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Tetragonula (Tetragonula) malaipanae , new species Figures 1���3 DIAGNOSIS: The species generally agrees with the subgeneric characterization as presented by Sakagami (1978). Worker dark but paler than Tetragonula laeviceps (Smith, 1857), T. pagdeni (Schwarz, 1939), and T. pagdeniformis (Sakagami, 1978) (figs. 1A, 1B), clypeus yellowish (fig. 1C), legs partly or wholly red brown or testaceous (fig. 1A). Size similar to that of T. testaceitarsis (Cameron, 1901) (likely senior synonym of T. valdezi [Cockerell, 1918]), larger than T. laeviceps (as restricted by Rasmussen and Michener, 2010), T. pagdeni, and T. pagdeniformis (see Discussion, below). Mesoscutum with pale pubescence not or weakly banded (fig. 1B), sublateral bare bands not or scarcely recognizable. Setae of body largely pale, but those of vertex, especially behind ocelli, blackish or sometimes dusky. Mean metatibial length 1.81 mm, mean forewing length including tegula 4.58 mm, mean head width 1.90 mm. Male generally colored as in worker (fig. 2), with S5 lacking gradulus (fig. 3C), apical process of S6 long and parallel sided (fig. 3D), S7 and T7 sharply pointed apically (figs. 3B, 3E); genitalia with penis valves and gonostyli directed posteriorly (figs. 3F, 3H), sclerotization probably representing gonobase across base (figs. 3F, 3H). DESCRIPTION: Worker. Body length 3.92 mm (3.50���4.70 mm); forewing length, including tegula, 4.58 mm (4.40���4.70 mm); head length from anterior margin of clypeus to summit of vertex, in facial view 1.42 mm (1.40���1.50 mm); head width 1.91 mm (1.75���2.00 mm); metatibia length 1.82 mm (1.63���1.88 mm); forewing diagonal from base of vein M to base of crossvein r-rs at margin of pterostigma 1.31 mm (1.23���1.38 mm). Scape (excluding basal bulb) more than half as long as compound eye; flagellomere IV slightly broader than long. Ocelloccipital distance over half OD, ocellocular distance less than interocellar distance. Genal area widest at about upper third of compound eye, and narrower than compound eye in profile. Metatibia and metabasitarsus shaped as usual in this group (see Sakagami, 1978: fig. 2); basal sericeous area at base of inner surface of metabasitarsus well defined, reaching to middle of metabasitarsus [variable, not attaining middle or extending beyond middle]. Metasoma short, about as wide as head, widest at T2 and T3, remaining terga progressively much narrower and largely telescoped into preceding segments. Cuticular surface polished and shining with few minute punctures or setal bases. Head black, the following parts yellow testaceous [to reddish testaceous or pale yellowish brown]: clypeus except dusky upper and upper lateral margins [entire clypeus commonly yellowish], supraclypeal area [sometimes dusky or blackish], mandible except blackish apex and extreme base, and labrum. [Lower genal area and hypostomal area behind mandibular base sometimes weakly brownish, grading into black background]. Antenna yellowish [to reddish] testaceous, flagellum blackish [or dusky] on upper surface, on lower surface part of each flagellomere dusky [or not]. Thorax black except pronotum dark brown [to blackish], pronotal lobe dusky testaceous [to black], mesoscutellum brownish [or black], [mesepisternum sometimes brownish black], anterolateral part of mesepisternum brown [to black]. Legs (including coxae and trochanters) reddish brown, meso- and metatibiae and tarsi dark reddish brown [or rarely legs testaceous or legs blackish brown, sometimes only mid and hind legs dark, inner surface of metatibia and metatarsus and often basal part of metafemur reddish brown]. Tegula dusky testaceous. Wings clear, faintly dusky, veins and pterostigma blackish. Metasoma with T1 dusky brown [brown], anterior surface largely testaceous; T5 and T6 [sometimes on T6] testaceous [brown or dusky]; T2 and T4 [sometimes also T5] brownish black, posterior margins paler translucent [posterior part of T1 sometimes similarly dark]. [Sometimes, especially specimens from nests 6 and 9, with all terga testaceous, posterior ones darker than anterior ones]. Metasomal sterna testaceous [brown, sometimes slightly dusky]. Pubescence decumbent and pallid except as otherwise noted, mostly ochraceous; clypeus without erect simple setae, even lower margin without such setae. Setae of face short (0.5 OD), plumose, rather dense but not hiding surface, particularly sparse and thin on lower paraocular area, whitish, those of frons [or upper half of frons] brownish to yellowish and hence inconspicuous in certain views; erect setae of scape absent except for exceedingly short ones; longer setae (1 OD) of vertex, especially behind ocelli, blackish or dusky; short pale setae of mesoscutum weakly [or not recognizably] banded. When recognizable, median asetose band (���G1��� of Sakagami, 1978) narrow, extending full length of mesoscutum except anterior extremity, G2 similarly narrow anteriorly but broadening and diverging posteriorly, G3 (lateral asetose band) poorly defined, especially on anterior half of mesoscutum [not recognizable except in some fresh, clean specimens]; longer, erect setae of mesoscutum (2 OD) pale or in certain illuminations dusky; fringe of long setae (up to 3 OD) on posterior margin of mesoscutellum ochraceous, with longest setae (3���4 OD) sometimes appearing dusky; metepisternum, side of propodeum, and anterior half of mesepisternum covered with dense white setae hiding surface [those of mesepisternum sometimes ochraceous]; some setae of outer surfaces of meso- and metatibiae and basitarsi dusky [commonly most setae of these parts, including large simple discal setae of corbicula, black or blackish]; anterior fringe of corbicula with a few [nearly all] setae dusky or blackish. T1 smooth and asetose; T2 to T5 smooth and asetose except for preapical very sparse bands, that of T2 consisting of only a few minute ochraceous setae, those of following terga with progressively more and longer setae, a few of those of T4 reaching beyond apical margin of tergum, those of T5 and T6 still longer, but all small and not conspicuous; setae of sterna denser and longer, almost white. Male. Agrees with description of worker except for variation noted above in brackets, in measurements provided, and as follows (except for measurements, description based on male collected from nest 4 on 17 September 2007): body length 4.50 mm; forewing length, including tegula, 4.45 mm (4.40���4.50 mm); head length (measured as for worker) 1.40 mm; head width 1.78 mm; metatibia length 1.46 mm (1.43���1.48 mm); forewing diagonal (measured as for worker) 1.26 mm (1.25���1.27 mm). Scape (excluding basal bulb) slightly less than half as long as compound eye. Flagellomere 4 (also 2���10) longer than broad. Ocelloccipital distance less than half OD; ocellocular distance about half of interocellar distance. Metatibia widest at about apical third, narrowed at apex, without plumose setae (fig. 3A). Metasomal T7 without Y-shaped ridge, apex broadly rounded (fig. 3B); S5 short, without noticeable gradulus, with shallow, broad, midapical emargination (fig. 3C); apical process of S6 slender, parallel sided (fig. 3D); terminalia as illustrated in figures 3D���3H; unbroken, weakly sclerotized band in position of gonobase (fig. 3F); long membranous midventral genital process present (figs. 3G, 3H), presumably fragmentum from base of penis; penis valves and gonostyli directed posteriorly (figs. 3F, 3H); gonostylus arising near outer distal margin of gonocoxite (figs. 3F, 3H), flattened, so that in lateral view it is rather broad and subapically constricted (fig. 3H), in dorsal view very slender apically. Pale areas of head yellowish brown, upper median part of supraclypeal area black. Genal and hypostomal areas black. Flagellum black. Pronotum including pronotal lobe, mesoscutellum, and mesepisternum black. Legs yellowish brown, distal parts of femora, basal parts of tibiae, and distal parts of meso- and metatibia infuscated. Metasoma with T1 testaceous, with broad preapical dusky band; remaining terga cleared for study but apparently dusky testaceous (note that when dissected the metasoma appears pale testaceous). Pubescence generally thinner and sparser than in female, particularly on clypeus, mesoscutellum, and mesepisternum; pubescence of head, mesosoma, and legs largely ochraceous. Mesoscutum almost without banding of setae; setae of mesepisternum and sides of propodeum white, somewhat less dense than in worker; dense setae of anterior part of mesepisternum dusky. HOLOTYPE: ���, Thailand, Kanchanaburi [Province], Thong Pha Phum [District], 17.ix.2007 [17 September 2007], Y. Boontop, nest 2 (SEMC). The holotype is from nest number 2 in Boontop���s records, a site 549 m above sea level at coordinates 14.42�� N, 98.28�� E. PARATYPES (2♂♂, 340������): 1♂, same data as holotype except nest 4 (SEMC); 1♂, same data as holotype except 12.iii.2008 [12 March 2008] and nest 11 (SEMC); 24������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 1 (16 SEMC, 2 CSCA, 2 DZUP, 2 EMBT, 2 ICNC); 11������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 1 (SEMC); 18������, same data as holotype (16 SEMC, 2 EMBT); 17������, same data as holotype except 12. iii.2008 [12 March 2008] (9 SEMC, 4 AMNH, 2 BBSL, 2 CNIN); 24������, same data as holotype except 27.vii.2009 [27 July 2009] (16 SEMC, 2 NHML, 2 SANC, 2 USNM, 2 EMBT); 8������, same data as holotype except nest 3 (SEMC); 13������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 3 (SEMC); 14������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 3 (10 SEMC, 2 ZMHB, 2 ZISP); 2������, same data as holotype except nest 4 (SEMC); 12������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 4 (10 SEMC, 2 KSMA); 11������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 4 (SEMC); 2������, same data as holotype except nest 5 (SEMC); 14������, same data as holotype except 12. iii.2008 [12 March 2008] and nest 5 (11 SEMC, 3 EMBT); 15������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 5 (11 SEMC, 2 KUEC, 2 EMBT); 5������, same data as holotype except 31.vii.2006 [31 July 2006] and nest 6 (SEMC); 9������, same data as holotype except nest 6 (SEMC); 15������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 6 (13 SEMC, 1 FSCA, 1 LIPI); 21������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 6 (19 SEMC, 1 FSCA, 1 LIPI); 5������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 7 (SEMC); 12������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 7 (SEMC); 5������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 8 (SEMC); 9������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 9 (SEMC); 5������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 10 (SEMC); 4������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 10 (SEMC); 14������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 11 (SEMC); 3������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 11 (SEMC); 6������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 12 (SEMC); 3������, same data as holotype except 27.vii.2009 [27 July 2009] and nest 12 (SEMC); 8������, same data as holotype except 12.iii.2008 [12 March 2008] and nest 13 (SEMC); 2������, same data as holotype except 31.vii.2006 [31 July 2006] and nest 19 (SEMC); 2������, same data as holotype except 17.ix.2007 [17 September 2007] (SEMC). ADDITIONAL MATERIAL: A further six workers are remarkably similar to T. malaipanae and we tentatively include them here, pending further work on these more southern populations, particularly utilizing morphometrics, males, and DNA data: 4������, Thailand, Nakhonsithammarat, Khao Laung, 3.v.2007 [3 May 2007], Y. Boontop, nest 11 [a site 359 m above sea level at coordinates 8.49�� N, 99.78�� E] (SEMC); 2������, Thailand, Suratthani, Khao sok, 3.v.2007 [3 May 2007], Y. Boontop, nest 15 [a site 421 m above sea level at coordinates 8.91�� N, 98.52�� E] (SEMC). COMMENTS: Probably variation in coloration of males is more or less comparable to that of workers. The male taken 12 March 2008 at nest number 11 differs from the male described above as follows: integument of head and mesosoma (including clypeus and tegula) black; scape and pedicel reddish brown; legs dusky pale brown; meso- and metatarsi and distal parts of tibiae blackish, palest areas on outer and posterior surfaces of protibia and basal parts of meso- and metatibiae; metasoma reddish brown with vague transverse dusky areas, apical terga darker; membranous midventral genital process not evident (likely a result of damage during dissection). ETYMOLOGY: The specific epithet honors Savitree Malaipan of the Department of Agriculture, Kasetsart University, Bangkok, beloved mentor of Y.B. and a prominent Thai melittologist., Published as part of Engel, Michael S., Michener, Charles D. & Boontop, Yuvarin, 2017, Notes on Southeast Asian Stingless Bees of the Genus Tetragonula (Hymenoptera: Apidae), with the Description of a New Species from Thailand, pp. 1-17 in American Museum Novitates 2017 (3886) on pages 5-11, DOI: 10.1206/3886.1, http://zenodo.org/record/5062278, {"references":["Sakagami, S. F. 1978. Tetragonula stingless bees of the continental Asia and Sri Lanka (Hymenoptera, Apidae). Journal of the Faculty of Science, Hokkaido University, Series 6, Zoology 21 (2): 165 - 247, + 1 pl. [pl. v].","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 (6): 42 - 88, + 2 pls.","Schwarz, H. F. 1939. The Indo-Malayan species of Trigona. Bulletin of the American Museum of Natural History 76 (3): 83 - 141.","Cameron, P. 1901. On the Hymenoptera collected during the \" Skeat Expedition \" to the Malay Peninsula, 1899 - 1900. Proceedings of the Zoological Society of London 2 (1): 16 - 44.","Cockerell, T. D. A. 1918. Descriptions and records of bees - LXXX. Annals and Magazine of Natural History (9) 2 (10): 384 - 390.","Rasmussen, C., and C. D. Michener. 2010. The identity and neotype of Trigona laeviceps Smith (Hymenoptera: Apidae). Journal of the Kansas Entomological Society 83 (2): 129 - 133."]}

Tetragonula (Tetragonula) sarawakensis (Schwarz, 1937) Trigona sarawakensis Schwarz, 1937: 313. Cotype workers (AMNH, examined by M.S.E. and C.D.M.). Trigona (Tetragona) sarawakensis var. sarawakensis Schwarz; Schwarz, 1939: 106. Tetragonula sarawakensis (Schwarz); Moure, 1961: 210. Trigona (Tetragonula) sarawakensis (Schwarz); Sakagami, 1978: 210. Since this is not a well-known species, we include below notes based on a series of workers in the AMNH, two labeled as cotypes, and include comparative information on T. malaipanae in brackets. DESCRIPTIVE NOTES: Setae of clypeus and lower face minute, minutely if at all plumose, not at all hiding smooth surface [broadly plumose, partially hiding surface which shows some punctation]; tegula ferruginous [dark testaceous, sometimes grading to dusky especially near inner margin]; mesoscutellum with posterior extremity (overhanging propodeum) with transverse yellow mark (sometimes ferruginous, perhaps artificially darkened in such instances owing to method of collection or changes resulting from long-term preservation) [no such mark or if mark recognizable, testaceous or dusky]; legs ferruginous [usually dull ferruginous or brown with dusky areas at least on meso- and metatibiae]; setae of legs whitish or yellowish, sometimes a few dusky setae on tibiae and basitarsi [some setae on tibiae and basitarsi dusky or blackish, large simple setae arising from corbicular surface black or dusky although marginal setae largely or partly yellowish]; metasoma entirely ferruginous [dusky brown to testaceous]. MATERIAL EXAMINED (7������): 1��� [cotype], Sarawak, Mt. Dulit, 4000 ft, moss forest, 18.x.1932 [18 October 1932], Oxford Univ. Exp. B.M. Hobby & A.W. Moore (AMNH); 1��� [cotype], same data as preceding specimen except 19.x.1932 [19 October 1932] (AMNH); 5������, Mt. Tibang [Mount Tibang, Sarawak, near border with Kalimantan], 1300 m, Borneo, Nov. 1925, Eric Mj��berg (AMNH)., Published as part of Engel, Michael S., Michener, Charles D. & Boontop, Yuvarin, 2017, Notes on Southeast Asian Stingless Bees of the Genus Tetragonula (Hymenoptera: Apidae), with the Description of a New Species from Thailand, pp. 1-17 in American Museum Novitates 2017 (3886) on pages 11-12, DOI: 10.1206/3886.1, http://zenodo.org/record/5062278, {"references":["Schwarz, H. F. 1937. Results of the Oxford University Sarawak (Borneo) expedition: Bornean stingless bees of the genus Trigona. Bulletin of the American Museum of Natural History 73 (3): 281 - 328, + 6 pls. [pls. ii-vii].","Schwarz, H. F. 1939. The Indo-Malayan species of Trigona. Bulletin of the American Museum of Natural History 76 (3): 83 - 141.","Moure, J. S. 1961. A preliminary supra-specific classification of the Old World meliponine bees (Hymenoptera, Apoidea). Studia Entomologica 4 (1 - 4): 181 - 242.","Sakagami, S. F. 1978. Tetragonula stingless bees of the continental Asia and Sri Lanka (Hymenoptera, Apidae). Journal of the Faculty of Science, Hokkaido University, Series 6, Zoology 21 (2): 165 - 247, + 1 pl. [pl. v]."]}

Tetragonula (Tetragonula) malaipanae , new species Figures 1–3 DIAGNOSIS: The species generally agrees with the subgeneric characterization as presented by Sakagami (1978). Worker dark but paler than Tetragonula laeviceps (Smith, 1857), T. pagdeni (Schwarz, 1939), and T. pagdeniformis (Sakagami, 1978) (figs. 1A, 1B), clypeus yellowish (fig. 1C), legs partly or wholly red brown or testaceous (fig. 1A). Size similar to that of T. testaceitarsis (Cameron, 1901) (likely senior synonym of T. valdezi [Cockerell, 1918]), larger than T. laeviceps (as restricted by Rasmussen and Michener, 2010), T. pagdeni, and T. pagdeniformis (see Discussion, below). Mesoscutum with pale pubescence not or weakly banded (fig. 1B), sublateral bare bands not or scarcely recognizable. Setae of body largely pale, but those of vertex, especially behind ocelli, blackish or sometimes dusky. Mean metatibial length 1.81 mm, mean forewing length including tegula 4.58 mm, mean head width 1.90 mm. Male generally colored as in worker (fig. 2), with S5 lacking gradulus (fig. 3C), apical process of S6 long and parallel sided (fig. 3D), S7 and T7 sharply pointed apically (figs. 3B, 3E); genitalia with penis valves and gonostyli directed posteriorly (figs. 3F, 3H), sclerotization probably representing gonobase across base (figs. 3F, 3H). DESCRIPTION: Worker. Body length 3.92 mm (3.50–4.70 mm); forewing length, including tegula, 4.58 mm (4.40–4.70 mm); head length from anterior margin of clypeus to summit of vertex, in facial view 1.42 mm (1.40–1.50 mm); head width 1.91 mm (1.75–2.00 mm); metatibia length 1.82 mm (1.63–1.88 mm); forewing diagonal from base of vein M to base of crossvein r-rs at margin of pterostigma 1.31 mm (1.23–1.38 mm). Scape (excluding basal bulb) more than half as long as compound eye; flagellomere IV slightly broader than long. Ocelloccipital distance over half OD, ocellocular distance less than interocellar distance. Genal area widest at about upper third of compound eye, and narrower than compound eye in profile. Metatibia and metabasitarsus shaped as usual in this group (see Sakagami, 1978: fig. 2); basal sericeous area at base of inner surface of metabasitarsus well defined, reaching to middle of metabasitarsus [variable, not attaining middle or extending beyond middle]. Metasoma short, about as wide as head, widest at T2 and T3, remaining terga progressively much narrower and largely telescoped into preceding segments. Cuticular surface polished and shining with few minute punctures or setal bases. Head black, the following parts yellow testaceous [to reddish testaceous or pale yellowish brown]: clypeus except dusky upper and upper lateral margins [entire clypeus commonly yellowish], supraclypeal area [sometimes dusky or blackish], mandible except blackish apex and extreme base, and labrum. [Lower genal area and hypostomal area behind mandibular base sometimes weakly brownish, grading into black background]. Antenna yellowish [to reddish] testaceous, flagellum blackish [or dusky] on upper surface, on lower surface part of each flagellomere dusky [or not]. Thorax black except pronotum dark brown [to blackish], pronotal lobe dusky testaceous [to black], mesoscutellum brownish [or black], [mesepisternum sometimes brownish black], anterolateral part of mesepisternum brown [to black]. Legs (including coxae and trochanters) reddish brown, meso- and metatibiae and tarsi dark reddish brown [or rarely legs testaceous or legs blackish brown, sometimes only mid and hind legs dark, inner surface of metatibia and metatarsus and often basal part of metafemur reddish brown]. Tegula dusky testaceous. Wings clear, faintly dusky, veins and pterostigma blackish. Metasoma with T1 dusky brown [brown], anterior surface largely testaceous; T5 and T6 [sometimes on T6] testaceous [brown or dusky]; T2 and T4 [sometimes also T5] brownish black, posterior margins paler translucent [posterior part of T1 sometimes similarly dark]. [Sometimes, especially specimens from nests 6 and 9, with all terga testaceous, posterior ones darker than anterior ones]. Metasomal sterna testaceous [brown, sometimes slightly dusky]. Pubescence decumbent and pallid except as otherwise noted, mostly ochraceous; clypeus without erect simple setae, even lower margin without such setae. Setae of face short (0.5 OD), plumose, rather dense but not hiding surface, particularly sparse and thin on lower paraocular area, whitish, those of frons [or upper half of frons] brownish to yellowish and hence inconspicuous in certain views; erect setae of scape absent except for exceedingly short ones; longer setae (1 OD) of vertex, especially behind ocelli, blackish or dusky; short pale setae of mesoscutum weakly [or not recognizably] banded. When recognizable, median asetose band (“G1” of Sakagami, 1978) narrow, extending full length of mesoscutum except anterior extremity, G2 similarly narrow anteriorly but broadening and diverging posteriorly, G3 (lateral asetose band) poorly defined, especially on anterior half of mesoscutum [not recognizable except in some fresh, clean specimens]; longer, erect setae of mesoscutum (2 OD) pale or in certain illuminations dusky; fringe of long setae (up to 3 OD) on posterior margin of mesoscutellum ochraceous, with longest setae (3–4 OD) sometimes appearing dusky; metepisternum, side of propodeum, and anterior half of mesepisternum covered with dense white setae hiding surface [those of mesepisternum sometimes ochraceous]; some setae of outer surfaces of meso- and metatibiae and basitarsi dusky [commonly most setae of these parts, including large simple discal setae of corbicula, black or blackish]; anterior fringe of corbicula with a few [nearly all] setae dusky or blackish. T1 smooth and asetose; T2 to T5 smooth and asetose except for preapical very sparse bands, that of T2 consisting of only a few minute ochraceous setae, those of following terga with progressively more and longer setae, a few of those of T4 reaching beyond apical margin of tergum, those of T5 and T6 still longer, but all small and not conspicuous; setae of sterna denser and longer, almost white. Male. Agrees with description of worker except for variation noted above in brackets, in measurements provided, and as follows (except for measurements, description based on male collected from nest 4 on 17 September 2007): body length 4.50 mm; forewing length, including tegula, 4.45 mm (4.40–4.50 mm); head length (measured as for worker) 1.40 mm; head width 1.78 mm; metatibia length 1.46 mm (1.43–1.48 mm); forewing diagonal (measured as for worker) 1.26 mm (1.25–1.27 mm). Scape (excluding basal bulb) slightly less than half as long as compound eye. Flagellomere 4 (also 2–10) longer than broad. Ocelloccipital distance less than half OD; ocellocular distance about half of interocellar distance. Metatibia widest at about apical third, narrowed at apex, without plumose setae (fig. 3A). Metasomal T7 without Y-shaped ridge, apex broadly rounded (fig. 3B); S5 short, without noticeable gradulus, with shallow, broad, midapical emargination (fig. 3C); apical process of S6 slender, parallel sided (fig. 3D); terminalia as illustrated in figures 3D–3H; unbroken, weakly sclerotized band in position of gonobase (fig. 3F); long membranous midventral genital process present (figs. 3G, 3H), presumably fragmentum from base of penis; penis valves and gonostyli directed posteriorly (figs. 3F, 3H); gonostylus arising near outer distal margin of gonocoxite (figs. 3F, 3H), flattened, so that in lateral view it is rather broad and subapically constricted (fig. 3H), in dorsal view very slender apically. Pale areas of head yellowish brown, upper median part of supraclypeal area black. Genal and hypostomal areas black. Flagellum black. Pronotum including pronotal lobe, mesoscutellum, and mesepisternum black. Legs yellowish brown, distal parts of femora, basal parts of tibiae, and distal parts of meso- and metatibia infuscated. Metasoma with T1 testaceous, with broad preapical dusky band; remaining terga cleared for study but apparently dusky testaceous (note that when dissected the metasoma appears pale testaceous). Pubescence generally thinner and sparser than in female, particularly on clypeus, mesoscutellum, and mesepisternum; pubescence of head, mesosoma, and legs largely ochraceous. Mesoscutum almost without banding of setae; setae of mesepisternum and sides of propodeum white, somewhat less dense than in worker; dense setae of anterior part of mesepisternum dusky. HOLOTYPE: ⚲, Thailand, Kanchanaburi [Province], Thong Pha Phum [District], 17.ix.2007 [17 September 2007], Y. Boontop, nest 2 (SEMC). The holotype is from nest number 2 in Boontop’s records, a site 549 m above sea level at coordinates 14.42° N, 98.28° E. PARATYPES (2♂♂, 340⚲⚲): 1♂, same data as holotype except nest 4 (SEMC); 1♂, same data as holotype except 12.iii.2008 [12 March 2008] and nest 11 (SEMC); 24⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 1 (16 SEMC, 2 CSCA, 2 DZUP, 2 EMBT, 2 ICNC); 11⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 1 (SEMC); 18⚲⚲, same data as holotype (16 SEMC, 2 EMBT); 17⚲⚲, same data as holotype except 12. iii.2008 [12 March 2008] (9 SEMC, 4 AMNH, 2 BBSL, 2 CNIN); 24⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] (16 SEMC, 2 NHML, 2 SANC, 2 USNM, 2 EMBT); 8⚲⚲, same data as holotype except nest 3 (SEMC); 13⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 3 (SEMC); 14⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 3 (10 SEMC, 2 ZMHB, 2 ZISP); 2⚲⚲, same data as holotype except nest 4 (SEMC); 12⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 4 (10 SEMC, 2 KSMA); 11⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 4 (SEMC); 2⚲⚲, same data as holotype except nest 5 (SEMC); 14⚲⚲, same data as holotype except 12. iii.2008 [12 March 2008] and nest 5 (11 SEMC, 3 EMBT); 15⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 5 (11 SEMC, 2 KUEC, 2 EMBT); 5⚲⚲, same data as holotype except 31.vii.2006 [31 July 2006] and nest 6 (SEMC); 9⚲⚲, same data as holotype except nest 6 (SEMC); 15⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 6 (13 SEMC, 1 FSCA, 1 LIPI); 21⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 6 (19 SEMC, 1 FSCA, 1 LIPI); 5⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 7 (SEMC); 12⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 7 (SEMC); 5⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 8 (SEMC); 9⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 9 (SEMC); 5⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 10 (SEMC); 4⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 10 (SEMC); 14⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 11 (SEMC); 3⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 11 (SEMC); 6⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 12 (SEMC); 3⚲⚲, same data as holotype except 27.vii.2009 [27 July 2009] and nest 12 (SEMC); 8⚲⚲, same data as holotype except 12.iii.2008 [12 March 2008] and nest 13 (SEMC); 2⚲⚲, same data as holotype except 31.vii.2006 [31 July 2006] and nest 19 (SEMC); 2⚲⚲, same data as holotype except 17.ix.2007 [17 September 2007] (SEMC). ADDITIONAL MATERIAL: A further six workers are remarkably similar to T. malaipanae and we tentatively include them here, pending further work on these more southern populations, particularly utilizing morphometrics, males, and DNA data: 4⚲⚲, Thailand, Nakhonsithammarat, Khao Laung, 3.v.2007 [3 May 2007], Y. Boontop, nest 11 [a site 359 m above sea level at coordinates 8.49° N, 99.78° E] (SEMC); 2⚲⚲, Thailand, Suratthani, Khao sok, 3.v.2007 [3 May 2007], Y. Boontop, nest 15 [a site 421 m above sea level at coordinates 8.91° N, 98.52° E] (SEMC). COMMENTS: Probably variation in coloration of males is more or less comparable to that of workers. The male taken 12 March 2008 at nest number 11 differs from the male described above as follows: integument of head and mesosoma (including clypeus and tegula) black; scape and pedicel reddish brown; legs dusky pale brown; meso- and metatarsi and distal parts of tibiae blackish, palest areas on outer and posterior surfaces of protibia and basal parts of meso- and metatibiae; metasoma reddish brown with vague transverse dusky areas, apical terga darker; membranous midventral genital process not evident (likely a result of damage during dissection). ETYMOLOGY: The specific epithet honors Savitree Malaipan of the Department of Agriculture, Kasetsart University, Bangkok, beloved mentor of Y.B. and a prominent Thai melittologist.

Engel, Michael S., Michener, Charles D., Boontop, Yuvarin (2017): Notes on Southeast Asian Stingless Bees of the Genus Tetragonula (Hymenoptera: Apidae), with the Description of a New Species from Thailand. American Museum Novitates 2017 (3886): 1-17, DOI: 10.1206/3886.1, URL: http://www.bioone.org/doi/10.1206/3886.1

Key to Species of Astibomelissa 1.Mesoscutum with numerous appressed to decumbent, plumose setae (usually missing on central disc, presumably due to abrasion), setae longer, not squamiform; outer surface of probasitarsus with fine, erect to suberect setae with distinctly sinuate or wavy apices (= type IV of Pasteels and Pasteels, 1974); inner setae of metabasitarsus white to pale yellow and similar to setae of scopa; metasomal tergum I without transverse carina at angle of anterior- and dorsal-facing surfaces; smaller species, 5.5���7.5 mm in total length (Saudi Arabia; northeastern Africa).................................................... 2 ���Mesoscutum tomentose, with dense covering of appressed, minute (typically about 0.50�� ocellar diameter), squamiform setae; outer surface of probasitarsus with fine, subdecumbent setae tapering to straight apices; inner setae of metabasitarsus fuscous, contrasting with surrounding white to pale yellow setae of scopa; metasomal tergum I with transverse carina at angle of anterior- and dorsal-facing surfaces; larger species, 7.9���8.8 mm in length [Iran]............................................... T. persica (Warncke) 2.Female with yellow facial maculation confined to labrum, clypeus, and supraclypeal area (fig. 3C); metasomal scopa dense, with setae greatly elongate and subdecumbent to suberect; small species, approximately 5.5���6.5 mm in length (male unknown) [Egypt, Sudan].................................................... T. mimetes (Cockerell) ���Female with yellow facial maculation extending to paraocular area and scape (fig. 6C); metasomal scopa not so dense, with setae long and largely decumbent to subdecumbent; moderate-sized species, approximately 6.7���7.5 mm in length (central Saudi Arabia)........................................................... T. kindahensis Engel, n. sp., Published as part of Engel, Michael S., Michener, Charles D. & Boontop, Yuvarin, 2017, Discovery of the Bee Tribe Tarsaliini in Arabia (Hymenoptera: Apidae), with the Description of a New Species, pp. 1-28 in American Museum Novitates 2017 (3877) on page 12, DOI: 10.1206/3877.1, http://zenodo.org/record/4598486, {"references":["Pasteels, J. J., and J. M. Pasteels. 1974. Etude au microscope electronique a balayage des scopas abdominales chez de nombreuses especes d'abeilles (Apoidea, Megachilidae). Tissue and Cell 6 (1): 65 - 83."]}

Genus Tarsalia Morawitz, 1895 Tarsalia Morawitz, 1895: 9. Type species: Tarsalia hirtipes Morawitz, 1895, by monotypy. Friese, 1896: 211; Baker, 1998: 837; Michener, 2000: 666; Michener, 2007: 686. Ancyla (Tarsalia) Morawitz; Warncke, 1977: 58; Warncke, 1979: 192. DIAGNOSIS: As for tribe (above). COMMENTS: The genus currently includes two rather distinct groups, one with three species in northeastern Africa, Arabia, and Iran, while the other has five species in Western and Central Asia, south into India, and west on the Mediterranean islands of Cyprus and Sardinia (table 1). 2017 ENGEL ET AL.: ARABIAN TARSALIA 9 10 AMERICAN MUSEUM NOVITATES NO. 3877 Key to Subgenera of Tarsalia 1.Metasoma and mesosoma dark reddish brown to black (fig. 1); metatibial scopa open (fig. 2A); subtriangular patch of distinctive, squamiform setae on outer surface of metatibia apical to metabasitibial plate (Sardinia, Cyprus, Western and Central Asia, India)......................................................................... Tarsalia s. str. ���Metasoma and majority of mesosoma light castaneous to testaceous (figs. 3, 4); metatibial scopa densely closed (fig. 2B); outer surface of metatibia apical to metabasitibial plate without patch of distinctive setae, area similar to remainder of scopal setae (northeastern Africa, Arabia, Iran)...................................... Astibomelissa, n. subgen. closed (fig. 2B); outer surface of metatibia apical to metabasitibial plate without patch of distinctive setae, area similar to remainder of scopal setae. Metasomal tergum I with or without carina at angle of anterior- and dorsal-facing surfaces (all species of Tarsalia s. str. with carina, albeit sometimes weakly so)., Published as part of Engel, Michael S., Michener, Charles D. & Boontop, Yuvarin, 2017, Discovery of the Bee Tribe Tarsaliini in Arabia (Hymenoptera: Apidae), with the Description of a New Species, pp. 1-28 in American Museum Novitates 2017 (3877) on pages 8-10, DOI: 10.1206/3877.1, http://zenodo.org/record/4598486, {"references":["Morawitz, F. F. 1895. Beitrag zur Bienenfauna Turkmeniens. Horae Societatis Entomologicae Rossicae 29: 1 - 76.","Friese, H. 1896. Die Bienen Europa's (Apidae europaeae) nach ihren Gattungen, Arten und Varietaten auf vergleichend morphologisch-biologischer Grundlage. Theil II. Solitare Apiden. Genus Eucera. Berlin: Friedlander & Sohn, 216 pp.","Baker, D. B. 1998. Taxonomic and phylogenetic problems in Old World eucerine bees, with special reference to the genus Tarsalia Morawitz, 1895 (Hymenoptera: Apoidea: Anthophoridae). Journal of Natural History 32 (6): 823 - 860.","Michener, C. D. 2000. The bees of the world. Baltimore: Johns Hopkins University Press, xiv + [i] + 913 pp., + 16 pls.","Michener, C. D. 2007. The bees of the world, 2 nd ed. Baltimore: Johns Hopkins University Press, xvi + [i] + 953 pp., + 20 pls.","Warncke, K. 1977. Ideen zum naturlichen System der Bienen (Hymenoptera, Apoidea). Mitteilungen der Munchner Entomologischen Gesellschaft 67: 39 - 63.","Warncke, K. 1979. Beitrage zur Bienenfauna des Iran: 10. Die Gattung Ancyla Lep., mit einer revision der Bienengattung Ancyla Lep. Bolletino del Museo Civico di Storia Naturale di Venezia 30: 183 - 195."]}

Engel, Michael S., Michener, Charles D., Boontop, Yuvarin (2017): Discovery of the Bee Tribe Tarsaliini in Arabia (Hymenoptera: Apidae), with the Description of a New Species. American Museum Novitates 2017 (3877): 1-28, DOI: 10.1206/3877.1, URL: http://www.bioone.org/doi/10.1206/3886.1

Tarsalia (Astibomelissa) kindahensis Engel, new species Figures 4–6 DIAGNOSIS: Tarsalia kindahensis can be recognized by combination of largely pale castaneous to light testaceous integument on metasoma, legs, and sometimes large portions of mesosoma although nota and much of pleura darker (figs. 4, 6); outer surface of female probasitarsus with dense, long, suberect setae with sinuate or wavy apices; metasomal tergum I without transverse dorsal carina; female metasomal scopa with long, subdecumbant setae with sinuate or wavy apices; male tergum VII with medioapical produced margin weakly bilobate (fig. 5A); male sternum VI apically truncate and with medial tubercle (fig. 5B); male hidden sterna asymmetrical and of characteristic form (fig. 5C); left penis valve hypertrophied (figs. 5E, 5G); and unique possession of yellow facial maculation on female paraocular area and scape, in addition to labrum, clypeus, and supraclypeal area (fig. 6C). DESCRIPTION: MALE: Total body length 6.72 mm (7.14 mm); forewing length 4.70 mm (5.18 mm). Head wider than long, length 1.69 mm (1.94 mm), width 2.28 mm (2.34 mm); inner margins of compound eyes comparatively straight, parallel in lower two-thirds, slightly diverging in upper third, upper interorbital distance 1.31 mm (1.37 mm), lower interorbital distance 1.16 mm (1.22 mm); apex of clypeus slightly below lower tangent of compound eyes; clypeus weakly protuberant and convex; ocelli at upper tangent of compound eyes; lateral ocelli separated from median ocellus by about one-half median ocellar diameter; lateral ocellus separated from eye by about 1.6× lateral ocellar diameter; lateral ocellus separated from posterior of head by about its diameter; scape short, extending at most to lower border of median ocellus; FI slightly longer than apical width, longer than FII. Intertegular distance 1.41 mm (1.53 mm). Basal area of propodeum weakly defined, about as long as metanotum. Legs without special modifications; mesobasitarsus elongate, straight, spur extending to slightly basad basitarsal midlength; metafemur slightly thicker than mesofemur; metabasitarsus elongate. Forewing with basal vein basad cu-a by 2–3 times vein width; 1m-cu entering second submarginal cell; second submarginal cell shortest, narrowed toward anterior wing margin; 2rs-m arcuate. Hind wing with nine distal hamuli. Metasomal tergum I without carina at angle of anterior- and dorsal-facing surfaces (similar to T. mimetes); tergum VII gradulus medially pointed posteriorly, without lateral gradular teeth (sensu Baker, 1998), but lateral area instead angulate, apical margin projected, deflexed medially, margin shallowly and broadly emarginate (fig. 5A); sternum V with medioapical margin produced with broad, apically blunt lobe; sternum VI narrowed apically, medioapical margin truncate, with broadly concave, smooth ovoid areas laterally along margin producing narrow medial disc bearing a medial tubercle (fig. 5B); hidden sterna VII and VIII asymmetrical and as in figures 5C and 5D; genitalia as in figures 5E–G, with penis valves markedly asymmetrical (figs. 5E, 5G). Integument somewhat shining. Labrum with coarse, shallow, contiguous punctures, otherwise smooth. Clypeus with coarse, faint, shallow punctures separated by less than a puncture width, except slightly more spaced along borders and more oblique, approximately separated by a puncture width or sometimes slightly more, integument between punctures smooth. Supraclypeal area with smaller, more well-defined punctures, separated by a puncture width, except medially slightly more separated, integument otherwise smooth. Face below level of antennae with small, faint, scattered punctures, separated by less than a puncture width along border with clypeus and subantennal sulcus, then becoming sparser toward inner margin of compound eye such that paraocular margin impunctate, integument between punctures smooth. Face above level of antennae with small, well-defined punctures separated by less than a puncture width, integument between punctures faintly imbricate; punctures sparse in short, broad triangular patch immediate anterior to median ocellus and absent in paraocular margin where integument is strongly imbricate; ocellocular space medially as on upper face but with punctures absent along outer border of lateral ocellus and in paraocular margin bordering compound eye, former patch smooth, latter strongly imbricate; vertex as on upper face; gena as on upper face except punctures smaller and becoming gradually more widely spaced ventrally such that on lower gena punctures separated by 2–3 times a puncture width, integument between punctures smooth; postgena with minute punctures similar to those on lower gena except sparse, integument otherwise smooth. Pronotum with sparse, minute punctures, integument otherwise smooth. Mesoscutum with small, well-defined punctures separated by less than a puncture width, with some posteromedially separated by about a puncture width, integument between punctures; tegula generally smooth to faintly imbricate, with sparse, minute punctures; mesoscutellum with punctures as on mesoscutum except somewhat larger and coarser. Metanotum finely imbricate with scattered, shallow, faint punctures. Pleura with small, well-defined punctures, integument otherwise smooth except more imbricate in area of mesepisternal border with metepisternum; punctures on hypoepimeral area becoming oblique ventrally, nearly contiguous, except sparse along scrobal sulcus; punctures on preepisternal area smaller, becoming shallower, fainter, and oblique toward anterior margin; punctures on mesepisternum below scrobal sulcus becoming fainter along posterior border and on ventral-facing surface; metepisternum imbricate with sparse punctures. Basal area of propodeum largely obscured by setae, integument apparently imbricate and coarsely punctate; posterior and lateral surfaces of propodeum faintly imbricate to smooth, with scattered, small punctures. Metasomal terga with minute, well-defined punctures separated by less than a puncture width, except anterior-facing depression of tergum I largely smooth and narrow apical margins of terga smooth, integument between punctures smooth; sterna as on terga except punctures coarser and integument otherwise more imbricate. 2017 ENGEL ET AL.: ARABIAN TARSALIA 17 Coloration of head generally testaceous to black (lighter in holotype male, darker in paratype male), except labrum entirely yellow; mandible largely yellow except dark reddish brown to black apically; clypeus and supraclypeal area entirely yellow; paraocular area yellow below level of antennae and broadly along compound eye in lower half of upper face; paraocular area in upper half of face above antennae dark brown to testaceous; scape yellow, pedicel and flagellomeres I and II yellowish brown; remainder of flagellum orange. Pronotum dark reddish brown to testaceous (testaceous in holotype male, dark testaceous along borders blending to dark reddish brown posteriorly in paratype male). Mesoscutum and mesoscutellum dark testaceous to black; tegula semitranslucent; metanotum testaceous to dark reddish brown; pleura testaceous to dark reddish brown; basal area of propodeum dark testaceous to black, remainder of propodeum testaceous to dark reddish brown. Legs testaceous. Wing membranes hyaline, faintly parchment colored; veins yellowish brown to dark brown. Metasoma testaceous. 18 AMERICAN MUSEUM NOVITATES NO. 3877 Pubescence white and often obscuring integument on frons, vertex, gena, mesosoma, tibiae, tarsi, and metasomal terga where such setae usually appressed to decumbent and plumose (setae with numerous, minute branches along length of rachis, giving feathered appearance: similar to short-branched setal forms described by Saunders, 1878, and Braue, 1913), setae of mesoscutum not squamiform, typically as long as or slightly longer than ocellar diameter (setae abraded from large areas of mesosoma and anterior metasomal terga on paratype male, allowing view of integumental sculpture); labrum with transverse, medial patch of short, erect, branched setae; setae below level of antennae typically sparse, suberect to erect, long, and minutely branched, similar setae on postgena; mesosoma with setae less prominent on preepisternal area and posteroventrally on mesepisternum and anteroventrally on lateral propodeal surface; tergal setae largely short and appressed; sternal setae longer, suberect, and posteriorly directed. FEMALE: As described for male, except as follows: total body length 7.50 mm; forewing length 5.00 mm. Head wider than long, length 1.81 mm, width 2.44 mm; inner margins of compound eyes comparatively straight, slightly convergent below, upper interorbital distance 1.51 mm, lower interorbital distance 1.33 mm; lateral ocelli separated from median ocellus by about one-half median ocellar diameter; lateral ocellus separated from eye by about 2.0× lateral ocellar diameter; lateral ocellus separated from posterior of head by about its diameter; scape short, extending to slightly less than distance to lower border of median ocellus; FI elongate, length about twice apical width, about twice as long as FII. Intertegular distance 1.56 mm. Coloration as in darker male (above) (fig. 6). Pubescence as in male except for sex differences: probasitarsus with fine, erect to suberect setae with distinctly sinuate or wavy apices; metatibial and metabasitarsal scopa composed of dense, elongate, plumose setae; metasomal sterna II–V with rows of elongate, subdecumbent setae with sinuate or wavy apices, forming distinct scopa; sternum VI with medioapical patch of dense, short, erect setae. HOLOTYPE: ♂, Saudi Arabia: Riyadh, al-Amariah [al-‘ Amāriah, a locality about 12 km west of Riyadh], Majra [Mazra‘ah] al-Gasim [al-Qasim, farm], 23.v.2011 [23 May 2011], M.A. Hannan (KSMA). PARATYPES: 1♀, Saudi Arabia: Riyadh, al-Amariah [al-‘Amāriah], Majra [Mazra‘ah] alGasim [al-Qasim, farm], 19.vi.2011 [19 June 2011], M.A. Hannan & I. Naser (SEMC); 1♂, Saudi Arabia: Qassim [al-Qassim], Unizah [Unayzah], al-Watania Farm, Rowdah, 29.v.2013 [29 May 2013], M.A. Hannan; Pulicaria undulate [sic] [Pulicaria undulata] (SEMC). FLORAL RECORD: The paratype male was taken while visiting flowers of Pulicaria undulata (L.) C.A. Mey (Asteraceae: Asteroideae: Helianthodea: Inuleae), a rather common annual herb in the region, known as false fleabane or regionally as “Jethjath” and used in medicinal tea. Regional beekeepers consider P. undulata as a good source for pollen and nectar during the spring. ETYMOLOGY: The specific epithet refers to the ancient kingdom of Kindah, which occupied, among other areas, large portions of the Najd, encompassing the type localities. 2017 ENGEL ET AL.: ARABIAN TARSALIA 19 COMMENTS: There is variation in color of the head and mesosoma between the two males, the one from Qassim being darker and similar to the female from al-‘Amāriah, while the male from al-‘Amāriah is distinctly lighter. The lighter male is more testaceous in those areas where the Qassim male is dark testaceous to dark reddish brown, and dark testaceous to reddish brown in those places where the female and male from Qassim are dark reddish brown to black. However, despite these color differences, the integumental, structural, and genitalic traits are identical between the males. The male of T. kindahensis superficially resembles to some degree the male of the sympatric Tetraloniella (Tetraloniella) persiciformis Alqarni et al. (2012), a eucerine that has also been taken visiting flowers of P. undulata.

Astibomelissa Engel, new subgenus TYPE SPECIES: Tarsalia kindahensis Engel, new species. DIAGNOSIS: Body length approximately 5.5���9.0 mm; integument pale castaneous to testaceous on metasoma and majority of mesosoma (figs. 3���5, 6). Female metatibial scopa densely, Published as part of Engel, Michael S., Michener, Charles D. & Boontop, Yuvarin, 2017, Discovery of the Bee Tribe Tarsaliini in Arabia (Hymenoptera: Apidae), with the Description of a New Species, pp. 1-28 in American Museum Novitates 2017 (3877) on page 10, DOI: 10.1206/3877.1, http://zenodo.org/record/4598486

Astibomelissa Engel, new subgenus TYPE SPECIES: Tarsalia kindahensis Engel, new species. DIAGNOSIS: Body length approximately 5.5–9.0 mm; integument pale castaneous to testaceous on metasoma and majority of mesosoma (figs. 3–5, 6). Female metatibial scopa densely

Tarsalia (Astibomelissa) kindahensis Engel, new species Figures 4���6 DIAGNOSIS: Tarsalia kindahensis can be recognized by combination of largely pale castaneous to light testaceous integument on metasoma, legs, and sometimes large portions of mesosoma although nota and much of pleura darker (figs. 4, 6); outer surface of female probasitarsus with dense, long, suberect setae with sinuate or wavy apices; metasomal tergum I without transverse dorsal carina; female metasomal scopa with long, subdecumbant setae with sinuate or wavy apices; male tergum VII with medioapical produced margin weakly bilobate (fig. 5A); male sternum VI apically truncate and with medial tubercle (fig. 5B); male hidden sterna asymmetrical and of characteristic form (fig. 5C); left penis valve hypertrophied (figs. 5E, 5G); and unique possession of yellow facial maculation on female paraocular area and scape, in addition to labrum, clypeus, and supraclypeal area (fig. 6C). DESCRIPTION: MALE: Total body length 6.72 mm (7.14 mm); forewing length 4.70 mm (5.18 mm). Head wider than long, length 1.69 mm (1.94 mm), width 2.28 mm (2.34 mm); inner margins of compound eyes comparatively straight, parallel in lower two-thirds, slightly diverging in upper third, upper interorbital distance 1.31 mm (1.37 mm), lower interorbital distance 1.16 mm (1.22 mm); apex of clypeus slightly below lower tangent of compound eyes; clypeus weakly protuberant and convex; ocelli at upper tangent of compound eyes; lateral ocelli separated from median ocellus by about one-half median ocellar diameter; lateral ocellus separated from eye by about 1.6�� lateral ocellar diameter; lateral ocellus separated from posterior of head by about its diameter; scape short, extending at most to lower border of median ocellus; FI slightly longer than apical width, longer than FII. Intertegular distance 1.41 mm (1.53 mm). Basal area of propodeum weakly defined, about as long as metanotum. Legs without special modifications; mesobasitarsus elongate, straight, spur extending to slightly basad basitarsal midlength; metafemur slightly thicker than mesofemur; metabasitarsus elongate. Forewing with basal vein basad cu-a by 2���3 times vein width; 1m-cu entering second submarginal cell; second submarginal cell shortest, narrowed toward anterior wing margin; 2rs-m arcuate. Hind wing with nine distal hamuli. Metasomal tergum I without carina at angle of anterior- and dorsal-facing surfaces (similar to T. mimetes); tergum VII gradulus medially pointed posteriorly, without lateral gradular teeth (sensu Baker, 1998), but lateral area instead angulate, apical margin projected, deflexed medially, margin shallowly and broadly emarginate (fig. 5A); sternum V with medioapical margin produced with broad, apically blunt lobe; sternum VI narrowed apically, medioapical margin truncate, with broadly concave, smooth ovoid areas laterally along margin producing narrow medial disc bearing a medial tubercle (fig. 5B); hidden sterna VII and VIII asymmetrical and as in figures 5C and 5D; genitalia as in figures 5E���G, with penis valves markedly asymmetrical (figs. 5E, 5G). Integument somewhat shining. Labrum with coarse, shallow, contiguous punctures, otherwise smooth. Clypeus with coarse, faint, shallow punctures separated by less than a puncture width, except slightly more spaced along borders and more oblique, approximately separated by a puncture width or sometimes slightly more, integument between punctures smooth. Supraclypeal area with smaller, more well-defined punctures, separated by a puncture width, except medially slightly more separated, integument otherwise smooth. Face below level of antennae with small, faint, scattered punctures, separated by less than a puncture width along border with clypeus and subantennal sulcus, then becoming sparser toward inner margin of compound eye such that paraocular margin impunctate, integument between punctures smooth. Face above level of antennae with small, well-defined punctures separated by less than a puncture width, integument between punctures faintly imbricate; punctures sparse in short, broad triangular patch immediate anterior to median ocellus and absent in paraocular margin where integument is strongly imbricate; ocellocular space medially as on upper face but with punctures absent along outer border of lateral ocellus and in paraocular margin bordering compound eye, former patch smooth, latter strongly imbricate; vertex as on upper face; gena as on upper face except punctures smaller and becoming gradually more widely spaced ventrally such that on lower gena punctures separated by 2���3 times a puncture width, integument between punctures smooth; postgena with minute punctures similar to those on lower gena except sparse, integument otherwise smooth. Pronotum with sparse, minute punctures, integument otherwise smooth. Mesoscutum with small, well-defined punctures separated by less than a puncture width, with some posteromedially separated by about a puncture width, integument between punctures; tegula generally smooth to faintly imbricate, with sparse, minute punctures; mesoscutellum with punctures as on mesoscutum except somewhat larger and coarser. Metanotum finely imbricate with scattered, shallow, faint punctures. Pleura with small, well-defined punctures, integument otherwise smooth except more imbricate in area of mesepisternal border with metepisternum; punctures on hypoepimeral area becoming oblique ventrally, nearly contiguous, except sparse along scrobal sulcus; punctures on preepisternal area smaller, becoming shallower, fainter, and oblique toward anterior margin; punctures on mesepisternum below scrobal sulcus becoming fainter along posterior border and on ventral-facing surface; metepisternum imbricate with sparse punctures. Basal area of propodeum largely obscured by setae, integument apparently imbricate and coarsely punctate; posterior and lateral surfaces of propodeum faintly imbricate to smooth, with scattered, small punctures. Metasomal terga with minute, well-defined punctures separated by less than a puncture width, except anterior-facing depression of tergum I largely smooth and narrow apical margins of terga smooth, integument between punctures smooth; sterna as on terga except punctures coarser and integument otherwise more imbricate. 2017 ENGEL ET AL.: ARABIAN TARSALIA 17 Coloration of head generally testaceous to black (lighter in holotype male, darker in paratype male), except labrum entirely yellow; mandible largely yellow except dark reddish brown to black apically; clypeus and supraclypeal area entirely yellow; paraocular area yellow below level of antennae and broadly along compound eye in lower half of upper face; paraocular area in upper half of face above antennae dark brown to testaceous; scape yellow, pedicel and flagellomeres I and II yellowish brown; remainder of flagellum orange. Pronotum dark reddish brown to testaceous (testaceous in holotype male, dark testaceous along borders blending to dark reddish brown posteriorly in paratype male). Mesoscutum and mesoscutellum dark testaceous to black; tegula semitranslucent; metanotum testaceous to dark reddish brown; pleura testaceous to dark reddish brown; basal area of propodeum dark testaceous to black, remainder of propodeum testaceous to dark reddish brown. Legs testaceous. Wing membranes hyaline, faintly parchment colored; veins yellowish brown to dark brown. Metasoma testaceous. 18 AMERICAN MUSEUM NOVITATES NO. 3877 Pubescence white and often obscuring integument on frons, vertex, gena, mesosoma, tibiae, tarsi, and metasomal terga where such setae usually appressed to decumbent and plumose (setae with numerous, minute branches along length of rachis, giving feathered appearance: similar to short-branched setal forms described by Saunders, 1878, and Braue, 1913), setae of mesoscutum not squamiform, typically as long as or slightly longer than ocellar diameter (setae abraded from large areas of mesosoma and anterior metasomal terga on paratype male, allowing view of integumental sculpture); labrum with transverse, medial patch of short, erect, branched setae; setae below level of antennae typically sparse, suberect to erect, long, and minutely branched, similar setae on postgena; mesosoma with setae less prominent on preepisternal area and posteroventrally on mesepisternum and anteroventrally on lateral propodeal surface; tergal setae largely short and appressed; sternal setae longer, suberect, and posteriorly directed. FEMALE: As described for male, except as follows: total body length 7.50 mm; forewing length 5.00 mm. Head wider than long, length 1.81 mm, width 2.44 mm; inner margins of compound eyes comparatively straight, slightly convergent below, upper interorbital distance 1.51 mm, lower interorbital distance 1.33 mm; lateral ocelli separated from median ocellus by about one-half median ocellar diameter; lateral ocellus separated from eye by about 2.0�� lateral ocellar diameter; lateral ocellus separated from posterior of head by about its diameter; scape short, extending to slightly less than distance to lower border of median ocellus; FI elongate, length about twice apical width, about twice as long as FII. Intertegular distance 1.56 mm. Coloration as in darker male (above) (fig. 6). Pubescence as in male except for sex differences: probasitarsus with fine, erect to suberect setae with distinctly sinuate or wavy apices; metatibial and metabasitarsal scopa composed of dense, elongate, plumose setae; metasomal sterna II���V with rows of elongate, subdecumbent setae with sinuate or wavy apices, forming distinct scopa; sternum VI with medioapical patch of dense, short, erect setae. HOLOTYPE: ♂, Saudi Arabia: Riyadh, al-Amariah [al-��� Amāriah, a locality about 12 km west of Riyadh], Majra [Mazra���ah] al-Gasim [al-Qasim, farm], 23.v.2011 [23 May 2011], M.A. Hannan (KSMA). PARATYPES: 1♀, Saudi Arabia: Riyadh, al-Amariah [al-���Amāriah], Majra [Mazra���ah] alGasim [al-Qasim, farm], 19.vi.2011 [19 June 2011], M.A. Hannan & I. Naser (SEMC); 1♂, Saudi Arabia: Qassim [al-Qassim], Unizah [Unayzah], al-Watania Farm, Rowdah, 29.v.2013 [29 May 2013], M.A. Hannan; Pulicaria undulate [sic] [Pulicaria undulata] (SEMC). FLORAL RECORD: The paratype male was taken while visiting flowers of Pulicaria undulata (L.) C.A. Mey (Asteraceae: Asteroideae: Helianthodea: Inuleae), a rather common annual herb in the region, known as false fleabane or regionally as ���Jethjath��� and used in medicinal tea. Regional beekeepers consider P. undulata as a good source for pollen and nectar during the spring. ETYMOLOGY: The specific epithet refers to the ancient kingdom of Kindah, which occupied, among other areas, large portions of the Najd, encompassing the type localities. 2017 ENGEL ET AL.: ARABIAN TARSALIA 19 COMMENTS: There is variation in color of the head and mesosoma between the two males, the one from Qassim being darker and similar to the female from al-���Amāriah, while the male from al-���Amāriah is distinctly lighter. The lighter male is more testaceous in those areas where the Qassim male is dark testaceous to dark reddish brown, and dark testaceous to reddish brown in those places where the female and male from Qassim are dark reddish brown to black. However, despite these color differences, the integumental, structural, and genitalic traits are identical between the males. The male of T. kindahensis superficially resembles to some degree the male of the sympatric Tetraloniella (Tetraloniella) persiciformis Alqarni et al. (2012), a eucerine that has also been taken visiting flowers of P. undulata., Published as part of Engel, Michael S., Michener, Charles D. & Boontop, Yuvarin, 2017, Discovery of the Bee Tribe Tarsaliini in Arabia (Hymenoptera: Apidae), with the Description of a New Species, pp. 1-28 in American Museum Novitates 2017 (3877) on pages 16-19, DOI: 10.1206/3877.1, http://zenodo.org/record/4598486, {"references":["Baker, D. B. 1998. Taxonomic and phylogenetic problems in Old World eucerine bees, with special reference to the genus Tarsalia Morawitz, 1895 (Hymenoptera: Apoidea: Anthophoridae). Journal of Natural History 32 (6): 823 - 860.","Saunders, E. 1878. Remarks on the hairs of some of our British Hymenoptera. Transactions of the Entomological Society of London 1878 (2): 169 - 172, + 1 pl.","Braue, A. 1913. Die Pollensammelapparate der beinsammelnden Bienen. Jenaische Zeitschrift fur Naturwissenschaft 50 (1): 1 - 96, + 4 pls.","Alqarni, A. S., M. A. Hannan, and M. S. Engel. 2012. A new wild, pollinating bee species of the genus Tetraloniella from the Arabian Peninsula (Hymenoptera, Apidae). ZooKeys 172: 89 - 96."]}

Key to Species of Astibomelissa 1.Mesoscutum with numerous appressed to decumbent, plumose setae (usually missing on central disc, presumably due to abrasion), setae longer, not squamiform; outer surface of probasitarsus with fine, erect to suberect setae with distinctly sinuate or wavy apices (= type IV of Pasteels and Pasteels, 1974); inner setae of metabasitarsus white to pale yellow and similar to setae of scopa; metasomal tergum I without transverse carina at angle of anterior- and dorsal-facing surfaces; smaller species, 5.5–7.5 mm in total length (Saudi Arabia; northeastern Africa).................................................... 2 —Mesoscutum tomentose, with dense covering of appressed, minute (typically about 0.50× ocellar diameter), squamiform setae; outer surface of probasitarsus with fine, subdecumbent setae tapering to straight apices; inner setae of metabasitarsus fuscous, contrasting with surrounding white to pale yellow setae of scopa; metasomal tergum I with transverse carina at angle of anterior- and dorsal-facing surfaces; larger species, 7.9–8.8 mm in length [Iran]............................................... T. persica (Warncke) 2.Female with yellow facial maculation confined to labrum, clypeus, and supraclypeal area (fig. 3C); metasomal scopa dense, with setae greatly elongate and subdecumbent to suberect; small species, approximately 5.5–6.5 mm in length (male unknown) [Egypt, Sudan].................................................... T. mimetes (Cockerell) —Female with yellow facial maculation extending to paraocular area and scape (fig. 6C); metasomal scopa not so dense, with setae long and largely decumbent to subdecumbent; moderate-sized species, approximately 6.7–7.5 mm in length (central Saudi Arabia)........................................................... T. kindahensis Engel, n. sp.

Engel, Michael S., Alqarni, Abdulaziz S., Hannan, Mohammed A., Hinojosa-Díaz, Ismael A., Michener, Charles D. (2014): Allodapine Bees in the Arabian Peninsula (Hymenoptera: Apidae): A New Species of Braunsapis from the Sarawat Mountains, with an Overview of the Arabian Fauna. American Museum Novitates 2014 (3801): 1, DOI: 10.1206/3801.1, URL: http://www.bioone.org/doi/abs/10.1206/3801.1

Engel, Michael S., Gonzalez, Victor H., Hinojosa-Díaz, Ismael A., Michener, Charles D. (2013): Introducing the Journal of Melittology: An outlet for disseminating bee research and raising melittological awareness. Journal of Melitology 2013 (1): 1-3, DOI: 10.17161/jom.v0i1.4433, URL: https://doi.org/10.17161/jom.v0i1.4433

Engel, Michael S., Grimaldi, David A., Gonzalez, Victor H., Hinojosa-Díaz, Ismael A., Michener, Charles D. (2012): An Exomalopsine Bee in Early Miocene Amber from the Dominican Republic (Hymenoptera: Apidae). American Museum Novitates 2012 (3758): 1-16, DOI: 10.1206/3758.2, URL: http://www.bioone.org/doi/abs/10.1206/3758.2

Anthophorula (Anthophorula) persephone Engel, new species Figures 1���4 DIAGNOSIS: The new species can be most readily distinguished from other Anthophorula proper by the broader pterostigma in which its inner breadth is greater than that of its marginal veins (fig. 4). In addition, the species can be characterized by the combination of a uniformly punctate mesoscutum and mesoscutellum, similarly punctate pleura, absence of distinct setal bands on the metanotum, and the pygidial fimbria composed of dense fuscous setae with a reddish tint. DESCRIPTION: Female. Total body length (as preserved) 6.1 mm; forewing length 4.6 mm; integument dark (fig. 1), apparently black throughout except legs dark brown, with spurs and pretarsal claws lighter brown, maculations absent; wing membranes hyaline, veins dark brown. Head broader than long (width as preserved 2.08 mm, length 1.30 mm) (fig. 2), with inner orbits of compound eyes only slightly diverging above; clypeus weakly convex; gena rounded, narrower than compound eye; lateral ocellus separated from occipital margin by approximately 1.5 times its diameter and from compound eye margin by about three times its diameter (values approximate as the ocelli are slightly distorted, making it difficult to determine their proper diameter in life); toruli situated well below midlength of face. Intertegular distance 1.46 mm; mesoscutellum more than twice length metanotum; metabasitibial plate short, broadly rounded, with planar surface, without carinate rim or covering of fine setae (fig. 3); scopa on metatibia and metabasitarsus composed of dense, plumose setae (fig. 3); metabasitarsus with apical process and penicillus. Forewing marginal cell obliquely truncate and bent away from costal margin (fig. 4); marginal cell apex somewhat pointed (fig. 4), length of cell slightly greater than distance from its apex to wing tip; pterostigma three times as long as prestigma, its inner breadth greater than that of bordering marginal veins; second medial cell shorter than second cubital cell; other venational details in figure 4. Anteriorfacing surface of first metasomal tergum depressed and apparently only slightly shorter than length of dorsal-facing surface; first metasomal tergum with distinct, transverse carina demarcating anterior- and dorsal-facing surfaces; pygidial plate black, apparently broad basally with relatively straight, apically tapering lateral margins to narrowly rounded apex (very difficult to discern). Face with small, well-defined punctures separated by a puncture width or less, integument between smooth and shining; frontal line weakly impressed, disappearing approximately two ocellar diameters before median ocellus. Mesoscutum uniformly with punctures separated by a puncture width or less, often only slightly less, integument between punctures smooth and shining; mesoscutellum sculptured as on mesoscutum; pleura sculptured as on mesoscutum. Metasomal terga with ill-defined, coarse, shallow punctures separated by less than a puncture width (except anterior-facing surface of first tergum apparently impunctate), such punctures slightly more defined along lateral margins, integument between finely imbricate and shining. Pubescence appearing largely fuscous or lightly fuscous with reddish tint except those setae of face whitish; setae of mesosoma branched and numerous, not dense or obscuring integument. Scopal setae elongate, fuscous, plumose throughout and along entire rachis. Metasomal terga without dense transverse bands of setae, setae most prominent laterally on terga; pygidial fimbria dense, composed of fuscous branched setae with reddish tint. Male. Unknown. HOLOTYPE: Female (fig. 1), DR-KL1; Dominican Republic; Early Miocene amber (Burdigalian?), specific mine unknown (AMNH). ADDITIONAL MATERIAL: Fragmentary female, same amber piece as holotype (AMNH DR-KL1). This individual is largely represented by the metasoma, fragments of the hind and midlegs, a very few fragments of one foreleg, and the wings which are folded over the metasoma and somewhat fractured (fig. 4). ETYMOLOGY: The specific epithet is from Greek mythology. Persephone, daughter of Zeus and Demeter, was abducted by Hades, who was later forced to return her. Before she left, Hades tricked her into eating four pomegranate arils thereby dooming her eternally to the underworld. Persephone was therefore bound to spend four months of each year with Hades as his consort., Published as part of Engel, Michael S., Grimaldi, David A., Gonzalez, Victor H., Hinojosa-D��az, Ismael A. & Michener, Charles D., 2012, An Exomalopsine Bee in Early Miocene Amber from the Dominican Republic (Hymenoptera: Apidae), pp. 1-16 in American Museum Novitates 2012 (3758) on pages 5-7, DOI: 10.1206/3758.2, http://zenodo.org/record/5360852