Revue suisse de Zoologie 114 (3): 471-505; septembre 2007 A review ofthe catfish genus Hara, with the description of four new species (Siluriformes: Erethistidae) Heok Hee NG1 & Maurice KOTTELAT2 1 Fish Division, Museum ofZoology, University ofMichigan, 1109 Geddes Avenue, AnnArbor, Michigan 48109-1079, USA. Current address: Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore, 6 Science Drive 2, #03-01, Singapore 117546. Email: [email protected] 2 Case postale 57, Cornol, CH-2952, Switzerland. Email: [email protected] A review of the catfish genus Hara, with the description of four new species (Siluriformes: Erethistidae). - Species of the erethistid catfish genus Hara are reviewed in this study. Eight species are recognized: Hara filamentosa, H. hara, H. horai, H.jerdoni, H. longissima, H. mesembrina, H.minuscula,andH. spinulus,ofwhich the latterfourare described as new herein.Erethistesmaesotensisis ajuniorsynonymofHarafilamentosa,and H. saharsai and H. serrata junior synonyms of H. hara. A neotype is designated forH.filamentosa Blyth, 1860 and a lectotype is designated for Hara horai Misra, 1976. Keywords: Sisoroidea - Bangladesh - India - Myanmar - Thailand. INTRODUCTION The Erethistidae are a small family of sisoroid catfishes inhabiting hillstreams in the subhimalayan region and western Indochina. The erethistid catfish genus Hara consistsofsmalltomediumsizedcatfishes withabrowncrypticcolorationandheavily tuberculate skin. Species ofHara are diagnosed from other members ofthe family in having an anteroventral flange on the ventral surface ofthe complex centrum and the parapophysis ofthe fifth vertebra with a vertical lamina (de Pinna, 1996). The genus has been recorded from the Ganges-Brahmaputra river drainages eastwards to the Salween River drainage in western Thailand (Hora, 1950; Kottelat, 1989; Ukkatawewat & Vidthayanon, 1998). Recently, the authors obtained material from southern Thailand that consider- ablyextends the southernmostextentofthe distributionofHara. Ourattempts to iden- tify this material prompted us to investigate the identities ofspecimens collected from India,Myanmar andThailand. The following study presents the results ofthis investi- gation, which reviews the species ofHara. We report here eight species ofHara, of which four are described as new. WereceivedThomson & Page's (2006) list ofgeneric and species names in the families Erethistidae and Sisoridae after the present work was completed. A compila- Manuscript accepted07.03.2007 472 H.H.NG &M. KOTTELAT tion ofsuch a nature would make sense as the completion ofa revision ofthe various genera and species (but not before). Thomson & Page treatHara as ajunior synonym ofErethistes. In the discussion below, we mention the results of a phylogenetic ana- lysis based on both morphological and molecular data that make it premature to formally synonymise these genera. This is discussed in greater detail in HHN's doctoral dissertation (Ng, 2006). Thomson & Page's list ofspecies and synonymies of Hara are apparently compilations, and do not affect our conclusions below (based on our studies ofaconsiderable number ofspecimens). MATERIALS AND METHODS Measurements weremade pointtopoint withdial callipers anddatarecordedto 0.1 mm. Counts and measurements were made on the left side ofspecimens whenever possible. Subunits ofthe head are presented as proportions ofhead length (HL). Head length itself and measurements of body parts are given as proportions of standard length (SL). Measurements and counts were made following Ng & Kottelat (1998) withthefollowingaddition: the lengthoftheposteriorprocessoncoracoidismeasured from the anterioredge ofthe coracoid where the process is situated to the posteriortip ofthe process. Fin rays were counted under a binocular dissecting microscope using trans- mitted light. Vertebrae were counted from radiographs obtained with a Hewlett Packard Faxitron 43855B cabinet x-ray machine. Vertebral counts and terminology follow those of Roberts (1994). Material for this study is deposited in the following collections: Natural History Museum, London (BMNH), California Academy of Sciences, San Francisco (CAS), collection of the second author, Cornol (CMK), Muséum d'histoire naturelle Genève (MHNG), Naturhistoriska Riksmuseet, Stockholm (NRM), Museum of Zoology, University of Michigan, Ann Arbor (UMMZ), and the Zoological Reference Collection of the Raffles Museum of Biodiversity Research, Singapore (ZRC). RESULTS Harafilamentosa Blyth, 1860 Fig. 1 Harafilamentosa Blyth, 1860: 152 (type locality: Tenasserim) Erethistes conta (non Hamilton): Vinciguerra, 1890: 250. Erethistes maesotensis Kottelat, 1983: 71, Fig. 1 (type locality: Mae Nam Moei, 5 km west of Mae Sot.Thailand) Material examined: CMK 11961 (2), 48.9-53.1 mm SL; CMK 14689 (9), 42.1-44.4 mmSL;Thailand:Takprovince,MaeNamMoeiatBanWaLe.CMK 12175 (2),44.6-47.0mm SL; ZRC 42223 (1),46.3 mm SL; Thailand: Tak province, stream at km 57 on road from Mae SottoWaLe (5 kmbeforeWaRei).ZRC41269 (1),44.3 mmSL;Thailand: Takprovince,Mae sNias)m, 1M9o.7eimmat S1L6;1T7h'a4il7a.n1d":NT9a8k°4p2r'o4v4i.nc9e",E.MaMeHNNaGm2M1o5e0i.,55(1kmpaWratoyfpeMaofeESroett.hiZsRteCs5ma0e6s4o7te(1n)-, 43.1 mm SL; Thailand: Tak province, Mae Nam Moei at Mae Sot (border with Myanmar), CMK mm MHNG 16°41'13.2"N 98°30'54.0"E. 14640 (11),32.1-48.6 SL; 2680.076 (3),40.2- mm CMK mm MHNG mm 48.1 SL; 15102 (17), 20.6-30.1 SL; 2680.075 (5), 24.1-27.2 SL; Thailand: Tak province. Huai Mae Lamao, about 2 km N ofroad 105 on road from Ban Mae Lamao to Ban Mae Ramat. CMK 5529 (1), 48.3 mm SL; Thailand: Tak province. Huai Mae Lamao at Ban Mae Lamao. UMMZ 197583 (1), 47.4 mm SL; Thailand: Tak province, Mae AREVIEWOFTHECATFISH GENUS HARA 473 Fig. 1 Harafilamentosa,neotype,MHNG2687.038,43.7 mmSL; Myanmar: MegathatChaung.Scale bar: 25 mm. 474 H H NG&M KOTTELAT - - - Lamao, tributary of Salween River. CMK 14769 (1), 55.2 mm SL; Thailand: Mae Hong Son province,HuaiHai,about5kmfromMaeHongSontoPairoad,onroadturningoffnorthatkm 44.CMK 16186(5),41.4-52.4mmSL;CMK 17951 (2),44.8-62.7mmSL;Thailand:MaeHong Songprovince,SalweenRiveratMae SamLeap.CMK 16198 (1),45.3 mmSL;Thailand: Mae Hong Son province, Nam Mae Kong Ka, 4-8 km upstrean ofconfluence with Salween River. CMK 16227 (1),46.9 mm SL; Thailand: Mae Hong Son province, Salween River about 1 km upstream ofBan Mae Sam Leap. BMNH 1893.2.16.12-14 (3),49.4-59.1 mm SL. - Myanmar: Meetan (=Mitan Chaung, a rivulet flowing S from the summit of Mulayet Taung, 16°H'N MHNG mm CMK 98°32'E,intheSalweenbasin). 2687.038,neotype,43.7 SL; 17786(4),46.8- 59.7mmSL;CMK 17951 (2),45.1-61.2mmSL;Myanmar: Kayinstate,AtaranRiverdrainage, stream"Chon Son"betweenKyondaw andPhadaw,about20kmNWofPayathouzu (atborder with Thailand), 15°25'N 98°15'E. UMMZ 243692 (6), 46.4-50.0 mm SL; Myanmar: from aquariumtrade.UMMZ245962 (1),53.7 mmSL; Myanmar: Bagodivision,Pyutownship,Pyu stream(tributaryofSittangRiver) ca. 229kmfromYangon, 18°29'N96°26'E.UMMZ245964 (9),56.7-67.1 mmSL; Myanmar: Kayin state,hillstreams inAtaranRiverdrainage inthevicin- ity ofPayathonzu. Diagnosis: Hara filamentosa can be distinguished from congeners (except H. mesembrina and H. longissima) in sometimes having (vs. always lacking) a fila- mentous extension to the first principal ray ofthe upper caudal-fin lobe; it is also dis- tinguished from all congeners exceptH. hara andH. longissima in having a shortpos- terior process on coracoid (reaching to midway ofdistance between bases ofpectoral spine and first pelvic-fin ray vs. two thirds orbeyond; length 19.9-22.8% SLvs. 23.5- 38.3). ItdiffersfromH. longissima inhaving ashorter(length 14.9-17.8% SLvs. 18.8- 21.3) and deeper(depth 6.1-7.2% SLvs. 5.0-6.1) caudal peduncle (depth 2.1-2.9times in its length for H.filamentosa vs. 3.1-3.9 times in H. longissima) and deeper body (depth 13.9-19.2% SL vs. 12.5-14.5). Harafilamentosa can be further distinguished from H. hara in having a longer adipose-fin base (15.7-21.7% SL vs. 11.8-15.6), deeper caudal peduncle (depth 6.1-7.2% SLvs. 5.0-5.8) and lacking (vs. presence) of serrations on the anterior edge of the dorsal spine, from H. horai in having a deeper caudal peduncle (depth 6.1-7.2% SLvs. 5.2-6.2) and a shorteradipose-finbase (length 15.7-21.7% SLvs. 10.0-13.7) andhead (length26.5-29.1% SLvs. 30.3-34.2),from//. jerdoni in having a shorter pectoral spine (length 23.3-29.9% SLvs. 42.2-51.7), more soft pectoral-fin rays (total number of soft fin rays modally 7 vs. 5), more vertebrae (31-34 vs. 27-29), the supraoccipital spine not reaching (vs. reaching) the anterior nuchal plate and adeeply forked (vs. emarginate) caudal fin. Itdiffers fromH.mesem- brina in having a shorter caudal peduncle (length 14.9-17.8% SLvs. 17.2-18.9), from H. minuscula in having more soft pectoral-fin rays (total number of soft fin rays modally 7 vs. 5) and the supraoccipital spine not reaching (vs. reaching) the anterior nuchal plate, and fromH. spinulus in having a longerdorsal spine (length 20.4-27.6% SL vs. 14.7-19.8), adipose-fin base (length 15.7-21.7% SL vs. 12.9-16.1) and caudal peduncle (length 14.9-17.8 % SLvs. 11.8-15.0). Description: Morphometric data in Table 1. Head and body moderately compressed. Dorsal profile curved from tip of snout to level of nares, then evenly sloping to origin of dorsal fin; evenly sloping ventrally to origin of adipose fin and gentlyconcavefromoriginofadipose fintoendofcaudalpeduncle.Ventralprofile flat to pelvic-fin base; sloping gently dorsally to end of anal-fin base and gently concave from end of anal-fin base to end of caudal peduncle. Anus and urogenital openings AREVIEWOFTHECATFISH GENUS HARA 475 Table 1. Morphometric data forHarafilamentosa (n=25: 43.7-67.1 mm SL). Range Mean±SD %SL Predorsal length 42.1-46.7 44.3±1.44 Preanal length 69.1-72.9 70.4±1.14 Prepelvic length 49.0-52.3 51.0+1.04 Prepectoral length 21.3-27.8 24.0±1.66 Length ofdorsal finbase 17.2-20.7 18.9±1.16 Dorsal spine length 20.4-27.6 23.6±2.48 Anal fin length 13.0-17.2 14.8+1.10 Pelvic fin length 16.0-19.6 17.2+1.20 Pectoral fin length 25.6-32.4 28.8±2.16 Pectoral spine length 23.3-29.9 27.0±1.99 Caudal fin length 20.9-30.3 26.7±2.80 Length ofadipose finbase 15.7-21.7 18.7+1.99 Dorsalto adipose distance 5.1-11.0 7.2±1.96 Post-adiposedistance 14.1-18.3 16.5±1.50 Caudal peduncle length 14.9-17.8 16.7±0.88 Caudal peduncle depth 6.1-7.2 6.8±0.37 Bodydepth atanus 13.9-19.2 16.9+1.65 Head length 26.5-29.1 28.0±0.91 Head width 21.0-27.0 24.7±1.75 Headdepth 19.1-24.3 21.5+1.54 Lengthofposteriorprocess on coracoid 19.9-22.8 21.3+1.04 %HL Snoutlength 50.0-60.7 54.9+3.60 Interorbital distance 32.2-41.4 37.3±3.04 Eye diameter 12.5-15.5 13.7+1.21 Nasal barbel length 12.6-22.1 17.0±3.59 Maxillarybarbel length 57.5-94.4 85.0±10.66 Innermandibularbarbel length 31.0-45.4 38.1±4.47 Outermandibularbarbel length 37.4-61.3 52.5±8.00 located at vertical through posterior three quarters ofadpressed pelvic fin. Skin tuber- culate, tubercles distributed evenly throughout head and body. Lateral line complete andmidlateral. Vertebrae 14+17=31 (1), 14+18=32 (2), 15+17=32 (2), 14+19=33 (1), 15+18=33 (5), 16+17=33 (4), 15+19=34 (2) or 16+18=34 (7). Snout narrow; profile dorsally rounded when viewed laterally and acutely triangular when viewed from above. Snout with pronounced steepening of curvature anteriorlyatlevelofposteriornareswhenviewedlaterally.Anteriorandposteriornares large and separated by flap of skin comprising base of nasal barbel. Gill openings narrow, extending from immediately ventral to posttemporal to immediately lateral to pointimmediately ventraltobaseofpectoral spine. Bony elements ofdorsal surface of head covered with thin, tuberculate skin and very prominent. Supraoccipital spine not reaching anterior nuchal plate. Weberian lamina well developed, approximately same length as supraoccipital spine andextending parallel toeither side ofspine. Eye ovoid, horizontal axis longest; located entirely in dorsal halfofhead, with faint dorsal ridge on frontal bone dorsal to eye. Orbit with free margin. 476 H.H. NG & M. KOTTELAT Barbels in fourpairs.Maxillarybarbel long and slender,extending to middle of pectoral-spine base. Nasal barbel short, subtended by flap ofskin at base and extend- ing to midway between posterior edge ofposterior nares and anterior margin oforbit. Inner mandibular-barbel origin close to midline, extending to middle of distance between its base and that of pectoral spine. Outer mandibular barbel originating posterolateralofinnermandibularbarbel,extending to aboutthreequarters ofdistance between its base and that ofpectoral spine. Mouth inferior, premaxillary tooth band not exposed when mouth is closed. Upperlip with indistinctplicae. Oral teeth small and villiform, in irregularrows on all tooth-bearing surfaces. Premaxillary teeth in single crescentic band across midline. Palate edentulous. Dentary teeth in two narrow crescentic bands separated at midline. Dorsal fin located above anterior third of body, with 1,5,i (25) rays; dorso- posterior fin margin straight. Dorsal spine moderately long and gently curved; with smooth anteriormargin and posteriormargin with4-7 serrae.Adipose fin with anterior margin straight and posterior margin angular. Caudal fin strongly forked, with i,7,7,i (1), i,7,8,i (23) ori,8,7,i (1) principal rays; upper lobe slightly longerthan lower. First principal ray of upper caudal-fin lobe produced into a filament extending posteriorly forlengthofuppercaudal lobeproper.Procurrentrays symmetricalandextendingonly slightly anteriorto fin base.Anal-fin base approximately in line with adipose-fin base. Anal fin with convex margin anteriorly and straight margin posteriorly, with iii,6,i (3), iv,6 (2) iv,6,i (8), iii,7 (2), iii,7,i (5),v,6 (1), iv,7 (1) or iv,7,i (3) rays. Pelvic-fin origin anteriorto vertical through posteriorend ofdorsal-fin base. Pelvic fin with pointed tip and straightposteriormargin,withi,5 (25) rays; tipofadpressed finjustreaching anal- fin origin. Pectoral fin with 1,5,i (3) or 1,6,i (22) rays; fin margin gently convex posteriorly; anterior spine margin with 14-23 small distally directed serrations, posterior margin with 9-13 large medially directed serrations. Coracoid with well developed posterior processes, extending to midway between base of posteriormost pectoral-fin ray and pelvic-fin origin. Colour: Head and body dark chestnut brown to light chocolate brown, with a mottling caused by numerous smaller, irregular darker brown patches. Belly and ven- tral surfaces of head light brown. Two thin, irregular light brown bars on body: first chevron-shapedandlocatedbetweendorsal-andadipose-finbases,secondthickerthan first and located on caudal peduncle. Dorsal fin dark mottled brown, except for thin hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and either a small ovoid hyaline spot or a crescentic median hyaline band on middle third of fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. Adipose fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, with irregularW-shaped band subdistally and with small dark brown flecks randomly distributed throughout fin. Two small light brown patches present at bases ofcaudal- finlobes in some individuals.Maxillary andmandibularbarbels lightbrown,withdark brown annuii. Distribution.HarafilamentosaisknownfromtheAtaran,Salween andSittang riverdrainages (Fig. 2).Inthe Salween,ithasbeencollectedinthe Salweenproperand the Mae Nam Moei. in Tak and Mae Hong Son Provinces ofThailand. AREVIEWOFTHECATFISH GENUS HARA All •O Harafilamentosa +* Harahara ** Harahorai Harajerdoni D Haralongissima V Haramesembrina AA Haraminuscula X» Haraspinulus 5 a 80 500 1000 kilometers Fig. 2 Map showingcollecting localities ofHara examined.Outlined symbols are type localities. Hara hara (Hamilton 1822) Fig.3 PimelodusharaHamilton, 1822: 190 (type locality: Nathpur,India) Pimelodus asperus M'Clelland, 1844: 404, pi. 24 fig. 2 (type locality: Hooghly River S of Ranaghat) 478 H. H. NG & M. KOTTELAT AREVIEWOFTHECATFISH GENUS HARA 479 Hara saharsai Datta Munshi & Srivastava, 1988: 265, fig. 5 (type locality: Kosi Belt, northern Bihar,India) Hara serratus Vishwanath & Kosygin, 2000: 143, figs 2-5 (type locality: Jiri River at Jiribam, Manipur,India) UMMZ mm W Material examined: 244697 (neotype), 47.2 SL; India: Bengal, Hooghly River S ofRanaghat. UMMZ 208748 (44), 33.0-50.0 mm SL; Bangladesh: Gowain RiveratGowainghat. Diagnosis: Hara hara can be distinguished from congeners in possessing (vs. lacking) serrations on the anterior edge of the pectoral spine. It further differs from H.filamentosa, H. mesembrina and H. longissima in always lacking (vs. sometimes having) a filamentous extension to the first principal ray ofthe upper caudal-fin lobe; itis also distinguished from all congeners except H.filamentosa and H. longissima in having a short posteriorprocess on coracoid (reaching to midway ofdistance between bases of pectoral spine and first pelvic-fin ray vs. two thirds or beyond; length 19.9- 24.8% SL vs. 23.5-38.3). Hara hara further differs from H filamentosa in having a . shorter adipose-fin base (11.8-15.6% SLvs. 15.7-21.7) and slenderer caudal peduncle (depth 5.0-5.8% SL vs. 6.1-7.2), from H. horai in having a longer caudal fin (25.8- 28.0% SL vs. 20.5-23.5), slenderer body (11.5-14.3% SL vs. 15.1-17.9) and a shorter head (length 27.2-30.9% SL vs. 30.3-34.2), from H.jerdoni in having a shorter pec- toral spine (length 26.9-33.3% SL vs. 42.2-51.7), more soft pectoral-fin rays (total numberofsoftfinraysmodally7vs.5),morevertebrae(31-34vs.27-29),the supraoc- cipital spine notreaching (vs. reaching) the anterior nuchal plate, and a deeply forked (vs.emarginate) caudal fin.Itfurtherdiffers fromH.mesembrina inhaving aslenderer caudal peduncle (depth 5.0-5.8% SL vs. 6.0-7.4), from H. minuscula in having a narrower head (width 22.4-24.3% SL vs. 24.5-27.2), more soft pectoral-fin rays (total numberofsoft fin rays modally 7 vs. 5) and the supraoccipital spine notreaching (vs. reaching) the anterior nuchal plate, and from H. spinulus in having a longer dorsal spine (22.7-27.4% SL vs. 14.7-19.8), narrower head (width 22.4-24.3% SL vs. 25.9- 30.0) and slenderercaudal peduncle (depth 5.0-5.8% SLvs. 6.7-8.5). Description: Morphometric data in Table 2. Exactly as in Harafilamentosa, except for the following. Vertebrae 14+18=32 (7), 15+17=32 (4), 14+19=33 (1), 15+18=33 (9) or 15+19=34(3).Dorsalfinwith1,5,i(24)rays.Dorsal spinemoderately long and gently curved; with serrated anterior margin and posterior margin with 8-13 serrae.Caudal fin with i,7,6,i (24) principalrays.Firstprincipalray ofuppercaudal-fin lobe notproduced into a filament.Anal fin with iv,6,i (24) rays. Pelvic fin with i,5 (24) rays. Pectoral fin with 1,6,i (24) rays; anterior spine margin with 17-24 small distally directed serrations, posterior margin with 12-17 large medially directed serrations. Coracoid with well developed posteriorprocesses,extending to midway between base ofposteriormost pectoral-fin ray and pelvic-fin origin. Colour: Head and body chestnut brown or light brown, with a mottling caused by numerous smaller, irregular darker brown patches. Belly and ventral surfaces of head light brown. Two thin, irregular light brown bars on body: first chevron-shaped andlocatedbetweendorsal- andadipose-finbases,secondthickerthan firstandlocated Fig. 3 Harahara,neotype,UMMZ244697,47.2mmSL;India:HooghlyRiveratRanaghat.Scalebar: 25 mm. 480 H. H. NG & M. KOTTELAT Table2.Morphometric dataforHara hara (n=24: 33.0-50.0 mmSL). Range Mean±SD %SL Predorsal length 44.0-45.5 44.6±0.64 Preanal length 66.9-70.9 68.7+1.54 Prepelvic length 47.6-53.6 51.1+2.28 Prepectoral length 21.2-27.3 24.4±2.22 Lengthofdorsal finbase 16.7-21.0 18.6±1.60 Dorsal spine length 22.7-27.4 24.5±1.88 Anal fin length 14.6-16.8 15.6±0.82 Pelvic fin length 15.8-18.2 16.8±0.91 Pectoral fin length 30.0-36.2 32.5+2.40 Pectoral spine length 26.9-33.3 30.0±2.32 Caudal fin length 25.8-28.0 27.0±1.01 Length ofadipose finbase 11.8-15.6 14.0±1.46 Dorsal to adiposedistance 10.8-12.7 12.0±0.71 Post-adipose distance 15.7-18.6 17.2+1.27 Caudal peduncle length 14.6-19.4 17.7+1.85 Caudal peduncle depth 5.0-5.8 5.4±0.34 Body depth atanus 11.5-14.3 12.8+1.28 Head length 27.2-30.9 29.6±1.41 Headwidth 22.4-24.3 23.5±0.91 Headdepth 18.6-20.1 19.3+0.61 Length ofposteriorprocess on coracoid 19.9-24.8 22.0±1.79 %HL Snout length 50.0-54.7 51.8+1.87 Interorbital distance 31.5-38.0 35.4±2.53 Eye diameter 9.4-10.9 10.1±0.65 Nasal barbel length 14.8-21.9 18.0±3.07 Maxillary barbel length 80.8-108.2 93.4±9.78 Innermandibularbarbel length 34.2-49.6 43.6±6.13 Outermandibularbarbel length 52.1-67.2 60.8±6.13 on caudal peduncle. Dorsal fin dark mottled brown, except for thin hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and either a small ovoid hyaline spot or a crescentic median hyaline band on middle third of fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. Adipose fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, with irregular W- shaped band subdistally and with small dark brown flecks randomly distributed throughout fin. Two small light brown patches present at bases ofcaudal-fin lobes in some individuals. Maxillary and mandibular barbels light brown, with dark brown annuii. Distribution: Hara hara is known from the Brahmaputra and Ganges river drainages (Fig. 2). Hara horai Misra, 1976 Fig.4 Hara horai Misra, 1976: 245, PL 9 figs 1-3 (type locality: Terai and Duars, northern Bengal, India) Materialexamined:ZSIFF955 (lectotype),58.8mmSL;India:WestBengal,Teraiand Duars.UMMZ245633 (2),56.2-73.1 mmSL;ZRC50578 (2),41.8-62.7mmSL; India:Assam,