Tormus helmsi Sharp, 1884

Fikáček, Martin, Minoshima, Yûsuke, Vondráček, Dominik, Gunter, Nicole & Leschen, Richard A. B., 2013, Morphology of adults and larvae and integrative taxonomy of southern hemisphere genera Tormus and Afrotormus (Coleoptera: Hydrophilidae), Acta Entomologica Musei Nationalis Pragae 53 (1), pp. 75-126 : 96-110

Tormus helmsi Sharp, 1884

( Figs 1 , 7 , 9 C–N, 10–16)

Tormus helmsi Sharp, 1884: 474 . Type locality: New Zealand, Buller, Greymouth.

= Stygnohydrus nitidus Broun, 1893b , syn. nov. Type locality: New Zealand, Westland, Boatmans. Trasferred to Tormus by HANSEN (1991: 183).

= Tormus nitidus Broun, 1893c: 1402 , syn. nov. Secondary homonym of Stygnohydrus nitidus . Type locality: New Zealand, Westland, Capleston. Synonymized with Stygnohydrus nitidus by HANSEN (1997: 360).

= Stygnohydrus basalis Orchymont, 1937: 155 , syn. nov. Replacement name for Tormus nitidus Broun, 1893 . Synonymized with Stygnohydrus nitidus by HANSEN (1997: 360).

= Stygnohydrus femoralis Broun, 1910: 12 , syn. nov. Type locality: New Zealand, Taupo, Raurimu.

Type material examined. Tormus helmsi : HOLOTYPE: unsexed specimen ( BMNH): Tornus [sic!] / helmsi / Type D.S. / Greymouth N. Zd / Helms [handwritten on the label with the specimen] // Type / H. T. [rounded label with red margin] // Greymouth / New Zealand / Helms // Sharp Coll. / 1905-313.’

Stygnohydrus nitidus : HOLOTYPE: unsexed teneral specimen ( BMNH): Syn- / type [rounded label with blue margin] // Type [rounded label with red margin] // New Zealand / Broun Coll. / Brit. Mus. / 1922-482 // Westland // 2343 // Stygnohydrus nitidus’

Tormus nitidus : LECTOTYPE (designated by HANSEN (1997)):unsexed specimen ( BMNH): Type [round label with red margin] // 2445 // Capleston / Westland // New Zealand / Broun Coll. / Brit. Mus. / 1922-482 // Tormus / nitidus // LECTOTYPE / Tormus nitidus Broun / M. Hansen des. 1996’

Stygnohydrus femoralis : HOLOTYPE: 1 ♀ ( BMNH): [female symbol] // Type [round label with red margin] // Raurimu / Jany. 1909 // 3032. // Stygnohydrus / femoralis. // New Zealand / Broun Coll. / Brit. Mus. / 1922-482. ’

Additional adult specimens examined (445 specimens). NEW ZEALAND: North Island : COROMANDEL: 1 ♀ ( NZAC): Maumaupaki Track , 36°58.2′S 175°34.6′E, 17.xi.2004, lgt. A. C. Eyles & J. I. Townsend. BAY OF PLENTY: 4 spec. ( NZAC): Horohoro state forest, Mamaku Plat , elev. 550 m [38°16′23″S 176°7′45″E], 24.vii.1976, lgt. J. S. Dugdale (76/43); 26 spec. ( FMNH, NMPC, YMC): Kaimai-Mamaku Forest , Mount Te Aroha , upper end of Tui Mine Track near summit road, elev. 775 m, 37°31.658′S 175°44.684′E, 19.xi.2005, lgt. Newton & Thayer (FMHD#2005-021); 3 spec. ( NZAC): Mount Te Aroha [37°32′29″S 175°42′39″E], 21.x.1967, lgt. J. C. Watt; 1 ♀ ( NZAC): Mount Te Aroha 37°32.5′S 175°42.7′E, 27.ii.1992, lgt. J. S. Dugdale (92/6); 1 ♀ ( NZAC): same locality, 12.xii.2004, lgt. R. Leschen & S. Williford (RL890). TARANAKI: 1 spec. ( LUNZ): Mount Egmont National Park , North Egmont Road [ca. 39°16.2′S 174°5.1′E], 24.v.1986, lgt. C. L. Lyal (CL479); 6 spec. ( LUNZ): Mount Egmont National Park , Potaema picnic area, elev. 650 m [39°18.8′S 174°8.8′E], 24.xii.1985, lgt. R. M. Emberson & P. T. Syrett; 4 spec. ( LUNZ): Mt. Egmont , elev. 823 m [ca. 39°17′S 174°7′E], 29.xi.1970, lgt. A. W.Don; 9 spec. ( NMPC, NZAC): Mt. Egmont NP, Mangaroaka Walk at Egmont road, elev. 600 m, 39°14.5′S 174°6.8′E, 1.xii.2012, lgt. Fikáček (NZ29); 15 spec. ( NMPC, DBCP, NHMW, NZAC, ZMUC): Mt. Egmont NP, Potaema Walk 6.8 km W of Pembroke, elev. 650 m, 39°21.3′S 174°8.8′E, 28.xi.2012, lgt. Becker, Fikáček & Hájek (NZ16); 1 spec. ( NMPC): Mt. Egmont NP, walk from The Plateau to Manganui Lodge, elev. 1230 m, 39°18.4′S 174°5.5′E, 28.xi.2012, lgt. Becker, Fikáček & Hájek (NZ18); 2 spec. ( LUNZ): North Egmont , Holly Hut, elev. 950 m [39°15.9′S 174°2.8′E], 27.xi.1975, lgt. A. K. Walker; 1 spec. ( LUNZ): Pouakai Range [39°14.5′S 174°1.2′E), 9.–13.i.1978, lgt. J. C. Watt; 2 spec. ( LUNZ): Pouakai Road, Ahukawakawa Track, elev. 1067 m [39°15.3′S 174°2.5′E], 11.i.1978, lgt. J. C. Watt (78/10). TAUPO: 1 spec. ( NZAC): Ohakune Mount Rd. Tk. [39°25.1′S 175°24′E], 24.xi.1965, lgt. J. I. Townsend; 2 spec. ( NZAC): same locality, 27.xi.1985, lgt. R. C. Craw (85/65); 2 ♀♀ ( BMNH): Waimarino [39°26′S 175°8′E], i.1910, without collector; Taranaki, 1 ♀ ( JNIC): Mt. Egmont , Patea walk, 39°18.5′S 174°6.8′E, 10.xii.1995, lgt. J. T. Nunn. WAIKATO: 2 spec. ( NZAC): Mt. Pirongia , Track 2 to Wharauroa [37°59.5′S 175°5.9′E], 16.i.1977, lgt. A. K. Walker (77/2); 1 ♂ ( NZAC): same locality, 9.vi.1977, lgt. B. A. Holloway (77/61). WELLINGTON: 1 ♀ ( NZAC): Mt. Holdsworth Track [40°52.4′S 175°25′E], 3.ix.1965, lgt. J. I. Townsend.

South Island: NELSON: 1 ♀ ( NMPC): 1000 acre Plateau , site 1, elev. 1022 m, 41°35.0′S 172°12.4′E, 8.ii.2011, lgt. L. Dunning & L. Shield (TB505) ; 1 spec. ( NZAC): Blue Duck Creek, Gouland Downs ,elev. 762 m [41°50′S 172°21.9′E], 5.ii.1966, lgt. J. I. Townsend ; 1 ♀ ( NMPC): Charming Creek walkway at Watsons Mill , 3.3 km E of Hector, elev. 70 m, 41°36.2′S 171°54.9′E, 5.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ35) ; 1 ♀ ( NZAC): Frazer Stream, S. Paturad [42°18.7′S 172°16.6′E], 22.viii.1967, lgt. F.Alack ; 1 ♂ ( NZAC): Glen Hope [41°21.3′S 173°9.3′E], 20.vi.1915, lgt. T. Hall ; 23 spec. ( NZAC): Gourland Downs [= Gouland Downs, 40°53.5′S 172°21.2′E], 5.ii.1965, lgt. J. I. Townsend ; 1 ♀ ( NMPC): Hanlon Lake , 5.2 km S of Te Namu on Karamea Highway, elev. 80 m, 41°25.5′S 172°6.1′E, 6.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ39) ; 1 spec. ( NZAC): Hope [41°21.3′S 173°9.3′E], 21.xii.1915, lgt. T. Brookes ; 1 ♀ ( NZAC): Lake Rotoiti [41°50.5′S 172°50.6′E], iii.1949, lgt. A. E. Brookes ; 1 ♀ ( NZAC): Mt. Arthur [41°13.1′S 172°40.9′E], 19.xi.1969, lgt.J.I. Townsend ; 2 spec. ( NZAC): Mt.Augustus, Granity ,elev. 853 m [41°40.5′S 171°51.4′E], 4.iii.1970, lgt. J. S. Dugdale ; 3 spec. ( NZAC, NMPC): Oparara Arches, 14.8 km NNE of Karamea, end of McCallum Mill Road at Box Canyon Cave , elev. 230 m [41°8.1′S 172°11.5′E], 6.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ37) ; 2 spec. ( NZAC): Punakaiki, Bullock Creek [42°12.1′S 171°24.5′E], 12.x.1970, lgt. J. I. Townsend ; 1 ♀ ( NZAC): Surveyors Creek, Karamea Saddle, elev. 609 m [41°18.3′S 173°24.8′E], 13.x.1970, lgt. J. I. Townsend. BULLER: 1 ♂ ( NZAC): Bullock Creek , elev. 20 m [ca. 42°6′S, 171°22″E], 6.xii.1982, lgt. C. A. Muir; 5 spec. ( NZAC): E of Mt. Dewar , elev. 1127 m [ca. 42°4.9′S, 171°32.7′E], 12.xii.1969, lgt. J. S. Dugdale ; 1 ♂ ( NZAC): Flagstaff Biological Reserve, Hochstetter [ca. 42°26.9′S 171°39.3′E], 20.ix.1972, lgt. J. S. Dugdale (72/174) ; 2 spec. ( NZAC): Lake Rotoroa near camp ground [ca. 41°51.9′S 172°38.8′E], 9.i.1993, lgt. J. W.M. Marris ; 9 spec. ( NZAC): Mawhera state forest [42°29.5′S 171°31′E], 10.xi.1971, lgt. J. McBurney (71/144) ; 1 spec. ( NZAC): Mawhera state forest, S of Ngahere [42°23.7′S 171°26.8′E], 27.i.1972, lgt. J. S. Dugdale (72/98) ; 5 spec. ( NZAC, ANIC, NMPC): Mawhera state forest, Kangaroo Creek , 5 km SE of Ngahere [42°25.4′S 171°29.6′E], 27.i.1972, lgt. J. S. Dugdale (72/86) ; 3 spec. ( NZAC): Mount Dewar , elev. 1000 m [42°4.9′S 171°32.7′E], xii.1969, lgt. J. I. Townsend (69/234) ; 1 spec. ( NZAC): same locality, xii.1969, lgt. J. I. Townsend (69/239) ; 2 spec. ( NZAC): same locality, xii.1969, lgt. J. I. Townsend ; 4 spec. ( NZAC, NMPC): same locality, 2.xii.1969, lgt. J. G. McBurney ; 3 spec. ( NZAC): same locality, 2.xii.1969, lgt. J. I. Townsend ; 1 ♂ ( NZAC): same locality, 5.xii.1969, lgt. J. C. Watt ; 5 spec. ( NZAC): same locality, 10.xii.1969, lgt. J. C. Watt ; 1 ♂ ( NZAC): Paparoa National Park, Mt. Bovis Tarn , elev. 1000 m [42°6.9′S 171°29.3′E], 9.xi.1990, lgt. P. Syrett & R. M. Emberson ; 3 spec. ( NZAC): Paparoa Range, Croesus Track , elev. 850 m [42°15.6′S 171°21.8′E], 19.i.1982, lgt. R. M. Emberson & J. W. Early ; 2 spec. ( NZAC): Punakaiki Scenic Reserve, Porarari River , elev. 20 m [42°6.5′S 171°19.6′E], 9.vi.1985, lgt. J. W. Early (LCNZ 85/7) ; 6 spec. ( NZAC, ANIC, NMPC): Stoney Creek , Inangahua valley [ca. 41°51.5′S 171°56.9′E], 19.ix.1972, lgt. J. S. Dugdale (72/182) ; 6 spec. ( NZAC): 6 km SW of Rotokohu [42°0.7′S 171°51.1′E], 25.i.1972, lgt. J. S. Dugdale (72/99) ; 2 spec. ( NZAC): 9 miles E of Reefton [42°6.9′S 172°2.4′E), 10.ii.1965, lgt.N.A. Walker ; 5 spec. ( LUNZ): Ananui Creek, Charleston state forest [41°56.9′S 171°30.3′E], 7.viii.1977, lgt. R. M. Emberson ; 6 spec. ( NZAC, NMPC): Boatmans Creek [42°2.6′S 171°53.7′E], 4.x.1971, lgt. G. Kuschel (71/117); 5 spec. ( NZAC): Capleston [42°4′S 171°54.9′], 8.iii.1972, lgt. J. C. Watt (72/111); 12 spec. ( NZAC): Capleston env., Beetle reserve , (42°4′3.69″S, 171°54′57.66″E), 6.iv.1973, lgt. J. C. Watt; 1 ♀ ( NZAC): Cave Creek Monument , 42°6′S 171°24′E, 20.i.2005, lgt. R. Leschen, T. Buckley, P. Lambert (RL928); 20 spec. ( NZAC, ANIC, NMPC): Central Val , 4.5 km SE of Cronadun [42°2′S 171°53′E], 28.i.1972, lgt. J. C. Watt; 1 ♂ ( NZAC): Inangahua [41°51.3′S 171°56.9′E], 23.i.1957, lgt. E.S.Gourlay; 1 spec. ( NZAC): Inangahua state forest, Fletcher Creek [41°59.3′S 171°50.6′E), xi.1971, lgt. J. G. McBurney; 4 spec. ( NZAC): same locality, 25.i.1971, lgt. J. S. Dugdale; 10 spec. ( NZAC): same locality, 9.xi.1971, lgt. J. S. Dugdale; 1 spec. ( NZAC): same locality, 26.i.1972, lgt. J. S. Dugdale; 16 spec. ( NZAC): same locality, 19.ix.1972,lgt.J. S. Dugdale; 1♂ ( NZAC): Maruia Saddle, Murchinson district [42°2.1′S 172°17.5′E], 2.vi.1965, lgt. J. I. Townsend (65/338); 1 ♂ ( NZAC): Matiri Tops [41°38′S 172°14.9′E], 17.ii.1964, lgt. J. I. Townsend; 1 ♂ ( NZAC): Mt. Murchinson , Buller valley, elev. 1341–1432 m [41°43.8′S 172°29.9′E], 21.xi.1971, lgt. J. S. Dugdale; 3 spec. ( NZAC, NMPC): N of Capleston, Italian Creek [42°2.9′S 171°55.3′E], 21.iv.1972, lgt. J. S. Dugdale; 5 spec. ( NZAC): same locality, 12.i.1973, lgt. J. C. Watt; 1 ♂ ( NMPC): Paparoa NP, beginning of Cave Creek Track close to Bullock Creek Road , 5.6 km NEE of Punakaiki, elev. 115 m, 42°6.2′S 171°24.1′E, 4.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ34) ; 3 spec. ( LUNZ): Paparoa range, Buckland Peaks , elev. 1250 m [41°53.3′S 171°38.3′E], 15.xi.1987, lgt. R. M.Emberson & P.T. Syrett ; 3 spec. ( NZAC): Rahu saddle, elev. 600–670 m [42°18.8′S 172°7′E], 1.vi.1965, lgt. J. I. Townsend (65/334) ; 1 ♂ ( LUNZ):same locality, 16.x.1984, lgt.J. W. Early (LCNZ 84/7) ; 10 spec. ( NZAC): Redman Creek, Capleston [42°3.6′S 171°52.8′E], 10.i.1973, lgt. J. C. Watt (73/6) ; 1 spec. ( NZAC): Tawhai state forest [42°9′S 171°47′E), x.1972, without collector (72/190 A) ; 24 spec. ( NZAC): Tawhai state forest, 3 km S of Reefton [42°9′S 171°47′E], 9.xi.1971, lgt. J. S. Dugdale (71/128); 1 ♂ ( NZAC): Tutaki valley, saddle near Rotoroa, Braeburn Track [41°51′S 172°27.9′E], 9.iii.1965, lgt. J. I. Townsend & G. Kuschel. NORTH CANTENBURY: 1 ♀ ( NZAC): Lewis Pass [42°22.7′S 172°23.9′E], 19.iii.1965, lgt. G. W. Ramsay & J. I. Townsend (65/265). WESTLAND: 1 spec. ( NZAC): Mt. Greenland , elev. 228 m [42°57.2′S 170°49.8′E], i.–ii.1940, lgt. E. S. Gourlay; 2 ♂♂ ( NZAC): same locality, 10.vi.1983, lgt. H. P. McColl (8/83); 1 ♂ ( NZAC): Mt. Tuhua, E. side of lake Kaniere [42°49.3′S 171°11.4′E], 20.xi.1984, lgt. C. F. Butcher (84/74); 1 spec. ( NMPC): Pleasant Flat , elev. 120 m, 44°0.8′S 169°2.9′E, 13.i.2006, lgt. R. Leschen, T. Buckley, R. Hoare (RL1066) ; 25 spec. ( NMPC, DBCP, NZAC, NHMW, ZMUC): same locality, 11.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ49) ; 4 spec. ( NMPC, NZAC):along Three Mile Lagoon Track , 1.8–3.0 km SWW of Okarito, elev. 100–140 m, 43°14′S 170°8.5′E, 8.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ43) ; 1 ♂ ( NMPC): Cross Creek on Haast Pass Highway , 18.3 NNE of Makarora,elev. 500 m, 44°6′S 169°36.5′E, 11.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ51) ; 1 spec. ( NZAC): Denniston [41°44.2′S 171°47.7′E], i.–ii.1965, lgt. J. I. Townsend ; 1 spec. ( NZAC): same locality, i.–ii.1965, lgt. J. I. Townsend ; 4 spec. ( NZAC): same locality, 7.x.1969, lgt. J. I. Townsend ; 2 ♀♀ ( NZAC): Haast Pass [44°6.4′S 169°21.3′E], 20.iii.1968, lgt. R. A. Cumber ; 2 spec. ( NMPC): same locality, 19.i.2010, lgt. Jäger & Nunn ; 12 spec. ( NZAC): Nile river valley [42°1.4′S 171°30.9′E], 1.ix.1971, lgt. R. M. Emberson ; 4 spec. ( LUNZ): Okuku state forest, 10 km E of Kumara [42°37.8′S 171°18.5′E], 28.viii.1977, lgt. R. M. Emberson ; 1 spec. ( NZAC): Paparoa Range, Lochnagar Ridge , camp area, elev. 1060 m [42°6.5′S 171°33.1′E], xii.1969, lgt. J. S. Dugdale ; 2 spec. ( NZAC): same locality, xii.1969, lgt. J. I. Townsend ; 4 spec. ( NZAC): Ross Mikonui Track , 43°S 170°51′E, 9.xi.2005, lgt.R. Leschen & S.Nomura (RL1015) ; 2 spec. ( NZAC): 2 miles S of Fox Glacier [43°29.5′S 180°1.5′E], 8.ii.1965, lgt. N.A. Walker ; 2 spec. ( NZAC): Alex Knob [43°25.6′S 170°9.1′E], 3.xi.1965, lgt. A. C. Eyles ; 3 spec. ( NZAC):same locality, 23.ii.1966, lgt.J. I. Townsend ; 1♂ ( LUNZ): Canavans Knob env.[43°22.9′S 170°9.8′E], 4.ii.1982, lgt. A.B.Miller ; 2 spec. ( NZAC): Fox Glacier [43°28′S 170°1.2′E], 17.ii.1965, lgt.T. Wood ; 1 spec. ( NZAC): same locality, 11.xi.1965, lgt. J. I. Townsend ; 1 spec. ( NZAC): same locality, 12.xi.1965, lgt. J. I. Townsend ; 1 spec. ( NZAC): Fox Glacier, Minnehaha Track , [43°28′S 170°1.2′E], 8.ii.1965, lgt. N. A. Walker ; 2 spec. ( NZAC): Franz Josef [43°23.3′S 170°11′E], 2.xi.1960, lgt. J. I. Townsend ; 1 ♂ ( LUNZ): same locality, 1.iv.1988, lgt. R. M. Emberson & P. T. Syrett ; 3 spec. ( NZAC): Gillespies Beach [43°25′S 169°49.3′E], 12.xi.1968, lgt. J. I. Townsend (68/173) ; 2 spec. ( LUNZ): Lake Moeraki [43°43.7′S 169°16.3′E], 12.xii.1994, lgt. C. J. Vink ; 2 spec. ( NZAC): Lake Paringa , southern end [43°43.2′S 169°24.4′E], 7.ii.1960, lgt. N. A. Walker ; 12 spec. ( LUNZ, NMPC): Lake Wombat Track [43°24.2′S 170°10.1′E], 13.i.1986, lgt. J. W. Early ; 1 ♂ ( NZAC): Matheson Lake track, 5 km NWW of Fox Glacier village, elev. 120 m, 43°26.3′S 169°58′E, 9.xi.2007, lgt. R. Leschen & C. Carlton (RL1318) ; 1 spec. ( NMPC): same locality, 9.xii.2012, lgt. Fikáček, Hájek & Leschen (NZ45) ; 2 spec. ( LUNZ): Mt. Aspiring National Park, Haast Pass highway, Depot Creek [43°57.9′S 169°14.6′E], 11.xi.1995, lgt. R. M. Emberson & P. T.Syrett ; 2 spec. ( NZAC): Upper Callery [43°25.4′S 170°12.4′E], 2.xi.1965, lgt. J. I. Townsend & A. C. Eyles; 5 spec. ( LUNZ, NMPC): Westland National Park, Gillespies, Cook River road, elev. 120 m [43°28.9′S 169°56.3′E], 20.x.1981, lgt. R. M. Emberson (LCZN 81/16). OTAGO LAKES: 1 ♂ ( LUNZ): Mt.Aspiring National Park , Cameron Flat , Makarora river [44°9.5′S 169°18′E], 2.i.1985, lgt. R. M. Emberson & P. T. Syrett .

Larval specimens examined. NEW ZEALAND: North Island: BAY OF PLENTY: 19 L1, 23 L2, 20 L3 ( FMNH, YMC, NMPC): Kaimai-Mamaku Forest, Mount Te Aroha, upper end of Tui Mine Track near summit road, elev. 775 m, 37°31.658′S 175°44.684′E, 19.xi.2005, lgt. Newton & Thayer (FMHD#2005-021). Larvae were associated with adults using the cox1 molecular markers, see the Genetic and morphological diversity above and the results of the phylogenetic analyses in Fig. 8 .

Redescription of adults. Body length 2.7–3.9 mm (holotype 3.5 mm), body width 1.8–2.2 mm (holotype 2.1 mm). Head and pronotum with sparsely arranged but rather coarse punctures, without microsculpture on interstices; elytra anteriorly with sparse punctuation consisting of punctures ca. as large as on pronotum or distinctly smaller., interstices smooth or with very weak microsculpture. Elytra more or less evenly convex, oblique to nearly vertical posteriorly in lateral view. Parameres of aedeagus variable in shape, but never with hook-like apices ( Figs 9 C–N).

Description of larvae. General morphology. Third instar. Body moderately thick, nearly parallel-sided, widest between abdominal segments 2–4 ( Fig. 1D ). Color yellowish brown, sclerotized parts slightly darker ( Fig. 1D ).

Head. Head capsule ( Fig. 14A ) semicircular; cervical sclerites small. Frontal lines visible in basal two-thirds, almost straight and divergent anteriad, reaching base of head capsule without fusing; coronal line absent. Surface of head capsule smooth. Six stemmata on each anterolateral corner of head capsule; each stemma more or less protuberant. Clypeolabrum ( Fig. 14B ) somewhat asymmetrical. Nasale asymmetrical with three teeth; right two teeth closely aggregated. Epistomal lobes slightly asymmetrical, projecting as far as or slightly further than nasale; right lobe projecting slightly further than left one. Lateral margin of each epistomal lobe membranous; inner margin with fringe of short, trichoid cuticular projections.

Antenna ( Fig. 15 A–B) 3-segmented, short, rather stout.Antennomere 1 as long as antennomeres 2 and 3 combined, anterior two-thirds of inner face including intersegmental membrane between antennomeres 1 and 2 and large portion of ventral surface bearing cuticular projections; projections on inner face trichoid, long on apical part, short on basal part; projections on ventral surface fine, rather densely arranged. Antennomere 2 narrower than antennomere 1, bearing rather short, trichoid cuticular projections on basal part and lateral half of dorsal and lateral surfaces; intersegmental membrane between antennomeres 2 and 3 with few rather short, trichoid cuticular projections on inner face. Antennomere 3 the narrowest and shortest, as long as antennal sensorium SE1.

Mandibles ( Figs 15 C–D) moderately wide, symmetrical, with three inner teeth each. Distal two inner teeth large, basal one small, distal one shorter than middle one.

Maxilla ( Figs 15 E–F) 6-segmented, longer than antenna. Cardo moderate in size, irregularly shaped. Stipes the longest, longer than palpomeres 1–4 combined; dorsal surface bearing densely arranged, short, trichoid cuticular projections on inner and outer parts; apical part of outer face including intersegmental membrane between stipes and palpomere 1 bearing rather long, trichoid cuticular projections. Maxillary palpus rather stout, 4-segmented. Palpomere 1 widest and longest, completely cylindrically sclerotized, bearing densely arranged, short, trichoid cuticular projections on median part of dorsal surface to lateral surface, including intersegmental membrane between palpomeres 1 and 2. Palpomere 2 narrower than palpomere 1 but wider than palpomere 3, as long as or slightly longer than palpomere 3, bearing densely arranged, short, trichoid cuticular projections on apical part of inner face including intersegmental membrane between palpomeres 2 and 3 and median part of dorsal surface to lateral face including intersegmental membrane; fine cuticular projections present ventrally on basal part. Palpomere 3 much wider than and as long as or slightly shorter than palpomere 4, bearing densely arranged, short, trichoid cuticular projections on median part of dorsal surface to lateral surface, including intersegmental membrane between palpomeres 3 and 4. Palpomere 4 bearing short, trichoid cuticular projections dorsally; inner process sclerotized, slightly shorter than palpomere 1.

Labium ( Figs 15 G–J) well developed but ligula reduced. Submentum (see Fig. 10B ) fused to head capsule, rather small, subpentagonal, wider than mentum. Mentum subquadrate in dorsal view ( Fig. 15G ), subtrapezoidal in lateral view ( Fig. 15H ). Dorsal surface with rather densely arranged, strong cuticular teeth; lateral face of apical part including intersegmental membrane between mentum and prementum bearing densely arranged short to long trichoid cuticular projections; remaining part of dorsal surface of intersegmental membrane bearing short, trichoid cuticular projections. Prementum subrectangular, slightly transverse, apical part wider than basal part; dorsal and lateral surfaces including membranous area of prementum bearing densely arranged short and narrow trichoid cuticular projections; median part of membrane (on place of the missing ligula) bearing very densely arranged, very long, trichoid cuticular projections; ventral surface of membrane bearing a few short to rather long, trichoid cuticular projections. Labial palpi moderately short, palpomere 1 larger than palpomere 2, bearing a few short, trichoid cuticular projections laterally; dorsal and lateral face of intersegmental membrane between palpomere 1 and 2 bearing a transverse row of very long to rather long, trichoid cuticular projections.

Thorax. Prothorax wider than head capsule ( Fig. 1D ). Proscutum ( Figs 1D , 14 C–D) formed by one large plate subdivided by fine sagittal line, anterior part rather weakly sclerotised; median part weakly convex transversely on each side, this part bare except few setae; anterolateral portion weakly convex; whole sclerite bearing densely arranged small, conical cuticular projections and sparsely arranged rather short lanceolate setae and a few long trichoid setae; anterior margin of sclerite bearing fringe of stout, lanceolate setae ( Fig. 14D ); setae composing a transverse row. Prosternal sclerite divided into two plates, bearing setae along anterior margin ( Fig. 14E ). Mesonotum with two sclerites on each side ( Fig. 14F ); anterior ones small, transverse, subtriangular, bearing fine cuticular projections; posterior ones large, subtriangular, bearing a few rather short, lanceolate, rod-like or spiniform setae of variable length, without cuticular projections; posterior mesonotal sclerites narrowly touching each medially, whereas anterior ones not. Two pairs of weakly convex tubercles present dorsolaterally; median ones posterolaterally to posterior mesonotal sclerite, lateral ones behind spiracles; tubercles bearing a few rod-like setae; lateroventral surface with two weakly convex projections. Metanotum with one sclerite on each side (e.g., Fig. 14F ); metanotal sclerites transverse, oval, bare but with a few setae of variable length and shape; sclerites not touching each other medially. Six rather large setiferous tubercles bearing a few rather short, rod-like setae (e.g., Fig. 12C ) present dorsally on posterior part, composing a transverse row; median two pairs behind metanotal sclerites, remaining ones more laterally; lateroventral surface with two weakly convex projections. Legs (see Fig. 12B ) short, stout, slightly or not visible in dorsal view, 5-segmented; all three pairs similar in shape.

Abdomen. Abdomen 10-segmented, tapering posteriad, covered with fine cuticular projections densely arranged, with tubercles bearing several rather short, rod-like setae (e.g., Fig. 12C ); dorsal sclerites on segments 1–7 absent. Segments 1 to 6 similar in shape and size, segment 7 smaller than others. Segments 1–6 with two transverse rows of tubercles dorsally to laterally; transverse fold subdividing segment, present between rows of tubercles. Anterior row composed of six large tubercles, spiracle present between lateral two tubercles, close to second one from the outest, lateral pair present on lateral face; posterior row composed of five small tubercles, mesal one slightly transverse, lateral pair on lateral face; lateroventral surface with two weakly convex projections; lateral part of ventral surface with closely aggregated four weakly convex projections. Segment 7 similar to segment 1–6 but posterior row of tubercles composed of three tubercles smaller and more weakly projected; projections of ventral surface indistinct, with a pair of weakly convex areas laterally.

Spiracular atrium modified ( Fig. 14G ). Segment 8 with irregularly shaped, oval dorsal plate, and with two pairs of tubercles on posterolateral part; dorsal plate bearing densely arranged, small cuticular projections and sparsely arranged, rather short rod-like or slightly lanceolate setae; posterior margin of segment 8 with two rather short and four long trichoid setae. One pair of membranous procerci present on lateral part of posterior margin of segment 8, bearing two rather short setae and one long seta. Segment 9 unilobed and bifurcated (see also Discussion); median lobe and each lateral lobe of spiracular atrium fused into one lobe, median part of dorsal surface weakly sclerotised, margin of sclerite unclear; urogomphi small, sclerotized, one segmented. Acrocercus and prostyli absent. Ventral surface of spiracular atrium with two projections.

Second instar. Very similar to third instar larva ( Fig. 1C ).

Head. Frontal lines clearly visible, almost straight and divergent anteriad, very slightly lyriform, reaching base of head capsule without fusing (as in Fig. 10B ).

Antenna, maxilla and labium proportionally slightly stouter than in third instar ( Fig. 13 ).

Thorax and abdomen. Cuticular projections on membrane finer and narrower than in third instar.

First instar. Very similar to second instar ( Fig. 1B ).

Head. Antenna, maxilla and labium proportionally slightly stouter than in second instar ( Fig. 11 ).

Thorax and abdomen. Cuticular projections on membrane slightly narrower than in second instar, but the size of projection similar to second instar, thus proportionally longer than in second instar; arrangements of cuticular projections sparser than in third instar.

Chaetotaxy of head. First instar. Frontale ( Figs 10A, C ). Central part with three pairs of sensilla (FR1–3) divergent posteriad; FR1 short seta, rather close to frontal line; FR2 pore-like, situated more anteriorly and more mesally to FR1; FR3 short seta, situated anteriorly to FR2. Pore-like sensillum FR4 and setae FR5–6 situated posteromesally to antennal socket, forming triangular group, FR5 rather short seta, FR6 moderately long seta; FR4 mesally to FR5–6, FR5 posteriorly to FR6; FR6 laterally to FR4. FR7 rather long seta situated on inner face of antennal socket. Pore-like sensillum FR15 and moderately short seta FR8 situated on mesal part of clypeolabrum, behind nasale; FR15 anteriorly to FR8. FR9–14 forming oblique group on basal and inner part of epistome anteromesally to antennal socket, FR11 and FR13–14 pore-like, FR9–10 rather long setae, FR9 longer than FR10, FR12 short and slightly stout seta; FR11 situated mesally to remaining sensilla, close to lateral-most seta of gFR1; FR12–13 close to FR11, between FR11 and FR9 or anteriorly to line connecting FR11 and FR9; FR10 and FR14 close to antennal socket, FR14 laterally to remaining sensilla, FR10 posteromesally to FR14; FR9 between FR14 and FR13. Nasale with a group of six stout and moderately short setae and with two minute setae (gFR1); lateral-most stout ones on emargination between nasale and epistomal lobe, remaining stout ones on median part of nasale; minute ones present ventrally on median tooth of nasale (not seen in Fig. 10C , see Fig. 14B ). Each epistomal lobe with a group of four stout setae on median part of anterior margin (gFR2), each seta bearing two subapical teeth on inner face; all setae nearly equidistant.

Parietale ( Figs 10 A–B). Dorsal surface with a group of five sensilla (PA1–5) forming irregularly longitudinal row in posterior part of parietale; PA1–2 and PA4–5 very short setae, PA3 pore-like. PA6 pore-like, located posteromesally, moderately close to posterior end of frontal lines. Two long setae PA7–8 situated along frontal line, PA7 more distant from frontal line than PA8; PA8 behind antennal socket, close and anterolaterally to mesal-most stemma of anterior low of stemmata; PA7 between PA6 and PA8, at ca. midlength between PA6 and PA8. Long seta PA9 situated very close and posterolaterally to antennal socket. Pore-like sensillum PA10 situated between mesal-most stemmata of anterior and posterior rows; rather long seta PA11 between lateral two stemmata of anterior row, slightly posteriorly to these. PA12 rather short seta close to and behind mesal-most stemma of posterior row, at midlength between PA7 and PA13; PA13 long seta, behind median stemma of posterior row. PA14–17 and PA26 forming irregular transverse row on anterior third of lateral surface ordered as follows from dorsal to ventral: PA14, PA15, PA16, PA17, PA26; PA26 more distant from PA17 than PA16; PA14 very long seta, PA15 and PA17 pore-like, PA26 long seta. PA18 long seta situated on median part of lateroventral surface, posteriorly to PA15–17, close and anterolaterally to PA30. PA19–22 closely aggregated, forming slightly irregular transverse row on anterior corner of head capsule, order from dorsal to ventral: PA19, PA20, PA21, PA22; PA19 porelike, PA20 rather short seta, PA21 very long seta, PA22 long seta. Pore-like sensilla PA23–25 situated close to ventral mandibular articulation; PA23 laterally to PA24–25, PA25 on inner margin, PA24 between PA23 and PA25. PA27–28 situated mesal third of ventral surface, at ca. midlength of ventral parietale; PA27 pore-like, anteriorly to PA28; PA28 very long seta. Pore-like sensillum PA29 situated posteriorly to PA28, on basal third of ventral parietale; pore-like sensillum PA30 on posterior two-fifths of lateroventral surface, behind PA18.

Antenna ( Figs 11 A–B). Antennomere 1 with five pore-like sensilla (AN1–5); AN1 situated dorsally on posterior two-fifths, in lateral part of dorsal surface; AN2 dorsally on anterior fifth of sclerite, between AN1 and AN4 more distant from AN1 than AN4; AN3 subapically on dorsal part of lateral face, AN4 on subapically on dorsal part of inner face, AN5 ventrally on median portion of anterior margin of sclerite.Antennomere 2 with one pore-like sensillum (AN6) situated dorsally on subapical part of sclerite, laterally to midwidth of antennomere 2; setae AN7–8 and AN10–11 and sensorium SE1 on intersegmental membrane between antennomeres 2 and 3, AN9 absent; AN7–8 and SE1 on lateral face, AN7–8 minute, behind SE1, SE1 long and moderately stout, rounded apically, about as long as antennomere 3; AN10–11 on inner face, AN10 long, AN11 short, both setae close to each other. Antennomere 3 with one long additional seta situated on midlength of inner face, with apical sensilla (gAN) in apical membranous area.

Mandibles ( Figs 11 C–D). Mandible with two setae (MN1 and MN5) and three pore-like sensilla (MN2–4) situated apically to midlength; MN6 probably absent. MN1 rather short, situated laterally on mandibular midlength. MN2–4 situated anteriorly to MN1 and posteriorly to MN5, laterally to apical inner tooth; MN3 close to base of apical inner tooth, MN2 on ca. at midlength between MN1 and MN3, MN3 anterolaterally to MN2; MN5 minute, on apical fifth of lateral mandibular face.

Maxilla ( Figs 11 E–F). Cardo with one rather long ventral seta ( MX 1). Stipes with an irregular row of five setae ( MX 7–11) situated dorsally along inner face; MX 7 rather short and stout but shorter and narrower than MX 8–11, situated on basal margin of sclerite; MX 8–11 rather short, stout, almost equidistant from each other; MX 8 more distant from MX 7 than from MX 9. Pore-like sensilla MX 2–3 situated ventrally, MX 2 on midlength of inner face, MX 2 medially on posterior two-fifth; MX 4–6 ventrally on subapical part of lateral face of sclerite, MX 4 pore-like, MX 5 very long seta, MX 6 long seta; MX 4 basally to MX 6; MX 5 mesally to MX 4; MX 6 close to borderline between sclerite and intersegmental membrane. Dorsal surface of palpomere 1 with one rather long, stout seta ( MX 16) situated basally on inner face. Three sensilla ( MX 12–14) situated on median to lateral part of ventral surface; MX 12 pore-like, MX 13 very long seta, MX 14 rather long seta; MX 12 and MX 14 close to distal margin of sclerite, MX 14 mesally to MX 12; MX 13 on ca. anterior one-third, behind MX 12 and MX 14. Pore-like sensilla MX 15 and MX 17 situated on membrane below inner appendage, MX 17 dorsally, MX 15 ventrally. Inner appendage with a few short setae apically (gAPP). Palpomere 2 with two pore-like sensilla ( MX 18 and MX 19) and one minute seta ( MX 27); MX 18 situated ventrally on median and anterior part of sclerite; MX 19 dorsally on inner part of intersegmental membrane between palpomeres 2 and 3; MX 27 basally on lateral face. Palpomere 3 with two long setae ( MX 21 and MX 23) and two pore-like sensilla ( MX 20 and MX 22) situated on or slightly anteriorly to midlength of sclerite; MX 21 and MX 22 closely aggregated, situated on inner face, MX 22 dorsally to MX 21; MX 20 on lateral part of ventral surface, close to MX 23; MX 23 on lateral face. Palpomere 4 with one rather short and slightly stout seta ( MX 24) situated basally on inner face, and with digitiform sensillum ( MX 25) and pore-like sensillum ( MX 26) apically on outer face of sclerite; MX 25 dorsally, MX 26 ventrally. Apical membranous area of palpomere 4 with several minute setae (gMX).

Labium ( Figs 10B , 11 G–I). Submentum with two pairs of setae (LA1–2) on anterolateral portion; LA1 very long situated posteriorly to LA2, LA2 short. Ventral surface of mentum with one pair of long setae (LA3) and one pair of pore-like sensilla (LA4) situated medially close to distal margin of sclerite; LA3 mesally to LA4. Prementum and its anterior membranous area with five pairs of sensilla (LA8–12) on dorsal surface and with three pairs of sensilla (LA5–7) on ventral surface. LA5–7 situated on lateral part of sclerite, LA5 very short seta, LA6 very long seta, LA7 pore-like; LA5 basally, LA6–7 apically close to distal margin of sclerite, LA7 laterally to LA6. Pore-like sensilla LA8 situated medially on borderline between sclerite and membrane; LA9–12 close to each other, on anterior membranous area of prementum, LA9 and LA11–12 pore-like, LA10 long seta; LA9–10 behind LA11–12, LA10 laterally to LA9; LA12 mesally to LA11. Palpomere 1 with two sensilla (LA13–14); LA13 minute seta, situated ventrally on basal part; LA14 pore-like, dorsally on intersegmental membrane between palpomeres 1 and 2. Palpomere 2 with pore-like sensillum LA15 situated subapically on outer face. Apical membranous area of palpomere 2 with several setae of variable length and shape (gLA).

Second instar. Primary sensilla of second instar larva similar to that of first instar; secondary chaetotaxy of second instar similar to third instar.

Parietale with five secondary sensilla. Two rather short secondary setae situated close to PA8; one posteriorly to PA8, one anteriorly to PA8; one rather short seta and one pore-like sensillum between PA9 and PA19, seta close to PA9, pore-like one close to PA19; one rather short seta between PA14 and PA15.

Mandible and maxilla ( Figs 13 C–F). See mandible and maxilla of third instar (some setae are missing from examined specimen, but judging from the arrangements of secondary sensilla, secondary chaetotaxy of second instar larva almost same as that in third instar).

Labium ( Figs 13 G–H). Dorsal surface of mentum with two or three stout secondary setae on each side, situated on distal margin of sclerite; lateral-most ones very long, median one or ones rather short.

Third instar. Very similar to second instar.

Mandible ( Fig. 15 C–D) with four to six secondary setae situated on outer face posteriorly to MN1; one minute close to MN1, one to two minute posteriorly to MN1, two to three rather short on basal part.

Maxilla ( Figs 15 E–F). Stipes bearing two secondary setae situated on ventral part of outer face; rather short one close to MX 5, remaining rather long one on basal part.

Comments on the synonymy. As we explain above in the Species-level taxonomy section, we adopt here the conservative approach and distinguish only two reciprocally allopatric species of Tormus which are both diagnosable morphologically. The reciprocal allopatry allows us to establish synonymies of T. helmsi on the basis of the position of type localities of the respective species within the known range of T. helmsi ( Fig. 16 ). Three species described by BROUN (1893b,c, 1910), Tormus nitidus , Stygnohydrus nitidus and Stygnohydrus femoralis , hence become synonyms of T. helmsi . Of these, T. nitidus and S. nitidus belong to the same genetically distinct clade as T. helmsi (which is here hence called the ‘ helmsi s.str. ’ clade), whereas S. femoralis represents the North Island clade called here for that reason the ‘ femoralis ’ clade. Remaining two clades of T. helmsi do not correspond to any of the original species described by historical authors, and are therefore called ‘Okarito’ and ‘Haast’ clades following their known geographic ranges.

Distribution. The species occurs in the central and southern part of the North Island (but is likely absent from the northern and eastern parts of the island) and along the western coast and in the Southern Alps Mts. of the South Island, reaching southwards to southern Westland (Haast). We have found a single male specimen localized Reef Point, Antipodes Is., 28.ii.1969, lgt. G. Kuschel (69/70)’ in the collection of NZAC. As Antipodes Islands are however more or less bare rocks without vegetation ca. 700 km southeast from New Zealand (whereas Tormus is shown here to inhabit forests) and as Tormus was never recorded from these islands before ( ORDISH 1974, MARRIS 2000), we consider this specimen as mislabeled and do not list it in the material examined above.

Figures

Fig. 1. Habitus of Tormus helmsi Sharp, 1884. A – adult; B–D – larva, first (B), second (C), and third (D) instar.

Fig. 7. Morphology of Tormus helmsi Sharp, 1884.A – prothoracic leg; B – mesothoracic leg; C – metathoracic leg; D – metafurca; E– female genitalia; F – male sternite 9.

Fig. 9. Variability of the aedeagus morphology in Tormus posticalis (Broun, 1917) (A–B) and T. helmsi Sharp, 1884 (C–N). Data on the specimens: A – Westland, Jackson Bay (voucher COL1770); B – Fiordland: Milford Sound (voucher D2); C – Taranaki, Mt. Egmont NP (voucher MF500); D – Bay of Plenty, Mt. Te Aroha (voucher D5); E – Waikato, Mt. Pirongia; F – Buller, Bullock Creek; G – Nelson, Oparara Arches (voucher MF499); H – Buller, Paparoa NP (voucher MF503); I – Buller, Italians Creek; J – Westland, Three Mile Lagoon (voucher MF506); K – Westland, Pleasant Flat; L – Westland, Cross Creek (voucher MF498); M – Westland, Gillespies; N – Westland, Matheson Lake (voucher COL1802).

Fig. 8. Results of phylogenetic analyses of Tormus Sharp, 1884 and its genetic diversity. A – map of localities of the sequenced specimens. B – Bayesian analysis with posterior probabilities above/below branches (outgroup and values lower than 50 omitted).C – maximum likelihood analysis with bootstrap support values above/below branches (outgroup and values below 50 omitted) and mapped clade specific morphological character and conserved amino acid changes. D – haplotype network with grey numbers indicating the number of substitutions on each branch. Color marking indicates the clades/clusters, numbers in color circles indicate different localities.

Fig. 14. Tormus helmsi Sharp, 1884, second (F) and third (A–E, G) instar larva.A – head, dorsal view; B – detail of anterior margin of head capsule, dorsal view; C – anterior part of dorsal surface of pronotum; D – detail of seta on anterior margin of pronotum; E – prosternum, ventral view; F – meso- and metanotal sclerites, dorsal view; G – spiracular atrium, dorsal view.

Fig. 15. Head appendages of Tormus helmsi Sharp, 1884, third instar larva.A–B – antenna, dorsal (A) and ventral (B) view; C–D – mandibles, dorsal view; E–F – maxilla, dorsal (E) and ventral (F) view; G–J – labium, dorsal (G), lateral (palpus omitted) (H), ventral (I) view, and detail of anteromedian surface of prementum, anterior view (J) (palpomere 2 of right palpus missing).

Fig. 10. First instar larva of Tormus helmsi Sharp, 1884. A–B – head capsule, dorsal (A) and ventral (B) view; C – detail of anterior margin of head capsule, dorsal view.

Fig. 12. First instar larva of Tormus helmsi Sharp, 1884. A – prosternum, ventral view; B – mesothoracic leg, anterior view; C – tubercle on dorsal surface of abdomen.

Fig. 13. Head appendages of Tormus helmsi Sharp, 1884, second instar larva.A–B – antenna, dorsal (A) and ventral (B) view; C–D – mandibles, dorsal view; E–F – maxilla, dorsal (E) and ventral (F) view; G–H – labium, dorsal (G) and ventral (H) view.

Fig. 11. Head appendages of Tormus helmsi Sharp, 1884 first instar larva. A–B – antenna, dorsal (A) and ventral (B) view; C–D – mandibles, dorsal view; E–F – maxilla, dorsal (E) and ventral (F) view; G–I – labium, dorsal view (G), detail of dorsal surface of labial palpus (H) and ventral view (I).

Fig. 16. Summary of known distribution of Tormus Sharp, 1884, combining the distribution data of the museum specimens (black and grey symbols), known distribution of genetically distict clades based on sequenced specimens (color symbols: colors and locality numbers correspond to those in Fig. 8) and the position of the type localities of all five described species of Tormus and Stygnohydrus Broun, 1893.