Description of four new species of Lepidotrigona (Hymenoptera: Apidae: Meliponini) from north-east India Author Viraktamath, Shashidhar Department of Entomology, University of Agricultural Sciences, GKVK, Bengaluru 560065 Author Thangjam, Rojeet 0000-0002-5352-4700 Department of Entomology, College of Horticulture, Central Agricultural University (Imphal), Thenzawl, Mizoram 796186 rojeetthangjam @ gmail. com https: // orcid. org / 0000 - 0002 - 5352 - 4700 text Zootaxa 2022 2022-08-15 5175 1 1 30 journal article 123828 10.11646/zootaxa.5175.1.1 9941acef-9d69-485b-875c-fafef9933fd9 1175-5326 7003134 396127FC-097F-42F6-BDB3-18CAF3FDD9AD Lepidotrigona thenzawlensis Viraktamath and Rojeet sp. n. ( Figs. 1 , 2 , 3 , 4 : a s4–s6, 5: a1–a3, 6 a, b) Diagnosis. Lepidotrigona thenzawlensis is a member of the ventralis species group with male and female bees measuring a mean of 5.18 and 4.79 mm long with forewing length of 4.66 and 4.52 mm , respectively ( Table 1 ). The species can be distinguished from L. rajithae sp. n. , L. amruthae sp. n. and L. sikkimensis sp. n. by the following characters: In male bees, gradulus of sternum 4 does not touch antecosta in L. thenzawlensis ( Fig. 4a s-4) (touches briefly in L. rajithae ( Fig. 4 b s-4) and L. sikkimensis ( Fig. 4 d s-4), does not touch in L. amruthae ( Fig. 4 c s-4); apical margin with a distinct emargination medially whereas, it is weakly emarginate in L. rajithae , produced into a broad lobe in L. amruthae ; gently convex in L. sikkimensis ; gradulus of sternum 5 touches antecosta extensively in L. thenzawlensis ( Fig. 4 a s-5), (touches briefly in L. rajithae (4 b s-5) and L. amruthae ( Fig. 4 c s-5), touches extensively in L. sikkimensis ( Fig. 4 d s-5)); distance between two apicosubmedian lobes is very short in L. thenzawlensis ( 0.15 mm ) and L. rajithae ( 0.18 mm ) but great in L. amruthae ( 0.29 mm ) and L. sikkimensis ( 0.31 mm ); apicolateral lobes longer than wide in L. thenzawlensis , L. rajithae and L. sikkimensis but as long as wide in L. amruthae . Antecosta in sternum 6 is weakly bisinuate in L. thenzawlensis ( Fig 4 a s-6), straight in L. rajithae ( Fig. 4 b s-6), weakly convex in L. amruthae ( Fig.4c s-6) and L. sikkimensis ( Fig. 4 d s-6); the apicomedial lobe 0.29 mm long, broadly pointed in L. thenzawlensis compared to L. rajithae ( 0.32 mm long, bluntly rounded), L. amruthae ( 0.33 mm long, acutely pointed) and L. sikkimensis ( 0.35 mm long, acutely pointed), the apodemal lobes widely separated in L. thenzawlensis moderately separated in L. rajithae , L. amruthae and L. sikkimensis ( Fig. 4 a-d s-6). The penis valve is shorter in L. thenzawlensis ( 0.62 mm ), moderately long in L. rajithae ( 0.80 mm ) and very long in L. amruthae ( 0.87 mm ) and L. sikkimensis ( 0.89 mm ) ( Table 2 ). Males of L. thenzawlensis also differ from other members of the ventralis species group namely L. flavibasis (Cockerell) , L. satun Attasopa and Banziger , L. doipaensis (Schwarz) and L. ventralis in the following characters. Mesoscuta of L. thenzawlensis , L. flavibasis . and L. doipaensis are not bordered by scale like hairs but bordered in L. satun ; in sternum 4, the apicomedial margin in L. thenzawlensis is with a distinct emargination while angularly emarginated in L. satun , slightly concave in L. doipaensis and convex in L. flavibasis , distinctly convex in L. ventralis ; apicosubmedian lobes of sternum 5 slender and pointed in L. thenzawlensis while they are rounded in L. satun , slender and pointed in L. flavibasis , L. doipaensis and triangular and pointed in L. ventralis ; gonostylus in L. thenzawlensis is shorter ( 0.82 mm ) than that of L. flavibasis ( 0.91 to 0.96 mm ) and L. ventralis ( 1.25 mm ) longer than in L. satun ( 0.75–0.78 mm ), L. doipaensis ( 0.67-0.71 mm );gonostylus expanded apically in all the species of ventralis group except L. ventralis ; spinulation in the apical expanded part of gonostylus gradually becomes denser towards apex in L. thenzawlensis , L. rajithae , L. amruthae and L. sikkimensis and L. satun while it is equally dense throughout the expanded part in L. flavibasis and L. doipaensis . Female bees of L. thenzawlensis differ from L. rajithae , L. amruthae , L. sikkimensis and L. arcifera , in the following respects. Mesoscutum of L. thenzawlensis is not bordered by scale-like plumose hairs but bordered in L. rajithae , L. amruthae and L. sikkimensis and L. arcifera ( Figs. 2 b , 8 b , 11 b , 14 b ); wings in all these new species with blackish brown tinge compared to clear and iridescent wings in L. arcifera ; metasomal first tergum black and no semi-circular band in L. thenzawlensis while in L. rajithae , L. amruthae , L. sikkimensis , and L. arcifera the tergum is light yellow or brown with light or dark brown or black semi-circular band, while in L. ventralis a dark spot on each side of pale tergum ( Sakagami 1975 ); malar space length is greater in L. thenzawlensis ( 0.15 mm ) as against 0.12, 0.10, 0.10 and 0.13 mm in L. rajithae , L. amruthae , L. sikkimensis and L. arcifera , respectively; ratio of interocellar to ocellocular distance, scape length to eye length, hind tibial length to forewing diagonal length and hind basitarsus width to hind tibial width are greater in all these four new species than those in L. arcifera ( Table 3 ). Description Males Coloration. Head, mesosoma and metasoma light brown to black; labrum reddish-brown; clypeus black; scape black except the basal bulb reddish-brown; pedicel, flagellomeres dark reddish-brown; ocelli transparent, shiny, light brown to reddish-brown; compound eyes black ( Fig. 1 a-c). Tegula, pterostigma, veins dark brown; legs light brown to reddish-brown, tarsi and hind tibia light brown. Mesoscutum with three very narrow, shiny longitudinal stripes arising from the anteromedial margin, extending up to the middle of mesoscutum. Metasomal terga dark reddish-brown approaching to black; tergum 1 black without any band ( Fig. 1 b ); sterna light brown with dark reddish-brown patches. Pilosity. Labrum, apical margin of clypeus fringed with long plumose white hairs; basal part of clypeus, face with white, short plumose hairs ( Fig. 1 c ); vertex with grey erect hairs; post genal margin fringed with long white hairs ( Fig. 1 a ). Mesoscutum without border of plumose scale-like hairs; integument with short greyish hairs; mesoscutellum fringed with short pale hairs; all legs with short white hairs; anterior and posterior margins, upper surface of the hind tibia with similar short white hairs. Pronotal lobe, mesepisternum and metepisternum with yellowishwhite long plumose hairs, the density and length of hairs increasing on the lower parts; the anterolateral surface of propodeum with thick white short plumose hairs ( Fig. 1 a ). Metasomal terga and sterna shiny, with white fine hairs, the density and length of hairs progressively increasing towards the terminal segments and medial regions of the sterna. Morphometry. Mean body length 5.18 mm , head width including compound eyes 1.83 mm ( Table 1 ); head length 1.26 mm ; eye 1.22 mm long 0.51 mm wide; upper interocular distance 3.09× greater than interocellar distance ( 0.35 mm ); median ocellus 19× greater than malar space length ( 0.01 mm ); scape 3.60× longer than its width ( 0.15 mm ); mandible 0.44 mm long, 0.22 mm wide; forewing 4.66 mm long 1.63 mm wide; forewing diagonal length 1.33 mm ; hind tibia 2.75× longer than its width ( 0.55 mm ) while the hind basitarsus 1.79× longer than its width ( 0.38 mm ). The ratio of head length to width 0.69; scape length to eye length 0.44; forewing length to width 2.86; hind tibial length to head width 0.83; hind basitarsus width to hind tibial width 0.69 ( Table 3 ). Metasomal sterna and genitalia. The following description is based on dissection of five males . Gradulus of sternum 4 transverse medially and not in touch with antecosta ( Fig. 3 a ); apico-medial margin with a distinct emargination, weakly angulate on both lateral sides. Gradulus of sternum 5 in touch with antecosta extensively; apical margin with two sub-median and two lateral lobes ( Fig. 3 b ); sub-median lobes longer ( 0.31 mm ), slender, pointed, terminating into 2–3 short spine-like structures; both lobes separated by 0.15 mm distance with a deep inverted U-shaped emargination; apicolateral lobes longer than wide ( 0.15 mm long, 0.07 mm wide), pointed, separated from sub-median lobe by a shallow inverted U-shaped emargination. Antecosta of sternum 6 weakly bisinuate in the middle with 0.29 mm long, triangular, broadly pointed apicomedian lobe. Apodemal lobes widely separated from each other ( Fig. 3 c ). Genitalia reddish-brown, asymmetrical (except gonostyli) with well sclerotized gonocoxae and penis valves while only basal 1/3 rd of gonostyli sclerotized ( Fig. 3 d, e ); left gonocoxa larger ( 0.23 mm long, 0.45 mm wide) than the right gonocoxa ( 0.23 mm long, 0.39 mm wide); penis valves dark reddish-brown; each penis valve 0.62 mm long, 0.32 mm wide at the base; terminal half of penis valve sharply curved laterally at a right angle and later narrowing terminally with a bluntly pointed apex; curvature of both penis valves asymmetrical, sometimes overlapping on each other. Each gonostylus arises laterally with a delicate attachment to the gonocoxa; 0.82 mm long, slender and tubular; sinuate at about the mid-length; 0.04 mm wide at the base, then widening to 0.06 mm in the middle and later slightly widening to 0.08 mm at the apical 1/4 th ; series of short and long setae arise from the medial and lateral surface of terminal expanded part with higher density near the apex ( Fig. 3 e ). Females Coloration. Head, mesosoma, metasoma black ( Fig. 2 a-c). Labrum, clypeus, compound eyes black. Scape, pedicel, flagellomeres black except basal bulb of scape reddish-brown ( Fig. 2 c ). Tegula black; wings with brownish-black tinge; pterostigma, veins dark brown; all the legs black except tarsi lighter. Mesoscutum with three very narrow, shiny longitudinal stripes arising from the anteromedial margin, extending up to the middle of mesoscutum. Metasomal terga and sterna black with inter-segmental lines light brown; tergum 1 black without any semi-circular band ( Fig. 2 b ). In some females, the depressed area of propodeum reddish- brown. Pilosity. Labrum fringed with short intermixed with long white hairs; clypeus, face, gena, post-gena, lower part of occiput clothed with short white hairs; vertex with brownish short erect hairs ( Fig. 2 a-c). Mesoscutum without a border of scale-like plumose hairs; mesoscutellum fringed with yellowish mixed with reddish-brown hairs ( Fig. 2 b ) . Pronotal lobe, anterior upper part of mesepisternum, metepisternum, anterolateral areas of propodeum with thick felt of white plumose hairs obscuring underlying integument; lower part of mesepisternum covered with long white hairs ( Fig. 2 a ). Anterior and posterior margins of hind tibia fringed with dark brown intermixed with pale hairs; upper surface shiny and bare. Basal margin of terga of metasoma with a transverse row of short white hairs; terminal terga fringed with long white hairs; sterna with long white hairs with high density on the medial region and terminal sterna. Morphometry. Female paratypes measure a mean of 4.79 mm in body length and 1.90 mm in head width ( Table 1 ). Head length, eye length, malar space length 1.35, 1.16, 0.15 mm , respectively; upper interocular distance 1.24 mm ; interocellar distance 1.17× greater than ocellocular distance ( 0.30 mm ); mandible 0.75 mm long, 0.28 mm wide; forewing 2.86× longer than its width ( 1.58 mm ); wing diagonal length 1.35 mm ; hind tibial length 2.23× longer than the hind basitarsus length ( 0.70 mm ); hind tibial width 1.45× greater than hind basitarsus width ( 0.40 mm ) The ratio of head length to width 0.71 ( Table 3 ); interocellar to ocellocular distance 1.17; scape length to eye length 0.61; forewing length to width 2.76; hind tibial length to head width 0.80; hind tibial length to forewing diagonal length 1.12; hind basitarsus width to hind tibial width 0.68. Nest. A colony of this species was found in the tree trunk. The entrance tube was 1.5 cm long, soft, brownish, with a wide opening of 1.10 cm . The brood cells were arranged in a single thick multi-layered comb ( Fig. 6 a, b ). FIGURE 1. Lepidotrigona thenzawlensis n. sp. Male holotype: a. lateral habitus, b. head, mesosoma and basal part of metasoma dorsal view, c. head frontal view FIGURE 2. Lepidotrigona thenzawlensis n. sp. Female paratype: a. lateral habitus, b. head, mesosoma and basal part of metasoma dorsal view, c. head frontal view FIGURE 3. Lepidotrigona thenzawlensis n. sp. Male paratype: a. Metasomal sternum 4, b. sternum 5, c. sternum 6, d. genitalia, e. gonostylus terminal part TABLE 1. Morphometry of four new species of Lepidotrigona and holotype of L. arcifera

Parameter/ ↓ Species→	Lepidotrigona thenzawlensis sp. n.		Lepidotrigona rajithae sp. n.		Lepidotrigona amruthae sp. n.		Lepidotrigona sikkimensis sp. n.		Lepidotrigona arcifera
Male N-11	Female N-10	Male N-11	Female N-10	Male N-11	Female N-10	Male N-09	Female N-10	Holotype Female*
Length of body	5.18 ± 0.35	4.79 ± 0.13	6.00 ± 0.22	4.15 ± 0.20	5.08 ± 0.39	4.09 ± 0.27	4.71 ± 0.16	4.16 ± 0.07	4.70
Width of head including eyes	1.83 ± 0.02	1.90 ± 0.02	1.82 ± 0.06	1,78 ± 0.02	1.78 ± 0.03	1.77 ± 0.02	1.77 ± 0.02	1.83 ± 0.03	1.89
Length of head	1.26 ± 0.03	1.35 ± 0.01	1.26 ± 0.02	1.27 ± 0.03	1.20 ± 0.02	1.26 ± 0.02	1.23 ± 0.02	1.29 ± 0.05	1.51
Length of eye	1.22 ± 0.03	1.16 ± 0.02	1.23 ± 0.02	1.09 ± 0.03	1.13 ± 0.04	1.12 ± 0.02	1.14 ± 0.01	1.11 ± 0.05	1.13
Width of eye	0.51 ± 0.02	0.42 ± 0.02	0.52 ± 0.02	0.42 ± 0.03	0.49 ± 0.03	0.38 ± 0.01	0.48 ± 0.01	0.39 ± 0.02	0.47
Upper inter ocular distance	1.08 ± 0.02	1.24 ± 0.02	1.07 ± 0.02	1.16 ± 0.03	1.05 ± 0.03	1.17 ± 0.02	1.05 ± 0.02	1.19 ± 0.04	1.23
Diameter of median ocellus	0.19 ± 0.01	0.17 ± 0.01	0.18 ± 0.01	0.16 ± 0.01	0.19 ± 0.01	0.15 ± 0.00	0.17 ± 0.01	0.15 ± 0.00	0.08
Inter ocellar distance	0.35 ± 0.01	0.35 ± 0.01	0.35 ± 0.01	0.34 ± 0.01	0.38 ± 0.01	0.35 ± 0.01	0.39 ± 0.02	0.35 ± 0.01	0.36
Ocello-ocular distance	0.19 ± 0.01	0.30 ± 0.01	0.21 ± 0.01	0.29 ± 0.01	0.20 ± 0.01	0.28 ± 0.01	0.20 ± 0.01	0.30 ± 0.01	0.32
Length of clypeus	0.41 ± 0.01	0.41 ± 0.04	0.43 ± 0.02	0.35 ± 0.03	0.38 ± 0.02	0.35 ± 0.01	0.37 ± 0.02	0.35 ± 0.01	0.36
Maximum width of clypeus	0.69 ± 0.03	0.86 ± 0.03	0.69 ± 0.02	0.78 ± 0.04	0.66 ± 0.02	0.73 ± 0.01	0.73 ± 0.02	0.74 ± 0.02	0.71
Malar space length	0.01 ± 0.00	0.15 ± 0.01	0.01 ± 0.00	0.12 ± 0.02	0.04 ± 0.02	0.10 ± 0.00	0.05 ± 0.01	0.10 ± 0.00	0.13
Length of scape	0.54 ± 0.01	0.69 ± 0.03	0.54 ± 0.01	0.65 ± 0.02	0.50 ± 0.01	0.65 ± 0.00	0.51 ± 0.20	0.65 ± 0.04	0.60
Width of scape	0.15 ± 0.01	0.11 ± 0.01	0.14 ± 0.01	0.12 ± 0.01	0.14 ± 0.01	0.10 ± 0.00	0.15 ± 0.01	0.10 ± 0.01	0.13
Length of pedicel + flagellum	1.83 ± 0.05	1.37 ± 0.03	1.82 ± 0.06	1.32 ± 0.05	1.66 ± 0.05	1.26 ± 0.05	1.79 ± 0.05	1.33 ± 0.04	1.35
Length of flagellomere 1	0.10 ± 0.01	0.12 ± 0.01	0.09 ± 0.01	0.10 ± 0.01	0.08 ± 0.02	0.10 ± 0.01	0.07 ± 0.01	0.13 ± 0.01	0.10
Length of flagellomere 2	0.15 ± 0.00	0.13 ± 0.00	0.16 ± 0.01	0.11 ± 0.01	0.15 ± 0.01	0.12 ± 0.00	0.15 ± 0.01	0.13 ± 0.01	0.10
Length of flagellomere 3	0.15 ± 0.00	0.13 ± 0.01	0.16 ± 0.01	0.11 ± 0.01	0.15 ± 0.01	0.12 ± 0.00	0.15 ± 0.01	0.13 ± 0.01	0.12
Width of flagellomere 3	0.14 ± 0.00	0.13 ± 0.01	0.15 ± 0.01	0.12 ± 0.01	0.14 ± 0.01	0.13 ± 0.00	0.14 ± 0.01	0.13 ± 0.00	0.14
Length of mandible	0.44 ± 0.02	0.75 ± 0.00	0.47 ± 0.02	0.66 ± 0.02	0.40 ± 0.01	0.70 ± 0.01	0.43 ± 0.01	0.68 ± 0.03	0.65
Width of mandible	0.22 ± 0.01	0.28 ± 0.01	0.23 ± 0.01	0.28 ± 0.02	0.19 ± 0.01	0.24 ± 0.01	0.19 ± 0.01	0.27 ± 0.03	0.21

......continued on the next page TABLE 1. (Continued)

Parameter/ ↓ Species→	Lepidotrigona thenzawlensis sp. n.		Lepidotrigona rajithae sp. n.		Lepidotrigona amruthae sp. n.		Lepidotrigona sikkimensis sp. n.		Lepidotrigona arcifera
Male N-11	Female N-10	Male N-11	Female N-10	Male N-11	Female N-10	Male N-09	Female N-10	Holotype Female*
Length of forewing + tegula	4.66 ± 0.01	4.52 ± 0.09	4.78 ± 0.14	4.32 ± 0.10	4.39 ± 0.12	4.24 ± 0.16	4.63 ± 0.10	4.49 ± 0.16	4.60
Width of forewing	1.63 ± 0.07	1,58 ± 0.09	1.69 ± 0.05	1.47 ± 0.05	1.46 ± 0.05	1.47 ± 0.03	1.49 ± 0.08	1.55 ± 0.07	1.65
Length of pterostigma	0.72 ± 0.04	0.73 ± 0.02	0.77 ± 0.02	0.72 ± 0.04	0.70 ± 0.02	0.72 ± 0.02	0.74 ± 0.02	0.73 ± 0.04	0.71
Length of marginal cell	1.52 ± 0.04	1.51 ± 0.01	1.59 ± 0.04	1.32 ± 0.05	1.40 ± 0.07	1.31 ± 0.01	1.48 ± 0.05	1.39 ± 0.10	1.40
Width of marginal cell	0.35 ± 0.01	0.36 ± 0.01	0.35 ± 0.01	0.33 ± 0.02	0.32 ± 0.01	0.33 ± 0.02	0.33 ± 0.02	0.35 ± 0.03	0.32
Wing diagonal length	1.33 ± 0.06	1.35 ± 0.03	1.36 ± 0.04	1.24 ± 0.02	1.27 ± 0.05	1.23 ± 0.02	1.27 ± 0.02	1.23 ± 0.04	1.33
Hamuli	6.00 ± 0.00	6.00 ± 0.00	6.00 ± 0.00	6.00 ± 0.00	6.33 ± 0.05	6.00 ± 0.00	6.00 ± 0.00	6.00 ± 0.00	6.00
Length of mesoscutum	1.25 ± 0.06	1.03 ± 0.03	1.27 ± 0.04	1.02 ± 0.03	1.16 ± 0.05	0.97 ± 0.01	1.21 ± 0.02	1.05 ± 0.04	1.08
Maximum width of mesoscutum	1.33 ± 0.05	1.33 ± 0.03	1.35 ± 0.06	1.20 ± 0.04	1.28 ± 0.03	1.23 ± 0.04	1.33 ± 0.04	1.23 ± 0.03	1.26
Length of hind tibia	1.51 ± 0.04	1.56 ± 0.04	1.54 ± 0.02	1.45 ± 0.03	1.45 ± 0.06	1.47 ± 0.02	1.41 ± 0.02	1.47 ± 0.03	1.43
Width of hind tibia	0.55 ± 0.00	0.58 ± 0.02	0.57 ± 0.02	0.54 ± 0.02	0.52 ± 0.02	0.56 ± 0.01	0.53 ± 0.02	0.56 ± 0.02	0.57
Length of hind basitarsus	0.68 ± 0.02	0.70 ± 0.04	0.70 ± 0.03	0.67 ± 0.02	0.62 ± 0.04	0.68 ± 0.02	0.55 ± 0.02	0.64 ± 0.02	0.62
Width of hind basitarsus	0.38 ± 0.01	0.40 ± 0.02	0.38 ± 0.02	0.40 ± 0.02	0.37 ± 0.02	0.41 ± 0.01	0.35 ± 0.01	0.36 ± 0.02	0.26

All measurements in mm (mean ± standard deviation) except hamuli, *Based on Rasmussen (2013) Materials examined. Holotype : Male : Mizoram : Thenzawl ( 23.2808° N , 92.7741° E , Altitude 783 m . a.s.l.), 20.x.2020 , leg. Rojeet T . deposited at UASB . Paratypes : Eight males, 30 females with the same collection data deposited at UASB ; one female paratype will be deposited at ZSIK . Etymology. This species is named after the place Thenzawl from where the samples were collected.