotype ♂ (A-type), Kakamega Equatorial Forest, 11.x.2002, S. Prigent leg. (MNHN); 1♂ paratype, ditto, except 11–23.x.2002 (MNHN). Distribution. Kenya.

3.2. Phortica (Ashima) glabra species subgroup, new Diagnosis. Palpus concave on subapicolateral surface (ch.9-1; Fig. 9C); abdominal sternite III widest among sternites (ch.23-1; Fig. 9E); sternite V with dense setulae on medial surface (ch.28-1; Fig. 9E); hypandrium asymmetric (ch.40-1; Fig. 9I); pregonite (Fig. 9G) with neither basal extension (ch.49-0) nor sensilla on basal portion (ch.53-0); aedeagus without spiny ornamentation on outer membrane (ch.55-1; Fig. 9G); aedeagal sheath with only 1 anterior or 1 posterior, strongly sclerotized postgonite on each side (Fig. 9H,I). Shared characters. Dorsomedial, tentorial apodeme Included species. Phortica (Ashima) glabra; P. (A.) idiasta; P. (A.) zhangyuanae; and P. (A.) takehiroi sp. nov.

3) Phortica (Ashima) eseta Chen, 2012 Phortica (Ashima) eseta Chen in Chen & Máca, 2012: 495. Diagnosis. Palpus concave on subapicolateral surface (ch.9-1); cibarial, medial sensilla 65–69 in 3–4 irregular rows per side; postgonites forming large, somewhat triangular plate (in ventral view); two pieces of aedeagal median rod not bifurcated; aedeagal free sclerite C-shaped, larger than median rod. Supplementary description. Supracervical setae approximately 20–21. Dorsomedial, tentorial apodeme 4/5 as long as basal, parallel portion of dorsolateral, tentorial apodeme. Arista with arb = 6/4–5. Cibarial, posterior sensilla approximately 5 per side. All tarsi entirely yellow. Specimens examined. Malaysia: holotype ♂, Poring, Sabah, 30.xii.1998, M.J. Toda leg. (SEHU); 2♂ paratypes, 1♂, Ulu Senagang, Crocker Range, Sabah, 18.x.1999, M.J. Toda leg. (SEHU); 2♂, Tawau, Sabah, 17.iii.2002, M.J. Toda leg. (SEHU); 3♂, Deramakot, Sabah, 22–31.ii.2004, K. Akutsu leg. (SEHU); 1♂ paratype, Kubah, Sarawak, 21.i.1999, M.J. Toda leg. (FRCK). Distribution. Borneo (Sarawak, Sabah). Remarks. This species is peculiar in the diagnostic characters among the species of the afoliolata group.

Subgenus Ashima Chen, 2011 Ashima Chen in Cao et al., 2011: 682 (as subgenus of Phortica); Gong et al., 2018: 438. Type species: Phortica xyleboriphaga Senior-White, 1921. Diagnosis. Projections at anterolateral corners of the cibarium longer than 1/2 width of cibarial, anterior margin (ch.10-1; Fig. 2C); mid-dorsally constricted epandrium (ch.29-1; Figs 3B,D, 4B, 6D, 7E, 8G, 9F); less sclerotized, broad lobe-like surstylus (ch.34-1; Figs 3E, 4B, 5C, 6D, 7E, 9F); hypandrium with a pair of anterolateral, inward extensions articulated to base of pregonite (ch.42-1; Figs 3C,F, 4D, 5E, 6G, 7F, 8J, 9I). Shared characters. Characters described as commonly seen across this subgenus (the Phortica foliiseta species complex at that time) by Chen et al. (2005b) are not repeated here. Occipital setae on lower portion shorter than postocular setae (ch.2-0; Figs 7A, 9A). Pedicel with only 1 thick seta (ch.4-0; Figs 2A, 4A, 5A, 6A, 7C, 8A, 9B). Clypeus medially pale (ch.8-1; Fig. 8F). Lateral membrane of proboscis scale-like patterned (ch.13-1; Fig. 2D). Prominent postpronotal seta 1. Epandrium not extended at anteroventral corner (ch.30-1; Figs 3D, 4B, 6D, 7E, 8G, 9F). Surstylus completely or largely covered with epandrium (ch.36-1; Figs 4B, 7E, 8G, 9F), with pubescence (ch.38-0; Figs 3E, 4B, 5C, 6D, 7E). Cercus longer than twice of width (ch.39-0; Figs 3B,D, 4B, 5C, 6D, 7E, 8G, 9F). Pregonite articulated to tip of aedeagal guide (ch.54-1; Figs 3G, 4I, 5G, 6E, 7G, 8D, 9C), without sensilla on apical portion (ch.51-0; Figs 3G,H, 4C, 5D, 6E, 7G, 8I, 9G). Phallapodeme vertically flat plate, with posterior, rod-like process apically articulated to aedeagal median rod (ch.60,61-1; Figs 3G,H, 4C, 5D, 6E, 7G, 8I). Aedeagal sheath with more or less sclerotized postgonite(s) (ch.64-1; Figs 3 F���H, 4D, 5D,E, 6F,H, 7F, 8I���K, 9G���I). Key. The digital, multiple-entry key ��� Ashima (Diptera: Drosophilidae: Phortica)��� to all species of this subgenus has been constructed largely based on the character matrix of Appendix 1, and its database package including software instructions is available from the following website: https://drive.google.com/open?id=10ht68FhP1DJDg h2T33kZn7I2ro8YGGX0., Published as part of Toda, Masanori J., B��nziger, Hans, Sati, Pradeep C., Fartyal, Rajendra S., Suwito, Awit & Katoh, Toru, 2020, Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species, pp. 1-54 in Zootaxa 4789 (1) on pages 9-10, DOI: 10.11646/zootaxa.4789.1.1, http://zenodo.org/record/3884610, {"references":["Cao, H. L., Wang, X. L., Gao, J. J., Prigent, S. R., Watabe, H., Zhang, Y. P. & Chen, H. W. (2011) Phylogeny of the African and Asian Phortica (Drosophilidae) deduced from nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 61, 677 - 685. https: // doi. org / 10.1016 / j. ympev. 2011.08.002","Gong, L., Zhu, L. & Chen, H. W. (2018) Six new species of the subgenera Alloparadisa and Ashima (Diptera: Drosophilidae: Phortica) from Yunnan, China, with DNA barcoding information. Annales de la Societe entomologique de France, New Series, 54, 434 - 446. https: // doi. org / 10.1080 / 00379271.2018.1491327","Senior-White, R. A. (1921) New Ceylon Diptera. Spolia Zeylanica, 11, 381 - 396. https: // doi. org / 10.5962 / bhl. part. 12654","Chen, H. W., Toda, M. J. & Gao, J. J. (2005 b) The Phortica (s. str.) foliiseta species-complex (Diptera, Drosophilidae) from China and its adjacent countries. Acta Zootaxonomica Sinica, 30, 419 - 429."]}

2.2. Phortica (Ashima) tanabai species subgroup, new Diagnosis. Prementum with a pair of patches of pubescence (ch.12-1; Fig. 6B); aedeagal, outer membrane ornamented with somewhat long, conical spines (ch.55-0, ch.56-2; Figs 6E,G, 7G,H). Shared characters. Some interfrontal setulae as long as lowest setulae on ocellar triangle (ch.3-0; Fig. 7B). Arista apically with leaf-like expansion (ch.7-1; Figs 6A, 7C). Setulae just before prescutellar acrostichal setae somewhat longer than other acrostichal setulae (ch.15-1). Fore and mid tibiae entirely yellow (ch.17,18-0). Abdominal sternites III–V wider than long (ch.22,24,25-1; Figs 6C, 7D); III widest among sternites (ch.23-1; Figs 6C, 7D), with dense setulae on medial surface (ch.26-1; Figs 6C, 7D). Pregonite not concave on dorsosubapical margin of apical knob (ch.48-0; Figs 6E, 7G), without basal extension (ch.49-0; Figs 6E, 7G). Posterior, rod-like process of phallapodeme not longer than aedeagal median rod (ch.62-0; Fig. 6E). Aedeagal sheath without lateral lobes bearing strongly sclerotized postgonites (ch.65-0; Figs 6F, 7F). Included species. Phortica (Ashima) huiluoi Cheng & Chen *; P. (A.) mediarista; P. (A.) tanabei; P. (A.) akutsui sp. nov.; and P. (A.) halimunensis sp. nov. * The assignment was judged from the original description by Cheng et al. (2008).

Lepidotrigona palavanica (Cockerell, 1915) Type material examined: NHMUK 013379686; worker holotype; PHLIPPINES; labelled as follows: “Type”, “ Trigona / palavanica/ CKll. TYPE.”, “B.M. TYPE HYM. 17B.1120”, “3839”, “P. Princess/ Palawan/ Baker”, “Brit: Mus 1933-567.”., Published as part of Attasopa, Korrawat, Packer, Laurence, Suttiprapan, Piyawan, Thanoosing, Chawatat & Bänziger, Hans, 2020, Description of the male of Lepidotrigona nitidiventris (Smith, 1857), redescription of the female holotype and additional morphological data on the workers (Hymenoptera: Apidae: Meliponini), pp. 119-128 in Revue suisse de Zoologie 127 (1) on page 126, DOI: 10.35929/RSZ.0012, http://zenodo.org/record/5743375, {"references":["Cockerell T. D. A. 1915. Descriptions and records of bees. -- LXVIII. Annals and Magazine of Natural History 16: 1 - 9."]}

Lepidotrigona latipes (Friese, 1900) Type material examined: ZMB; worker holotype; MALAYSIA [Malacca, not Singapore, see discussion]; labelled as follows: “India/ Singapore/ 1890”, “ Trigona / latipes/F. /1909 Friese det.”, “Type”, “Coll Friese”, “HOLOTYPE/ Trigona / latipes Friese / Examined C Rasmussen ’07”., Published as part of Attasopa, Korrawat, Packer, Laurence, Suttiprapan, Piyawan, Thanoosing, Chawatat & Bänziger, Hans, 2020, Description of the male of Lepidotrigona nitidiventris (Smith, 1857), redescription of the female holotype and additional morphological data on the workers (Hymenoptera: Apidae: Meliponini), pp. 119-128 in Revue suisse de Zoologie 127 (1) on page 126, DOI: 10.35929/RSZ.0012, http://zenodo.org/record/5743375, {"references":["Friese H. 1900. Neue Arten der Bienengattungen Melipona Ill., und Trigona Jur. Termeszetrajzi Fuzetek 23: 381 - 394."]}

Lepidotrigona nitidiventris (Smith, 1857) Figs 1-17 Holotype Worker (female): Figs 1-3 Type material examined: OUMNH; worker holotype; MALAYSIA; labelled as follows: “Holo-/ type”, “M. OPHIR/ 79”, “ Trigona nitidiventris./ Smith” “ Tr. nitidiventris / Smith J. Pr. L. / Soc. T. 2 p 50/ Malacca M. Ophir” (see inset to Fig. 3). Diagnosis: Lepidotrigona nitidiventris is the type species of the genus and also the nominate species of the species group which comprises the largest species of the genus. Workers differ from those of L. ventralis and the L. terminata groups by a combination of body size (more than 5.5 mm) and apically expanded metatibia (spoon-like: as wide as approximately half its length, more than 0.46 times). In the other two groups the body length is usually less than 5.5 mm, and the metatibial ratio less than 0.45 times. Leipidotrigona nitidiventris also differs in the faintly yellow-tinged wings which are pale brownish grey in the other two groups. Lepidotrigona nitidiventris has robust hairs on the anterior and posterior margins of the mesoscutum and mesoscutellum, whereas the hairs are not robust on those areas in the L. ventralis group. Lepidotrigona nitidiventris can be differentiated from L. latipes by their brown tegula and black T2-T6, whereas they are yellowish brown and reddish brown, respectively, in L. latipes. The pale yellowish-brown vertex hairs in L. nitidiventris separate it from L. trochanterica, where those hairs are all black. L. palavanica differs from L. nitidiventris in being much smaller in body size and metatibial ratio. Re-description of the species: Structure: Body length 6.62 mm. Head wider than long (width 2.57 mm, length 1.95 mm); eye width (in side view) 0.67 mm; gena width 0.32 mm; interocellar distance 0.44 mm; ocellocular distance 0.38 mm; inner orbits slightly converging below, upper interorbital distance 1.51 mm, lower interorbital distance 1.46 mm; scape length excluding basal bulb 0.99 mm, with basal bulb 1.09 mm; 1st flagellomere length 0.16 mm; 2nd flagellomere length 0.24 mm, width 0.17 mm; 3rd flagellomere shorter than 2nd; malar space length 0.13 mm; length of longest hair on vertex 0.2 mm. Length of longest hair on mesoscutum 0.16 mm; length of longest hair on mesoscutellum 0.11 mm; forewing length 5.84 mm, width 2.32 mm; forewing diagonal 1.89 mm, length of 1st submarginal cell 0.41 mm, length of 2nd submarginal cell 0.87 mm; 1st recurrent vein before mid-length of posterior margin of 2nd submarginal cell. Metatibia length 2.51 mm, width 1.27 mm; metabasitarsus length 0.95 mm, width 0.66 mm. Coloration of integument: Head black except as follows: apex of mandible dark brown; basal bulb and extreme base of scape, and base of F1 yellow; rest of flagellum dark brown. Mesosoma black except tegula, pronotal lobe, and sides of metanotum all brown; all legs black except all trochanters, procoxa, metacoxa ventrally, profemur ventrally, and posterior portion of metabasitarsus brown, tarsomeres 3 to 5 yellowish brown; wings hyaline to pale yellow, wing veins yellowish brown except C, R, M, and Rs brown. Metasomal terga black except basal depression and posterior margin of T1 whitish brown; metasomal sterna yellowish brown. Pubescence: Facial hairs mostly plumose, densely appressed and whitish yellow; hairs of lower edge of labrum and mandible pale brown, hairs of vertex robust and pale yellowish-brown, longest hairs on vertex approximately 1/5 times as long as scape. Mesoscutum and mesoscutellum hairs minute yellowish white; hairs of anterior and posterior areas of mesoscutum and mesoscutellum robust, yellowish brown; mesoscutellum with some dark brown hairs intermixed, longest hairs on mesoscutum and mesoscutellum approximately 1/6th and 1/9th as long as scape, respectively; margins of mesoscutum and mesoscutellum with distinct band of short, plumose, scale-like yellow hairs, bands on mesoscutum wider anteriorly and posteriorly; mesepisternum, metepisternum, and pronotal lobe with whitish-yellow, plumose hairs, intermixed with long white hairs on mesepisternum ventrally; hairs on lateral surface of metanotum and propodeum plumose, yellowish white; metapostnotum bare; coxae and trochanters as well as ventral surface of profemur with yellowish-white hairs, rest of profemur with brown hairs intermixed with black hairs distally; tibiae with black hairs except metatibia with whitish-brown keirotrichia posteriorly; protarsi with brown hairs except probasitarsus with black hairs intermixed; meso- and metatarsi with black hairs intermixed with few brown hairs. T1 with minute, sparse, pale whitish-brown hairs except basal depression and posterior margin lacking hairs. Posterior margin of T2- T6 as well as visible portions of T5 and T6 with minute pale brown hairs intermixed with dark brown hairs on posterior half of T5 and T6; S1-S6 minute with sparse yellowish-brown hairs. Surface sculpture: Head and mesosoma with fine, dense punctures, slightly larger on mesoscutellum; metapostnotum with larger crowded punctures all over, diameters 1.5-2 times those of head; all legs with sparse, shallow and fine punctures; lateral part of T1, posterolateral parts of T1-T4 and complete T5 and T6 with sparse shallow, fine punctures; posterior margin of T2-T4 and complete T5 and T6 with minute, shallow punctures; S1-S6 with fine shallow punctures all over. Worker (female): Figs 4-6 Material examined: BCMU-LNW01; nest UP45; 2 workers; THAILAND, Tak Province., Umphang District; 5.V.2011; leg. prep. H. Bänziger. – All remaining specimens with the same labels except as follows: BCMU-LNW 02; nest UP45; 2 workers; 19.VI.2012. – BCMU-LNW03; nest UP45; 2 workers; 21.III.2016. – BCMU-LNW04; nest UP88; 3 workers; 7.V.2016. – MHNG-LNW05; nest UP88; 1 worker; 7.V.2016. Variation: N=10, measured as range (average±SD). As in holotype except as follows: Structure: Body length 5.69-6.56 (6.17±0.22) mm. Head width 2.58-2.87 (2.77±0.08) mm; head length 1.84- 2.03 (1.96±0.05) mm; eye width (in side view) 0.63- 0.68 (0.66±0.01) mm; gena width 0.3-0.42 (0.37±0.04) mm; interocellar distance 0.42-0.48 (0.45±0.02) mm; ocellocular distance 0.38-0.42 (0.4±0.01) mm; upper interorbital distance 1.51-1.61 (1.56±0.03) mm, lower interorbital distance 1.46-1.56 (1.52±0.03) mm; scape length excluding basal bulb 0.99-1.05 (1.03±0.02) mm, with basal bulb 1.09-1.2 (1.17±0.03) mm; 1st flagellomere length 0.16-0.18 (0.17±0.01) mm; 2nd flagellomere length 0.2-0.23 (0.21±0.01) mm, width 0.16-0.18 (0.17±0.004) mm; malar space length 0.14- 0.15 (0.14±0.005) mm; length of longest hairs on vertex 0.2-0.25 (0.21±0.02) mm. Length of longest hairs on mesoscutum 0.17-0.22 (0.19±0.01) mm; length of longest hairs on mesoscutellum 0.13-0.18 (0.15±0.02) mm; forewing length 5.73-6.27 (6.05±0.18) mm, width 2.35-2.52 (2.21±0.74) mm; forewing diagonal 1.94- 2.03 (1.98±0.03) mm, length of 1st submarginal cell 0.44-0.48 (0.46±0.01) mm, length of 2nd submarginal cell 0.84-0.95 (0.89±0.03) mm. Metatibia length 2.51- 2.67 (2.6±0.05) mm, width 1.23-1.32 (1.27±0.02) mm; metabasitarsus length 0.95-1.07 (1±0.04) mm, width 0.67-0.75 (0.7±0.02) mm. Coloration of integument: Tegula, wing veins M, Rs, and stigma slightly paler yellow than those of the holotype; mesoscutellum dark brown, sometimes with a brown spot posteromedially; tarsomeres 3 to 5 sometimes dark brown. T1 laterally with two large black triangular spots which occasionally are in contact with each other, rest of T1 sometimes light brown; metasomal sterna vary in darkness of brown. Pubescence: Hairs of vertex slightly darker than those of the holotype, sometimes intermixed with dark brown hairs. Metasomal sternal hairs sometimes paler than those of the holotype. Surface sculpture: Punctures on metasomal terga vary in density, otherwise as in holotype. Male: Figs 7-17 Material examined: BCMU-LNM01; nest UP88; 7 males; THAILAND, Tak Prov., Umphang Distr; 7.V.2016; leg. prep. H. Bänziger; caught leaving the nest [not swarming]. – MHNG-LNM02; nest UP88; 1 male; Tak Prov., Umphang Distr; 7.V.2016; leg. prep. H. Bänziger; caught leaving the nest [not swarming]. Male terminalia and associated sterna: Dissected from five males (BCMU-LNM 01) with the following codes: GP 3200, GP 3201, GP 3205, GP 3365, GPKA 88. Diagnosis: Males differ from the males of L. terminata and L. ventralis group by their overall larger size, shape of the metatibia and of S5 and S6 as follows: male of L. nitidiventris with expanded, triangular metatibia with apical margin straight, with a black comb near apicodorsal area (missing in other species where males are known); metatibial hairs dark brown to black (Figs 7, 8). Metatibia of other species simple, rounded apically, without comb, with hairs grey to brown or intermixed with few dark brown hairs. S5 of L. nitidiventris has long protruding apical-lateral lobes, but no subapical lobes (Fig. 11), whereas S5 of other known males of the genus have short to very short lateral lobes, two distinct subapical lobes, and strong apical setae. S6 of L. nitidiventris with two deep subapical emarginations with long, narrow apicalmedial lobes (Fig. 12), S6 of other known males of the genus do not have or have only weakly produced subapical emarginations, and short apical-medial lobes. Genital capsule of L. nitidiventris bilaterally symmetrical (Fig. 16), asymmetrical in the known species that belong to the L. ventralis group. Description: N=8, measured as range (average±SD). As in the worker holotype except as follows: Structure: Body length 6.19-6.98 (6.56±0.3) mm. Head width 2.4-2.62 (2.55±0.07) mm; head length 1.81- 2 (1.89±0.06) mm; eye width (in side view) 0.62-0.72 (0.68±0.04) mm; gena width 0.28-0.37 (0.31±0.03) mm; interocellar distance 0.4-0.55 (0.49±0.05) mm; ocellocular distance 0.3-0.33 (0.33±0.01) mm; inner orbits converging below, upper interorbital distance 1.49-1.56 (1.52±0.02) mm, lower interorbital distance 1.04-1.11 (1.07±0.02) mm; scape length excluding basal bulb 0.75-0.78 (0.78±0.01) mm, with basal bulb 0.88-0.95 (0.93±0.02) mm; 1st flagellomere length 0.14- 0.16 (0.15±0.01) mm; 2nd flagellomere length 0.28- 0.3 (0.3±0.01) mm, width 0.16-0.17 (0.16±0.004) mm; malar space length 0.02-0.03 (0.03±0.003) mm; length of longest hairs on vertex 0.33-0.43 (0.37±0.04) mm. Length of longest hairs on mesoscutum 0.2-0.24 (0.22±0.01) mm; length of longest hairs on mesoscutellum 0.16-0.18 (0.17±0.01) mm; forewing length 5.6-6.05 (5.86±0.14) mm, width 2.05-2.18 (2.12±0.04) mm; forewing diagonal 1.78-1.97 (1.86±0.06) mm, length of 1st submarginal cell 0.43-0.48 (0.45±0.02) mm, 2nd submarginal cell length 0.77-0.83 (0.8±0.02) mm. Metatibia length 2.48- 2.66 (2.57±0.07) mm, width 1.08-1.2 (1.16±0.03) mm; metabasitarsus length 0.7-0.77 (0.73±0.02) mm, width 0.48-0.53 (0.51±0.02) mm. Apex of S1 and S2 entire, S3 with small narrow emargination apicomedially; apical margin of metatibia flat diagonally (not rounded) forming a triangular-shaped metatibia, and with a black metatibial comb on the apex near the dorsal margin. Coloration of integument: Yellowish-brown area on F1 slightly darker than in holotype. Mesosoma black except tegula and pronotal lobe light brown, mesoscutellum seldom with yellowish-brown spot posteromedially; procoxa and proximoventral area of profemur brown, protrochanter and protarsus yellowish brown, rest of fore leg black; mid and hind legs black except mesotrochanter, and meso- and metatarsomeres 4-5 yellowish-brown, metatrochanter and ventral area of meso- and metafemurs brown, anterodistal and posterior area of metatibia and metabasitarsus sometimes brown, hyaline wings pale yellowish, slightly more brownish tinged than in workers, wing veins yellowish brown except C and R, dark brown. Metasomal terga black except basal depression of T1 brownish white or vary to light brown, T6 and T7 sometimes dark brown; S1-S7 light brown. Pubescence: Hairs of vertex long whitish brown, longest hairs approximately half of scape length; scape with thin white hairs medioventrally. Hairs of mesoscutum and mesoscutellum mostly whitish brown with brown hairs intermixed on mesoscutellum posteriorly, the longest hairs on mesoscutum and mesoscutellum approximately 3/10 and 2/9 as long as scape, respectively; posterolateral margin of mesoscutum with thinly scattered whitishyellow plumose hairs, which vary in density, sometimes to the point that the plumose hairs are missing altogether laterally; pronotal lobe, mesepisternum, metepisternum, lateral surface of propodeum, and metanotum with short, plumose, yellowish-white hairs intermixed with yellowish-white simple hairs, the simple hairs longest on mesepisternum ventrally; coxae and trochanters with long white hairs; femurs with white hairs ventrally and with short dark brown hair dorsodistally; protarsi with light brown hairs intermixed with few dark brown hairs; pro-, mesotibiae, and metatarsus with dark brown hairs intermixed with few whitish-brown hairs; metatibia with black or sometimes dark brown hairs, and with a black comb restricted to the apical angle, as well as with yellowish-white keirotrichia posteriorly. Metasomal tergal hairs as those of the holotype except lateral and posterior area of T4-T5, T6 and T7 posteriorly with black hairs; S1-S3 with short brown hairs. Surface sculpture: As for the holotype but punctures smaller and denser on mesoscutum. Terminalia and associated sterna: S4 emarginate apicomedially forming an apex with two broad shallow lobes, S4 gradulus almost touching anterior margin of sternum anteromedially; area posterior to gradulus with short hairs, the hairs denser on lateral lobes, and with a brown spot laterally (Fig. 10). S5 with long protruding apicolateral lobe posterolaterally oriented, and with deep emargination apicomedially, gradulus transverse medially, touching anterior margin of sternum; S5 with hairs on posterior area from gradulus, longer on posterior margin and apicolateral lobe; S5 brown marked laterally, darker on the apicolateral lobe (Fig. 11). S6 biconcave with long apicomedial lobe, narrowing distally with spatulate apex, apical emarginations with long hairs (Fig. 12). S7 convex apicomedially and bisinuate apicolaterally, with small asymmetrical subapicolateral lobes, the left lobe appearing to be slightly more protruding than the right one (Fig. 13). Gonostylus longer than penis valve, tip expanded widest at approximately apical 1/6, genital capsule and apical 1/3 of gonostylus light brown except penis valve and the remainder of gonostylus black. Penis valve narrow, first weakly curved ventrally mainly at mid-length, slightly curved lateroventrally towards apex (Figs 14-17)., Published as part of Attasopa, Korrawat, Packer, Laurence, Suttiprapan, Piyawan, Thanoosing, Chawatat & Bänziger, Hans, 2020, Description of the male of Lepidotrigona nitidiventris (Smith, 1857), redescription of the female holotype and additional morphological data on the workers (Hymenoptera: Apidae: Meliponini), pp. 119-128 in Revue suisse de Zoologie 127 (1) on pages 120-126, DOI: 10.35929/RSZ.0012, http://zenodo.org/record/5743375, {"references":["Smith F. 1857. Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca; and at Singapore, by A. R. Wallace. Journal of the Proceedings of the Linnean Society, Zoology 2: 42 - 88, pls 1 - 2."]}

Lepidotrigona trochanterica (Cockerell, 1920) Type material examined: NHMUK 013379685; worker holotype; MALAYSIA, labelled as follows: “Type”, “B.M. TYPE HYM. 17B.1102”, “ Trigona / trochanterica/ CKll. TYPE.”, “Sadakan/ Borneo/ Baket”, “Brit Mus./ 1933-567.”., Published as part of Attasopa, Korrawat, Packer, Laurence, Suttiprapan, Piyawan, Thanoosing, Chawatat & Bänziger, Hans, 2020, Description of the male of Lepidotrigona nitidiventris (Smith, 1857), redescription of the female holotype and additional morphological data on the workers (Hymenoptera: Apidae: Meliponini), pp. 119-128 in Revue suisse de Zoologie 127 (1) on page 126, DOI: 10.35929/RSZ.0012, http://zenodo.org/record/5743375

L. hoosana: Type material: Holotype (worker, AMNH); Type locality: TAIWAN; labelled as follows: “Formosa/ Taihorin. III.10 / H.Sauter S.G.”, “ T. ventralis hoosana / 1925 Friese det.”, “AMNH_IZC 00324480”, “acc 35178”, “Holotype”, “ Trigona / ventralis / var. hoosana / H.F. Schwarz”. [Examined]., Published as part of Attasopa, Korrawat, Bänziger, Hans, Disayathanoowat, Terd & Packer, Laurence, 2018, A new species of Lepidotrigona (Hymenoptera: Apidae) from Thailand with the description of males of L. flavibasis and L. doipaensis and comments on asymmetrical genitalia in bees, pp. 63-82 in Zootaxa 4442 (1) on page 79, DOI: 10.11646/zootaxa.4442.1.3, http://zenodo.org/record/1301390

L. arcifera: Type material: Holotype (worker, BMNH); Type locality: INDIA; labeled as follows: “ Type ”, “B.M. TYPE / HYM./ 17B.1081”, “ Trigona / arcifera Ckll / TYPE ”, “ Trigona / ventralis Sm / (worker) from Sladen.”, “Teesta/ Bridge/ Himalayas/ India / 10.1.97 ”, “T.D.A. Cockerell./ B.M.1936-415.”. [Images examined (Fig. 2 and Table 1 of Rasmussen, 2013)]., Published as part of Attasopa, Korrawat, Bänziger, Hans, Disayathanoowat, Terd & Packer, Laurence, 2018, A new species of Lepidotrigona (Hymenoptera: Apidae) from Thailand with the description of males of L. flavibasis and L. doipaensis and comments on asymmetrical genitalia in bees, pp. 63-82 in Zootaxa 4442 (1) on page 80, DOI: 10.11646/zootaxa.4442.1.3, http://zenodo.org/record/1301390, {"references":["Rasmussen, C. (2013) Stingless bees (Hymenoptera: Apidae: Meliponini) of the Indian subcontinent: Diversity, taxonomy and current status of knowledge. Zootaxa, 3647 (3), 401 - 428. https: // doi. org / 10.11646 / zootaxa. 3647.3.1"]}

Male of L. flavibasis Lepidotrigona flavibasis (Cockerell, 1929) , measured as range (Average±SD, n=4). Figs. 2: B1–4, 3: B1–4 & 8 Diagnosis. Lepidotrigona flavibasis differs from the other two species as follows: S4 apicolaterally bisinuate with medial convexity (Fig. 2: B1) (angularly emarginate in L. satun, weakly concave in L. doipaensis (Fig. 2: A1, C1)); S5 with apicosubmedial lobes pointed apically (as in L. doipaensis, rounded in L. satun), with one or two very short setae (generally only one very long seta and a much smaller seta in L. doipaensis but 4–7 setae of intermediate length in L. satun); and S5 gradulus mostly diagonal except short transverse section touching the antecosta medially (Fig. 2: B2) (gradulus parallel to and in touch with most of antecosta medially in L. doipaensis, but no contact in L. satun (Fig. 2: A2, C2)). L. flavibasis also differs in having only greyish yellow hairs on metatibia dorsally (Fig. 8: B) (dark brown intermixed with greyish yellow hairs in L. doipaensis, but variably brown, dark brown or greyish yellow hairs in L. satun). The ratio of the forewing diagonal to the length of the 2nd submarginal cell separates L. flavibasis from L. satun (see diagnosis of L. satun above and Table 1). The gonostyli and penis valves are also clearly different in the three species (see descriptions and Fig. 3). Description. As for the holotype of L. satun n. sp. except as follows: Structure: Body length 4.8–4.94 (4.91±0.07) mm. Head width 1.76–1.78 (1.78±0.01) mm; head length 1.34– 1.37 (1.35±0.01) mm; eye width 0.56–0.57 (0.56±0.005) mm; gena width 0.18–0.2 (0.19±0.01) mm; interocellar distance 0.34–0.35 (0.35±0.01) mm; ocellocular distance 0.2–0.22 (0.21±0.01) mm; upper interorbital distance 1.1–1.11 (1.1±0.01) mm, lower interorbital distance 0.75–0.78 (0.77±0.01) mm; scape length excluding basal bulb 0.51–0.52 (0.51±0.005) mm, with basal bulb 0.57–0.59 (0.58±0.01) mm; 1 st flagellomere length 0.08–0.13 (0.11±0.02) mm; 2nd flagellomere length 0.17–0.18 (0.18±0.005) mm, width 0.14–0.15 (0.14±0.004) mm; 3rd slightly shorter than 2nd; 4th longer than broad; malar space length 0.03–0.04 (0.03±0.005) mm. Forewing length 4.39–4.48 (4.42±0.04) mm, width 1.6–1.61 (1.6±0.01) mm; forewing diagonal 1.23–1.29 (1.26±0.03) mm, length of 1 st submarginal cell 0.3–0.33 (0.31±0.01) mm, 2nd submarginal cell length 0.58–0.61 (0.6±0.01) mm. Metatibia length 1.4–1.42 (1.41±0.01) mm, width 0.54–0.56 (0.55±0.01) mm; metabasitarsus length 0.58–0.59 (0.59±0.01) mm, width 0.37–0.39 (0.38±0.01) mm. S4 convex apicomedially, gradulus of S4 weakly concave medially, with small apicolateral angulation; S5 apical margin with two long, narrow processes bearing one or two very short thick setae, processes delimit an emargination ~0.54 times as deep as length of sternum and as long as wide, apicolateral lobe well developed, bearing long curved setae, gradulus touching antecosta medially; S6 with acute triangular apicomedial process; S7 approximate twice as wide as long, apical margin weakly emarginate. Genital capsule dark brown to black except base of penis valve, apical half to 2/3 of gonostylus and gonocoxa anteriorly pale brown to brown; the right penis valve overarches the left one and has two right angles directed to the right, so that the tip of the valve ends to the right of the midline of the bee, and generally posterior and often dorsal to the tip of the left penis valve. In the left penis valve the first angle is less acute and the second angle is missing but replaced by a curvature directed to the right, hence also bringing the tip of the left valve to the right of the midline, the tip is anterior and often ventral to the tip of the right penis valve. In addition, the final fourth or fifth of the valves is sinuous, the tips diverging from each other. Gonostylus sclerotized basally, approximately 12 times longer than wide (length 0.91–0.96 mm, width 0.073–0.080 mm) hairs equally dense throughout the apically expanded portion. Coloration of integument: Head black except as follows: yellowish brown on basal bulb and extreme apex of scape, labrum, and apical half of mandible; F2–F11 dark brown. Mesosoma black except pronotal lobe and tegula dark brown; fore and mid legs dark brown except medio- and distitarsi yellowish brown; hind leg black except tarsus yellowish brown (sometimes metabasitarsus dark brown). Metasomal terga black except basal depression of T1 transparent brown and apex of T6 and T7 brown. S1–S3 dark brown except posterior half of S1, posterior margin of S2, and apex of S3 whitish translucent; S4 brown except apical margin and most of disc medially translucent; S5 with apical and apicolateral lobes brown; S6 with anterolateral brown spot; S7 translucent except gradulus narrowly brown. Pubescence: Face with short plumose dense appressed white hairs; vertex with erect long pale whitish brown hairs. Mesoscutum with long and very short pale whitish brown hairs; mesoscutum, pronotal lobe and mesepisternum lacking distinct band of short plumose scale-like yellow hairs on margins; mesoscutellum with erect long pale whitish brown hairs longest on posterior margin; mesepisternum, metepisternum and metanotum with brownish white hairs; coxa and trochanter of all legs and ventral surface of all femora with long pale brownish white hairs, rest of femora with brownish white hairs intermixed with pale brown hairs; pro- and mesotibiae with dark brown hairs intermixed with brownish white hairs; metatibia with short greyish yellow hairs, coloration and density approximately the same on dorsal and ventral margins; all tarsi with whitish brown hairs intermixed with yellowish brown hairs, metatibia with short sparse brown keirotrichia. Posterolateral margin of T3–T7 with fine brownish white hairs progressively longer from T3–T7. Surface sculpture: All legs with fine dense punctures. T1 with sparse fine shallow punctures except impunctate on basal depression; metasomal sterna with coarse shallow crowded punctures intermixed with sparse coarse punctures. Material examined. Type material: Holotype (worker, AMNH); Type locality: THAILAND; labelled as follows: “Doi Sutep./Siam/ Feb. 9 / Alice Mackie”, “ Trigona / flavibasis / TYPE. CKll”, “acc 35740”. [Images examined (provided by AMNH, 2017)]. Additional material: Measured male specimens included in description and Table 1 (collected from wild nests): THAILAND, Chiang Mai Prov., Doi Suthep: 6.iv.2017, H. Bänziger (Nest SHQ2, 4 males, caught leaving nest [not swarming]). Measured worker specimens included in Table 1 (collected from wild nests): Chiang Mai Prov.: Doi Suthep, 26.xii.2015, K. Attasopa (Nest PHPL1, 2 workers); Doi Suthep-Pui, 1080 m, 6.iv.2017, H. Bänziger (Nest SHQ2, 2 workers). Non-measured specimens (collected from wild nests): Chiang Mai Prov.: Doi Inthanon, 17.i.2016, K. Attasopa (Nest MYAL1, 3 workers; MKLL3, 3 workers); Doi Suthep, 26.xii.2015, K. Attasopa (Nest DSTL1, 2 workers; nest PHPL1, 3 workers); Doi Suthep-Pui, 1080 m, 1.iv.2017, H. Bänziger (Nest SHQ2, 6 workers), 6.iv.2017, H. Bänziger (Nest SHQ2, 16 males, 9 workers, caught leaving nest [not swarming]). Nonmeasured specimens (collected by H.B. while they sucked his sweat): Chiang Mai Prov., Doi Inthanon, above Mae Ya Noi, 1700 m, 22.ii.1993 (4 workers). Genitalia examined: GP 3296, 3297, 3298, 3299, 3300, 3301, 3302, nest SHQ2 of type locality, Doi Suthep, 6.iv.2017., Published as part of Attasopa, Korrawat, Bänziger, Hans, Disayathanoowat, Terd & Packer, Laurence, 2018, A new species of Lepidotrigona (Hymenoptera: Apidae) from Thailand with the description of males of L. flavibasis and L. doipaensis and comments on asymmetrical genitalia in bees, pp. 63-82 in Zootaxa 4442 (1) on pages 77-79, DOI: 10.11646/zootaxa.4442.1.3, http://zenodo.org/record/1301390, {"references":["Cockerell, T. D. A. (1929) Descriptions and records of bees. - CXX. Annals and Magazine of Natural History, Series 10, 4, 584 - 594."]}

Lepidotrigona satun Attasopa and Bänziger n. sp. Figs. 2: A1–4, 3: A1–A4 & 4–6 Diagnosis. Lepidotrigona satun is a member of the “ ventralis ” species group based primarily on size: body and forewing length each less than 5 mm. It is the only species in the group known from the lower peninsula of Thailand. Males can be differentiated from those of the other two species of the “ ventralis ” group confirmed as occurring in Thailand (L. flavibasis and L. doipaensis) based upon external morphology as follows: margin of mesoscutum of L. satun with plumose, scale-like, yellow hairs (Fig. 4: A) (no such hairs in the other two species). S4 of L. satun is angularly emarginate apicomedially (Fig. 2: A1) (convex medially and bisinuate laterally in L. flavibasis, slightly concave in L. doipaensis). The apicosubmedial lobes of S 5 in L. satun are apically rounded each with 4–7 thick, long setae (Fig. 2: A2) (the lobes are pointed in the other two species and bear only 1–2 setae which are very short in L. flavibasis (Fig. 2: B2) or with one very long and, if present, a second much shorter in L. doipaensis (Fig. 2: C2)). S5 gradulus does not touch the antecosta in L. satun (Fig. 2: A2) whereas it does touch it in the other two species, briefly in L. flavibasis (Fig. 2: B2) and extensively in L. doipaensis (Fig. 2: C2). Males can also be differentiated from those of these other two species because of the very short malar space of L. satun, at most 0.16 times as long as the length of the first flagellomere whereas in the other two species it is at least 0.27 times as long as the first flagellomere. The ratio of the forewing diagonal to the length of the 2nd submarginal cell also varies among the three: it is largest (ḵ2.3) in L. satun, but at most 2.11 in the other two species (shortest - 1.94 to 2.05 - in L. doipaensis and intermediate - 2.05 to 2.11 - in L. flavibasis) (see Table 1). The genitalia are also very different in the three species (see below and Fig. 3). Workers of L. satun can be differentiated from all others in the L. ventralis species group, except L. flavibasis, due to the coloration of the pubescence. Lepidotrigona satun has dark hairs on the tibiae and pale yellowish brown to pale brown hairs on the vertex and mostly pale hairs (except posteriorly) on the mesoscutellum; no other member of the ventralis species group (except L. flavibasis) has this combination: L. hoozana and L. ventralis have some or entirely (respectively) whitish hairs on the metatibia, whereas the other four species have brown to black hairs; L. arcifera has pale hairs on the pro- and mesotibiae whereas L. satun, L. flavibasis and L. doipaensis have dark hairs; L. doipaensis has black hairs on the vertex and mesoscutellum, whereas L. flavibasis and L. satun do not have these hairs black but yellowish grey or variously pale to dark brown, respectively; the thickest of these hairs are about 0.007 mm, 0.003 mm and 0.005 mm wide, respectively. Workers of L. satun and L. flavibasis can be readily differentiated by the ratio of lengths of the metatibia and 2nd submarginal cell, at least 1.21 in L. satun, at most 0.98 in L. flavibasis and intermediate 1.05–1.09 in L. doipaensis. There are additional morphometric differences among the three species (Table 1). Description. Holotype Male. Structure: Body length 4.67 mm. Head wider than long; head width 1.77 mm; head length 1.31 mm; eye width (in side view) 0.51 mm; gena width 0.18 mm; interocellar distance 0.31 mm; ocellocular distance 0.17 mm; inner orbits strongly converging below, upper interorbital distance 0.98 mm, lower interorbital distance 0.68 mm; scape length excluding basal bulb 0.48 mm, with basal bulb 0.57 mm; 1 st flagellomere length 0.09 mm; 2nd flagellomere length 0.17 mm, width 0.13 mm; 3rd subequal to 2nd; 4th slightly longer than broad; malar space length 0.01 mm; mandible with subapical tooth. Forewing length (measured from humeral angle to apex) 4.03 mm, width 1.52 mm; forewing diagonal 1.1 mm, length of 1 st submarginal cell 0.35 mm, 2nd submarginal cell length 0.47 mm, vein Rs reaching stigma, 1 st recurrent vein at approximately mid-length of 2nd submarginal cell. Metatibia length 1.34 mm, width 0.52 mm; metabasitarsus length 0.51 mm, width 0.34 mm. Metasomal sterna and terminalia are described from paratypes in a separate section below. Coloration of integument: Head black except as follows: dorsal surface of scape, most of pedicel, proximal half of F1, and small marks towards base and apex of mandible dark brown; supraclypeal area and clypeus brown; labrum, most of mandible, ventral surface of scape, apical mark on pedicel and the rest of F1 yellowish brown. Mesosoma brown except pronotal lobe, mesoscutum, metepisternum and sides of metanotum black and metanotum medially yellowish brown; fore and mid legs pale brown except coxae, trochanters proximally and femora dorsally brown; hind leg brown except tarsus yellowish brown; wings hyaline to pale brown, wing veins and stigma pale brown except veins C, R, M, Rs, and R1 brownish black. Metasomal terga amber brown except T1 with semicircular brown band darker towards sides on disc, basal depression translucent, posterior marginal zone translucent amber, T4–T5 brown; S1 and S2 anteriorly whitish translucent, S3 with lateral brown spot. Pubescence: Most of face from lower margin of vertex to apex of clypeus with short plumose dense appressed yellowish white hairs; labrum, ventral scape, most portion of genal area, and lower edge of mandible with pale yellowish brown hairs longest at apex of labrum and lower mandible; ventral surface of scape with very short pale brown hairs; vertex with erect long pale yellowish brown hairs, longest hairs of vertex approximately one half as long as scape. Mesoscutum with long and very short pale yellowish brown hairs, the longest hairs approximately one quarter as long as scape; margins of mesoscutum with distinct band of short plumose scale-like yellow hairs interrupted anteromedially and posteriorly; mesoscutellum with pale yellowish brown hairs, progressively longer from anterior to posterior; mesepisternum, metepisternum and metanotum with pale brown hairs, somewhat darker anteroventrally on mesepisternum; posteroventral area of mesepisternum with long white hairs, pronotal lobe and mesepisternum anterodorsally with yellow plumose scale-like hairs; metapostnotum without hairs, lateral surface of propodeum with minute dense white hairs; coxa and trochanter of all legs with long pale brownish white hairs, ventral surface of all femora with pale brown hairs, remaining femora with pale brown hairs intermixed with dark brown hairs, protibia with pale brown hairs intermixed with few short dark brown hairs, the dorsal and anterior mesotibia with dark brown hairs, rest of mesotibia with pale yellowish brown hairs, metatibia dorsally with hairs variable, mostly brown, sometimes dark brown or greyish yellow, or intermixed; protarsus with yellowish brown hairs, meso- and metatarsi with yellowish brown hairs intermixed with few dark brown hairs, posterior surface of metatibia with short yellowish brown keirotrichia. Posterolateral margin of T1 and posterior and lateral parts of T2–T4 with minute sparse pale brownish white hairs; T3–T6 laterally with fine pale brown hairs progressively longer from T3–T6; T7 with pale brown hairs most dense on posterior margin, the longest hairs as long as the longest hairs of T6. Surface sculpture: Head and most of mesosoma with fine dense punctures; metapostnotum with coarse crowded sharp-edged punctures, diameters 2–2.5 times those of mesoscutum; all legs with fine sparse punctures; metasomal terga shining, T1 with few coarse punctures on posterolateral brown band, T2–T7 with few fine shallow punctures restricted to the posterior margin; metasomal sterna somewhat shining with sparse fine shallow punctures. Description. Paratype Males , measured as range (Average±SD, n=10). As in the holotype except as follows: Structure: Body length 4.33–4.85 (4.58±0.15) mm. Head width 1.72–1.77 (1.75±0.02) mm; head length 1.24– 1.34 (1.32±0.03) mm; inner orbits converging below, upper interorbital distance 0.94–1.01 (0.97±0.03) mm, lower interorbital distance 0.67–0.73 (0.69±0.02) mm; eye width 0.48–0.54 (0.52±0.02) mm; gena width 0.14–0.18 (0.17±0.02) mm; interocellar distance 0.28–0.31 (0.29±0.01) mm; ocellocular distance 0.1 8–0.22 (0.2±0.01) mm; scape length excluding basal bulb 0.47–0.52 (0.49±0.01) mm, with basal bulb 0.56–0.59 (0.58±0.01) mm); 1 st flagellomere length 0.09–0.11 (0.1±0.01) mm; 2nd flagellomere length 0.15–0.17 (0.16±0.01) mm, width 0.12– 0.14 (0.13±0.004); 3rd subequal to 2nd; 4th slightly longer than broad; malar space length 0.01–0.02 (0.01±0.002) mm. Forewing length 3.94–4.09 (4.01±0.05) mm, width 1.47–1.52 (1.49±0.02); forewing diagonal 1.09–1.18 (1.15±0.03) mm, length of 1 st submarginal cell 0.3–0.34 (0.32±0.01) mm, 2nd submarginal cell length 0.46–0.52 (0.49±0.01) mm. Metatibia length 1.34–1.43 (1.39±0.03) mm, width 0.47–0.53 (0.5±0.02) mm. Metabasitarsus length 0.47–0.55 (0.5±0.03) mm, width 0.29–0.33 (0.31±0.01) mm. Coloration of integument: Ventral scape usually yellowish brown rarely brown on distal half, yellowish brown area on pedicel and 1 st flagellomere varies in size, position, and sometimes brown; pale marking on disc of tegula varies in size, seldom absent; mesepisternum sometimes dark brown; propodeum sometimes dark brown or more or less black; all coxae sometimes brown, femora and tibiae vary in darkness of brown; semicircular brown band of T1 varies slightly in width, sometimes darker than other terga; T2–T7 sometimes darker than in the holotype. Pubescence: Long hairs on vertex, mesoscutum and mesoscutellum sometimes slightly paler and longer than the holotype; yellow plumose scale-like hairs on mesepisternum vary in density; minute hairs on visible portion of metasomal terga vary in length and density. Surface sculpture: Fine shallow punctures on T2–T7 vary in density. Description. Paratype Workers , measured as range (Average±SD, n=10). As in the holotype except for usual sexual dimorphism in flagellomeres, metasomal segment number, presence of corbicula, terminalia, and as follows: Structure: Body length 3.94–4.39 (4.16±0.15) mm. Head wider than long; head width 1.76–1.85 (1.81±0.03) mm; head length 1.38–1.45 (1.41±0.02) mm, inner orbits slightly diverging for upper half, converging below, upper interorbital distance 1.08–1.13 (1.11±0.02) mm, lower interorbital distance 0.97–1.03 (0.99±0.02) mm; eye width 0.41–0.48 (0.45±0.02) mm; gena width 0.2–0.27 (0.25±0.02) mm; interocellar distance 0.2 7–0.32 (0.29±0.02) mm; ocellocular 0.26–0.3 (0.28±0.01) mm; scape length excluding basal bulb 0.62–0.67 (0.65±0.01) mm, with basal bulb 0.72–0.75 (0.74±0.01) mm; 1 st flagellomere length 0.11–0.13 (0.12±0.01) mm; 2nd flagellomere length 0.1– 0.13 (0.11±0.01) mm, width 0.11–0.13 (0.13±0.01); 3rd approximate subequal to 2nd; 4th as long as broad; malar space length 0.08–0.11 (0.09±0.01) mm; mandible with two subapical teeth. Forewing length 3.97–4.09 (4.04±0.04) mm, width 1.51–1.54 (1.52±0.01) mm; forewing diagonal 1.15–1.19 (1.17±0.01) mm, length of 1 st submarginal cell 0.3–0.34 (0.32±0.01) mm, 2nd submarginal cell length 0.46–0.52 (0.49±0.01) mm, 1 st recurrent vein at approximately before middle-length of 2nd submarginal cell. Metatibia length 1.45–1.51(1.49±0.02) mm, width 0.6–0.63 (0.62±0.01) mm; metabasitarsus length 0.57–0.66 (0.61±0.03), width 0.38–0.41 (0.4±0.01). S1 emarginate posteromedially, posterior margin of S2–S5 with shallow broad emargination, S6 entire. Coloration of integument: Labrum, scape basal bulb, distal and basal marks on scape and 1 st flagellomere medially light brown; rest of antenna dark brown (scape and pedicel sometimes black). Mesosoma black except mesoscutellum and tegula dark brown with paler brown disc; all legs black to dark brown, probasitarsus dark to yellowish brown, apex of metatibia and metabasitarsus sometimes brown, tarsi of all legs light brown except slightly darker posteriorly. Semicircular brown band on disc of T1 usually wider laterally than in male; T2–T5 black, sometimes pregradular area of T2 brown; T6 usually black except apex pale, seldom wholly pale brown. Pubescence: Plumose appressed hairs on face absent from upper frons to vertex, occiput and upper genal area; scape ventrally with few very short pale brown hairs; vertex with erect pale brown hairs shorter and slightly darker than those of male, mesoscutum with sparse short pale brown hairs, medial interruption absent on distinct yellow band; mesoscutellum with long pale yellowish grey hairs variously intermixed with pale to dark brown (but not black) hairs, the thickest hairs about 0.005 mm wide (about 1/10 of the gonostylar width of the male), mesepisternum with fine brown hairs, with long white hairs ventrally, patches of yellow plumose scale-like hairs on mesepisternum anterodorsally and pronotal lobe usually larger than in males; metepisternum and lateral surface of propodeum with fine short pale yellowish white hairs (sometimes yellowish brown); all femora dorsally with dark brown hairs though paler posteriorly, with white (sometimes pale brown) hairs ventrally; pro- and mesotibiae with dark brown hairs intermixed with pale brown hairs posteriorly, metatibia with dark brown hairs dorsally, tips of hairs slightly paler, ventrally with paler hairs intermixed with a few dark hairs, posteriorly with whitish brown keirotrichia; all basitarsi with dark brown intermixed with yellowish brown hairs, remaining tarsomeres with yellowish brown hairs sometimes intermixed with few brown hairs; T1–T4 with minute hairs slightly darker than in male; T3–T6 towards sides with dark brown hairs; S1–S6 with minute pale brown hairs progressively longer from anterior to posterior on each sternum. Surface sculpture: Punctures on mesoscutellum, metanotum, and metepisternum slightly larger than on rest of mesosoma; all legs mostly with fine punctures intermixed with few coarse punctures, all femora with punctures similar to those of metapostnotum though slightly smaller; anterior surface of metatibia usually shining for ~apical 2/3, shining area varies in size. Male Metasomal Sterna and Genitalia. In the following, description of genitalic features is based upon dissections of paratype males from all four nests (see material examined). S4–S7 weakly sclerotized except S4 posterolaterally, S5 laterally and posteriorly brown (Fig. 2: A1–4); S4 angularly emarginate apicomedially, depth of emargination ~0.33 times length of sternum, emargination separating two broadly rounded lobes, more strongly sclerotized area of S4 with fine dense hairs, longest and densest lateral to apex of lobe; gradulus of S4 weakly convex close to anterior margin of S4 (Fig. 2: A1); S5 with four apical lobes, long submedian lobes separated by deep U-shaped emargination, depth of emargination ~0.63 times medial length of S5, short lateral lobe separated from medial lobe by shallow concavity, medial lobes bearing ~4–7 thick setae apically, gradulus of S5 transverse medially, near anterior margin (Fig. 2: A2); S6 apical margin weakly produced medially, weakly concave laterally (Fig. 2: A3); S7 1.56 times as wide as long, apical margin convex throughout (Fig. 2: A4). Genital capsule light brown except black for apical 3/5 of penis valve; penis valve curved ventrally, somewhat hook-like, curvature strongest on basal third to half, then almost straight, the right valve tends to overlap the left one somewhat and its apex is slightly posterior to that of the left valve. Gonostylus not sclerotized, narrow, at least 12 times as long as wide (0.75–0.78 mm long and 0.044–0.058 mm wide), slightly expanded for apical 1/3, medial surface setose on apical half, hairs denser towards apex, lateral surface with few short hairs in apical half (Fig. 3: A1–4). Etymology. The specific epithet refers to the province in Thailand where the species was collected; it is a noun in apposition. Material examined. Holotype. (Nest TB 2, male, to be deposited in QSBG) labelled as follows: “Thaleban Nat. Park/ Satun Prov., 100 m / 27.iv.2017 / ex nest TB2/ S. THAILAND / H. Bänziger leg prep det”, “ HOLOTYPE ”. Paratypes. All bearing labels that state “ PARATYPE ” and the same locality as those of the holotype except dates and nests as follows: Measured paratypes (included in description and Table 1) labelled as follows: 26.iv.2017 (Nest TB1, 1 worker); 27.iv.2017 (Nest TB1, 1 male); 27.iv.2017 (Nest TB2, 3 males, 3 workers); 27.iv.2017 (Nest TB3, 3 males, 3 workers); 27.iv.2017 (Nest TB4, 3 males, 2 workers); 29.iv.2017 (Nest TB1, 1 worker). Non-measured paratypes: 27.iv.2017 (Nest TB1, 2 males), 26.iv.2017 (Nest TB1, 2 workers), 29.iv.2017 (Nest TB1, 2 workers), 27.iv.2017 (Nest TB2, 17 males, 14 workers; 27.iv.2017 (Nest TB3, 7 males, 2 workers); 27.iv.2017 (Nest TB4, 11 males and 2 workers). Genitalia examined: GP 3304, 3305, 3306, 3308, 3309, 3310, 3311, same nest as holotype; GP 3307, 3312, 3313, 3314, 3315, 3316, 3317, paratypes of nest TB4; GP 3318, 3319, 3320, 3321, 3322, paratypes of nest TB3; GP 3323, paratype of nest TB1.

Male of L. doipaensis Lepidotrigona doipaensis (Schwarz, 1939) , measured as range (Average±SD, n=4). Figs. 2: C1–4, 3: C1–4 & 7 Diagnosis. Males of L. doipaensis can be differentiated from the other two Lepidotrigona species of the L. ventralis species group known from Thailand as follows (see also diagnosis of L. satun above). S4 slightly concave apicomedially (Fig. 2: C1) (strongly concave and convex in L. satun and L. flavibasis respectively). The apicosubmedial lobes of S5 pointed apically (as in L. flavibasis, rounded in L. satun), with one very long seta and often a much shorter one (two very short setae in L. flavibasis, 4–7 setae of intermediate length in L. satun). S4 gradulus touches the antecosta (Fig. 2: C1) (at least slightly separated from the antecosta in the other two species (Fig. 2: A1, B1)). S5 gradulus mostly transverse and touching antecosta medially (Fig. 2: C2) (not touching the gradulus in L. satun and doing so only briefly in L. flavibasis). Hairs on dorsal surface of metatibia dark brown intermixed with greyish yellow ones (Fig. 7: B) (only greyish yellow hairs in L. flavibasis, but variably brown, dark brown or greyish yellow in L. satun). The gonostylus and genital capsule are also clearly different in the three species (see descriptions and Fig. 3). The ratio of the forewing diagonal to the length of the 2nd submarginal cell separates all three species (Table 1) and see diagnosis of L. satun above. Description. As for the holotype of L. satun n. sp. except as follows: Structure: Body length 4.85–5(4.89±0.07) mm; head width 1.9–1.95 (1.93±0.02) mm; head length 1.42–1.48 (1.45±0.02) mm; eye width 0.55–0.58 (0.57±0.01) mm; gena width 0.18–0.19 (0.19±0.01) mm; interocellar distance 0.39–0.42 (0.4±0.01) mm; ocellocular distance 0.25–0.27 (0.26±0.01) mm; upper interorbital distance 1.08–1.11 (1.09±0.01) mm, lower interorbital distance 0.78–0.8 (0.79±0.005) mm; scape length excluding basal bulb 0.52–0.58 (0.55±0.02) mm, with basal bulb 0.61–0.67 (0.64±0.02) mm; 1 st flagellomere length 0.11–0.13 (0.12±0.01) mm; 2nd flagellomere length 0.2–0.22 (0.21±0.01) mm, width 0.13–0.14 (0.14±0.001) mm; 3rd shorter than 2nd; 4th longer than broad; malar space length 0.03–0.04 (0.04±0.001) mm. Forewing length 4.55– 4.76 (4.66±0.08) mm, width 1.7–1.76 (1.73±0.02) mm; forewing diagonal 1.28–1.34 (1.3±0.03) mm, length of 1 st submarginal cell 0.3–0.32 (0.31±0.01) mm, 2nd submarginal cell length 0.62–0.67 (0.65±0.02) mm. Metatibia length 1.61–1.73 (1.67±0.04) mm, width 0.62–0.66 (0.64±0.01) mm; metabasitarsus length 0.57–0.61 (0.59±0.02) mm, width 0.4–0.41 (0.41±0.005) mm. S4 broadly emarginate apicomedially, depth of emargination 0.15 times length of sternum medially; gradulus of S4 slightly concave medially; S5 apical margin with two long, narrow, acute lobes that delimit an emargination ~0.45 times as deep as length of sternum, emargination approximately as long as wide, submedial lobes each bearing a long, occasionally also a short, thick apical seta, S5 gradulus transverse touching antecosta medially; S6 apical margin convex medially with short acute process, S7 ~twice as wide as long, apical margin weakly emarginate, bearing some distinct hairs apicomedially. Genital capsule dark brown to black except base of penis valve, apical half to 2/3 of gonostylus, and gonocoxa anteriorly which are pale brown to brown. Penis valve curved ventrally, curvature strongest on basal third to half, then almost straight, valves normally in contact with each other (exceptionally not so as in Fig. 3: C1–4) and the right valve overarching the left valve. Gonostylus sclerotized basally, medial surface setose on apical ~3/7, approximately 10 times longer than wide (length 0.67–0.71 mm, width 0.062–0.073 mm). Coloration of integument: Head black except as follows: yellowish brown on basal bulb and extreme apex of scape (sometimes ventral surface of scape entirely yellowish brown); apical 2/3 of mandible; pedicel, and all flagellomeres dark brown. Mesosoma black except tegula; all coxae and femora black; trochanters brown; tibiae brown to black; all tarsi yellowish brown. Metasomal terga dark brown except basal depression and posterior margin of T1 yellowish brown; S1–S3 brown, except S3 laterally dark brown; S4 with large lateral dark brown spot; S5 dark brown apically, paler medially; S6 pale brown except paler on disc; S7 translucent with large lateral mark and gradulus brown. Pubescence: Face with plumose dense appressed white hairs; frons, vertex, lower genal area, and lower edge of mandible with long pale whitish brown hairs; longest hairs of vertex ~3/4 times as long as scape. Mesoscutum and mesoscutellum with long and very short pale whitish brown hairs, the longest hairs slightly shorter than those of vertex; mesoscutum lacking distinct band of short plumose scale-like yellow hairs; coxa, trochanter, and femora of all legs with pale brownish white hairs; all tibiae with pale brown to greyish yellow hairs, intermixed with short dark brown hairs; metatibia with short dense brownish white keirotrichia. Surface sculpture: As for male L. flavibasis. Material examined. Type material: Holotype (worker, NMNH); Type locality: THAILAND; labelled as follows: “Doi Pa/ Mai Deng/ Siam 750m / 12-29-32 ”, “HughMSmith/ Coll”, “TypeNo./ 53563/ U.S.N.M.”, “Holotype”, “ Trigona ventralis / var. doipaensis / H.F. Schwarz”, “7080102”. [Examined and measured in Table 1]. Paratypes (2 workers, AMNH), labelled as follows: “Doi Pa/ Mai Deng/ Siam 750m / 12-29-32 ”, “HughMSmith/ Coll”, “PARATYPE”, Trigona ventralis / doipaensis / H.F. Schwarz”, “acc 36579”. [Examined and measured in Table 1]. Additional material: Measured male specimens included in description and Table 1 (collected from wild nests): THAILAND, Chiang Mai Prov.: Doi Inthanon, 17.i.2016, K. Attasopa (Nest MYAL5, 2 males); Doi Suthep, Chang Khian stream area, 1020 m, 11.iv.2016, H. Bänziger (Nest CK 4, 2 males, caught leaving nest [not swarming]). Measured worker specimens included in Table 1 (collected from wild nests): Chiang Mai Prov.: Doi Inthanon, 17.i.2016, K. Attasopa (Nest MKLL1, 1 worker; nest MKLL4, 1 worker; nest MYAL3, 1 worker); Doi Suthep, Chang Khian, 1020 m, 29.ix.2010, H. Bänziger (Nest CK4, 1 worker); Doi Suthep, 26.xii.2015, K. Attasopa (Nest DSTL3, 1 worker). Non-measured specimens (collected from wild nests): Chiang Mai Prov.: Chaiprakan, 710 m, 20.ii.2010, H. Bänziger (Nest CP9, 13 workers), 11.iv.2011 (Nest CP26, 4 males, 6 workers, swarming), 16.iv.2016 (Nest CP26, 11 males, 2 workers, caught as leaving nest); Doi Suthep, Chang Khian stream area, 1020 m, 19.iii.2009, H. Bänziger (Nest CK4, 12 workers), 5.viii.2009 (Nest CK4, 4 workers), 19.viii.2009 (Nest CK4, 4 workers), 29.ix.2010 (Nest CK4, 5 workers), 11.iv.2016 (Nest CK4, 10 males, 2 workers, caught as leaving nest); Doi Suthep, Chang Khian stream area, 1040 m, 4.xii.2011, H. Bänziger (Nest CK11, 7 workers), 11.iv.2016 (Nest CK11, 11 males, 3 workers, caught as leaving nest); Doi Inthanon, 17.i.2016, K. Attasopa (Nest MKLL1, 2 workers; nest MKLL4, 2 workers; nest MYAL3, 2 workers); Doi Suthep, 26.xii.2015, K. Attasopa (Nest DSTL3, 2 workers); Doi Suthep, W of temple, 1070 m, 1.iv.2017, H. Bänziger (Nest SHQ1, 8 workers); Doi Suthep northern sector, QSBG area, 25.iii.2010, H. Bänziger (Nest, 13 workers); Hill opposite QSBG, 25.iii.2010, H. Bänziger (Nest in tree trunk, 1 worker); Pa Pä, Ban Huay Phra Chao, 29.vi.2008, H. Bänziger (Nest, 12 workers); North of Chiang Dao Dist., 6.xi.2008, H. Bänziger (Nest TL26, 8 workers), 8.ii.2009, H. Bänziger (Nest TL26, 6 workers). Genitalia examined: MYAL 5-01, MYAL 5-02, nest MYAL 5, Doi Inthanon, 17.i.2016; GP 3182, 3183, 3191, nest CK4, Chang Khian, Doi Suthep, 11.iv.2016; GP 3179, 3180, 3192, 3258, 3259, 3260, nest CK11, Chang Khian, Doi Suthep, 11.iv.2016. Comments. Based upon the colour of pubescence, the male of L. doipaensis might be misidentified as L. flavibasis because it has pale hairs on the vertex and mesoscutellum typical for the worker of the latter species. However, due to the association of the two sexes collected together in the field, the males described above are certainly L. doipaensis despite the fact that the hairs are dark in workers of this species.

L. ventralis: Type material: Holotype (worker, BMNH); Type locality: MALAYSIA: ���Borneo (Sarawak)���. [Not examined]. Additional material: MALAYSIA, Borneo, Sarawak, Gunung Buda, 20���28.xi.1996, A. D. Mudge (2 workers, PCYU); INDONESIA, W. Kalimantan, Gunung Palung Nat. Pk. 17���29.vi.1991, C. Darling, U. Rosichon, H. Sutrisno (3 workers, PCYU); W. Kalimantan, Gunung Palung Nat. Pk. 15.vi���15.vii.1991, C. Darling, U. Rosichon, H. Sutrisno (9 workers, ROM)., Published as part of Attasopa, Korrawat, B��nziger, Hans, Disayathanoowat, Terd & Packer, Laurence, 2018, A new species of Lepidotrigona (Hymenoptera: Apidae) from Thailand with the description of males of L. flavibasis and L. doipaensis and comments on asymmetrical genitalia in bees, pp. 63-82 in Zootaxa 4442 (1) on page 79, DOI: 10.11646/zootaxa.4442.1.3, http://zenodo.org/record/1301390