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Piraonapicha, Kanyakorn, Sangpradub, Narumon, Jaitrong, Weeyawat, Attasopa, Korrawat, Liu, Xingyue (2023): Frist record of Protohermes stigmosus Liu, Hayashi & Yang (Megaloptera Corydalidae: Corydalinae) in Thailand with the first description of female. Zootaxa 5256 (1): 87-93, DOI: 10.11646/zootaxa.5256.1.7, URL: http://dx.doi.org/10.11646/zootaxa.5256.1.7

Protohermes stigmosus Liu, Hayashi & Yang, 2007 (Figs 1–5) Protohermes stigmosus Liu, Hayashi & Yang, 2007: 38 figs 20, 123–127; Yang & Liu, 2010: 180, fig. 78; Martins et al, 2022: 29. Types. Holotype. Male, China (Xishuangbanna Yunnan Province), 31.III.2004, reared from larva, Liang Tang leg., deposited in the Shanghai Normal University (SNU, examined), China. Non-type material examined. Thailand. One male, Bo Kluea District, Nan Province, 19°11U13.35UN, 101°10U5.91UE, 1006 m a.s.l., 4.III.2020, K. Piraonapicha & L. Khaton leg. leg. (THNHM, THNHM-I-24146, pinned); 1 female, same locality, date, collectors (THNHM, THNHM-I-24147, in alcohol). Diagnosis. Head without dark markings; pronotum with two pairs of blackish markings at anterolateral corner and three pairs of blackish markings near posterolateral corners; fused gonocoxites 10 medially with a pair of large subtriangular processes, and laterally with a pair of digitiform gonostyli 10, leaving very short lateral arms; female fused gonocoxites 8 with feebly convex posterior margin, medially without any notch; no lateral sac-like lobes between abdominal segments 8 and 9. Description. Male (Figs 1–3). Measurements (n=1). Total body length 35.1 mm (including mouthparts); head width 6.2 mm, head length 4.8 mm (excluding labrum and mandibles); prothorax length 4.3 mm; prothorax width 4.4 mm; right forewing length 53.2 mm; right forewing width 16.7 mm; right hindwing length 47.6 mm; right hindwing width 18.3 mm; wing spans 101.5 mm. Head yellow with obtuse postocular spine; compound eyes dark, located anterior to mid-length of head; ocelli located between compound eyes, close to antennal socket, inner margin brown; antennae subserrate, scapes and pedicel yellowish brown, scapes almost twice as long as pedicel, flagellum dark; labrum yellowish brown, subtriangular, shorter than broad, anterior margin distinctly convex, and covered with sparse short setae; mandibles yellow to reddish brown (anterior half reddish brown and posterior half yellow), with long and sharp apical tooth, followed by broad and subtriangular preapical tooth, small subtriangular prebasal tooth, and subtriangular basal tooth (denticles reddish brown); occiput subtriangular, yellow, posterior margin roundly convex in dorsal view. Pronotum yellow, subquadrate, with two pairs of blackish markings at anterolateral corner and three pairs of blackish markings at posterolateral corner (Fig. 4); mesonotum and metanotum subrectangular, yellow, both clearly broader than long; legs yellow, covered with short dense setae; distal half of protibiae and all protarsi black, meso- and metatarsi with second to fifth tarsomeres black; tarsal claws reddish brown; forewing pale grayish brown, with a subtriangular yellowish brown marking near base, yellowish brown markings at basal 1/3, and with an irregular markings at apical 1/3 and several extremely small yellowish markings on crossveins on apical portion; costal cellules with distinct grayish brown stripes, hindwing entirely hyaline; veins yellow except veins in dark regions of forewing and apical portion of hindwing pale brown; RA 8-branched, with 9 crossveins between RA and RP, MP 1+2 4-branched, MP 3+4 2-branched. Abdomen yellowish brown. Genitalia. Tergum 9 subrectangular, clearly broader than long, with anterior margin concave medially as U-shaped and slightly concave posteriorly. Sternum 9 in ventral view subrectangular, clearly broader than long, with median portion apparently inflated; posterior margin of sternum 9 in ventral view widely concave (as U-shaped), and with posterolateral corners forming short, acutely angled digitiform processes. Gonostylus 9 in ventral view slender, unguiform, and curved inward. Ectoproct subcylindrical, short; in ventral view with a feebly developed tufted tubercle located anterior to posterolateral corners on inner margin; tip of ectoproct slightly incised and covered with dense short setae. Callus cerci in dorsal view suboval, connected to posterolateral corners of tergum 9. Fused gonocoxites 10 medially with a pair of large subtriangular processes, and laterally with a pair of digitiform gonostyli 10, leaving very short lateral arms (Fig. 2). Female. (Figs 3–5). Measurements (n=1). Total body length 27.2 mm; head width 5.3 mm, head length 4.3 mm (excluding labrum and mandibles); prothorax length 3.3 mm; prothorax width 3.8 mm; right forewing length 41.1 mm; right forewing width 13.8 mm; right hindwing length 39.1 mm; right hindwing width 14.8 mm. Morphologically similar to male, except these following characters:1) fused gonocoxites 8 in lateral view subtrapezoid, slightly protruding posteriad, in ventral view subrectangular, posterior margin medially slightly convex, and covered with dense short setae; 2) gonocoxite 9 in lateral view broadly subtriangular, slightly incised posteroventrally near tip and with a rather small lobe at tip; 3) upper part of ectoproct in lateral view digitiform, and lower part suboval; 4) callus cerci in lateral view large and suboval, completely fused with ectoprocts (Fig. 5). DNA barcode data. The mitochondrial COI sequences of our adult specimens are identical for both sexes showing genetic distance = 0. Phylogenetic analysis based on maximum likelihood estimation indicated that an unknown female specimen is conspecific with male adult of P. stigmosus. In addition, P. stigmosus, P. triangulates, and P. furcatus are genetically separated by high interspecific variations of COI sequences (Fig. 6). Distribution. China (Yunnan); Thailand (Nan Province, New record). Remarks. This species can be distinguished from the other members of the P. costalis group by the ten small dark spots (five pairs) on the pronotum in both sexes, and by the configuration of male fused gonocoxites 10 with a pair of large subtriangular lobes medially. Among the Thai species of Protohermes, P. stigmosus somewhat resembles P. triangulatus by the more complicated wing patterns. In this study, the female of P. stigmosus was collected and described for the first time. It is notable that the female genitalia of this species lacks the sac-like lateral lobes between abdominal segments 8 and 9. This trait is typical in the P. costalis group, although there are several species lacking this lobe, such as Protohermes niger Yang & Yang, 1988 and Protohermes basimaculatus Liu, Hayashi & Yang, 2007 (Liu et al. 2007; Chang et al. 2013). Whether the lack of the female sac-like lateral lobes indicates close relationships between above species (including P. stigmosus) or exclusion of these species from the P. costalis group needs further investigation. Protohermes stigmosus was first described based on a single adult male (reared from a larva) from Yunnan, China (Liu et al. 2007). Prior to the current record, no additional specimen had been found since 2007, suggesting possible rareness of this species. The locality of the present record is approximately 1,000 kilometers far from the type locality. Based on the few specimens that have been collected, this species inhabits mountainous areas with elevations ca. 1000 m. The holotype as well as the two males and one female from Thailand were collected in March, thus mating flight of the species might be in the dry season., Published as part of Piraonapicha, Kanyakorn, Sangpradub, Narumon, Jaitrong, Weeyawat, Attasopa, Korrawat & Liu, Xingyue, 2023, Frist record of Protohermes stigmosus Liu, Hayashi & Yang (Megaloptera Corydalidae: Corydalinae) in Thailand with the first description of female, pp. 87-93 in Zootaxa 5256 (1) on pages 87-93, DOI: 10.11646/zootaxa.5256.1.7, http://zenodo.org/record/7745407, {"references":["Liu, X. Y., Hayashi, F. & Yang, D. (2007) Systematics of the Protohermes costalis species-group (Megaloptera: Corydalidae). Zootaxa, 1439 (1), 1 - 46. https: // doi. org / 10.11646 / zootaxa. 1439.1.1","Yang, D. & Liu, X. Y. (2010) Fauna Sinica, Insecta. Vol. 51. Megaloptera. Science Press, Beijing, 457 pp. [in Chinese with English summary]","Martins, C. C., Ardila-Camacho, A., Rivera-Gasperin, S. L., Oswald, J. D., Liu, X. & Contreras-Ramos, A. (2022) A world checklist of extant and extinct species of Megaloptera (Insecta: Neuropterida). European Journal of Taxonomy, 812, 1 - 93. https: // doi. org / 10.5852 / ejt. 2022.812.1727","Chang, W., Hayashi, F., Liu, X. Y. & Yang, D. (2013) Discovery of the female of Protohermes niger Yang & Yang (Megaloptera: Corydalidae): Sexual dimorphism in coloration of a dobsonfly revealed by molecular evidence. Zootaxa, 3745 (1), 84 - 92. https: // doi. org / 10.11646 / zootaxa. 3745.1.7"]}

Protohermes stigmosus Liu, Hayashi & Yang, 2007 (Figs 1–5) Protohermes stigmosus Liu, Hayashi & Yang, 2007: 38 figs 20, 123–127; Yang & Liu, 2010: 180, fig. 78; Martins et al, 2022: 29. Types. Holotype. Male, China (Xishuangbanna Yunnan Province), 31.III.2004, reared from larva, Liang Tang leg., deposited in the Shanghai Normal University (SNU, examined), China. Non-type material examined. Thailand. One male, Bo Kluea District, Nan Province, 19°11U13.35UN, 101°10U5.91UE, 1006 m a.s.l., 4.III.2020, K. Piraonapicha & L. Khaton leg. leg. (THNHM, THNHM-I-24146, pinned); 1 female, same locality, date, collectors (THNHM, THNHM-I-24147, in alcohol). Diagnosis. Head without dark markings; pronotum with two pairs of blackish markings at anterolateral corner and three pairs of blackish markings near posterolateral corners; fused gonocoxites 10 medially with a pair of large subtriangular processes, and laterally with a pair of digitiform gonostyli 10, leaving very short lateral arms; female fused gonocoxites 8 with feebly convex posterior margin, medially without any notch; no lateral sac-like lobes between abdominal segments 8 and 9. Description. Male (Figs 1–3). Measurements (n=1). Total body length 35.1 mm (including mouthparts); head width 6.2 mm, head length 4.8 mm (excluding labrum and mandibles); prothorax length 4.3 mm; prothorax width 4.4 mm; right forewing length 53.2 mm; right forewing width 16.7 mm; right hindwing length 47.6 mm; right hindwing width 18.3 mm; wing spans 101.5 mm. Head yellow with obtuse postocular spine; compound eyes dark, located anterior to mid-length of head; ocelli located between compound eyes, close to antennal socket, inner margin brown; antennae subserrate, scapes and pedicel yellowish brown, scapes almost twice as long as pedicel, flagellum dark; labrum yellowish brown, subtriangular, shorter than broad, anterior margin distinctly convex, and covered with sparse short setae; mandibles yellow to reddish brown (anterior half reddish brown and posterior half yellow), with long and sharp apical tooth, followed by broad and subtriangular preapical tooth, small subtriangular prebasal tooth, and subtriangular basal tooth (denticles reddish brown); occiput subtriangular, yellow, posterior margin roundly convex in dorsal view. Pronotum yellow, subquadrate, with two pairs of blackish markings at anterolateral corner and three pairs of blackish markings at posterolateral corner (Fig. 4); mesonotum and metanotum subrectangular, yellow, both clearly broader than long; legs yellow, covered with short dense setae; distal half of protibiae and all protarsi black, meso- and metatarsi with second to fifth tarsomeres black; tarsal claws reddish brown; forewing pale grayish brown, with a subtriangular yellowish brown marking near base, yellowish brown markings at basal 1/3, and with an irregular markings at apical 1/3 and several extremely small yellowish markings on crossveins on apical portion; costal cellules with distinct grayish brown stripes, hindwing entirely hyaline; veins yellow except veins in dark regions of forewing and apical portion of hindwing pale brown; RA 8-branched, with 9 crossveins between RA and RP, MP 1+2 4-branched, MP 3+4 2-branched. Abdomen yellowish brown. Genitalia. Tergum 9 subrectangular, clearly broader than long, with anterior margin concave medially as U-shaped and slightly concave posteriorly. Sternum 9 in ventral view subrectangular, clearly broader than long, with median portion apparently inflated; posterior margin of sternum 9 in ventral view widely concave (as U-shaped), and with posterolateral corners forming short, acutely angled digitiform processes. Gonostylus 9 in ventral view slender, unguiform, and curved inward. Ectoproct subcylindrical, short; in ventral view with a feebly developed tufted tubercle located anterior to posterolateral corners on inner margin; tip of ectoproct slightly incised and covered with dense short setae. Callus cerci in dorsal view suboval, connected to posterolateral corners of tergum 9. Fused gonocoxites 10 medially with a pair of large subtriangular processes, and laterally with a pair of digitiform gonostyli 10, leaving very short lateral arms (Fig. 2). Female. (Figs 3–5). Measurements (n=1). Total body length 27.2 mm; head width 5.3 mm, head length 4.3 mm (excluding labrum and mandibles); prothorax length 3.3 mm; prothorax width 3.8 mm; right forewing length 41.1 mm; right forewing width 13.8 mm; right hindwing length 39.1 mm; right hindwing width 14.8 mm. Morphologically similar to male, except these following characters:1) fused gonocoxites 8 in lateral view subtrapezoid, slightly protruding posteriad, in ventral view subrectangular, posterior margin medially slightly convex, and covered with dense short setae; 2) gonocoxite 9 in lateral view broadly subtriangular, slightly incised posteroventrally near tip and with a rather small lobe at tip; 3) upper part of ectoproct in lateral view digitiform, and lower part suboval; 4) callus cerci in lateral view large and suboval, completely fused with ectoprocts (Fig. 5). DNA barcode data. The mitochondrial COI sequences of our adult specimens are identical for both sexes showing genetic distance = 0. Phylogenetic analysis based on maximum likelihood estimation indicated that an unknown female specimen is conspecific with male adult of P. stigmosus. In addition, P. stigmosus, P. triangulates, and P. furcatus are genetically separated by high interspecific variations of COI sequences (Fig. 6). Distribution. China (Yunnan); Thailand (Nan Province, New record). Remarks. This species can be distinguished from the other members of the P. costalis group by the ten small dark spots (five pairs) on the pronotum in both sexes, and by the configuration of male fused gonocoxites 10 with a pair of large subtriangular lobes medially. Among the Thai species of Protohermes, P. stigmosus somewhat resembles P. triangulatus by the more complicated wing patterns. In this study, the female of P. stigmosus was collected and described for the first time. It is notable that the female genitalia of this species lacks the sac-like lateral lobes between abdominal segments 8 and 9. This trait is typical in the P. costalis group, although there are several species lacking this lobe, such as Protohermes niger Yang & Yang, 1988 and Protohermes basimaculatus Liu, Hayashi & Yang, 2007 (Liu et al. 2007; Chang et al. 2013). Whether the lack of the female sac-like lateral lobes indicates close relationships between above species (including P. stigmosus) or exclusion of these species from the P. costalis group needs further investigation. Protohermes stigmosus was first described based on a single adult male (reared from a larva) from Yunnan, China (Liu et al. 2007). Prior to the current record, no additional specimen had been found since 2007, suggesting possible rareness of this species. The locality of the present record is approximately 1,000 kilometers far from the type locality. Based on the few specimens that have been collected, this species inhabits mountainous areas with elevations ca. 1000 m. The holotype as well as the two males and one female from Thailand were collected in March, thus mating flight of the species might be in the dry season.

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"]}

Lepidotrigona satun Attasopa and B��nziger 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 (���2.3) 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. B��nziger 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., 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 pages 66-74, DOI: 10.11646/zootaxa.4442.1.3, http://zenodo.org/record/1301390

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.

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