Hydrobiologia 474: 1–39, 2002. © 2002 Kluwer Academic Publishers. Printed in the Netherlands. 1 A review of selected South- and East Asian Tanytarsus v.d. Wulp (Diptera: Chironomidae) Torbjørn Ekrem Museum of Zoology, University of Bergen, Muséplass 3, N-5007 Bergen, Norway Tel: 555-82903. Fax: 555-89677. E-mail: Torbjorn.Ekrem@zoo.uib.no Received 25 June 2001; in revised form 3 January 2002; accepted 6 February 2002 Key words: Tanytarsus, Chironomidae, Diptera, taxonomy, Asia Abstract Male and pupa of Tanytarsus calorifontis sp. n. are described. Diagnoses, hypopygium drawings for the examined species and distribution are given for: Tanytarsus akantertius Sasa & Kamimura, T. angulatus Kawai, T. atagoensis Tokunaga,T. bathophilus Kieffer, T. boninensis Tokunaga, T. formosae (Kieffer), T. formosanus Kieffer, T. infundibulus Chaudhuri & Datta, T. kikuchii Sasa, Kawai & Ueno, T. konishii Sasa & Kawai, T. mcmillani Freeman, T. mendax Kieffer, T. miyakobrevis Sasa & Hasegawa, T. monstrosus Chaudhuri et al., T. ogasaquartus Sasa & Suzuki, T. ogasatertius Sasa & Suzuki, T. okuboi Sasa & Kikuchi, T. oscillans Johannsen, T. ovatus Johannsen, T. oyamai Sasa, T. pollexus Chaudhuri & Datta, T. shouautumnalis Sasa, T. shoudigitatus Sasa, T. takahashii Kawai & Sasa, T. tamaundecimus Sasa, T. tonebeceus Sasa & Tanaka, T. tusimatneous Sasa & Suzuki, T. unagiseptimus Sasa, T. uraiensis Tokunaga, T. yakuheius Sasa & Suzuki and T. yunosecundus Sasa. Tanytarsus ikiefeus Sasa & Suzuki is a new junior synonym of T. konishii. Tanytarsus ikifegeus Sasa & Suzuki, T. miyakoflavus Sasa & Hasegawa,T. oyabepallidus Sasa, Kawai & Ueno, T. simantoopeus Sasa et al. and T. tusimatheius Sasa & Suzuki are new junior synonyms of T. okuboi. Tanytarsus nippogregarius Sasa & Kamimura is a new junior synonym of T. bathophilus. Tanytarsus cultellus Chaudhuri & Datta and T. sibafegeus Sasa et al. are new junior synonyms of T. oscillans.Tanytarsus insulus (Guha et al., 1985) is a new junior synonym of T. ovatus. Tanytarsus sakishimanus Sasa & Hasegawa, T. vinculus Chaudhuri et al., T. parvistylus Chaudhuri & Datta, T. fusciabdominalis Guha et al. and T. euformosanus Kikuchi & Sasa are new junior synonyms of T. formosanus. Tanytarsus tsutaprimus Sasa, T. tokarajekeus Sasa & Suzuki, T. tusimatlemeus Sasa & Suzuki, T. tusimatopeus Sasa & Suzuki and T. yakugeheuus Sasa & Suzuki are new junior synonyms of T. shouautumnalis. Tanytarsus togasiroidus Sasa & Okazawa is a new junior synonym of T. shoudigitatus. Tanytarsus tusimatjekeus Sasa & Suzuki and T. tusimatkeleus Sasa & Suzuki are new junior synonyms of T. akantertius, and Tanytarsus tusimatpequeus Sasa & Suzuki is a new junior synonym of T. tusimatneous. Virgatanytarsus toganiveus (Sasa & Okazawa), Cladotanytarsus utonaiquartus (Sasa) and Zavrelia tusimatijeus (Sasa & Suzuki), all previously placed in Tanytarsus, are new combinations. Introduction Research on chironomids from the East Asian subregion has been extensive during the last 30 years, and a large number of new species have been described. Most of these were described by Dr Manabu Sasa and co- workers, who have given numerous descriptions and species lists from most regions in Japan. In South Asia, most of the recent taxonomic work has been done by Prof. P. K. Chaudhuri and his Indian col- leagues. Consequently, many of the recently described species are found only in Japan or India although they probably have a wider distributional range. A travel grant enabled me to visit the Sasa type collection now located at the National Institute of Environmental Studies (NIES), Tsukuba, Japan in November 2000; these types together with those of Indian Tanytarsus species kindly provided by Dr Chaudhuri form the basis for the review of the Tanytarsus species presented here. This paper is part of a thesis for the 2 partial fulfilment of a Dr scient degree at the University of Bergen, Norway, entitled “A revision of the Tanytarsus eminulus, gregarius, lugens and mendax species groups (Diptera: Chironomidae)”. For this reason, the selected species reviewed in the present study are almost all potential members of the above groups and will be included in a phylogenetic analysis later. However, a few species diagnosed here clearly do not belong to the above species group. They are nevertheless included because the original description either argues their placement in the T. eminulus, T. lugens, T. gregarius or T. mendax species groups or has had an unclear description regarding group placement. Material, methods and morphology The terminology with abbreviations follows Sæther (1980), with the additions and corrections given by Sæther (1990). The term ‘taeniae’ (Langton, 1994) is used for the filamentous setae (LS) of the pupal abdomen, ‘frontal tubercles’ is used for the tubercles on the frons of the imago and ‘cephalic tubercles’ is used for the tubercles on the frontal apotome of the pupal cephalothorax. Material for light microscopy was mounted or remounted in Euparal according to the procedures described by Sæther (1969). Measurements were taken according to Schlee (1966). The shape of the median volsella lamellae is usually difficult to see and flattening or dissection of the hypopygium is often necessary to examine this character properly. The following abbreviations are used: Larva (L), pupa (P), pupal exuviae (Pex); length of antennal pedestal divided by length of larval head capsule (AHR); length of antennal pedestal divided by length of larval antennal segment 1 (AAR); length of Lauterborn organs including stems, divided by length of larval antennal segments 3–5 (LOR) and width of mentum divided by width of ventromental plates (MVR). The Australian National Insect Collection, Canberra (ANIC), Department of Natural Sciences, Bishop Museum, Hawaii, U.S.A. (Bishop), The Natural History Museum, London, U.K. (BMNH), Deutsches Entomologisches Institut, Eberswalde, Germany (DEI), Entomology collection, Hiroshima Imperial University, Japan (HIU), National Institute of Environmental Studies (NIES), Naturhistorisches Museum Wien, Vienna, Austria (NMW), Royal Belgian Institute of Natural Sciences (RBIN), Insect collections, Department of Zoology, University of Burdwan, India (UBI), Museum of Zoology, University of Bergen, Norway (ZMBN), Zoological Survey of India, Calcutta (ZSI), the Zoologische Staatssammlung München, Munich, Germany (ZSM) and the private collection of M. J. Bolton, Ohio, U.S.A. (Bolton). Tanytarsus akantertius Sasa & Kamimura (Figs 1– 2) Tanytarsus akantertius Sasa & Kamimura, 1987: 21, Fig. J. Holotype ♂ (NIES Type no. A11: 37B) Japan, Hokkaido, Akan National Park, Lake Akan, 14.VI.1982, leg. M. Yasuno & Y. Sugaya; 4 paratypes ♂♂ as holotype (NIES A100: 35, 36A, 38A, 38B) [all examined]. Sasa (1991a: 70), male. Tanytarsus tusimatjekeus Sasa & Suzuki, 1999a: 31, Fig. 36. Holotype ♂ (NIES Type no. A355: 97) Japan, Tsushima Island, Kamaigata, Nita Dam, sweeping net, 26.III.1998, leg. H. Suzuki [examined] Syn. n. Tanytarsus tusimatkeleus Sasa & Suzuki, 1999a: 31, Fig. 37. Holotype ♂ (NIES Type no. A355: 98) Japan, Tsushima Island, Kamaigata, Nita Dam, sweeping net, 26.III.1998, leg. H. Suzuki [examined] Syn. n. Diagnosis T. akantertius is separable from other Tanytarsus species by the following combination of characters. Small to minute frontal tubercles; wing hairy; AR about 1.0; LR1 about 2 or less; anal tergite bands well separated, thin, not reaching crests; minute microtrichiafree area on either side of anal point base; microtrichia mostly absent between crests of anal point, only present among 6 small median tergite setae; spinulae in regular row between well developed anal crests; anal point somewhat triangular, with rounded apex; superior volsella with concave median margin and median hook- shaped apex, 4–5 dorsal setae and 2 median setae, microtrichia absent; digitus well developed, clearly reaching beyond median margin of superior volsella; 1 seta on small tubercle below digitus on stem of superior volsella; median volsella short, at almost right angle to main axis of hypopygium, with closely set apparently setose lamellae; inferior volsella more or less straight with about 17 apical setae. Remarks T. akantertius keys to the T. chinyensis species group (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971), except being larger and having a somewhat higher AR. The species is quite similar to Tanytarsus chinyensis Goetghebuer, but separates by having a higher AR, lower LR1 and a larger median volsella with more lamellae. Sasa & Kamimura 3 Figures 1–8. Hypopygia. Figures 1–2. Tanytarsus akantertius; Figures 3–4.Tanytarsus atagoensis; Figures 5–6. Tanytarsus boninensis; Figures 7–8. Tanytarsus calorifontis sp. n. 4 (1987) argue the placement of T. akantertius in the T. chinyensis group, but the authors failed to do so in the discussion on T. tusimatjekeus and T. tusimatkeleus (Sasa & Suzuki, 1999a) where they placed the species in the T. mendax species group. Sasa & Suzuki (op. cit.) separate the two latter species only in differences in frontal tubercles and LR1 values. These differences are not found upon re- examination. Other differences shown in the original drawings (Sasa & Suzuki op. cit.) are due to mounting procedures. T. akantertius has been recorded from Hokkaido, Toyama and the Tsushima Islands, Japan (Sasa, 1988, 1991a: Sasa & Kamimura, 1987; Sasa & Suzuki op. cit.). Tanytarsus angulatus Kawai Tanytarsus angulatus Kawai, 1991: 168, Fig. 6. Holotype ♂ (HIU) Japan, Toyama Prefecture, Dokawa River near Matsuo Jinja, 24.V.1983, leg. K. Kawai [not examined]. Sasa & Kikuchi (1995), adult male; Spies (1998b), adult male, pupa, larva. Examined material: 1 L, P, ♂ (Bolton) U.S.A., Ohio, Franklin Co., Scioto Park, Spring, 6.–13.V.1989, leg. M. J. Bolton. Diagnosis The following combination of characters separates T. angulatus from other Tanytarsus species. Anal point broadest at apex, with small spinulae between well developed anal crests; superior volsella long and narrow, tapering toward apex which is hook- like bent medially; digitus almost parallel and as long as superior volsella; median volsella well developed with subulate lamellae reaching tip of inferior volsella. Gonostylus with somewhat abruptly constricted apex. Pupa with slender, bare thoracic horn shorter than 450 µm; precorneals arranged in almost straight line, anterior most seta slightly longer than other 2; developed cephalic tubercles, 0.15–0.35 × length of frontal setae; pedicel sheath tubercles obvious; posterior thoracic mound well developed; hook row about 0.5 × width of segment II; spine patches of tergites III, IV, large with long, anally directed spines; spine patches of tergites V and VI small; 1 lateral taenia on segments (V), VI, 2 on segment VII and 5 on segment VIII; anal fringe with 43–54 taeniae in 1 row. Larval antennal segment 2 with unsclerotized apex, segments 3, 4 and 5 well pigmented; antennal blade longer than segment 2; AR 1.9–2.3; AAR 0.67; LOR 1.5–2.0; no pigmented band on LO stylus; SI and probably chaetulae basales plumose; S II, some chaetae and chaetulae laterales simple; some chaetae with small teeth; mandible with 3 inner teeth, 1 apical ventral tooth and 1 apical dorsal tooth; seta subdentalis reaching slightly beyond apex of dorsal apical tooth; apical tooth of pecten mandibularis stronger than rest; narrow post occipital plate without incision. Mentum with dark lateral edges on median tooth, MVR 0.77; ventromental plates, broad, evenly convex with obvious striations; 7–8 anal setae. Remarks A thorough description and drawings of all stages are given by Spies (1998b). Pupae of T. angulatus key to the T. lugens species group, while males key to the closely related T. mendax species group due to the presence of a long digitus. However, as mentioned by Spies (1998b), other characters like the shape of median volsella and absence of median tergite setae are shared with species in the T. lugens species group. A certain placement of the species has to await a phylogenetic analysis of the above and associated species groups. The species has previously been recorded from Toyama, Japan and California in addition to the present new record from Ohio (Kawai, 1991; Sasa & Kikuchi, 1995; Spies, 1998b). Tanytarsus atagoensis Tokunaga (Figs 3–4) Tanytarsus atagoensis Tokunaga, 1938: 348, Fig. 24, 25. Holotype ♂, Japan, Hyogu Prefecture, Mt. Atagoyama, 31.V.1931, leg. M. Tokunaga [not examined]. 3 paratypes ♂♂ (Kyushu University) as holotype [examined]. Diagnosis The following combination of characters separates T. atagoensis from other Tanytarsus species: anal point with narrow, evenly rounded tip and several small spinulae between anal crests; anal tergite with median setae at some distance from anal point base, microtrichial pattern uncertain, but possibly small microtrichia- free patches on either side of anal point; superior volsella almost quadrate, but with medially elongated apex, 3 median setae; digitus well developed, thin, not reaching past median margin of superior volsella; median volsella short, medially directed, with several lamellae some of which are probably foliate. Remarks The three paratypes were the only specimens found in the Tokunaga collection. All specimens are obviously 5 Figures 9–15. Hypopygia. Figures 9–10. Tanytarsus formosae; Figures 11–13. Tanytarsus formosanus; Figures 14–15. Tanytarsus infundibulus. worn from long time in alcohol and several genital features were difficult to see properly. The species keys to the T. pallidicornis species group (Cranston et al., 1989) and fits the group diagnosis except having a somewhat shorter digitus (Reiss & Fittkau, 1971). T. atagoensis has been recorded only from its type locality on Hyogu, Japan. Tanytarsus bathophilus Kieffer (Figs 58, 66, 75, 82, 86, 91, 96, 101, 106) 6 Tanytarsus bathophilus Kieffer, 1911a: 49. Holotype ♂ (RBIN) Germany, Holzmaar, 9.VIII.1910, leg. A. Thienemann [not examined]. Kieffer (1911a, 1915), Goetghebuer (1928, 1937–1954), Reiss (1968), Shilova (1976), male adult; Bause (1913), Thienemann (1915), Shilova (1976), pupa; Shilova (1976), larva. Calopsectra luticola (Kieffer, 1922: 109). Holotype ♂ Germany, Mecklenburg, Kirchensee, leg. A. Thienemann [not examined]. Synonymised by Thienemann (1929). Microtendipes microsandalum (Kieffer, 1915: 71). Synonymised by Reiss & Fittkau (1971). Calopsectra tripunctata (Reiss, 1968: 205). Holotype ♂ Germany, Bodensee, Wasserburger Bucht, 1962, leg F. Reiss [not examined]. Synonymised by Reiss & Fittkau (1971) Tanytarsus nippogregarius Sasa & Kamimura, 1987: 20, Fig. I. Holotype ♂ (NIES Type no. A100:21) Japan, Hokkaido, Akan National Park, Lake Akan, 14.VI.1982, leg. M. Sasa & K. Kamimura; 6 paratypes ♂♂ (A100:22-27) as holotype [all examined]. Syn. n. Tanytarsus yunosecundus Sasa, 1984: 36, Fig. 16. Partim: pupa. Tanytarsus gregarius sensu Sasa, 1984: 40, Figs 22– 24. Additional material examined: 2 ♂♂ pharate pupae and 3 Pex, Germany, Amper, Pegel Stegen, 19.IV.1988, leg. Hieber; 1 ♂ Pex, Germany, Holzmaar (Locus Typicus), possibly 9.VIII.1910, leg. A. Thienemann; 2 ♂♂ Pex Sweden, Lappland, 1936, leg. A. Thienemann; 1 ♂ Pex and 1 & Pex Germany, Bavaria, Brunnsee, 10.VII–19.IX.1990, leg. N. Reiff; 1 ♂ Japan, Hokkaido, Akan National Park, Lake Panke, 15.VI.1982, leg. M. Sasa & K. Kamimura. Diagnosis The following combination of characters separates T. bathophilus from other Tanytarsus species. Male adult dark brown to black; second palpomere as long or somewhat shorter than third; small frontal tubercles, LR1 about 1.7, AR about 1.4; wing hairy with macrotrichia on most veins, but Cu bare; median anal tergite setae absent; large microtricha- free area around base of anal point; anal point with granulose apex and several scattered spinulae between well developed anal crests; superior volsella oval, broadest proximal, with about 8 dorsal and 3 median setae; digitus minute and often difficult to observe; median volsella relatively large, reaching apex of superior volsella, with several subulate lamellae which are about 2× longer than broad. Pupa with long, slender, bare thoracic horns; precorneals arranged in triangular pattern, anterior most seta being slightly longer than other 2; developed cephalic tubercles, about as broad as long; pedicel sheath tubercles obvious; large posterior thoracic mound; anterior thoracic mound present; spine patches of tergites III, IV, large with long, anally directed spines; spine patches of tergites V and VI small; 1 lateral taeniae on segments V–VI, 2 on segment VII and 5 on segment VIII; usually about 30–50 filamentous setae in 1 continuos row on anal lobe. Larva with AR about 2.0; antenna with segment 3–5 pigmented; Lauterborn organs small, on pedestals about 2× longer than length of antennal segments 3–5 with broad, basal band; mentum with 3 broad median teeth; mandible with 3 inner ventral teeth, 1 apical ventral, 1 apical dorsal and 1 large dorsal plate-like tooth; postoccipital plate narrow with distinct cleavage. Remarks Tanytarsus bathophilus belongs to the T. lugens species group (Reiss & Fittkau, 1971). Males of T. bathophilus are separated from the other species in the T. lugens group by having second palpomere as long or shorter than third, a minute digitus and moderately large median volsella with subulate lamellae which are about 2× longer than broad. Pupae of T. bathophilus can be separated from the other species of the lugens species group by the continuous anal fringe and posterior thoracic mound larger than 40 µm. Diagnostic characters for larvae of the T. lugens species group have not been established, but common for T. bathophilus and T. lugens is the large dorsal plate-like tooth on the mandibles. This character, however, is also present on larval mandibles of Corynocera ambigua Zetterstedt and C. oliveri Lindeberg (Hofmann, 1984), but these species can be distinguished by their reduced mentum and mandible (C. ambigua) and three dorsoapical mandibular teeth (C. oliveri). Hofmann, op. cit. mentions that the middle tooth and first lateral teeth of the mentum of Corynocera larvae constitutes a distal ventral plate and that the next lateral teeth are situated dorsally of their median neighbour teeth. This mentum construction is also present, although not as dominating, in many if not all Tanytarsus species, and caution should be taken by palaeoecologists who wish to determine worn mouth parts of Tanytarsini head capsules to species and even genus level! The type material of T. yunosecundus Sasa, 1984 was 7 Figures 16–19. Hypopygia. Figures 16–17. Tanytarsus kikuchii; Figures 18–19. Tanytarsus konishii. examined and some of the pupal exuviae were conspecific with T. bathophilus as were males and pupae described in the same paper as T. gregarius Kieffer, 1909 (Sasa, 1984). The tentatively associated pupa of T. yunosecundus is diagnosed below. T. bathophilus has so far been recorded from Central Europe, Lake Rybinsk, Russia and Honshu and Hokkaido, Japan (Reiss & Fittkau, 1971; Shilova, 1976; Sasa, 1984; Sasa & Kamimura, 1987). Tanytarsus boninensis Tokunaga (Figs 5–6) Tanytarsus boninensis Tokunaga, 1964: 606, Fig. 17c. Holotype ♂ (Bishop Type no. 3376) Bonin Island, Chichi Jima, Futamiko, 10.V.1956, leg. C. K. Clagg; 1 allotype & and 2 paratypes ♂♂ as holotype [all examined]. Diagnosis The following combination of characters separates T. boninensis from other Tanytarsus species. Large frontal tubercles; AR c. 0.8; LR1 c. 2.5; wing moderately hairy with few macrotrichia in cell m and several on Cu. Anal tergite bands of Y- type; anal tergite with square dorsal margin and a large microtrichia- free area around anal point; anal point short, stout, with broad, evenly rounded tip and 3 small trifid spinulae between low anal crests; small field of microtrichia between crests; anal tergite with small median setae; superior volsella oval with c. 6 dorsal and 3 closely set median setae; digitus well developed, thick, almost reaching apex of superior volsella; median volsella short, medially directed, with 4 spatulate and some setose lamellae; inferior volsella almost straight with 8 distinct dorsoapical lobe; gonostylus almost equally wide over its whole length, medially curved. Remarks T. boninensis keys to the T. norvegicus species group due to the distally broad gonostyli and the Y- shaped anal tergite bands (Cranston et al., 1989) and is probably not a member of the T. gregarius group which it superficially resembles. The short and broad anal point and thick digitus will separate the species from the other members in the T. norvegicus group (Reiss & Fittkau, 1971). T. boninensis has so far been recorded only from Bonin Island. Tanytarsus calorifontis sp. n. (Figs 7, 8, 59, 67, 76, 83) Type material: Holotype ♂ (ZMBN Type no. 364) Thailand, Krabi, Khao Nor Chuchi, Hot spring, 21.I.1997, leg. Department of Zoology, UiB, student expedition; 20 paratypes Pex as holotype. Etymology Calorifontis from Latin, meaning ‘warm spring’ referring to the species’ type locality. Diagnosis Tanytarsus calorifontis sp. n. can be separated from other described Tanytarsus species by the following combination of characters. Male imago AR c. 0.8, LR1 c. 2.4; hypopygium with separated anal tergite bands which end at some distance from anal point base, median tergite setae absent, large microtrichiafree area around base of anal point, two small teeth on posterior margin of anal tergite; anal point with large spinulae between well developed anal crests; superior volsella somewhat pear- shaped with posterior constriction and dorsolateral field of microtrichia; digitus absent; median volsella moderately long with some subulate and several setose lamellae; inferior volsella relatively straight with a dorsomedially directed apex. Pupal frontal apotome with low cephalic tubercles (c. 20 µm), well developed pedicel sheath tubercles; thoracic horn narrow (c. 15 µm) with a row of chaetae on mid 2/3 which are somewhat longer than width of horn; 3 precorneals in almost a straight row, anterior precorneal stronger but not longer than other 2; posterior thoracic mound absent; anterior pair of dorsocentrals thin c. 2× longer than thick setae; TII with anterior transverse field of shagreen and 2 triangular patches posteriorly; TIII with small field of shagreen anterior of a small round spine patch; TIV– VI with small anterior spine patches; TVIII and anal Table 1. Lengths (in µm) and proportions of legs for T. calorifontis p1 p2 p3 p1 p2 p3 fe ti ta1 ta2 ta3 ta4 ta5 425 450 468 241 403 504 583 241 - 274 130 - 223 108 - 154 72 - 76 58 - LR 2.42 0.60 - BR 3.5 4.4 - BV 1.72 2.97 - SV 1.14 3.54 - lobe with anterior patches of shagreen; segment VII with 2 lateral taeniae, segment VIII with 4-5 lateral taeniae; anal fringe with about 30 taeniae. Description Male imago (n =1). Total length 2.0 mm. Wing length 1.03 mm. Total length/wing length 2.0. Cleared specimens pale brownish with darker vittae, preepisternum, median anepisternum, epimeron II, scutellum, postnotum and eyes. Head. Antenna normally developed, AR 0.83. Thirteenth flagellomere 310 µm long. Distance between eyes c. 180 µm. Frontal tubercles small, 7 µm long. Temporal bristles 8; 2 inner verticals, 4 outer verticals, 2 postorbitals. Clypeus triangular, 65 µm long with 15 setae. Tentorium 83 µm long, 22 µm wide at sieve pore. Stipes 108 µm long, 11 µm wide. Width of cibarial pump 47 µm. Lengths of palp segments (in µm): 25, 25, 58, 76, 79. Thorax. Scutal tubercle absent; dorsocentrals 6; acrostichals 14; prealars 0; scutellars 4; halteres with 5 setae. Wing. VR 1.12. Sc bare, R with 10 setae, R1 with 8, R4+5 with 4, m1+2 with 20, m3+4 with 4, Cu, Cu1 and PCu with 0 and An with 8 setae; m bare, r4+5 with 36 setae, m1+2 with 31 including on false vein, m3+4 , cu and an bare. Legs. Spur on front tibia 32 µm long including scale. Spines of middle and hind tibiae 25 µm long including 14 µm long comb. Lengths and proportions in Table 1. Hypopygium (Figs 7–8). Tergite IX 90 µm long with no median and 12 apical setae; apical margin with 2 pairs of small teeth; microtrichia absent from a large area around base of anal point. Anal point 47 µm long, 14 µm wide at base and 7 µm wide at apex. Anal point 9 Figures 20–27. Hypopygia. Figures 20–21. Tanytarsus mcmillani; Figures 22–23. Tanytarsus miyakobrevis; Figures 24–25. Tanytarsus ogasaquartus; Figures 26–27. Tanytarsus ogasatertius. 10 with 3 strong spinulae between well developed anal crests. Anal tergite bands curved caudally, ending at some distance before anal point. Transverse sternapodeme 43 µm long, phallapodeme 86 µm long. Gonocoxite 104 µm long, gonostylus 79 µm long. Superior volsella (Fig. 8) pear- shaped, constricted apically, bearing 5 small setae dorsally and 3 setae medially; dorsolateral microtrichia present. Digitus absent; median volsella moderately long 45 µm long including c. 2-3 µm long, subulate and simple lamellae. Inferior volsella relatively straight, apex dorsomedially directed, 77 µm long, c. 12 apical setae. HR 1.32, HV 2.5. Pupa (n = 6, unless otherwise stated). Total length 2.3–3.3, 2.9 mm, abdomen 1.8–2.6, 2.2 mm. Colouration light brown with darker areas around shagreen and spine patches on tergites II through VI. Leg and wing sheaths brown, muscle marks dark brown. Cephalothorax (Figs 59, 67). Length of frontal setae 36–72, 56 µm, cephalic tubercles (Fig. 59) low, 14–25, 21 µm long; pedicel sheath tubercle well developed. Thoracic horn slender, 234–277, 254 µm long, 14–18 µm wide, with a long row of 18–29, 24 µm long, chaetae on mid 2/3. 3 precorneals in almost a straight line, anterior precorneal strong, 54–83, 75 µm long; middle precorneal weaker 54–80, 67 µm long; posterior precorneal 43–80, 64 µm long. 2 antepronotals visible, median antepronotal 40–61, 50 (4) µm long; lateral antepronotal 36–72, 51 (5) µm long. Two pairs of dorsocentrals, anterior pair 25–36, 32 (thick) and 36–72, 58 (thin) µm long; posterior pair 40–43, 42 (thick) and 43–47, 45 (thin) µm long. Only very little granulation dorsal of anterior dorsocentrals. Nose of wing sheath well developed, posterior thoracic mound absent. Abdomen (Fig. 76). Shagreen on tergite (T) II as 1 anterior, transverse band and an additional 2 posterior, triangular patches; shagreen on TIII in small patches anterior to spine patches. Pedes spurii B on TII present; hook row 79–133, 98 µm long, less than 1/3 width of TII; length of longest spines/spinules of TIII– TVI (in µm): 18–32, 25; 14–18, 16; 11–18, 13 and 10–14, 12. Spines of TIII–VI in longitudinal patches, lengths of patches (in µm): 30–54, 45; 58–97, 76; 47– 61, 52 and 36–54, 42. Segment II–VI with 3 lateral setae; segment VII with 1–2 lateral setae and 2–3 lateral taeniae; segment VIII with 2 dorsal setae, anterior taeniate 50–86, 69 µm long, posterior setiform 43– 58, 48 µm long, 1 taeniate ventral seta 47–86, 58 µm long, 4–5 lateral taeniae; anal lobe with 2 dorsal, 76– 137, 105 µm long taeniae, anal fringe with 26–32, 29 taeniae, c. 220–290 µm long; posterior lateral comb of segment VIII (Fig. 83) 22–40, 39 µm wide with 4– 9, 6 apical teeth. Lengths of genital sac: 144–223, 179 µm (♂, n=4) and 72–94 µm (♀, n=2). Remarks Males of T. calorifontis key to the T. mendax species group (Cranston et al., 1989) when ignoring the absence of a digitus, but does not completely fit the group diagnosis in being smaller, having a lower AR and an absent digitus (Reiss & Fittkau, 1971). The shape of the hypopygium and its other appendices, however, is very similar to other species in the T. mendax species group. The pupa will also key to the T. mendax group when ignoring the low cephalic tubercles (Pinder & Reiss, 1986) and is very similar to other species in this group when this character is ignored. Probably, the closest related species is T. uraiensis Tokunaga which is collected from a hot spring in Taiwan. The new species differs from T. uraiensis in the absence of median anal tergite setae, shorter anal point, lower AR, higher LR1 , smaller pupal cephalic tubercles, smaller spine patches on TIII–VI and less taeniae in anal fringe. T. calorifontis is reported only from its type locality, but the species may be found in other hot springs on the Malaysian peninsula and adjacent regions. Tanytarsus formosae (Kieffer) (Figs 9–10) Tanytarsus formosae (Kieffer, 1921: 592). Lectotype designation by Ekrem (2001a). Lectotype ♂ (DEI) Rheotanytarsus formosae, Formosa, Anpin, X.1912, leg. H. Sauter [examined]. Five paralectotypes ♂♂ as lectotype [examined]; 2 paralectotypes ♂♂ and 1 ♀ as lectotype [not examined]. Additional material examined: 1 ♂ (UBI) India, West Bengal, Sagar Island, 29.X.1991, leg. A. Mazumdar. Diagnosis The species can be separated from other described Tanytarsus species by the following characters. AR c. 1.0; wing with apical macrotrichia only, Cu, cu and an bare; anal tergite with 2 median setae and a small microtrichia- free area at anal point base; anal tergite bands separate, reaching well developed anal point crests; 1–5 small spinulae on anal point; superior volsella oval to slightly bean- shaped with 6–8 dorsal and 3 strong, median setae placed on small tubercles; dorsolateral microtrichia present on superior volsella; digitus absent. Median volsella moderately long with numerous thin, subulate and setose lamellae. Inferior volsella slightly S- shaped; gonostylus straight. 11 Figures 28–37. Hypopygia. Figures 28–31. Tanytarsus okuboi; Figures 32–33. Tanytarsus oyamai; Figures 34–35. Tanytarsus oscillans; Figures 36–37. Tanytarsus ovatus. 12 Remarks The identity of Tanytarsus formosae (Kieffer, 1921), was clarified by Ekrem (2001a). One of the paralectotypes actually belongs to the quite different species Tanytarsus gracistylus Chaudhuri & Datta described by Datta et al. (1992). This species, however, resembles the genus Rheotanytarsus even less than T. formosae (Kieffer, 1921), and is probably accidentally a part of the type series. The species keys to the T. lugens species group (Cranston et al., 1989) due to the absence of microtrichiae between anal crests and a digitus, but is apparently closer related to species in the T. gregarius groups due to the shape of the superior volsella and the anal point. The species fits the group diagnosis of the T. gregarius group (Reiss & Fittkau, 1971) except being smaller, having a lower AR, microtrichia absent from anal point and a smooth apex of anal point, but a placement has to await a phylogenetic analysis of the genus. T. formosae differs from the probably closest related species, T. gregarius and T. inaequalis, by completely lacking a digitus, absence of microtrichia between anal crests, having two strong median setae on the anal tergite and the presence of dorsolateral microtrichia on superior volsella. T. formosae (Kieffer, 1921) has only been recorded from Taiwan. Tanytarsus formosanus Kieffer (Figs 11, 12, 13, 60, 68, 73, 77, 87, 92, 97, 102) Tanytarsus formosanus Kieffer, 1912: 42. Lectotype 1 ♂ (DEI) Formosa, Tainan province, Anpin, X. 1908, leg. H. Sauter; 2 paralectotypes ♂♂ as lectotype [examined]. Tanytarsus formosae Kieffer, 1923: 38. Syntypes 2 ♂♂ (DEI) Formosa, Taihoku province, Daitotei, V.– VI.1914, leg. H. Sauter. [examined]; synonymised by Ekrem (2001a). Tanytarsus horni Goetghebuer, 1934: 39. Holotype ♂ (NMW) Iraq, Basra, 13–15.IV.1926, leg. H. Schmidt; 2 paratypes ♂♂ (BMNH) as holotype; synonymised by Ekrem (2001a). Reiss & Fittkau (1971: 121), Cranston & Judd (1989: 268) adult male, ecology, distribution. Langton (1991: 357), pupa. Verschuren (1997: 500), larva. Tanytarsus aculeus Chaudhuri et al., 1988: 239. Holotype ♂ (UBI Type no. 138), India, West Bengal, Burdwan University campus, 6.VI.1983, leg. Nandi [not examined]; 5 paratypes: 2 ♂♂, 3 Pex (UBI) India, West Bengal, Burdwan, 3.V.1979, leg. S. K. Nandi [examined]; synonymised by Ekrem (2001a). Tanytarsus fuscimarginalis Chaudhuri et al., 1984: 33, Figs 10–13. Holotype ♂ (UBI Type no. 65), India, West Bengal, Bolpur, 18.VIII.1977, leg. D. Chatterje [examined]; paratypes: 1 ♂ (UBI) as holotype [examined]; 1 ♂ (UBI) India, West Bengal, Burdwan, 3.VII.1978, leg. M. Gosh [examined]; synonymised by Ekrem (2001a). Tanytarsus nigrocinctus Freeman, 1957: 220. Holotype ♂ (BMNH) Uganda, Lake Victoria, 4.V.1952 (synonymised by Reiss & Fittkau, 1971) [examined]; 1 paratype ♂ (BMNH) Kenya, Kitui, 29.VI.1953 [examined]. Freeman (1958: 339), Dejoux (1968b: 449), adult male. Tanytarsus dycei Glover, 1973: 444, Fig. 44. Holotype ♂ (ANIC) N. Qld, Danbulla Rd., 5 miles Tinaroo Falls, at light, 26.IV.1967, D. H. Colless; 4 paratypes as holotype [all examined]. Cranston (1996, 2000), larva, pupa; synonymised by Ekrem (2001b). Tanytarsus sakishimanus Sasa & Hasegawa, 1988: 237. Holotype ♂ (NIES Type no. A65: 12 according to original publication) Japan, Ishigaki Island, Ishigaki City, water filtration pond, 1.II.1982, leg. M. Sasa & H. Hasegawa [not examined]; 6 paratypes: 5 ♂♂ as holotype; 1 ♂ as holotype except from pond on Miyako Island [examined]. Syn. n. Tanytarsus euformosanus Kikuchi & Sasa, 1990: 319, Fig. 25. Holotype ♂ (NIES Type no. A200: 13) Indonesia, Sumatra, Lake Toba, 1.I.1987, leg. M. Kikuchi [not examined]; 4 paratypes: 2 ♂♂ (NIES A199: 13, 20) as holotype; 2 paratypes ♂♂ (NIES A199: 21, 22) as holotype except collected at the side of Ashan dam, 03.I.1989 [examined]. Syn. n. Tanytarsus vinculus Chaudhuri et al., 1984: 34, Figs 14–16. Holotype ♂ (UBI Type no. 64), India, West Bengal, Bolpur, 4.VIII.1977, leg. D. Chatterjee [examined]. Syn. n. Tanytarsus fusciabdominalis Guha et al., 1985: 34, Fig. 9. Holotype ♂ (UBI Type no. 133) India, Andaman Islands, Old Wandoor, 7.VI.1982, leg. R. Sharma [not examined]; 1 paratype ♂ as holotype [examined]. Syn. n. Tanytarsus parvistylus Chaudhuri & Datta in Datta et al., 1992: 49, Figs 26–30. Holotype ♂ (UBI Type no. 201) India, West Bengal, Polempur, 23◦ N, 87◦ 9′ E, 25 m a.s.l., 26.XI.1988, leg. A. K. De [examined] Syn. n. Additional material examined: 1 ♂ (UBI) India, Sagar Islands, light trap, 17.X.1990. 1 ♂ (UBI) India, Bolpur, leg. D. Chatterje; 1 ♂ (UBI) India, Andaman Islands, Old Wandoor, 27.V.1982, leg. R. Sharma; 1 ♂ (UBI) India, BSN, 6.VI.1984, leg. T. K. Datta; 1 ♂ 13 Figures 38–48. Hypopygia. Figures 38–39. Tanytarsus pollexus; Figures 40–43. Tanytarsus shouautumnalis; Figures 44–46. Tanytarsus shoudigitatus; Figures 47–48. Tanytarsus takahashii. 14 (UBI) India, Sagar Islands, West Bengal, 18.X.1990, leg. A. Mazumdar; 1 pharate ♂, 1P, 14 Pex, 8L (ZSM) Indonesia, Middle Sumatra, 2.III.1929, leg. A. Thienemann. Diagnosis T. formosanus is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; usually strong thoracic colourations anteriorly on scutum, laterally on scutum under parapsidal suture, basally on scutellum, postnotum, median anepisternum II, epimeron II and on preepisternum; abdominal colouration characteristic and often strong but pale specimens also exist; wing with sparse setation (restricted to apical 1/3), Cu bare, cell r4+5 with a greater or equal amount of setae as cell m1+2 ; AR usually larger than 1. Spinulae in 1 row between well developed anal crests; superior volsella tapered towards apex which is somewhat medially elongated with a ventral lip, superior volsella with dorsolateral microtrichiae, 6–8 dorsal setae of which 3 are placed laterally, 3 median setae where 2 most apical larger than innermost seta; digitus fairly short; median volsella relatively short with 4–6 broad, distally pointed lamellae in addition to 15–20 setose lamellae (often difficult to see, and dissection of the median volsella might be necessary). Pupa with large conical cephalic tubercles; long evenly tapered thoracic horn with chaetae longer than width of horn in 1 row on mid 2/3; strong armament on tergite III and IV with long spines in separate, longitudinal rows occupying 1/3– 1/2 length of tergite; segment VII with 2 anterior setae and 2 posterior lateral taeniae, segment VIII with 5 lateral taeniae where the 3rd is placed more medially than the other 4. Larval head with antenna about 0.6 × length of head capsule; Lauterborn organ small, on short pigmented pedestal, reaching to apex of antennal segment 4; 1.35 < AR < 1.6; AAR 0.52–0.63; SI, chaetae and chaetulae basales plumose; SII, S3 and chaetulae laterales all apparently simple; mandible with 1 large and 1 small dorsal tooth; premandible well pigmented with 5 teeth; 4 large anal tubules. Remarks The holotype slide of T. vinculus has the date 4.IX.1977 which differs from the date (18.IX.1977) in the original publication of the species (Chaudhuri et al., 1984). I assume that the original publication contains a typographic error and that the specimen examined is the holotype. The paratypes of T. fusciabdominalis, T. sakishimanus and T. euformosanus are all consistent with their the original descriptions and do not differ from diagnostic characters of T. formosanus. The synonymy thus is certain although the holotypes have been unavailable for examination. Both pupae and male adults of T. formosanus keys to the T. mendax-group (Pinder & Reiss, 1986; Cranston et al., 1989). As discussed by Sasa & Hasegawa (1988) for T. sakishimanus, the species is probably most closely related to T. oyamai Sasa, but differs by having a different pigment pattern on the male abdomen, a differently shaped superior volsella, digitus, median- and inferior volsella. The pupa differs in having spines on the abdominal tergite IV while the larval LO is larger and with longer styli. Distribution: Africa, Middle East, Taiwan, Hubei Province, China, India, South France, South Spain, Sumatra, Thailand, Ryukyu Islands, Miyako Island, Okinawa Island, Southern Japan (Kieffer, 1912; Kieffer, 1923; Goetghebuer, 1934; Freeman, 1957; Freeman, 1958; Dejoux, 1968a, b, 1976a, b; Dejoux, 1984; Reiss & Fittkau, 1971; Petr, 1972; Hashimoto et al., 1981; Chaudhuri et al., 1984, 1992; Hare & Carter, 1987; Harrison, 1987; Kikuchi & Sasa, 1990; Langton, 1991; Laville & Reiss, 1992; Wang & Zheng, 1992; Sasa, 1993c; Sasa & Hasegawa, 1988; Sasa & Suzuki, 1993; Verschuren, 1997; Mazumdar et al., 1998). T. formosanus is a freshwater species, but often found in lakes with higher salinity (Dejoux, 1976b; Verschuren, 1997). The highest conductivity in which larvae of T. formosanus have been found is 868 µS cm−1 at Lake Awasha, Ethiopia (Kibret & Harrison, 1989). The larvae have been recorded from both sandy and muddy bottoms with coarse organic debris (Dejoux, 1976b; Hare & Carter, 1987; Verschuren, 1997), and have at least in one lake shown intolerance to daily oxygen fluctuations (Petr, 1972). In Japan, the species is known from sandy bottom in water filtration ponds and other small water bodies. Tanytarsus infundibulus Chaudhuri & Datta (Figs 14–15) Tanytarsus infundibulus Chaudhuri & Datta in Datta et al., 1992: 47, Figs 19–25. Holotype ♂ (UBI Type no. 200), India, West Bengal, Gangasagar, 21◦ 9′ N 88◦ E, 21.I.1987, leg. A. K. De; 1 paratype ♂ India, West Bengal, Jalpaiguri, 22.III.1988, leg A. K. De [both examined]. 15 Figures 49–57. Hypopygia. Figures 49–50. Tanytarsus tusimatneous; Figures 51–52. Tanytarsus unagiseptimus; Figure 53. Tanytarsus uraiensis; Figures 54–55. Tanytarsus yakuheius; Figures 56–57. Tanytarsus yunosecundus. 16 Diagnosis T. infundibulus can be separated from other Tanytarsus species by the following combination of characters. AR c. 1.0; wing moderately hairy with macrotrichia on Cu; frontal tubercles present. Anal tergite bands reaching anal crests; 2 small, median tergite setae at base of anal point. Anal point long, narrow, with 4–5 spinulae between well developed crests, apex evenly rounded, relatively large microtrichia- free area around base. Superior volsella teardrop- shaped with medially directed posterior apex, dorsolateral microtrichiae absent, about 5 dorsal setae of which 2–3 are placed on apex, 3 median setae; digitus absent or a minute lobe. Median volsella well developed with c. 5 recurved, broad lamellae and several setose lamellae; inferior volsella broad and straight. Remarks The paratype examined is not included in the list of the type material (Datta et al., 1992) and possibly not included in the description. The specimen does, however, carry a paratype label and is collected by A. K. De at Jalpaiguri as are paratypes used in the original description. The specimen available for examination is almost identical to the drawings by Datta et al. op. cit., and differs from the original description only in the AR value. Datta et al., op. cit. describes frontal tubercles as absent for T. infundibulus but regards them as present in the remarks section. T. infundibulus keys to the T. lugens species group (Cranston et al., 1989), but does not fit the group diagnosis (Reiss & Fittkau, 1971) in being smaller, having a lower AR, less spinulae between anal crests and differently shaped superior- and median volsellae. The species is probably closer to the T. gregarius or the T. mendax species groups, but the absence of a digitus will hinder the placement of T. infundibulus in these species groups. The species is apparently closely related to T. nichollsi, T. formosae (Kieffer, 1921), T. elisabethae and T. harei, but can be distinguished from all these species by the following combination of characters: absence of microtrichia between the anal crests and on superior volsella, setae on Cu and broad, recurved lamellae on median volsella. Datta et al. (1992) recorded T. nichollsi from the same locality as T. infundibulus. Their diagnosis of T. nichollsi, however, does not match that of the Australian specimens and could possibly be specimens of T. infundibulus. The species is only recorded from its type locality. Tanytarsus kikuchii Sasa, Kawai & Ueno (Figs 16– 17) Tanytarsus kikuchii Sasa et al., 1988: 38, Plate 7 Fig. B. Holotype ♂ (NIES Type no. A136: 69) Japan, Toyama, Oyabe River, site Y-2, 19.VIII.1987, leg. Sasa, Kawai & Ueno [examined]. Sasa & Kikuchi (1995), adult male. Tanytarsus sp. ‘Tokushima’ Sasa & Kikuchi, 1986: 29, Fig. 5B. Diagnosis The following combination of characters separates T. kikuchii from other Tanytarsus species. LR1 3.42 (holotype). Anal point with spinulae in 1 row between anal crests; anal tergite bands separate; anal tergite with 6 median setae and 2 small microtrichia- free areas on either side of anal point base; some microtrichiae present between crests of anal point; superior volsella oval; digitus absent; median volsella large with numerous simple and subulate lamellae reaching tip of inferior volsella. Remarks The species keys to the T. gregarius species group, and as mentioned by Sasa et al. (1988) is probably closest to T. gregarius. It differs, however, in having much longer median volsella, higher LR1 , lower AR and a broader superior volsella. T. kikuchii is also similar to T. yakuheius and T. yunosecundus but differs in the absence of a digitus, higher LR1 and smooth anal point apex (compared to T. yakuheius). T. kikuchii has been recorded from Tokushima and Toyama Prefectures, Japan. Tanytarsus konishii Sasa & Kawai (Figs 18–19) Tanytarsus konishii Sasa & Kawai, 1985: 19, Fig. 5. Holotype ♂ (NIES Type no. unknown, specimen was not found under the number from the original description) Japan, Toyama Prefecture, Toyama, Kameike Pond, 23.X.1984; 1 paratype ♂ as holotype [not examined]. Tanytarsus ikiefeus Sasa & Suzuki, 1999b: 150, Fig. 6. Holotype ♂ (NIES Type no. A372: 83) Japan, Iki Island, Oushimizu, 28.III.1998, leg. H. Suzuki; 1 paratype male (NIES A372: 84) as holotype [examined]. Syn. n. Diagnosis The following characters can be regarded as diagnostic. Large species with wing length of about 2.5 mm; frontal tubercles c. 65 µm long; AR about 1.5; 17 Figures 58–65. Pupal frontal apotomes. Figure 58. Tanytarsus bathophilus; Figure 59. Tanytarsus calorifontis; Figure 60. Tanytarsus formosanus; Figures 61–62.Tanytarsus mendax; Figure 63. Tanytarsus oyamai; Figure 64. Tanytarsus uraiensis; Figure 65. Tanytarsus yunosecundus. LR1 c. 2.0; wing with sparse setation, mainly on distal half; anal tergite bands separate, barely reaching crests of anal point; median tergite setae absent; microtrichia- free area around base of anal point; anal point with about 6 spinulae between well developed anal crests, without microtrichia; superior volsella almost sickle- shaped with blunt medially directed apex with 3 median and 5–8 dorsal setae, proximal median seta smaller than distal setae; digitus absent; median volsella moderately long with numerous, apparently simple lamellae almost reaching apex of inferior volsella; inferior volsella slightly club- shaped. Remarks Although the types of T. konishii have not been examined, the good original description of the species 18 makes the synonymy with T. ikiefeus certain. T. konishii will key to the T. lugens species group due to the missing digitus (Cranston et al., 1989), but is probably closer to species in the T. mendax species group due to the microtrichial pattern on the anal tergite and shape of the superior volsella. T. konishii seems to be closely related to T. calorifontis and T. oyamai, but will separate from T. calorifontis by having a higher AR and no microtrichia on the superior volsella and from T. oyamai by the absence of median tergite setae and a longer differently shaped median volsella. T. konishii has been recorded from Iki Island, Fukushima and Toyama Prefectures (Sasa & Kawai, 1985; Sasa & Ogata, 1999; Sasa & Suzuki, 1999b; Sasa et al., 2000). Tanytarsus mcmillani Freeman (Figs 20–21) Tanytarsus mcmillani Freeman, 1958: 341, Fig. IIe. Holotype ♂ (BMNH) Nigeria, Kankiya, XII.1956– I.1957, leg. B. McMillan; 3 paratypes: 2 ♂♂ as holotype and 1 ♂ (BMNH) Nigeria, Kaduna, 19.X.1956, leg. B. McMillan [all examined]. Tanytarsus atroxitarsus Chaudhuri & Datta, in Datta et al., 1992: 42, Figs 1–6. Holotype ♂ (UBI, Type no. 196) India, West Bengal, Jalpaiguri, 26◦ 16′ N 88◦ 25′ E, 68.5 m a.s.l., 22.III.1988, leg. A. K. De; 1 paratype ♂ (UBI) as holotype except collected in Birpara, 26◦ 43′ N 89◦ 8′ E, 117.8 m a.s.l., 14.X.1990, leg. T. K. Datta [both examined] synonymised by Ekrem (2001a). Additional material examined: 1 ♂ (BMNH) Saudi Arabia, Medina, W. J. 12.l, 26◦ 42′ N 37◦ 15′ E, 1000 m a.s.l., 14.XI.1984, leg. W. Buttiker, 1 ♂ (BMNH) Saudi Arabia, Makkah, W. Maraum, 22◦ 16′ N 39◦ 44′ E, 280 m a.s.l., 3–4.V.1984, leg. W. Buttiker, 1 ♂ (ZSM) Kamerun, Adamaoua, 31.IV.1979, leg. W. Flauke & P. Nagel. Diagnosis The species differs from other Tanytarsus species by the following combination of characters. Frontal tubercles well developed; AR c. 1.2; LR1 c. 3.0; wing hairy, c. 10–15 setae in cell m; hypopygium with 2– 8 median setae on anal tergite, anal point with 4–7 small spinulae between well developed anal crests; microtrichia present around base of anal point; superior volsella oval with parallel sides, c. 7 dorsal setae, microtrichia absent; small digitus present; median volsella moderately large with several foliate and simple lamellae. Remarks Based on the adult male morphology, Tanytarsus mcmillani keys to the T. lugens species group (Cranston et al., 1989), but does not fit the group diagnosis (Reiss & Fittkau, 1971) in having AR < 1.4, smooth apex of anal point and a digitus which is c. half the length of superior volsella. A definitive group placement has to await a phylogenetic analysis of the genus. T. mcmillani is quite similar to T. okuboi and T. pollexus, but can be separated from these species by the larger anal point with minute spinules and long median tergite setae. T. mcmillani has been recorded from West Bengal, India, Saudi Arabia and several areas in West Africa (Dejoux, 1974; Cranston & Judd, 1989; Datta et al., 1992; Ekrem, 2001a). Tanytarsus mendax Kieffer (Figs 61, 62, 69, 79, 84, 88, 93, 98, 103, 107) Tanytarsus mendax Kieffer, 1925: 223. Lectotype ♂ (RBIN) Germany, Holstein, Plön, leg. A. Thienemann [not examined]. Reiss & Fittkau (1971 for T. holochlorus), Spies (1998a), list of synonymies, morphology and ecology. Tanytarsus holochlorus Edwards, 1929: 414. Synonymised by Lindeberg (1976). Tanytarsus xanthus Sublette, 1964: 142. Synonymised by Sublette & Sublette (1979) as synonym of T. holochlorus, confirmed by Spies (1998a). Material examined: 1 L, P, ♂ (ZMBN) Russia, Lake Rybinsk, 28.VII.1969, leg, A. I. Shilova; 1 L, P, ♂ (ZSM) Germany, Bavaria, Lerchenauer See near Munich, 29.V.–4.VI.1991, leg. N. Reiff; 1 P, ♂ as previous; 1 L, P (ZMBN) Germany, Bavaria, Kreis Dachau, Karlsfelder See, 15.–19.VII.1999, leg. T. Ekrem; 1 L, P as previous but 15.–21.VII.1999; 1 L, P, ♂ as previous but 15.VII.1999; 1 P, ♀ as previous but 30.VI–4.VII.1999; 1 L, P, ♀ as previous but 30.VI–5.VII.1999; 2 L, P, ♀♀ (ZMBN) Germany, Bavaria, Munich, Nymphenburger Park, Kleiner See, 13.–16.VII.1999, leg. T. Ekrem; 2 L as previous but 24.VII.1999; 1 L, P, ♀ as previous but hatched 2.VIII.1999; 1 prepupa (Bolton) U.S.A., Ohio, Columbiana county, Beaver Creek, Chemline tributary at SR 518, 8.IX.1987, leg. M. J. Bolton; 1 L, P, ♂ (Bolton) U.S.A., Ohio, Geauga county, Beaver Run, Fern Lake Bog, 25.VIII–28.VIII.1993, leg M. J. Bolton; 1 L, P, ♂ (Bolton) U.S.A., Ohio, Portage county, south of Ravenna, Crystal Lake, 24.IX.– 30.IX.1990, leg. M. J. Bolton; 1 P (Bolton) U.S.A., Ohio, Franklin county, Sharon Woods Park, Schrock 19 Figures 66–74. Pupal thoraces and thoracic horns. Figure 66. Tanytarsus bathophilus; Figure 67. Tanytarsus calorifontis; Figures 68, 73. Tanytarsus formosanus; Figure 69. Tanytarsus mendax; Figure 70. Tanytarsus oyamai; Figures 71, 74. Tanytarsus yunosecundus; Figure 72. Tanytarsus uraiensis. Lake, 29.IV.1989, leg. M. J. Bolton; 10 ♂♂ (NIES) Japan, Hokkaido Prefecture, Lake Utonai, 13.VI.1986, leg. M. Sasa. Diagnosis Tanytarsus mendax can be distinguished by the fol- lowing combination of characters. Male wing hairy, macrotrichia on Cu; large frontal tubercles present; hypopygium with median setae and large microtrichiafree area around anal point; row of large spinulae on evenly rounded anal point; superior volsella tapering towards apex; digitus large, extending bey- 20 ond posteromedian margin of superior volsella; median volsella fairly short with some foliate lamellae; inferior volsella largely club- shaped. Pupa with large cephalic tubercles and longer frontal setae, well developed pedicel sheath tubercles for both sexes; thoracic horn long (400–972 µm long, 29–47 µm broad), chaetae longer than width of horn (50–80 µm) in fairly short or long row (1/5–2/3 length of horn); 3 precorneals in a row or very low triangular pattern, posterior seta shorter than 2 anterior ones; scutal suture with weak granulation; 1 dorsocentral in posterior pair much stronger and longer than other three. 2 visible antepronotals (but 3 seta marks), median antepronotal about 80–130 µm long; hook row about 2/3 width of tergite II; spinule patches of tergites III, IV, V and VI relatively small, elongate, on anterior parts of tergites, spinules about 9–14 µm long; 0–1 lateral taenia on segment VI, 2–4 lateral taeniae on VII, 5 lateral taeniae segment VIII; 31– 44 taeniae in anal fringe; posterolateral comb 40–61 µm wide with 6–9 marginal teeth. Pedes spurii B present on segment II. Larva with head 360–432 µm long, head length/width 1.11–1.30; antennal pedestal 82–120 µm; antennal segment 2 with unsclerotized apex, segments 3, 4 and 5 well pigmented; AR 1.95– 2.23, Lauterborn organs including stylus 50–79 µm long; LOR 1.65–1.98; AAR 0.51–0.62; AHR 0.23– 0.28; SI, chaetae and chaetulae basales plumose, S II and chaetulae laterales simple; mandible with 3 inner teeth, 1 apical ventral tooth and 1 apical dorsal tooth; seta subdentalis reaching apex of apical tooth; apical tooth of pecten mandibularis stronger than rest; broad postoccipital plate; dark sclerotised postoccipital margin and tentorium. Mentum with dark lateral edges on median tooth, MVR 0.79–0.91; ventromental plates evenly convex with obvious striations; 4 anal tubules about 80 µm long. Remarks The larva and the pupa of Tanytarsus mendax are difficult to distinguish from those of T. occultus, and can apparently only be distinguished by a less developed pedicel sheath tubercle of the pupa and a slightly lower LOR value in the larva. The colouration of larval gula is variable; the apical tooth of pecten mandibularis is often broken off and could be large also in other species. In Asia, T. mendax probably is most easily confused with T. takahashii and T. oyamai, but will separate from T. takahashii by having a shorter median volsella and a superior volsella without microtrichia, and from T. oyamai by the shape of the anal point and superior volsella. T. mendax has a Holarctic distribution (Spies, 1998a) with one record from Hokkaido, Japan (Sasa, 1988). Tanytarsus miyakobrevis Sasa & Hasegawa (Figs 22–23) Tanytarsus miyakobrevis Sasa & Hasegawa, 1988: 236, Fig. 2. Holotype ♂ (NIES Type no. A64: 65) Japan, Miyako Island, unpolluted stream in plantation, 2.II.1982, leg. M. Sasa & H. Hasegawa [examined]. Sasa & Kikuchi (1995), Sasa (1998) adult male, keys. Diagnosis The following combination of characters separates T. miyakobrevis from other Tanytarsus species. Large LR1 , about 2.9. AR about 1.35; wing hairy, about 10 macrotrichia in m. Anal point long and narrow with apex wider than rest and spinulae in 1 row between anal crests; anal tergite with about 3 long, median setae and 2 small microtrichia- free areas on either side of anal point base; few microtrichiae present between crests of anal point; superior volsella roughly oval with somewhat narrower apex and extensive areas of microtrichia dorsolaterally and ventromedially; digitus small, triangular; median volsella moderately large with some simple and subulate lamellae only reaching to about 1/2 length of inferior volsella. Remarks The species keys to the T. gregarius species group (Cranston et al., 1989), and separates from the other species in this group except T. ovatus by the shape of the anal point and by the extensive areas of microtrichia on the superior volsella. T. ovatus, however, can be separated by having a differently shaped anal point, a larger digitus, larger LR1 and pectinate lamellae on the median volsella. T. miyakobrevis has been recorded only from its type locality. Tanytarsus monstrosus Chaudhuri et al. Tanytarsus monstrosus Chaudhuri et al., 1992: 384, Figs 5–7. Holotype ♂ (UBI Type no. 147) India, West Bengal, Dubrajpur, 2.V.1983, leg. S. K. Das [not examined]; paratypes reared from egg masses from Burdwan, 2.V.1983, leg. P. K. Chaudhuri and from Dubrajpur, leg. S. K. Das [not examined]. Diagnosis The following characters used in the original description (Chaudhuri et al., 1992) can be used as diagnostic. 21 Figures 75–85. Pupal tergites, anal lobes and posterolateral combs. Figures 75, 82. Tanytarsus bathophilus; Figures 76, 83. Tanytarsus calorifontis; Figure 77. Tanytarsus formosanus; Figure 78. Tanytarsus oyamai; Figures 79, 84. Tanytarsus mendax; Figures 80, 85. Tanytarsus yunosecundus; Figure 81. Tanytarsus uraiensis. Adult male without frontal tubercles; wing hairy with several macrotrichia in m; dark, posterior transverse bands on tergites V–VI; anal tergite bands separate, almost reaching crests of anal point; median tergite setae absent; median, round, brown- pigmented patch at base of anal point; anal point with apically rounded apex and 5 spinulae between developed anal crests; superior volsella somewhat oval, with straight posterolateral and convex anteromedian margin, 3 closely set median setae; digitus small; median volsella relatively small, brush like with several lamellae; inferior volsella straight. Pupae with large cephalic tubercles; 22 pedicel sheath tubercles well developed; thoracic horn long with small chaetae evenly distributed on apical 2/3 of horn; 2 precorneal setae; shagreen in a πpattern on TII, TII with pedes spurii B; TIII with posterior, laterally curved patch of spines, anterior patch of shagreen; TIV–VI with anterior, oval spinule patches; segment VII and VIII with 1 and 5 lateral taeniae respectively. Larval head with small Lauterborn organ on short pedestal, reaching to apex of antennal segment 5; AR c. 1.9; SI and chaetae plumose; SII apparently simple; mandible with 1 dorsal tooth; premandible well pigmented with 5 teeth; 4 large anal tubules. Remarks Unfortunately, no type material was available for examination and many peculiar characters reported by Chaudhuri et al. (1992), such as absence of frontal tubercles and the median pigmented patch of the anal tergite, could not be verified. The species will key to the T. mendax group both as pupa and male imago (Pinder & Reiss, 1986; Cranston et al., 1989) and is probably closely related to T. formosanus. T. monstrosus differs, however, in the absence of frontal tubercles, by having a brown pigmented patch at base of male anal point, brown pigmented bands on male tergites V–VI only, evenly distributed chaetulae on apical 2/3 of pupal thoracic horn, one dorsal tooth on larval mandible and larval AR of c. 1.9. The status of the species might change when the type material is examined. Tanytarsus ogasaquartus Sasa & Suzuki (Figs 24– 25) Tanytarsus ogasaquartus Sasa & Suzuki, 1997: 322, Fig. 4. Holotype ♂ (NIES Type no. A307: 19) Japan, Bonin Islands, Chichijima, Sakaiura, sweep net, 14.IV.1996, leg. H. Suzuki; 1 paratype ♂ (NIES A307: 18) as holotype except 17.IV.1996 [both examined]. Diagnosis The following combination of characters separates T. ogasaquartus from other Tanytarsus species. Small LR1 , 1.7–2.5. AR about 0.6–0.9; wing hairy, about 5 macrotrichia in m; anal point somewhat triangular with 2 small spinulae between anal crests; anal tergite with about 4 median setae and 2 moderately large microtrichia- free areas on either side of anal point base; some microtrichiae present between crests at base of anal point; superior volsella oval; digitus well developed, broad, extending beyond median margin of superior volsella; median volsella moderately large with some simple and subulate lamellae not reaching 1/2 length of inferior volsella. Remarks The species keys to the T. eminulus species group (Cranston et al., 1989), and separates from the other species in this group by the low LR1 , low AR, broad digitus and smooth apex of the anal point. The apparently most closely related species is T. ogasatertius which has a much longer median volsella with apical, spatulate lamellae, T. tonebeceus which has a higher AR and thinner digitus and T. okuboi which has a more delicate digitus, larger median volsella and higher LR1 . T. ogasaquartus is only reported from its type locality. Tanytarsus ogasatertius Sasa & Suzuki (Figs 26–27) Tanytarsus ogasatertius Sasa & Suzuki, 1997: 322, Fig. 3. Holotype ♂ (NIES Type no. A307: 17) Japan, Bonin Islands, Chichijima, Sakaiura, sweep net, 14.IV.1996, leg. H. Suzuki [examined]. Diagnosis The following combination of characters separates T. ogasatertius from other Tanytarsus species. LR1 about 2.5; AR about 1; wing hairy, about 6 macrotrichia in m. Anal point somewhat triangular with 2 small spinulae between anal crests; anal tergite with about 6 median setae and 2 small microtrichia- free areas on either side of anal point base; some microtrichiae present between crests at base of anal point; superior volsella oval; digitus well developed, broad, extending beyond median margin of superior volsella; median volsella long and thin with some simple and 3–4 broad, spatulate lamellae towards apex almost reaching tip of inferior volsella. Remarks The species keys to the T. eminulus species group if one regards the few microtrichia on the anal point to constitute a ‘field’ between the anal crests (Cranston et al., 1989). The species separates from the other species in this group by the broad digitus, smooth apex of the anal point and the long median volsella with apical spatulate lamellae. Apparently, the most closely related species is T. ogasaquartus which has a considerably shorter median volsella with subulate lamellae and T. yunosecundus which has numerous subulate lamellae on the median volsella. Not regarding the microtrichia on the anal point to constitute 23 Figures 86–108. Larval mandibles, antennae, menta, premandibles and postoccipital plates. Figures 86, 91, 96, 101, 106. Tanytarsus bathophilus; Figures 87, 92, 97, 102. Tanytarsus formosanus; Figures 88, 93, 98, 103, 107. Tanytarsus mendax; Figures 89, 94, 99, 104, 108. Tanytarsus oyamai; Figures 90, 95, 100, 105. Tanytarsus uraiensis. 24 a ‘field’, however, will place T. ogasatertius in the quite different T. mendax species group (Cranston et al., 1989). Tanytarsus okuboi Sasa & Kikuchi (Figs 28–31) Tanytarsus okuboi Sasa & Kikuchi, 1986: 28, Fig. 4N. Holotype ♂ (NIES Type no. A98: 24) Japan, Tokushima, LT at rice paddies, 9.X.1984, leg. M. Sasa & M. Kikuchi; 3 paratypes ♂♂ (NIES A98: 24, 29, 30) as holotype [all examined]. Sasa (1998) key. Tanytarsus miyakoflavus Sasa & Hasegawa, 1988: 233, Fig. 1. Holotype ♂ (NIES Type no. A64: 63) Japan, Miyako Island, 2.II.1982, leg. M. Sasa & H. Hasegawa; paratypes: 2 ♂♂ (NIES A64: 64, 66) as holotype [all examined]. Syn. n. Tanytarsus oyabepallidus Sasa et al., 1988: 40, Fig. 6A. Holotype ♂ (NIES Type no. A136: 71) Japan, Toyama, Oyabe River, site C-4, 19.VIII.1987, leg. Sasa, Kawai & Ueno; 6 paratypes ♂♂ as holotype [all examined]. Syn. n. Tanytarsus simantoopeus Sasa et al., 1998: 63, Fig. 16. Holotype ♂ (NIES Type no. A358: 56) Japan, Shikoku Island, Shimanto River, Nakamura, LT, 26.IV.1998, leg. M. Sasa, H. Suzuki and T. Sakai; 5 paratypes ♂♂ (NIES A359: 60, 61, 67; A361: 15, 17) as holotype [all examined]. Syn. n. Tanytarsus tusimatheius Sasa & Suzuki, 1999a: 29, Fig. 33. Holotype ♂ (NIES Type no. A355: 87) Japan, Tsushima Island, Nitagawa River, sweep net, 26.III.1998, leg. H. Suzuki [examined]. Syn. n. Tanytarsus ikifegeus Sasa & Suzuki, 1999b: 151, Fig. 7. Holotype ♂ (NIES Type no. A357: 82) Japan, Iki Island, Katsumoto Dam, Sweeping, 28.III.1998, leg. H. Suzuki; 2 paratypes ♂♂ (NIES A357:83–84) as holotype [examined]. Syn. n. Diagnosis The following combination of characters separates T. okuboi from other Tanytarsus species. AR c. 1.0; LR1 c. 2.6. Anal point with granulose apex and spinulae in 1 row between anal crests; anal tergite with about 4 median setae and 2 small microtrichia- free areas on either side of anal point base; some microtrichiae present between crests of anal point; superior volsella oval; digitus present, about 1/2 length of superior volsella, usually barely extending beyond median margin of superior volsella; median volsella moderately large with some simple and subulate lamellae never reaching tip of inferior volsella. Pupa described by Sasa & Kikuchi (1986), but not possible to separate from pupae of other species in the T. eminulus group by the characters described. Remarks The species keys to the T. eminulus species group (Cranston et al., 1989) and separates from the other species in this group by having a much shorter median volsella. T. oyabepallidus and T. miyakoflavus were collected from quite different areas of Japan, namely Oyabe River (at about 36.5◦ N) and a rice paddy on Miyako Island (at about 25◦ N). T. okuboi is known from a rice paddy area in the suburbs of Tokushima city (about 34◦ N), while T. simantoopeus is known from southern Shikoku Island, T. ikifegeus from Iki Island and T. tusimatheius is known from Tsushima Island. The Miyako Island has a different climate compared with the other localities, but the habitat of the specimens collected on Miyako and near Tokushima is the same. T. tusimatheius and T. simantoopeus were both collected near larger rivers. Since only minute morphological differences are visible, I choose to synonymise the six species and conclude that the species is relatively tolerant to different climates and lentic and lotic habitats. T. okuboi is probably closest related to T. tonebeceus, T. tamaundecimus and T. ogasaquartus which apparently only differs in having a significantly different LR1 , smooth anal point apex and more setae on wing. In addition to the localities mentioned above, T. okuboi has been recorded from West Bengal, India (Datta et al., 1992). These specimens have, however, not been available for examination so this record still is questionable. Tanytarsus oscillans Johannsen (Figs 34–35) Tanytarsus oscillans Johannsen, 1932: 35. Holotype ♂ (BMNH) Indonesia, Java, Buitzenborg, Pond in the Botanical Garden, 15.IX.1928, leg. A. Thienemann [examined]. Tanytarsus cultellus Chaudhuri & Datta, in Datta et al., 1992: 44, Figs 9–11. Holotype ♂ (UBI Type no. 198), India, West Bengal, Alipurduar, 26◦ 25′ N 91◦ 5′ E, 60.9 m a.s.l., pond side, 15.IV.1987, leg. A. K. De; 1 paratype ♂ as holotype, except collected 15.IV.1990 [both examined] Syn. n. Tanytarsus sibafegeus Sasa et al., 1999: 189, Fig. 9. Holotype ♂ (NIES Type no. A316: 23) Japan, Ishikawa Prefecture, Shibayamagata Lake, Sumita, 16.VII.1998, leg. M. Sumita [examined]. Syn. n. Diagnosis Tanytarsus oscillans can be separated from other 25 Tanytarsus species by the following combination of characters. Wing hairy, with about 10 macrotrichia in m and 14 on Cu; AR about 1.1, frontal tubercles 10– 15 µm. Anal tergite bands separated, almost reaching anal crests; anal tergite with shoulders on posterior margin, 2 median setae and small microtrichia- free areas on either side on anal point base; anal point relatively long with pointed apex and 5 large spinulae between well developed crests, microtrichia extensively distributed in between; superior volsella oval, slightly narrower in middle, medially directed, with 5–7 dorsal and 3 closely set median setae; digitus relatively long, finger like, extending to or barely beyond apex of superior volsella; median volsella short, knoblike with 4 medially curved foliate lamellae and 4–5 setose lamellae; inferior volsella thin, somewhat medially curved with slightly enlarged apex. Remarks Although the examined paratype specimen of T. cultellus is not mentioned in the original publication, I believe it is a part of the original type series (Datta et al., 1992) since it is collected at the type locality by A. K. De, and bears a paratype label. The specimen is inseparable from the original description and it is probable that the absence of frontal tubercles and low AR observed by Datta et al. (1992) in the description of T. cultellus are due to misinterpretations as are several hypopygial features by Sasa et al. (1999) for T. sibafegeus. T. oscillans keys to the T. eminulus species group (Cranston et al., 1989) and is very similar to T. smolandicus Brundin, 1947 and T. unagiseptimus. It differs, however, from T. smolandicus by having only two median setae on the anal tergite, shorter frontal tubercles and lower AR and from T. unagiseptimus by having microtrichia extensively distributed between anal point crests and a narrow inferior volsella. T. oscillans has been recorded from India, Indonesia and Ishikawa Prefecture, Japan. Tanytarsus ovatus Johannsen (Figs 36–37) Tanytarsus ovatus Johannsen, 1932: 35. Holotype ♂ (BMNH) Indonesia, Middle Java, Sarangan, Lake Pasir, 8.XII.1928, leg. A. Thienemann [examined]. Tanytarsus insulus (Guha et al., 1985). Tanytarsus minimus Guha et al., 1985: 37, Fig. 11. Replacement name given by Mazumdar et al. (1998). Holotype ♂ (UBI Type no. 138), India, S. Andaman Island, Wright Myo, 26.V.1982, leg. R. Sharma [not examined]; 1 paratype ♂ as holotype (ZMBN), 1 paratype ♂ as holotype, except collected 27.V.1982 [both examined]. Syn. n. Additional material examined: 2 ♂♂ (UBI and ZMBN) India, Andaman Islands Wright Myo, 27.V.1982, leg. R. Sharma. Diagnosis Tanytarsus ovatus can be separated from all other Tanytarsus species by the following character combination. AR about 1.2, LR1 larger than 3.1. Hypopygium with separated anal tergite bands that almost reach anal crests, 5–6 median tergite setae at anal point base; anal point with a large field of microtrichia and several trifid spinulae in 1 row between crests; microtrichiafree areas on either side of anal point base. Superior volsella kidney- to inversely pear-shaped, apex medially directed, 6–7 dorsal and 3 median setae of which posterior is smaller and situated on ventral side of superior volsella, dorsolateral and ventromedian microtrichia present; digitus small, pointed; median volsella well developed with 2–3 broad, pectinate and several setose lamellae; inferior volsella club- shaped with obvious dorsomedian lobe. Remarks The holotype of T. insulus was not available for examination, and although the original drawing of the hypopygium (Guha et al., 1985: 37, Fig. 11b) is more similar to Tanytarsus formosanus Kieffer than to the paratypes examined, the description is more similar to the paratypes. Especially the LR1 value is much higher in T. insulus, and clearly different from that of T. formosanus (1.9–2.7). It is possible that the type series of T. insulus is heterogenous, and the holotype conspecific with T. formosanus. But until it is examined, I choose to synonymise T. insulus with T. ovatus. The three specimens collected 27.V.1982 are not specifically mentioned in the original description as part of the type material. One slide bears a paratype label, the two other slides are wrongly labelled Tanytarsus vinculus. T. ovatus keys to the T. gregarius species group if one regard the digitus to be inconspicious (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971) except having a smooth anal point apex, pectinate lamellae on the superior volsella and a somewhat larger digitus. The probably closest related species is T. miyakobrevis Sasa & Hasegawa, 1988 which also has quite extensive fields of microtrichia on the superior volsella, but T. ovatus separates by having a differently shaped anal point, a larger digitus, larger 26 LR1 and pectinate lamellae on the median volsella. T. ovatus has been recorded from West Bengal in addition to the Andaman Islands and Java (Mazumdar et al., 1998), but this record can not be confirmed in the present study. Tanytarsus oyamai Sasa (Figs 32, 33, 63, 70, 78, 89, 94, 99, 104, 108) Tanytarsus oyamai Sasa, 1979: 3, Fig. 5–7. Holotype ♂ (NIES Type no. A13: 51) Japan, Tsukuba, National Institute for Environmental Studies, 22.VIII.1978, leg. M. Sasa; 7 paratypes (NIES A13): 2 P, ♂, as holotype except 23.VIII.1978, 5 L as holotype except 21.VIII.1978 and 08.VIII.1978 [all examined]. Sasa & Kikuchi (1995), Sasa (1998) adult male, keys; Sasa (1980), Sasa (1989a), Sasa & Kikuchi (1986), Sasa et al. (1988), Sasa et al. (1998), Kawai et al. (1998), Kawai et al. (1999) additional records and ecology. Additional material examined: 1 P, ♂ (NIES A13: 11) as holotype, except 28.VI.1977; 1 ♂ (NIES A15: 01) as holotype, except 07.X.1976; 4 ♂♂ (NIES A137: 23) Japan, Toyama, Oyabe River, 19.VIII.1987, leg. M. Sasa, K. Kawai & R. Ueno. Diagnosis T. oyamai is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; usually strong thoracic colorations anteriorly on scutum, laterally on scutum under parapsidal suture, basally on scutellum, postnotum, median anepisternum II, epimeron II and preepisternum; abdominal tergites uniformly dark brown; wing with sparse setation (restricted to apical 1/4), Cu bare, r4+5 with a greater or equal amount of setae as m1+2 ; AR about 1.0; spinulae in 1 row between well developed anal crests; superior volsella abruptly tapered towards median elongated apex, about 6 dorsal setae of which 3 are placed laterally, 3 median setae where 2 most apical are larger than innermost seta; digitus short; median volsella relatively short and slender with some subulate and setose lamellae (often difficult to see, and dissection of the median volsella might be necessary); inferior volsella finger like, only weakly medially curved. Pupa with large conical cephalic tubercles; long evenly tapered thoracic horn with chaetae longer than width of horn in 1 row on mid 2/3; strong points in transverse, anterior band on TII not connected with the posterior field of weaker points; posterior, short, elongate spine patches on TIII, oval, anterior spinule patches on TIV–V, round on TVI; segment VII with 1–2 lateral taeniae, segment VIII with 5 lateral taeniae where 2 posterior most taeniae usually placed close together. Larval head with antennae moderately long, about 0.6 times length of head capsule; Lauterborn organ moderately large, almost as long as antennal segment 4+5 combined, on pigmented pedestal barely reaching past apex of antennal segment 5; AR about 1.6; SI and chaetulae basales plumose, SII, S3, chaetae and chaetulae laterales all apparently simple; mandible with 1 large dorsal tooth; premandible well pigmented with 5 teeth; 4 large anal tubules. Remarks T. oyamai keys to the T. mendax species group (Cranston et al., 1989) and is apparently most closely related to T. formosanus and T. mendax. Males of T. oyamai differs from those of T. formosanus by having a narrower, somewhat longer apex of anal point, an abruptly tapering superior volsella without lateral microtrichia, a small, tubercle- like digitus, a fingerlike inferior volsella and by having an uniformly dark brown coloured abdomen; pupae by having spinules instead of spines on TIV and a lower posterior thoracic mound; larvae by having somewhat longer styli of LO, larger LO and only one large dorsal tooth on mandible. Males of T. oyamai differ from those of T. mendax by having a differently shaped anal point and superior volsella; pupa by having long chaetae, 4× width of thoracic horn, in one row on middle 1/3 of horn and spines instead of spinules on TIII; larva by having shorter Lauterborn organ (including pedicel) and a lower AR. T. oyamai has been recorded from eutrophic lakes and rivers in Ibaraki, Tokyo, Kagoshima, Yamanashi, Tokushima, Shiga, Toyama, Nagasaki Hiroshima, Ishikawa and Kochi prefectures, Japan (Sasa, 1979, 1980, 1985a,b, 1987, 1993a; Sasa & Kikuchi, 1986; Sasa et al., 1988, 1989a, 1991b, 1998, 1999; Ueno et al., 1993; Kawai et al., 1998, 1999; Sasa & Ogata, 1999). Tanytarsus pollexus Chaudhuri & Datta (Figs 38– 39) Tanytarsus pollexus Chaudhuri & Datta in Datta et al., 1992: 51, Figs 31–35. Holotype ♂ (UBI Type no. 202), India, Mongpu, 27◦ N, 88◦ E, 2000m a.s.l., 8.IV.1984, leg. T. K. Datta [examined]. Diagnosis The following combination of characters separates 27 Tanytarsus pollexus from all other Tanytarsus species. AR c. 1.1; LR1 large, about 3.2; large frontal tubercles; anal tergite with separate tergite bands almost reaching anal point base, 4 median setae, about 14 apical setae, small microtrichia- free patches on either side of anal point base; anal point with about 6 small trifid spinulae in 1 row between well developed anal crests; superior volsella oval with about 7 dorsal and 3 median setae; digitus small, inconspicuous; median volsella with 4 broad, foliate and c. 10 setose lamellae; inferior volsella straight with medioproximal hump and dorsoapical, medially directed lobe. Remarks T. pollexus keys to the T. eminulus group due to the digitus being present and well discernible (Cranston et al., 1989), and fits the group diagnosis well, except for having a smaller digitus (Reiss & Fittkau, 1971). The probably closest related species is T. okuboi which only differs in having a lower LR1 , usually a larger digitus, a granulose anal point apex and somewhat differently shaped apodemes, and T. ikifegeus which has a significantly lower LR1 . T. pollexus is recorded only from India. Tanytarsus shouautumnalis Sasa (Figs 40–43) Tanytarsus shouautumnalis Sasa, 1989a: 39, Fig. 13. Holotype ♂ (NIES Type no. A152: 88) Japan, Toyama, Shou River, St. 4, 25.VIII–5.IX.1988, leg. M. Sasa; 1 paratype ♂ (NIES A152: 89) as holotype [both examined]. (Sasa & Suzuki, 1993), adult male. Tanytarsus tsutaprimus Sasa, 1991a: 78, Fig. 2.4. Holotype ♂ (NIES Type no. A202: 59) Japan, Toyama, Tsuta- onsen, 1.VII–21.VII.1989, leg. M. Sasa [examined]. Syn. n. Sasa & Suzuki (1999a) adult male. Tanytarsus tokarajekeus Sasa & Suzuki, 1995: 268, Fig. 10. Holotype ♂ (NIES Type no. A290:46) Japan, Kagoshima, Tokara Islands, Nakanoshima, LT, 20.V.1994, leg. H. Suzuki [examined]. Syn. n. Tanytarsus tusimatlemeus Sasa & Suzuki, 1999a: 32, Fig. 38. Holotype ♂ (NIES Type no. A353: 12) Japan, Tsushima Island, Izuhara, Azugawa River, sweep net, 23.III.1998, leg. H. Suzuki [examined]. Syn. n. Tanytarsus tusimatopeus Sasa & Suzuki, 1999a: 34, Fig. 41. Holotype ♂ (NIES Type no. A355: 70) Japan, Tsushima Island, Kamitsushima, Toyo, sweep net, 26.III.1998, leg. H. Suzuki [examined]. Syn. n. Tanytarsus yakugeheuus Sasa & Suzuki, 2000: 64, Fig. 17. Holotype ♂ (NIES Type no. A383: 93) Japan, Kyushu, Yakushima Island, Shirotani River, sweep net, 27.III.1999, leg. H. Suzuki [examined]. Syn. n. Diagnosis T. shouautumnalis is separable from other Tanytarsus species by the following combination of characters. Head with small frontal tubercles; wing hairy with more than 3 macrotrichia in m; AR about 1.0; LR1 > 2.4. Anal tergite bands thin and relatively short, not reaching anal crests; microtrichia- free areas present on either side of anal point base; microtrichia present between crests at base of anal point; about 6 long median tergite setae placed close to anal point base; spinulae in irregular or regular row between well developed anal crests; anal point normally developed, with truncate apex; superior volsella somewhat triangular in shape with dorsolateral microtrichia, 3 median setae where 2 most apical larger than basal seta; digitus long, clearly extending beyond median margin of superior volsella; median volsella relatively short and slender with 2 – 3 spatulate and some setose lamellae; inferior volsella finger like, weakly curved medially. Remarks T. shouautumnalis keys to the T. eminulus species group (Cranston et al., 1989), but does not resemble the European species in this group due to the shape of the superior volsella and the small frontal tubercles (Reiss & Fittkau, 1971). Although T. shouautumnalis and T. tsutaprimus syn. n. originally were described from a river and a hot spring respectively (Sasa, 1989a, 1991a), there are, however, no obvious morphological differences between the holotypes. The fact that T. tsutaprimus has been collected alongside Azugawa River on Tsushima island (Sasa & Suzuki, 1999a) and that T. shouautumnalis has been found in lakes with low PO4 concentration (Kawai et al., 1999) confirms that this species is probably not typical for spring fauna, but rather quite general in habitat choice. T. shouautumnalis also has been recorded from several other rivers in South and Southwestern Japan, mostly as the synonym species listed above. I suspect the authors did not detect these synonyms, although some of them even were collected at the same locality at the same date, because of distortion of the genitalia in the mounting process thus making comparison difficult (compare anal points in Figs 40, 41). T. shouautumnalis can be separated from similar Asian Tanytarsus species by the shape of the anal point and superior volsella. 28 Tanytarsus shoudigitatus Sasa (Figs 44–46) Tanytarsus shoudigitatus Sasa, 1989a: 40, Fig. 16. Holotype ♂ (NIES Type no. A154: 21) Japan, Toyama, Shou River, Kouzochinai, St. 2, 7.II.1989, leg. M. Sasa; 3 paratypes ♂♂ (NIES A154: 22–24) as holotype. Tanytarsus togasiroidus Sasa & Okazawa, 1991: 117, Fig. 7.9. Holotype ♂ (NIES Type no. A186: 50) Japan, Toyama, tributary of Toga River, 11.IX.1990, leg. M. Sasa & T. Okazawa [examined]. Syn. n. Diagnosis T. shoudigitatus is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; wing hairy with about 10 macrotrichia in m; AR about 1.0; LR1 > 3.0; anal tergite bands thin almost reaching anal crests; small microtrichia- free areas on either side of anal point base; microtrichia present between crests of anal point; about 4 short median tergite setae placed close to anal point base; small spinulae in irregular row between well developed anal crests; anal point normally developed, with rounded apex; superior volsella oval with distinct ventral tubercle near apex, 5 long, dorsal setae and 3 median setae where 2 most apical larger than innermost seta; digitus long, clearly extending beyond posteromedian margin of superior volsella; median volsella moderately large with 4–5 subulate, occasionally 1 pectinate and some setose lamellae; inferior volsella club- shaped, weakly curved medially. Remarks T. shoudigitatus keys to the T. eminulus species group (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971). It is separated from other members in this group by its high LR1 and the distinct ventral tubercle on superior volsella. The species is recorded only from Toyama and Ishikawa prefectures (Sasa et al., 1999). Tanytarsus takahashii Kawai & Sasa (Figs 47–48) Tanytarsus takahashii Kawai & Sasa, 1985: 22, Fig. 8. Holotype ♂ (HIU) Japan, Hiroshima Prefecture, Ohta River, rearing of bottom sample, St. 4, 28.VIII.1981, leg. K. Kawai [not examined]; 3 paratypes: 2 ♂♂ as holotype except collected at St. R5, 7.VI.1981; 1 ♂ as holotype, except collected at St. 12, 10.IX.1981 [examined]. Diagnosis T. takahashii is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; wing hairy; LR1 > 3; AR about 1.0. Anal tergite bands almost reaching anal crests; small microtrichia- free areas at base of anal point; only a few microtrichia present between crests of anal point; 2 – 3 minute to small median tergite setae at base of anal point; small spinulae in a row between well developed anal crests; anal point with parallel sides and rounded apex; superior volsella almost tear drop- shaped, abruptly tapering on apical half, with dorsolateral microtrichia, 5 – 6 dorsal setae and 3 median setae where 2 most apical larger than innermost seta; digitus large, mostly not covered by and extending beyond posteromedian corner of superior volsella; median volsella long, thin, with several long and thin subulate or setiform lamellae; inferior volsella straight. Remarks T. takahashii keys to the T. mendax species group (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971) if one does not regard the few microtrichia between the anal crests to constitute a ‘field of microtrichiae’. The actual shape of the superior volsella and the digitus is similar to several other species in this group (see T. calorifontis Fig. 8), but the long median volsella with long and thin lamellae will separate T. takahashii from all other members of this species group. Considering the few microtrichiae to constitute a field of microtrichia will put T. takahashii in the T. eminulus species group, while due to the shape of the median volsella it shows similarities to the T. lestagei aggregate. The high LR1 and shape of superior volsella will, however, make T. takahashii unmistakably different from other members in this aggregate. A phylogeny of these species groups may reveal where this species probably belongs. T. takahashii has been recorded from Hiroshima, Shiga, Gifu, Tokushima, Fukushima, Ishikawa and Toyama prefectures (Sasa, 1987, 1989b, c, d, 1990, 1993b; Sasa et al., 1999; Kawai & Sasa, 1985; Kawai et al., 1998) Tanytarsus tamaundecimus Sasa Tanytarsus tamaundecimus Sasa, 1980: 27, Figs 30– 32. Holotype ♂ (NIES Type no. A43: 91) Japan, Tokyo Prefecture, Minamiasakawa River, 17.VIII– 3.IX.1979, leg. Sasa et al. 5 paratypes: 1 ♂, 1 ♀ and 3 29 Pex as holotype [not examined]. Diagnosis The following characters from the original description (Sasa, 1980) can be regarded as diagnostic. Male wing moderately hairy, with few macrotrichia in m; AR about 0.8, frontal tubercles c. 30 µm long, LR1 about 3.3; anal tergite with separated anal tergite bands, c. 3 median setae close to anal point base; anal point somewhat triangular, apically rounded, with trifid spinulae in a row and microtrichia between well developed anal crests; superior volsella somewhat kidney- shaped, 4 dorsal, 3 median setae; digitus large, extending beyond median margin of superior volsella; median volsella relatively long, reaching past apex of superior volsella, with some medially directed lamellae; inferior volsella club- shaped. Pupal thoracic horn long with small chaetae distributed on apical half; TII with large square patch of shagreen, small bare area in middle; TIII with long, slightly laterally curved spine patches consisting of long spines; TIV with long spine patches, anterior end strongly medially curved, posterior end straight, consisting of long spines; TIV and TV with small rounded anterior patches of spinules; 2 lateral taeniae on segment VII, 5 on segment VIII; anal fringe with 38–42 taeniae. Remarks Unfortunately, box 43 of the collection was missing when I visited NIES and is probably still used by Prof. Sasa in Toyama. T. tamaundecimus keys to the T. eminulus species group (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971) except for the AR which is somewhat lower in T. tamaundecimus. The species is probably closely related to T. medius, T. yunosecundus, T. ogasaquartus and T. tonebeceus, but adults can be separated from T. medius and T. yunosecundus by having a lower AR and a much higher LR1 , and from T. ogasaquartus and T. tonebeceus by a higher LR1 . The pupa of T. tamaundecimus can be separated from those of T. medius by having spine patch of TIII only slightly curved, chaetae only on distal half of thoracic horn and more shagreen on TII, and from those of T. yunosecundus by having anterior spine patch on TIV strongly medially directed and chaetae only on apical half of thoracic horn. No pupa are described for T. ogasaquartus and T. tonebeceus. T. tamaundecimus is only recorded from Tokyo Prefecture (Sasa, 1980, 1983). Tanytarsus tonebeceus Sasa & Tanaka Tanytarsus tonebeceus Sasa & Tanaka, 2000: 1, Fig. 2. Holotype ♂ (NIES Type no. A391: 19) Japan, Gunma Prefecture, Taisho Bridge, 1.VII.1999 [not examined] Diagnosis The following characters from the original description (Sasa & Tanaka, 2000) can be regarded as diagnostic. AR c. 1.0; LR1 about 2.0; wing hairy with setae almost on entire surface; hypopygium with separate anal tergite bands, 4 median tergite setae close to base of anal point; anal point triangular with 4 spinulae and possibly microtrichia between crests; superior volsella oval, with 4 dorsal and apparently only 2 median setae (although probably 3 are present as a basal seta might have been overlooked); digitus moderately long, narrow, reaching median margin of superior volsella; median volsella moderately long, stem reaching to distal half of superior volsella, with several medially and posteriorly directed lamellae. Remarks If microtrichiae are present between the anal point crests, T. tonebeceus will key to the T. eminulus species group (Cranston et al., 1989). The shape of the anal point, superior and median volsella also suggests that the species should belong here. The probably closest related Asian species are T. okuboi and T. tamaundecimus which differ by having a higher LR1 and less setae on the wing surface, T. yunosecundus which has a longer median volsella and T. ogasaquartus which has a lower AR, more robust digitus and a shorter median volsella. Tanytarsus tusimatneous Sasa & Suzuki (Figs 49– 50) Tanytarsus tusimatneous Sasa & Suzuki, 1999a: 33, Fig. 40. Holotype ♂ (NIES Type no. A354: 95) Japan, Tsushima Island, Sumokawa, sweeping net, 25.III.1998, leg. H. Suzuki [examined]. Tanytarsus tusimatpequeus Sasa & Suzuki, 1999a: 35, Fig. 42. Holotype ♂ (NIES Type no. A353: 92) Japan, Tsushima Island, Uchiyama, sweeping net, 24.III.1998, leg. H. Suzuki. [examined]. Syn. n. Diagnosis T. tusimatneous is separable from other Tanytarsus species by the following combination of characters. Head with minute frontal tubercles; wing moderately hairy without macrotrichia in m; AR < 0.8; LR1 about 30 2.3. Anal tergite bands well separated, ending far from anal crests; microtrichial pattern on anal tergite uncertain, but minute microtrichia- free areas on each side of anal point probably exist; microtrichia absent between crests of anal point; about 8 median tergite setae between anal tergite bands and anal point base; about 5 small spinulae placed irregularly in rounded pit on anal point; anal point short, triangular with pointed apex; superior volsella almost circular, but with almost straight median margin, 5–8 dorsal setae, 3 median setae set close to each other, 2 most apical larger than innermost seta; digitus long, broad, extending beyond median margin of superior volsella; median volsella short, with some short, thin, subulate and setose lamellae reaching approximately half length of inferior volsella; inferior volsella slightly Sshaped, with about 11 setae. Remarks T. tusimatneous keys to the T. chinyensis species group (Cranston et al., 1989) and not to the T. mendax group as claimed be Sasa & Suzuki (1999a). The species fits the group diagnosis given by Reiss & Fittkau (1971), but does not have the basal most median seta of the superior volsella placed on the digitus suggested as diagnostic for the group by Ekrem (2001a). T. tusimatneous separates from other species in this group by having a pointed apex of the anal point and a more rounded superior volsella. The synonymy was probably not detected by Sasa & Suzuki, op. cit. due to mounting differences. The species description might change slightly in the future when more material is examined. Tanytarsus unagiseptimus Sasa (Figs 51–52) Tanytarsus unagiseptimus Sasa, 1985a: 47, Fig. 19, 20. Holotype ♂ (NIES Type no. A78: 51) Japan, Kyushu, Lake Unagi, bottom sediment rearing, 6.II.1981, leg. M. Sasa [examined]. Diagnosis T. unagiseptimus is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; wing hairy with several macrotrichia in m; AR > 1.0; LR1 2.4–2.7. Anal tergite bands thin almost reaching anal crests; small microtrichia- free areas on either side of anal point base; microtrichia present between crests of anal point; 1–3 short median tergite setae placed close to anal point base; large spinulae in regular row between relatively low anal crests; anal point parallel sided, with rounded apex; superior volsella oval with 5–8 short, dorsal setae and 3 median setae where 2 apical larger than basal most seta; digitus barely extending beyond posteromedian corner of superior volsella; median volsella small, knob-like with 3–5 subulate and several setose lamellae; inferior volsella with only slightly enlarged apex. Pupa with bare thoracic horn, TIII with about 20 µm long spines in patches which are about 1/3 length of tergite; shorter spines and spinules in anterior, oval patches on TIV; spinules in small, anterior, round patches on TV and TVI; 1 lateral taenia on segment VII, 5 on segment VIII. Remarks T. unagiseptimus keys to the T. eminulus species group (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971). The male can be separated from other members in this group by its evenly rounded anal point, low number of median setae on anal tergite and a small knob- like median volsella with three to five subulate lamellae. The pupa separates from the other species in the T. eminulus species group by a bare thoracic horn and the quite short spine patches on TIII–IV. The closest related Asian species apparently is T. oscillans which has extensive distribution of microtrichia between anal point crests and a pointed anal point apex. T. unagiseptimus is recorded from lowland mesotrophic and eutrophic lakes and river pools in central and southern parts of Honshu, Japan (Sasa, 1985a, 1987; Ueno et al., 1993; Sasa & Suzuki, 1995; Kawai et al., 1998; Kawai et al., 1999). Tanytarsus uraiensis Tokunaga (Figs 53, 64, 72, 81, 90, 95, 100, 105) Tanytarsus uraiensis Tokunaga, 1938: 350, Fig. 26, 27. Holotype ♂, Taiwan (Formosa), Urai, Hot spring, 14.IX.1924, leg. R. Takahashi [not examined]; 16 paratypes: 2 ♂, 2 P, ♀, 1 ♀, 1 Pex, 10 L (HIU) as holotype [examined]. Kikuchi & Sasa (1990), Sasa & Kikuchi (1995) adult male. Diagnosis T. uraiensis is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; wing hairy; AR about 1.0. Anal tergite bands almost reaching anal crests; microtrichiafree area around base of anal point; microtrichia absent between crests of anal point; 2–5 well developed median tergite setae placed at end of anal tergite bands; large spinulae in regular row between well developed anal crests; anal point parallel sided, with rounded 31 apex; superior volsella almost egg- shaped, evenly tapering towards apex, with ventral, apical lip, 4–5 dorsal setae and 3 median setae where 2 most apical larger than innermost seta; digitus apparently absent, but can also be small and hyaline; median volsella moderately long, narrow, with 1 or 2 thin, subulate and some setose lamellae; inferior volsella with only slightly enlarged apex and more than 12 setae. Pupal thoracic horn with chaetae longer than width of horn in 1 row extending about 3/4 length of thoracic horn; cephalic tubercles large, conical; TIII with about 80 µm long spines in patches which are about 1/3 length of tergite; shorter spines and spinules in anterior, oval patches on TIV; spinules in small, anterior, round patches on TV and TVI; 2 lateral taeniae on segment VII, 5 on segment VIII; about 37 taeniae in anal fringe. Larva of T. mendax group type with 1 large dorsal tooth on mandible, small LO on stylus reaching to apex of antennal segment 5; SII apparently simple although Tokunaga (1938: 353, Fig. 27o) reports labial appendages to be atrophied; AR about 1.2; relatively straight ventromental plates. Remarks T. uraiensis keys to the T. mendax species group (Cranston et al., 1989), and fits the group diagnosis (Reiss & Fittkau, 1971). It can be separated from other members in this group by its pale colour, evenly rounded anal point, presence of median setae on anal tergite, evenly tapered superior volsella, the apparent absence of a digitus and by having only 1 or 2 thin subulate lamellae on median volsella. The pupa separates from the other species in the T. mendax species group except T. formosanus by having chaetae longer than width of thoracic horn in 1 row over 3/4 length of horn, spines and spinules on TIV, 2 lateral taeniae on segment VII and about 37 taeniae in anal fringe. The pupa pictured in Tokunaga (1938) differs from the types examined by having chaetae over the apical 7/8 length of thoracic horn (compared to covering 2/3 of thoracic horn length, Fig. 72), a wide field of shagreen on TII (compared to well separated fields, Fig. 81), broad spine patches on TIII (compared to a row of spines, Fig. 81) and only 1 lateral taenia on segment VII (compared to 2, Fig. 81). It is both possible that Tokunaga’s drawings are inaccurate or that the pupae remaining in the type material are of a different species, namely T. formosanus. Larva very similar to larvae of T. formosanus and T. oyamai, but can apparently be separated from those of T. formosanus by a somewhat longer stylus of LO (reaching apex of an- tennal segment 5) and only 1 dorsal tooth on mandible. The larva of T. oyamai is, as far as I can see, not separable from that of T. uraiensis. T. uraiensis is also quite similar to T. calorifontis of which the males can be separated from T. uraiensis by the absence of median anal tergite setae and higher AR (c. 1.0 compared to < 0.9 for T. uraiensis). The pupa of T. uraiensis can be separated from those of T. calorifontis by having large cephalic tubercles and more taeniae in the anal fringe (c. 37 compared to c. 32). The holotype was not possible to locate. The paratype material examined had been in alcohol for 75 years, and was difficult to macerate in KOH. Only 1 male hypopygium was available for examination, and the other males including the holotype might be lost. Several characters, even with an interference contrast microscope, were impossible to see properly and the diagnosis above might change slightly if fresh material is examined in the future. Kikuchi & Sasa (1990) recorded the species from Lake Toba, Sumatra in 1989, but although similar, their description differs from Tokunaga (1938) by reporting a different LR1 (2.88 compared to 1.59 in Tokunaga) and from the examined paratype male in having more median setae and no microtrichia- free area on anal tergite and only two median setae on the superior volsella. However, separate front legs found in the vial containing the paratypes have a LR1 of 2.54, thus the measures of Tokunaga (1938) could have been inaccurate. In addition, the microtrichial pattern of the anal tergite is sometimes difficult to observe, and the microtrichiafree area has not been observed in Kikuchi & Sasa’s (1990) description of neither T. euformosanus nor T. uraiensis. Neither have the three median setae and dorsolateral microtrichia on the superior volsella been drawn, structures that are present on the holo- and paratypes (see diagnosis on T. formosanus). It is thus possible that Kikuchi found T. uraiensis at the shore of Lake Toba, but this has to be confirmed by examination of this material which was not available to me for the present study. Sasa & Kikuchi (1995) mentions that Hashimoto et al. (1981) has recorded T. uraiensis in Thailand. However, only the similarity of T. formosanus and T. uraiensis is discussed by Hashimoto et al., op. cit. and a new record is not mentioned. T. uraiensis is thus so far only with certainty known from its type locality on Taiwan. Tanytarsus yakuheius Sasa & Suzuki (Figs 54–55) Tanytarsus yakuheius Sasa & Suzuki, 2000: 64, Fig. 18. Holotype ♂ (NIES Type no. A384: 03) Japan, 32 Yakushima Island, Shirotani River, sweeping net, 27.III.1999, leg. H. Suzuki [examined]. Diagnosis T. yakuheius is separable from other Tanytarsus species by the following combination of characters. Head with large frontal tubercles; wing hairy with about 5 macrotrichia in m; AR about 1.0; LR1 about 2.9. Anal tergite bands ending far from anal crests; small microtrichia- free areas on each side of anal point; microtrichia present between crests of anal point; about seven median tergite setae between anal tergite bands and anal point base; small spinulae in a regular row between well developed anal crests; anal point short, triangular with rounded, slightly granulose apex; superior volsella oval, with several ventral, transverse ridges, 4 dorsal setae, 3 median setae where 2 most apical larger than innermost seta which is placed on small ventral tubercle; digitus triangular, not extending beyond median margin of superior volsella; median volsella long, with several long, thin, subulate and setose lamellae reaching beyond apex of inferior volsella; inferior volsella slightly club- shaped, with about 9 setae. Remarks T. yakuheius keys to the T. eminulus species group (Cranston et al., 1989), and fits the group diagnosis, except having a much shorter digitus (Reiss & Fittkau, 1971). This character, together with the short, triangular anal point, long median volsella and transverse, ventral ridges on the superior volsella distinguishes T. yakuheius from all other members of this group. The species is apparently most similar to T. kikuchii and T. yunosecundus due to the long median volsella, ventral, transverse ridges on superior volsella and a triangular anal point, but separates clearly from T. kikuchii by having a digitus, granulose anal point apex and lower LR1 value and from T. yunosecundus by having a shorter digitus, shorter median volsella, granulose anal point and only a slightly convex inner lobe of gonocoxite (opposed to an almost right angled lobe in T. yunosecundus). T. yakuheius is only reported from its type locality. Tanytarsus yunosecundus Sasa (Figs 56, 57, 65, 71, 74, 80, 85) Tanytarsus yunosecundus Sasa, 1984: 36, Fig. 14, 15, 17. Holotype ♂ (NIES Type no. A36:51) Japan, Honshu, Kanto, Nikko national park, Lake Yunoko, rearing of bottom sediment, 27.IV.–12.V.1979, leg. T. Iwakuma & Y. Sugaya; 5 paratypes: 1 ♂, 1 Pex (NIES A36: 53, 52) as holotype; 1 ♂ (NIES A36: 54) as holotype except emerged 13.VI.1979; 2 ♀♀ with Pex (NIES A36: 66, 68) as holotype, except emerged 14.VI.1979 and 26.V.1979. [all examined]. Sasa & Kikuchi (1986), Sasa (1988), Sasa & Kikuchi (1995), Sasa (1998), adult male, key. Diagnosis T. yunosecundus is separable from other Tanytarsus species except species in the T. lestagei aggregate by the following combination of characters. Head with large frontal tubercles; wing moderately hairy with cell m bare and Cu bare or with 2–3 macrotrichia; AR about 1.1; LR1 about 2.4. Anal tergite bands almost reaching anal crests; small microtrichia- free areas at base of anal point; microtrichia present between crests of anal point; about 3 small median tergite setae at anal point base; spinulae in a regular row between well developed anal crests; anal point triangular with rounded apex; superior volsella oval, with several ventral, transverse ridges, seven dorsolateral setae in a constricted area, 3 median setae with 2 most apical larger than innermost seta; digitus long, extending beyond posteromedian apex of superior volsella; median volsella long, curved, with several long, thin, subulate and setose lamellae reaching beyond apex of inferior volsella; inferior volsella slightly enlarged at apex, with about 11 setae. Pupal thoracic horn with small chaetulae evenly distributed on apical 4/5 of thoracic horn; cephalic tubercles minute; posterior thoracic mound very small; TI without shagreen patch; TIII with about 60 µm long spines in diagonal patches which are about 1/2 length of tergite, small round patch of shagreen anteriorly of spine patch; TIV with about 50 µm long spines anteriorly in straight, long patches, spines decreasing in length posteriorly and ending as spinules; spinules in small, anterior, oval patches on TV and TVI; TVII–IX with anterior, round shagreen patches; 2 lateral taeniae on segment VII, 5 on segment VIII; about 36 taeniae in anal fringe. Remarks The pupa described as T. yunosecundus by Sasa (1984) is wrongly associated (by mass rearing) and actually belongs to T. bathophilus Kieffer. This species is found at the same locality, but described as T. gregarius Kieffer (Sasa, 1984) (see T. bathophilus). The type series includes two different morphotypes of which one clearly belongs to the T. lestagei aggregate. Obviously, the present description of the pupa differs 33 from that of Sasa (1984), and it should be noted that the association although more probable, still is tentative. The larva of T. yunosecundus described by (Sasa, 1984) unfortunately was unavailable for examination, but this tentative association is probably correct since the mandible does not have the large, ventral, platelike tooth typical for T. bathophilus. In addition, the high AR and the long pedicel of LO recorded by Sasa is observed for several members of the T. lestagei aggregate and not in T. bathophilus. T. yunosecundus keys to the T. eminulus species group (Cranston et al., 1989), and is a clear member of the T. lestagei aggregate (Lindeberg, 1967; Reiss & Fittkau, 1971). It is difficult to separate from the European members in this aggregate, and until present I am not able to separate T. yunosecundus based on adult male characters. Sasa (1984) separates T. yunosecundus from T. lestagei Goetghebuer by the digitus placement medially of superior volsella, digitus shape and colour differences. These differences, however, can be due to mounting and seasonal variation and are not obvious species specific characters. The pupae of most species in the T. lestagei aggregate are to some extent separable, but the pupa of T. yunosecundus seems identical to those of T. decipiens. However, the synonymy will not be made until more material of the European members of the T. lestagei aggregate has been examined. The most similar Asian species are probably T. kikuchii and T. yakuheius (see remarks section for these species). T. yunosecundus has been reported from Toshigi, Yamanashi, Tokushima, Fukushima and Hokkaido Prefectures and Iki Island (Sasa, 1984, 1985a, 1988, 1993b; Sasa & Kikuchi, 1986; Iwakuma et al., 1993; Sasa & Suzuki, 1999b). New combinations, doubtful species and questionable distributions Holo- and paratypes of Virgatanytarsus toganiveus (Sasa & Okazawa, 1991) comb. n., Cladotanytarsus utonaiquartus (Sasa, 1988) comb. n. and Zavrelia tusimatijeus (Sasa & Suzuki, 1999a) comb. n. were examined and found to belong other genera than Tanytarsus. The following species are unrecognisable from their original or subsequent description and type material for examination has not been found. Nomina dubia already listed in the catalogue of the Chironomidae of the Oriental region (Sublette & Sublette, 1973) are not included. Tanytarsus hirtipes Kieffer, 1911b: 168. Type specimen in ZSI (small fragment only). Tanytarsus lasiopterus Kieffer, 1911b: 169. Type specimen in ZSI lost. Tanytarsus leptogastrus Kieffer, 1911b: 172. Type specimen in ZSI lost. Tanytarsus pictus (Doleschall, 1857) was first described in Chironomus, but later transferred to Tanytarsus Johannsen (1932). Unfortunately, the type material has not been located in Naturalis, Leiden nor in the Rijksmuseum in Amsterdam and according to the curators in these collections the types are probably lost (C. van Achterberg & H. de Jong pers. comm.). The drawing and description of the abdominal colouration of T. pictus is similar to the types of T. formosanus Kieffer, 1912, and the material collected by Thienemann on Java (larva, pupa, ♂) and used by Johannsen (1932) and Zavřel (1934) in their descriptions of T. pictus is identical to T. formosanus. However, it appears that neither Johannsen (1932) nor Zavřel (1934) have examined the types of T. pictus and although a synonymy with T. formosanus is quite possible, it has to await the localization and examination of the type material. Tanytarsus flaviradialis Guha et al., 1985: 32. No type material has been available for examination, and the original description is not sufficient to separate the T. flaviradialis from T. formosanus and related species. A male of Tanytarsus viridis Kieffer, 1911b (BMNH) collected at the type locality 18–19.I.1908 is found to belong to the genus Paratanytarsus. However, it is not a type, and although the unidentifiable syntype present at BMNH is collected only some days later (2.III.1908) it is not possible to determine if the two specimens are conspecific. I regard T. viridis to be a nomen dubium of Tanytarsus. Tanytarsus agraensis Singh & Kulshrestha, 1975: 421. The type material has neither been found in St. Johns College nor in ZSI (Chaudhuri pers. comm.), and the original description is insufficient to discriminate T. agraensis from T. formosanus. Tanytarsus ungituberculata Singh & Kulshrestha, 1975: 419. Same situation as for T. agraensis. Tanytarsus magnituberculus Guha et al., 1985: 37. Same situation as for T. flaviradialis. The following species have questionable reported distributions. Tanytarsus nichollsi Glover, 1973: 450, has been recorded twice from India (Datta et al., 1992; Mazumdar et al., 1998). One specimen collected on Sagar Island, 34 West Bengal was available for examination and belongs to the species T. formosae (Kieffer, 1921). Datta et al. (1992) records T. nichollsi from Jalpaiguri, West Bengal, but their specimens apparently differ from the diagnosis given by Glover (1973) and it is probable that these specimens belong to the closely related species T. infundibulus Chaudhuri & Datta in Datta et al., 1992 which is found at the same locality. Tanytarsus fuscithorax Skuse, 1889: 272, has been recorded from West Bengal, India by Chaudhuri et al. (1984), but apparently is not identical to the previous descriptions of the species (Chaudhuri et al., op. cit.). Chaudhuri et al., op. cit. did not examine the type specimens or species described by Glover (1973) and the Indian material has not been available for examination. Tanytarsus okuboi Sasa & Kikuchi, 1986: 28 has been recorded from West Bengal, India by Datta et al. (1992), but their specimens apparently differ some from the type material examined (Datta et al. op. cit.). Key to adult males Included in the key are potential South and East Asian species of the Tanytarsus eminulus, gregarius, lugens and mendax species groups. The species T. monstrosus is included in the key below although it is reported not to have frontal tubercles (Chaudhuri et al., 1992). This because the character is believed to be a misinterpretation in the original description. 1. Frontal tubercles well developed; anal tergite bands well separated; median tergite setae when present, never on tubercle; anal point well developed, straight or slightly triangular with several spinulae regularly arranged between anal crests; superior volsella oval to pear- shaped, with 3 setae on median margin, sometimes with apex medially directed; tubercle with seta never present ventrally of superior volsella; digitus absent or relatively straight, evenly tapered when long; median volsella usually with some foliate lamellae; gonostylus evenly tapered towards apex with widest width at 1/3–1/2 length . . . . . . . . . . . . . . . . . . . . . . 2 If anal tergite bands not separated and spinulae are present between crests of anal point, then digitus not straight and/or median tergite setae on tubercle and/or frontal tubercles minute or absent and/or superior volsella not oval to pearshaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . not keyed 2. Superior volsella apically tapered to a point, Fig. 7, 11, 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Superior volsella more or less oval, if slightly tapered, never to a point, Fig. 9, 40 . . . . . . . . . . 12 3. Median tergite setae absent, Fig. 7 . . . . . . . . . . . . 4 Median tergite setae present, although sometimes very small, Fig. 14 . . . . . . . . . . . . . . . . . . . . . . . . . . 7 4. Brown pigmented patch at base of anal point; posterior transverse bands on tergites V–VI, (Chaudhuri et al., 1992, Fig. 5) . . . . T. monstrosus No pigmented patch at base of anal point; colouration of abdominal tergites different . . . . . . . . . . .5 5. Digitus present, reaching apex of superior volsella (Spies, 1998b, Fig. 3) . . . . . . . . . . . . . T. angulatus Digitus absent, Fig. 7 . . . . . . . . . . . . . . . . . . . . . . . . 6 6. AR about 0.8–0.9; superior volsella with dorsolateral microtrichia, Fig. 8 . . . . . . . . . . . T. calorifontis AR about 1.5; superior volsella without dorsolateral microtrichia, Fig. 18 . . . . . . . . . . . . T. konishii 7. Anal point long, narrow, anal tergite bands reaching crests of anal point; digitus absent, Fig. 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. infundibulus Anal point shorter, if anal tergite bands end close to crests of anal point, digitus small but present, Fig. 36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Dorsolateral microtrichia on superior volsella, Fig. 12, 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Superior volsella without dorsolateral microtrichia, Fig. 33 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 9. Digitus reaching apex of superior volsella; median volsella reaching past inferior volsella, Fig. 47 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. takahashii Digitus not reaching apex of superior volsella; median volsella reaching 1/2 length of inferior volsella, Fig. 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Median tergite seta large, digitus well distiguishable, Fig. 11 . . . . . . . . . . . . . . . . . . . . . T. formosanus Median tergite seta small, digitus absent or hyaline, Fig. 53 . . . . . . . . . . . . . . . . . . . . . . T. uraiensis 11. Anal point with thick apical margin; median margin of superior volsella concave, Fig. 32T. oyamai Anal point with thin apical margin; median margin of superior volsella convex (Reiss & Fittkau, 1971, Fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. mendax 12. Median tergite setae absent, digitus minute or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. bathophilus Median tergite setae present, digitus present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Digitus absent or minute to small, Fig. 17, 18 . . 14 Digitus well developed, extending beyond medioapical margin of superior volsella, Fig. 35 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 35 14. Median volsella large, lamellae reaching apex of inferior volsella, Fig. 16 . . . . . . . . . . . . . . . . . . . . . 15 Median volsella shorter, lamellae never reaching apex of superior volsella, Fig. 9 . . . . . . . . . . . . . 16 15. Digitus absent, LR1 > 3.2, Fig. 16 . . . . T. kikuchii Digitus present, LR1 < 3.0, Fig. 54 . T. yakuheius 16. Microtrichia present on superior volsella, Fig. 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Superior volsella without microtrichia, Fig. 29 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 17. Digitus absent; microtrichia only present on dorsolateral margin of superior volsella; microtrichia absent from anal point, Fig. 9 . . . . . . . T. formosae Digitus minute to small; microtrichia also on ventromedian margin of superior volsella; microtrichia present between anal crests, Fig. 22 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18. Digitus minute; anal point parallel sided with apex wider than base; some lamellae of median volsella subulate, Fig. 22 . . . . . . . . . . . . . . . . T. miyakobrevis Digitus small; anal point slightly triangular; some lamellae of median volsella pectinate, Fig. 36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T. ovatus 19. Anal point large with parallel margins and minute spinulae; long median tergite setae; superior volsella long with parallel lateral margins, Fig. 20 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. mcmillani Anal point shorter, slightly triangular with small spinulae; shorter median tergite setae; slightly oval superior volsella, Fig. 30 . . . . . . . . . . . . . . . . 20 20. Anal point apex granulose; LR1 about 2.4–2.8, Fig. 28 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. okuboi Anal point apex smooth, LR1 about 3.2, Fig. 38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. pollexus 21. Median volsella long and thin, Fig. 26 . . . . . . . . 22 Median volsella shorter and wider, Fig. 44 . . . . 23 22. Median volsella with 3–4 broad, spatulate lamellae almost reaching apex of inferior volsella, Fig. 26 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. ogasatertius Median volsella with several thin lamellae reaching beyond apex of inferior volsella, Fig. 56 . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. yunosecundus 23. Superior volsella with ventral tubercle, Fig. 45, 46 . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. shoudigitatus Superior volsella without ventral tubercle . . . . . 24 24. Superior volsella almost triangular with dorsolateral microtrichia; anal point apex almost quadrate, Fig. 40–42 . . . . . . . . . . . . . . . . . . T. shouautumnalis Superior volsella more oval, without dorsolateral microtrichia; anal point apex pointed or evenly rounded, Fig. 34 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25. Anal point with large spinulae between parallel crests and median volsella short and knob- like, bearing medially directed, subulate lamellae, Fig. 51 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Anal point triangular with small spinulae and/or median volsella long with simple lamellae, Fig. 24 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 26. Microtrichia extensively distributed between crests of anal point; inferior volsella thin, medially curved towards apex, Fig. 34 . . . . . . . . T. oscillans Microtrichia not present between spinulae of anal point; inferior volsella thick, not medially curved towards apex, Fig. 51 . . . . . . . . . . T. unagiseptimus 27. Anal point apex granulose, Fig. 28 . . . . . T. okuboi Anal point apex smooth, Fig. 24 . . . . . . . . . . . . . 28 28. LR1 about 3.3, (Sasa, 1980, Fig. 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tamaundecimus LR1 < 2.6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 29. AR < 0.9; digitus broad, Fig. 24 . T. ogasaquartus AR about 1.0; digitus narrow, (Sasa & Tanaka, 2000, Fig. 2) . . . . . . . . . . . . . . . . . . . . . T. tonebeceus Concluding remarks As shown, the taxonomy of East and South Asian Tanytarsus species belonging to the T. eminulus, gregarius, mendax and lugens species groups has been somewhat confused with many synonyms and unclear descriptions. This is still the case for most of the Tanytarsus species not belonging to these groups and further revisionary work is needed. The robustness of the species groups created by Reiss & Fittkau (1971) for European Tanytarsus species has yet to be tested for species of other fauna regions, but some characters in the pupal morphology suggests that the groups might hold. Pupae of the species of which the adults key to the T. mendax species group tend to have large cephalic tubercles, chaetae in one row on thoracic horn and almost equal large spine patches on tergites III–VI with spinules or short spines only. Larvae have short Lauterborn organ pedicels and large postoccipital plates, but too few larvae are described to say anything with certainty. Pupae of species in the T. eminulus species group tend to have minute or no cephalic tubercles, quite short thoracic horn with minute chaetae evenly distributed on parts or whole surface of horn and large spine patches with long spines on tergites III–IV. Pupae of species in the T. gregarius and T. lugens species groups are observed also to have small or missing cephalic 36 tubercles and large spine patches with long spines on tergites III–IV, but their thoracic horn is longer and bare, and they have a band of spinules lateral to the spines of tergite IV. To few larvae are described to say anything about potential diagnostic larval characters for the T. gregarius, eminulus and lugens species groups. Hopefully, further descriptions of immature stages as well as molecular data will contribute to a stable phylogeny for these zoogeographically and ecologically interesting species. Acknowledgements I am indebted to Dr Manabu Sasa, Dr Seichii Nohara, Dr Ryuhei Ueno and Dr Kiyoshi Satake for their friendly help during my stay at NIES, Prof. P. K. Chaudhuri, Dr Frank Menzel and Dr John Chainey for providing me with type material. Thanks also to Prof. Ole A. Sæther, Prof. James E. Sublette, Angela Sanseverino and Dr Elisabeth Stur for kindly reading through different versions of the manuscript. My scientific visit to NIES, Tsukuba, Japan was financed by Nansenfondet and associated funds (ref. 164/2000). References Bause, E., 1913. Die Metamorphose der Gattung Tanytarsus und einiger verwandter Tendipedidenarten. Ein Beitrag zur Systematik der Tendipediden. Arch. Hydrobiol. Suppl. 2: 1–139. Brundin, L., 1947. Zur Kenntnis der schwedischen Chironomiden. Arkiv Zool. 39 A: 1–95. Chaudhuri, P. K., A. Ali & A. Majumdar, 1992. Life stages of Tanytarsus van der Wulp with description of a new species from India (Diptera, Chironomidae). Dtsch. ent. Z., N. F. 39: 381–396. 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