Iteaphila nitidula Zetterstedt

Sinclair, Bradley J. & Shamshev, Igor V., 2012, 3561, Zootaxa 3561, pp. 1-61 : 29-37

Iteaphila nitidula Zetterstedt

( Figs. 1C, 3B, 8B, 9, 10)

Iteaphila nitidula Zetterstedt, 1838: 541 . Type locality: Dalekarlia , Sweden [= Dalarna].

Hilara migrata Walker, 1849: 491 . Type locality: St. Martin’s Falls, Canada; Smith, 1971: 354.

Hilara obscura Meigen : Zetterstedt, 1849: 3018. Misidentification.

Hilara carbonella Zetterstedt, 1859: 5003 . Type locality: Wittangi , Sweden [after lectotype designation]. syn. nov.

Steleocheta stiriensis Becker, 1891: 284 . Type locality: Rolle-Pass [ Italy].

Steleocheta meridionalis Becker, 1892: 126 (new name for Steleocheta stiriensis Becker, 1891 ). Unnecessary name change.

Empis conjuncta Coquillett, 1900: 411 . Type locality: Alaska, USA. syn. nov.

Iteaphila fuscipennis Frey, 1913: 39 (as variation of Iteaphila obscura sensu Frey ). Type locality: Lt. Kola [ Russia: Murmansk Province]; Tuomikoski, 1958: 130.

Iteaphila styriensis: Melander, 1928: 104 , error or emendation.

Iteaphila americana Melander, 1946: 32 . Type locality: Yellostone Park, USA; Tuomikoski, 1958: 130; Smith, 1971: 354.

Type material examined. Iteaphila nitidula : HOLOTYPE ♀, labelled: “Dlc. [= Dalekarlia]”; “Bhm. [= Boheman]”; “ HOLOTYPE / Iteaphila / nitidula / Zetterstedt, 1838 / design. 2007 Shamshev & Sinclair [red; due to poor condition of pin, specimen was re-mounted]” ( NHRS).

Hilara carbonella : LECTOTYPE ♂ (here designated), labelled: “ H. obscura M.? / Wittangi. ♂ [white, Zetterstedt’s handwritting]”; “ Lectotypus / Hilara / carbonella / Z. 1859/ design. 2004/ I. Shamshev [red, handwritten]”; “ Empididae , Type no. 3034: 1–2, Zool. Mus. Lund Sweden ” ( MZLU) . PARALECTOTYPE: 1 ♂ from Finland (Wallengren Collection (292: 4), MZLU) .

Steleocheta stiriensis : HOLOTYPE ♀, labelled: “Rolle-Pass/ 20|6.90, 25693”; “meridio-/nalis B. [Becker's hand writting]”; “? Typus”; “Zool. Mus. Berlin ”; “ Holotype [red]”; “ Iteaphila nitidula Zetterstedt, 1838 / det. Shamshev & Sinclair, 2004 [wings and postpedicel on one antenna missing]” ( ZMHB).

Iteaphila fuscipennis : HOLOTYPE ♂, labelled: “Kola/ Palmén/ 469 [pink]”; “ HOLOTYPUS, Iteaphila obscura Zett. var. fuscipennis Frey, 1913: 39 , ident. by Chvála, 1980” ( ZMH).

Iteaphila americana : HOLOTYPE ♂, labelled: “YellstPark/ Thumb Sta/ 16 July [19]23/ ALMelander”; “ HOLOTYPE / Iteaphila / americana/ Mel.”; “ALMelander/ Collection/ 1961” ( USNM) . PARATYPES: CANADA. British Columbia: Vancouver , 13.v.1916 (2 ♀, USNM) . Quebec: Aylmer , 23, 28.v.1923, C.H. Curran (1 ♂, 2 ♀, USNM) . USA. Colorado: Monarch Pass , 8000 & 11400 ft., 21.vi.1940, ALM (2 ♀, USNM) . Idaho: Lake Waha, 9.vi.1918, ALM (1 ♂, 3 ♀, USNM); Moscow Mt. , 17.vi.1916, 26.vi.1920, ALM (1 ♂, 1 ♀, USNM) . Missouri: Atherton , 13.iv.1901 (1 ♂, USNM) . Montana: Glacier Park, Grinnell Lake , 24.vii.1935, ALM (1 ♀, USNM) . New Mexico: Las Vegas Mts. , 11,000 ft., vi.1901, Cockerell ( USNM) . Oregon: Mt. Hood , 5000 ft., 25.vi.1935, ALM (1 ♀, USNM) . Washington: Deer Park, 5.v.1912 (3 ♀, USNM); Ilwaco, 25.v.1917, ALM (1 ♀, USNM); Mt. Baker, Skyline Trail, 10.viii.1925, ALM (1 ♀, USNM = Anthepiscopus ); Mt. Rainier , Paradise, 28.vi.1935, ALM (2 ♂, USNM); Mt. Rainier , van Trump, 21.vii.1922, ALM (1 ♂, 1 ♀, USNM); Olympia, 24, 31.iii.1894 (2 ♂, USNM); Seattle, 17.iii.1896, 24.iii.1903, 9.iv.1899, 14.iv.1898, 15.iv.1898 (5 ♂, 1 ♀, USNM); Snoqualmie Pass, 29.vi.1924, ALM (1 ♀, USNM) . Wyoming: same data as holotype (5 ♀, USNM) .

Empis conjuncta : LECTOTYPE ♂ (here designated), labelled: “ Sitka Alaska / June 16 ’99”; “ Harriman Expedition’ 99/ T. Kincaid , Collector ”; “ Type / No. 5215/ U.S. N.M. [red label]”; “ Empis / conjuncta/ Coq. [Coquillett’s handwriting]”; “ Iteaphila / conjuncta/ Coq. [hand written]” ( USNM) . PARALECTOTYPES: same as lectotype (1 ♀, USNM); Orca, 27.vi.1899 (1 ♀, USNM) .

Taxonomic notes. Zetterstedt (1838) described I. nitidula after a single specimen (female), which he received for identification from Boheman and which was taken from Dalekarlia (mountain area in middle Sweden). In Zetterstedt's collection in Lund there is one specimen of this species (female), however, it is not the holotype because it was collected by Zetterstedt in Sweden Lapland. Probably when Zetterstedt described the new species, he returned the single specimen to Boheman in Stockholm, where among other specimens of I. nitidula there is a female with the same data as in Zetterstedt's original description ( Tuomikoski 1958). We examined all specimens of I. nitidula housed in Lund and Stockholm and added a holotype label to this female specimen. See Michelsen (1985) for discussion of Zetterstedt type depositories.

The authors of the last catalogues on Empididae ( Chvála & Wagner 1989; Yang et al. 2007) indicated that Hilara obscura was described by Zetterstedt (1849) as a separate species. However, our comparison of Zetterstedt’s papers and his collections indicate that it is a misleading conclusion. Zetterstedt (1842: 359) described Hilara minuta (= Ragas unica Walker ) and in the end of the description of this species he made a note that he has a male taken from “Wittangi Lapponiae Tornensis d. 14 Jun. 1821 inveni” that possibly is Hilara obcura Meigen. But because he had only one specimen in bad condition and had no female he did not discuss this species. However, Zetterstedt (1849: 3014 and 3018) included H. obcura under number 32 in his list of species of Hilara indicating Meigen as the author of the specific name. At the same time he noted a male from “Westrogothia valde rara, d. 6 Jun. a Prof. Boheman inventa” and did not note a specimen that he had indicated in 1842. In the Diptera Scandinavica Collection (Lund) at the end of material on the genus Hilara there is one specimen with the name H. obcura (“ H. obscura M?, Wittangi, ♂ ”), which is the specimen noted by Zetterstedt in 1842. Thus, it is evident that Zetterstedt never described the species with the name H. obscura . An additional supporting argument is that Zetterstedt (1859: 5003) described Hilara carbonella ( Diptera Scandinavica Collection, Lund; species No. 34 box with Hilara ) and indicated that the specimen from “Wittangi” noted by him in 1842 belongs to this species together with other specimens taken from Finland (“Alterum specimen masculum in Finlandia ad Enare d. 26 Jun. 1856 captum, bene conservatum, mecum communicatum D. Profess. E.J. Bonsdorff”). We examined both these specimens and designated a lectotype to clearly establish the identity of this species. Chvála (pers. comm. 2012) studied the types of Hilara carbonella at Lund in 1996 and identified it to be a species of Iteaphila , but mistakenly forgot to include this nomenclatural change in his book on Hilara of Fennoscandia ( Chvála 2005).

It should be noted that Frey (1913) indicated mistakenly Zetterstedt as the author of H. obscura , transferred this “species” to the genus Iteaphila and provided a re-description. However, a figure of the male genitalia given by Frey (1913, fig. 3) indicates clearly that it is I. nitidula (see also Tuomikoski 1958).

Becker (1891) described Steleocheta stiriensis (= S. meridionalis ) after a single specimen taken from Italy (Rolle-Pass). We added a red holotype label to fix formally the type status of this specimen. In addition, a determination label stating that this species is syntypic with I. nitidula was added.

Frey (1913) described Iteaphila fuscipennis as a variation of I. obscura Zetterstedt sensu Frey. We examined the holotype of this species and conclude with Tuomikoski (1958) that this species is a junior synonym of I. nitidula .

The synonym, Empis conjuncta , was designated in the unpublished thesis of Anderson (1973). We have examined the type series and confirm this conclusion.

Additional material examined. BULGARIA. Pirin, Banderica, 7.vii.1982, 1800 m, F. Kocourek (1 ♂, CHVC). CANADA. Alberta: Banff-Jasper Hwy, Sunwapta Pass, 6600', 5.vii.1955, R. Coyles (1 ♂, CNC); Banff NP, 14 mi W Banff, 4500', 7.vii.1955, on Ledium groenlandicum flowers, GES (1 ♂, 1 ♀, CNC); Banff NP, Mt. Eisenhower, 11, 16.v.1968, Mosquin & Seaborn (2 ♂, CNC); Dunvegan, N side Peace R, 22.v.-6.vi.1995, pan tp in grass, S. Boucher (1 ♂, CNC); Edmonton, 21.v.1925, O. Bryant (2 ♂, CAS); Edmonton, Ecol. Res., 29.vi.- 5.vi.1986, B. V. Brown (3 ♂, 9 ♀, CNC); Elkwater, 11.vi.1956, O. Peck (2 ♀, CNC); Jasper, 3.vii.1932, E. Hearle (1 ♀, CNC); Kananaskis, For Exp. Sta. Seebe, 21.vi.1968, H.J. Teskey (1 ♀, CNC); Waterton Lks NP, Cameron Lk, 17.vi.1956, E.E. Stern (1 ♀, CNC); Ptarmigan Trail, 6500', 20.vii.1928, O. Bryant (1 ♀, CAS). British Columbia: Alaska Hwy, 37 km W Ft. Nelson, MT, 12.vi.-5.ix.1984, S & J Peck (1 ♀, CNC); Alaska Hwy, milepost DC 520 mi (831 km), Whirpool Canyon, 9.vi.1996, PHA (2 ♂, 7 ♀, USNM); Alaska Hwy, mi 392, Summit Lk, 16.vi.- 4.vii.1959, R.E. Leech (5 ♂, 27 ♀, CNC); Alliford Bay, QC Islands, 29.v.1957, E.E. MacDougall (2 ♀, CNC); Atlin, 2200', 13.vi.1955, H. Huckel (3 ♂, CNC); Bevan, 18.vi.1955, J. R. McGillis (1 ♀, CNC); Bowen Is, 24.iv.1960, GJS (1 ♂, UBCZ); Bowser, 5-7.vi.1955, J. R. McGillis (1 ♂, 8 ♀, CNC); Burnaby Mt., 26.iv.1979, D. Gillspie (2 ♀, CNC); Campbell R, Goslin Lk, 25.v.1988, G. Hutchings (1 ♂, RBCM); Cathedral PP, Glacier Lk, 7.vii.1986, R.A. Cannings (1 ♂, RBCM); Cathedral PP, Quiniscoe Lk, 7.vii.1986, S.G. Cannings (1 ♂, UBCZ); Circle Hot Springs, 24.vi.1996, PHA (1 ♀, USNM); Cottonwood R, Hwy 97, 23.v.1981, L. Vasington & S.G. Cannings (1 ♀, UBCZ); Devonian Regional Pk, 1.5 km S Metchosin, Sherwood Ck, 48º21.9'N 123º32.1'W, 9.iv.2004, C.J. Borkent (1 ♀, CNC); Duncan, 9.vi.1955, GES (1 ♀, CNC); Galiano Is, 28.iv.1974, 12.v.1980, G.G.E. Scudder (2 ♀, UBCZ); Hart Hwy, Bear Lk Camp, 72 km from Prince George, 24.vi.1978, PHA (1 ♂, USNM); Hixon, 21.vi.1976, E. Dyer (1 ♀, CNC); Horne Lk, 4.vi.1955, R. Coyles (1 ♀, CNC); Juskatla, QC Islands, 12-14.vi.1957, E.E. MacDougall (2 ♀, CNC); Kamloops, 11.vi.1939, 10.vi.1945, GJS (2 ♀, UBCZ); 4 km E Kingfisher, 50º36.92'N 118º40.801'W, flowers of Shepherdia , 11.iv.2010, AB (2 ♂, CNC); King Salmon Lk, 58º43', 132º54', 14.vii.1960, WWM (1 ♂, CNC); Kitimat, 2.vi.1960, JGC (2 ♀, CNC); Kitsumkalum Lk, 20 mi N Terrace, 31.v., 16.vi.1960, JGC, R. Pilfrey (2 ♀, CNC); Klanawa, 30.v.1952 (1 ♀, CNC); Kootenay NP, Daer-Pitts Aspen Control/Burn 1–2, 2–9.vi., 3–17.vi., 17.vi.-2.vii., 25.vi.-9.vii., 2–16.vii.2000, MT, G. Gareau (3 ♂, 8 ♀, CNC); Lac la Hache, 27.vi.1964, L.H. McMullen (1♀, CNC); Lisadele Lk, 58º41', 133º04', 9.vii.1960, WWM (1 ♂, CNC); Manning Pk, Dry Ridge summit, 1830 m, 25.vi.1983, S.G. Cannings (1 ♂, UBCZ); Manning Pk, 1700 m, 27.vi.1984, R. Danielsson (1 ♂, MZLU); Moresby Is, Sandspit, 12 km W, 10.v.1984, G.G.E. Scudder (1 ♂, 10 ♀, UBCZ); Mt Revelstoke, 17.vii.1952, GJS (1 ♀, CNC); Pleasant Camp, 2.vi.1981, CSG (1 ♂, UBCZ); 4 mi S Prince George, Hwy 17, Southpark RV Pk, 2.vi.1996, PHA (2 ♀, USNM); Qualicum, 15.vi.1955, GES (1 ♀, CNC); Qualicum Bay, Horne Lk, 28.v.1955, J. R. McGillis (1 #, CNC); Queen Charlotte Islands, T’lell, 30.vi.1955, G. Holland (1 ♂, 1 ♀, CNC); QC Islands, T’lell, 25.v.1957, E.E. MacDougall (2 ♀, CNC); QC Islands, Alliford Bay, 29.v.1957, E.E. MacDougall (2 ♀, CNC); QC Islands, Laskeek Bay, Lost Is., 15.v.2001, N 52º52'39''W 131º31'07'', S. Allombert (15 ♂, 12 ♀, RBCM); Robson, 25.v., 3.vi.1968, H. R. Foxlee (2 ♀, UBCZ); Salmon Arm, 2.v.1991, AB (2 ♂, 1 ♀, CNC); 3 km E Salmon Arm, 4.v.1992, AB (1 ♀, CNC); Salmon Arm, 16.iv.-26.v.1989, 2.v.1991, 1.v.1992, AB (3 ♂, 1 ♀, CNC); 6 km E Sicamous, 30.iv.1992, AB (1 ♂, CNC); 8 km E Sicamous, 8.vi.1992, AB (1 ♂, 3 ♀, CNC); nr Terrace, Lakelse Lk, 27.vi.1960, R. Pilfrey (1 ♀, CNC); nr Terrace, Mt. Thornhill, 21.vi.1960, GES (1 ♂, 1 ♀, CNC); 5 mi S Terrace, 3.vi.1960, JGC (3 ♀, CNC); 32 mi SW Terrace, ex. Heracleum lanatum , 4–11.vi.1969, GES, CHM, JGC, R. Pilfrey, B. Heming (6 ♂, 19 ♀, CNC); Tunjony Lk, 58º26', 132º45', 18–19.vii.1960, WWM, R. Pilfrey (3 #, CNC); Vancouver Is, Point Grey, 1.v., 9.iv.1973, JRV (2 ♀, CNC). Labrador: Goose Bay, 25.vi.1948, W.E. Beckel (1 ♀, CNC). Manitoba: Churchill, 20.vi.1948, GES (1 ♂, CNC); nr Churchill, burn S Twin Lks, 58º36.92'N 93º49.33'W, 11–15.vii.2009, MT 3, J.M. Cumming (2 ♂, 3 ♀, CNC). New Brunswick: Kouchibouguac NP, 25.v.1977, B. Cooper, Code 6133K (1 ♂, CNC); ditto, 8.vi.1977, JRV, Code 5222 V (2 ♀, CNC). Northwest Territories: Aklavik, 16.vi.1931, O. Bryant (1 ♀, CAS); Norman Wells, 9.vi.1949, W. R.M. Mason (1 ♂, CNC). Nova Scotia: Cape Breton Highlands NP, MacKenzie Mtn, 400 m, 27.v.-8.vi.1984, BEC (6 ♂, 5 ♀, CNC); Cape Breton Highlands NP, Pleasant Bay, 7–11.vi.1984, BEC (2 ♂, 2 ♀, CNC); Cape Breton Highlands NP, North Mt., bog, 31.v.-3.vi.1984, BEC (6 ♂, 5 ♀, CNC); Cape Breton Highlands NP, Cheticamp R., 26–30.v.1984, BEC (3 ♀, CNC); Cape Breton Highlands NP, Beulach Ban Falls, 28.v.1984, BEC (1 ♀, CNC). Ontario: Algonquin PP, Swan Lake area, 13–15.v.1990, J.E. O’Hara (1 ♂, CNC); Bruce Co., Black Ck PP, 28.v.2000, Potentilla anserina flowers, J.J. Dombroskie (1 ♂, DEBU); Bruce Co., Crane R., 8.v.1997, SAM (1 ♂, DEBU); Fathom Five NP, Flowerpot Is., Marl Bed, Mossy Cedar, pan trp#2, 23.vi.-30.v.1995, T. Woodcock & SAM (1 ♂, DEBU); 7 mi E Griffith, 23.v.1988, 3.vi.1990, BEC (4 ♂, 1 ♀, CNC); Haldimond-Norfolk Co., Manestar Tr., 6 kmNNW St. Williams, 42°42'17''N 80°27'38''W, 8.v.2008, J.H. Kits (2 ♂, 1 ♀, DEBU); Low Bush, Lake Abitibi, 5.v.1925, N.K. Bigelow (1 ♀, CNC); Maynooth, 23.v.1970, J.F. McAlpine (1 ♀, CNC); Metcalfe, 10.v.1986, BEC (1 ♀, CNC); North Gower Rd 6 & Montague Boundary Rd, N 45º02' W 75º54', MT, 14–16.v.2004, A. Bennet & D. Barnes (1 ♀, CNC); Northumberland Co., Peter’s Woods PNR, N 44º7'26'' W 78º2'31'', forest MT, 19.v.-1.vi.2011, Brunke & Paiero (1 ♀, DEBU); Ottawa, 29.v.1997, JRV (1 ♀, CNC); Ottawa, Rockcliffe Pk, 14.v.1987, BEC (1 ♀, CNC); Shaw Forest, 6 km NNE Eganville, MT, N 45º38' W 77º04', 27.v.-3.vi.1993, H. Goulet, M. Sharkey (1 ♀, CNC); Tobermory, Dunks Bay, 15.vi.1996, SAM (2 ♀, DEBU); Wellington Co., Fergus, MT, 1.vi.1992, SAM & Cannings (1 ♀, RBCM). Quebec: Gatineau Pk, Harrington Lk, 30.v.1954, J.E.H. Martin (1 ♀, CNC); Gatineau Park, King Mtn, summit 45°29'20''N 75°51'45''S, 13.v.2009, J.M. Cumming (1 ♂, 2 ♀, CNC); Kingsmere (Gatineau Pk), 12.v.1958, JGC (1 ♀, CNC); Mont St. Hilaire, 45º32.6'N 73º9.1'W, MT, 10–28.v.2001, E. Fast, S. Hawkins (5 ♂, 2 ♀, LEM); Old Chelsea (Gatineau Pk), 26.v.1964, JRV (1 ♀, CNC); St. Flore, Lac Mondor, 17.v.1951, E.G. Munroe (1 ♂, CNC). Yukon: Alaska Hwy, Slims R. delta along Sheep Ck Rd, km 1706, 60º59'N 138º34'W, MT, 10–12.vi.1979, #791020a, ROM Fld.Pty (1 ♀, ROM); Dempster Hwy, km 49, 4.vi.2003, AB (1 ♂, CNC); Dempster Hwy, mi 51, 17–21.vi., 25–27.vi.1973, G. Wood & DMW (14 ♂, 7 ♀, CNC); Dempster Hwy, km 68, N Klondike R, 64º29'N 138º25'W, 17.vi.1981, CSG (1 ♂, UBCZ); Dempster Hwy, km 155, pan tp, 28.vi.-2.vii.1982, G. Wood & DMW (1 ♀, CNC); Klondike Hwy, km 12, Ethel Lk Rd, sweeping roadside, 63º19'N 136º21'W, 30.vi.1980, #800067c, ROM Fld.Pty. (1 ♀, ROM); Klondike Loop, J 243 mi (390 km), McQueston, 14.vi.1996, PHA (3 ♀, USNM); North Fork Crossing, mi 43, Peel Plateau Rd., 3500', 26.vi.-5.vii.1962, R.E. Leech & PJS (7 ♀, CNC); Old Crow, 6 mi upriver Porcupine R, MT, 67º34'N 139º41'W, 4–5.vii.1981, #810537d, R. Jaagumagi (1 ♀, ROM); South Carol Rd., km 160 at Lapie Lks, sweeping, 61º37'N 133º04'W, 15.vi.1980, #800034d, ROM Fld.Pty (1 ♀, ROM). CZECH REPUBLIC. Beskydy Mts, Lysá hora hilltop, MT, 1323 m, 49°32'46''N 18°26'53''E, 21.v.-7.vii.2004, MB (1 ♂, 8 ♀, BARC); Bílina, Štrbický vrch, edge of wood, YPWT 50.33.10N 13.50.40E, 400 m, 23–30.iv.1997, MB (2 ♂, 1 ♀, BARC); Bílina, Vršíček, mixed wood, 440 m, 50°33'12''N 13°49'57''E, MT, 2.iv.-14.v.1998, MB (1 ♂, BARC); Bohemia, Kleť Holubov-Křemže br., 23.v.1978, M. Chvála; Stellaria blossoms (1 ♂, CHVC); Bohemia, Lijmo-Luč, 6.v.1993, J. Máca (1 ♂, CHVC); Krkonoše, Liščí hora, MT, upper forest line, 1320 m, 50°42'04''N 15°40'35''E, 29.vi.-26.vii.2005, Vanĕk (1 ♂, BARC); MS Beskydy, Muřinkový, peat-bog, 49.31N 18.39E, 950 m, 13.vi.1987, MB (2 ♀, BARC); Prachatice, Libínské Sedlo, 48.59N, 13.59E, 800 m, 9.v.1987, MB (1 ♂, BARC); Spálenec-Prachatice, car net, 21.v.1992, MB (1 ♂, BARC); Šumava Mts. 1100 m, Rokytecká slať, peat-bog, 49°00'59''N 13°25'05''E, PT, 18–20.v.1999, MB, Kubik (3 ♂, 1 ♀, BARC); Šumava Mts. 1100 m, Rokytecká slať, peat-bog, 49°00'59''N, 13°25'05''E, MT, 18.v.-16.vi.1999, MB, Kubik (1 ♂, 2 ♀, BARC); Šumava, Zhůřské slatĕ, peat-bog, 1140 m, 17.vi.-21.vii.1999, MT 49°04'30''N 13°33'54''E, MB, Kubik (1 ♀, BARC) Šumava Mts. 1130 m, Zhůřské slatĕ, peat-bog, 49°04'N 13°34'E, PT, 18–20.v.1999, MB, Kubik (1 ♂, BARC); Šumava, 870 m Nová Hůrka, peat-bog, 49°09'N 13°20'E, PT, 18–20.v.1999, MB, Kubik (1 ♀. BARC); Šumava Mts. 870 m Nová Hůrka, peat-bog, 49°09'N, 13°20'E, MT, 18.v.- 16.vi.1999, MB, Kubik (1 ♀, BARC); MS Beskydy, H.Lomná—Hruška, small brook, Picetum, 43°30'29''N 18°36'56''E, 740 m, 23.v.-19.vi.1999, MT, MB (1 ♀, BARC); Šumava, Malá Niva, peat-bog, 750 m, MT, 29.iv.- 16.vi.1999, 48°54'35''N 13°49'21''E, MB & Kubik (1 ♀, BARC); Šumava, Boubin, PT, 1250–1350 m, 48°59'27''N 13°49'01''E, vi.2004, J. Farkač (20 ♂, 10 ♀, BARC); Šumava Mts, Trojmezná, Malaise trap, 16.v.-1.vi.2003, J. Farkač (3 ♀, BARC); Šumava, Boubín, mixed wood, 990 m, MT, 48°58'40''N 13°49'00''E, 15–28.v.2002, 18.v.- 12.vi.2002, K. Spitzer (2 ♂, 4 ♀, BARC); Šumava, Borová Lada, 2 km W, damp meadow nr. river, 49.00N 13.39E, 910 m, 4.vi.1996, MB (1 ♀, BARC); U. Janovice-Kostelec, car net, 8.v.1992, MB (1 ♀, BARC); Vraz u Pisku, mixed wood, 49.23N 14.08E, 400 m, 22–25.v.1995, MB (1 ♀, BARC). FINLAND. Fennia, Ab, Karislojo, Krogerus, 5.vii.1932 ( ZMUC); Kuusamo, 1549, R. Frey (1 ♀, CHVC); Utsjoki, 7157, Hellén, 7157 (1 ♂, CHVC). GERMANY. Bayerischer Wald, Spiegelau, 3 km N loc. 1, 48.57N 13.22E, 760 m, 8.vi.1995, MB (8 ♂, 8 ♀, BARC; 2 ♀, CHVC); Wölfelsgrund, Schles. 16.v.1921, L. Duda, Pseudoragas velutinus (1 ♂, ZMHB); Wustung, b. Habelschweldt, 6.v.1927, L. Duda; Pseudoragas velutinus (2 ♀, ZMHB); same locality, 31.v.1923 (1 ♀, ZMHB). JAPAN. Ehime: Shikoku: Matsuyama, 4.iv.1957, M. Takahashi (1 ♂, KUMF). Fukuoka: Kôrasan, Kurume, 21.iii.2001, S. Ide (29 ♂, 8 ♀, KUMF); ditto, 26.iii.2004, S. Ide (5 ♂, 12 ♀, KUMF). Hokkaido: Upepesanke, alpine zone, 20.vii.1967, TS (2 ♂, 5 ♀, KUMF); Sugataminoike, Taisetsuzan, 24.vi.1986, TS (2 ♂, KUMF); Takagi–S. (1 ♂, KUMF); 28.v.1957, M. Takahashi (1 ♀, KUMF); Nibushi, Kutcharo, Kushiro, 8.vi.1957, M. Takahashi (1 ♀, KUMF). Kumamoto: Haki, Izumimura, 21.iv.1980, K. Ohara (2 ♂, KUMF). Nagano: Okushiga-gohgen, 31.v.1975 (1 ♂, 2 ♀, KUMF). Yamanashi: Gozeishi, Hoozan, 2.v.1961, TS (1 ♀, KUMF). RUSSIA. Kamchatka: Karyak Autonomous Okrug, env. of Apuka, Pakhachinskiy Range, shrubby tundra, 200–300 m, 3.vii.1959, Gorodkov (4 ♂, ZIN). Karelia: Khumala & Polevoy, 2009. Khabarovsk Territory: Ozerpakh, estuary of Amur River [53°02'32''N, 141°14'45''E], 8–9.vi.1915, Chernavin (4 ♂, ZIN); same locality, 13.vi.1915, Chernavin (1 ♂, 3 ♀, ZIN); same locality, 15.iv.1915, Chernavin (10 ♂, 4 ♀, ZIN). Magadan Prov.: Indigirka R, mouth of Inyali river, border of larch forest near brook, 18.vi.1976, V. Kovalev (1 ♂, ZMMU). Murmansk Prov.: Vudyavr Lake, Khibiny, Kola Peninsula [67°37'50"N, 33°40'15"E], 28.vi.1930, E. Belyakova (1 ♀, ZIN). Tyumen' Prov. (Khanty-Mansy Autonomous Okrug): Khanty-Mansy AO, E Ural, 63.818°N 59.562°E, 622 asl, 6–8.vii.2010, K.Tomkovich (3 ♂, ZMMU). Yakutia: Mikhaylovka, 60 km NE of Amga, 17.vi.1986, Bagachanova (1 ♂, ZIN); Jakutskaja obl., Suntar-Chajata Mts. 900 m, vegetation, 26.vi.1988, Horák (1 ♀, BARC). SLOVAKIA. Malá Fatra, N slopes of Stoh 1300 m (3), 1.vi.1983, M. Chvála (2 ♂, CHVC); Vysoké Tatry, Hrebienok, 29.v.1968, M. Chvála (1 ♂, CHVC); Zadnå Polana, 1250 m, spruce wood, MT, 6.v.-3.vii.2006, 48°39'40''N, 19°29'50''E, Ševčík & Roháček (1 ♂, 5 ♀, BARC). SWEDEN. Abisko, To., leg. O. Ringdahl (2 ♂, 1 ♀, MZLU); T. Lpm. Abisko, loc. 1. Abiskodalen, 7.vii.1957, P.I. Persson (2 ♀, MZLU); T. Lpm. Abisko, loc. 4. Nuoljatunneln, 13.vii.1957, P.I. Persson (1 ♀, MZLU); T. Lpm. Stordalem, loc. 2, 6.vii.1957, P.I. Persson (1 ♂, MZLU); Sm. Växjö, S Åreda, 29.iv.1989, R. Danielsson (1 ♀, CHVC; 29 ♂, 3 ♀, MZLU). TAIWAN. Tonpogoe, 2500 m, Kagi-Keu, 29.iii.1967, T. Shirozu (1 ♀, KUMF); vicinity of Taataka-anbu, ca 2700 m, Kagi-Keu, 3.iv.1967, T. Shirozu (1 ♂, KUMF); Tungpu-Taataka, Lulinshan, Nantou Hsien, 10.iv.1965, TS (2 ♂, KUMF); Tungpu-Taataka, Lulinshan, Nantou Hsien (1 ♀, KUMF). USA. Alaska: Anchorage, Chugach Mt, 2600', Salix catkin, 30.v.1948, Sailer & Sommerman (1 ♂, USNM); 173 mi NE Anchorage, Glenn Hwy, 27.v.1951, R.I. Sailer (1 ♀, USNM); 11 mi S Anderson Jct, Rte 3, mi 270, 23.vi.-11.viii.1984, MT, S&J Peck (1 ♀, CNC); Chena R Rec Area, 30 mi E Fairbanks, 22.vi.-12.viii.1984, MT, S&J Peck (1 ♂, 3 ♀, CNC); Douglas, 16.vi.1949, W.C.F. (1 ♀, USNM); Eagle Summit, 3800', 25.vi.1948, R.I. Sailer (1 ♀, USNM); Haines, 19.vii.1954, W.C.F. (1 ♀, WSU); Igloo Ck, 24.vi.1954, W.C.F. (1 ♀, WSU); Ketchikan, 3-28.vi.1954, 26.vii.1955, 11.vi.1956, F. Baker (1 ♂, 4 ♀, WSU); Mt. McKinley NP, Savage R, 790 m, 17.vii.1978, PHA (1 ♀, USNM); Naknek, AFB, 9.v.1954, W.C.F. (1 ♂, WSU); Nome Area, mi 17 Kougarok Rd, N 64º42'22''W 165º17'50'', 21–23.vi.2005, J. & R. Skevington (2 ♀, CNC); Richardson Hwy, mi 206, beaver pond, 20.vi.1987, DMW (1 ♂, CNC); Richardson Hwy, mi 206, 2900', Isabel Pass, 13.vii.1962, PJS (1 ♀, CNC); Richardson Hwy, mi 213, 17.vi.1951, W. R.M. Mason & J. R. McGillis (2 ♀, CNC); Savage R, 19.vi.1954, W.C.F. (1 ♀, WSU); Sitka Nat. Mon., Baranof Is., 17.v.1954, R. Coleman (1 ♂, USNM); Taylor Hwy, Walker Fk Cpgd, 17.vi.1996, PHA (1 ♂, 2 ♀, USNM); Toklat R, 4.vii.1956, W.C.F. (1 ♂, 1 ♀, WSU); Unalakleet, 28.vi.1961, R. Madge (1 ♀, CNC). California: Alpine Co., West Fk Carson R, Snowshoe Springs Cpgd, 6600', 20.vi.1971, PHA (7 ♂, 5 ♀, USNM); Camp Angelus [Bernardino Co., N 34.145 W 116.982], 28.v.1947, ALM (1 ♀, USNM); Del Norte Co., Jedediah Smith Redwoods SP, Howland Hill Rd at Mill Ck, 16.v.1978, DDW (2 ♂, 1 ♀, CAS); El Dorado Co., Echo Lk, 31.v.2009, 2285 m, N 38º50'1.34'' W 120º2'34.79'', BJS (3 ♂, 1 ♀, CNC); Fresno Co., Dinkey Ck area, Laurel Ck, 4.vi.1987 (1 ♀, CNC); Lassen Co., Westwood, 16.v.1945, W.W. Worth (2 ♂, 1 ♀, USNM); Mono Co., Tioga Pass, 9.vii.1969, ex. Ribes montgenam, A.R. Moldenke (1 ♀, CAS); Mono Co., Twin Lakes, 8600', SW Mono Lks, PHA (1 ♀, USNM); Nevada Co., Sagehen Ck, nr Hobart Mills, 17.vi.1964, C. Slobodchikoff (1 ♀, CAS); Placer Co., Tahoe NF, Ward Ck, 4.8 km SW Tahoe City, 2073 m, 17.vii.1982, T.W. Davies (1 ♀, CAS); Plumas Co., Lassen NF, N Fk Feather R, 28.v.1977, DDW (10 ♂, 11 ♀, CAS); Siskiyou Co., El Capitan, 2.viii.1971, 6000ft, J. Kraemer (1 ♀, CAS); San Mateo Co., Purisima Ck Canyon, 4.iv.1985, PHA (1 ♂, USNM); Sierra Co., Webber Lk, 3.vii.1964, C. Slobodchikoff (1 ♀, CAS); Shasta Co., Lassen NP, 6.vii.1952, M. Cazier, W. Gertsch, R. Schrammel (1 ♀, AMNH); Shasta Co., Vista Pl, 4 mi SW Burney Co, 3000–4000 m 14.v.1951, S.D. Gaimari (1 ♀, CNC); Tulare Co., 15 km E Pinehurst, 29.v.1995, D. Burdick (3 ♂, CNC); Tuolumne Co., Yosemite NP, Tuolumne Meadows, 2630 m, 37º52'46''N 119º21'30''W, 14.vi.2003, J. & A. Skevington (1 ♀, CNC). Colorado: Boulder Co., Corona Pass, 10600', 6.vii.1961, JGC (1 ♀, CNC); Echo Lk, Mt. Evans, 10600', 13,20. vii.1961, CHM (2 ♀, CNC); Mt. Evans, 11300', 10.viii.1961, CHM (1 ♀, CNC); Mt. Evans, Timberline, 11600', 11,29. vii.1961, CHM (6 ♀, CNC); Mt. Evans, Timberline, 11700', 22.vii.1961, CHM (1 ♀, CNC); Gilpin Co., 2 mi E Rollins Pass, 3048 m, 16.viii.1973, DDW (1 ♂, 2 ♀. CAS); Gunnison Co., Schofield, 3538 m, 39º2'29''N 107º2'54''W, 7.vii.2006, T.A. Wheeler (4 ♀, LEM); Lake Co., Independence Pass, 12100', 31.vii.1961, BHP (1 ♂, 1 ♀, CNC); Loveland Pass, 9850', 28.vii.1961, JGC (1 ♀, CNC). Idaho: Latah Co., 3 mi S Helmer, Meadow Ck, 2800', 20.v.1976, ex. Lomatium orogenoides, N.E. Woodley (1 ♂, 1 ♀, USNM); Latah Co., 3 mi S Helmer, Meadow Ck, 2800', 17.v.1975, ex. Lomatium orogenoides, W.J. Turner (2 ♂, 2 ♀, CHVC); Moscow Mt., 8.vi.1921, ALM (1 ♂, USNM); Moscow Mt., 3.vi.1911 (2 ♀, USNM). Michigan: Ontonogan Co., 18.vi.1960, R. & K. Dreisbach (1 ♀, CNC). Montana: Gallatin Co., Hyalite Dam, 20 mi S Bozeman, 6800', 12.vi.1999, R. Hurley (1 ♂, MTEC). New Mexico: Torrance Co., New Canyon Cpgd, 9 mi W Manzano, MT, 7800', 25.v.1991, J.E. O’Hara (1 ♀, CNC). North Carolina: GSMNP, ATBI Plot, Andrews Bald, 35º32'20''N 83º29'39''W, 10–24.v., 24.iv.-10.v.2001, MT, I.C. Stocks et al. (2 ♂, 1 ♀, CNC). Oregon: Baker Co., Anthony Lk, SP, 8.vii.1977, R.S. & V.L. Zack (1 ♂, WSU); Clackamas Co., Mt. Hood NF, Still Ck Cpgd, 23.vi.1977, DDW, (1 ♂, 1 ♀, CAS); Crater NP, 10.vii.1968, B. V. Peterson (1 ♂, 1 ♀, CNC); Hood R Co., Mt. Hood NF, Mitchell Ck nr Sahalie Falls, 1400 m, 27.vi.1978, DDW (1 ♂, 3 ♀, CAS); Hood R Co., 20 mi SSW Mt. Hood, Mt. Hood Meadow Ski Area, east Fk Hood R nr. Umbrella Falls, 5200', 7.vii.1978, N. Herman (2 ♂, 4 ♀, AMNH); Hood R Co., Mt. Hood NF, Hood R Meadows, 1350 m, 27.vi.1978, DDW (4 ♀, CAS); Josephine Co., Bolan Lk, ex. pollen in gut, 16.vi.1988, J.A Downes (1 ♀, CNC); Lane Co., 1 mi S Hand Lk, 11.vii.1972, G. Steyskal (2 ♀, USNM); Lincoln Co., Boiler Bay, 24.v.1930, H.A. Scullen (2 ♀, USNM); Veronia, 1.iv.1938, ex. willow, J. Schuh, K. Gray (1 ♂, WSU). Utah: Garfield Co., 15 kmN Boulder, 12, 20.vi.1994, JFM (5 ♀, CNC); Mt. Timpanogos Lodge, 1925 m, MT, N 40°23'22.26'' W 111°35'8.13'', 7–9.vi.2011, I. Winkler (3 ♀, USNM). Virginia: Fairfax Co., Dead Run, 16.iv.1915, ex. Amelanchier canadensis, R.C. Shannon (1 ♀, USNM). Washington: Clallam Co., Olympic NP, Sunrise Ridge, Mt. Angeles, 1550 m, 24.vii.1978, DDW (1 ♀, CAS); Lewis Co., Big Ck Cpgrd, 4.4 km SE Ashford, 2.vii. 1974, 550 m, PHA (1 ♀, CAS); Mt. Rainier, Paradise Pk, viii.1917, viii.1921, ALM (5 ♀, USNM); Mt. Rainier, Sluiskin, 28.vii.1922, ALM (1 ♀, USNM); Mt. Rainier NP, E end Yakima Pk, creek, 1830 m, 3.viii.1977, DDW (1 ♀, CAS); Nahcotta, 24.v.1917, ALM (1 ♀, USNM); Pierce Co., Fort Lewis, 15.iv.1945, PHA (1 ♀, USNM); Pierce Co., Mt. Rainier NP, 14 km ENE Paradise, Owyhigh Tr, 46º45'44''N 121º39'26''W, 16.vi.2004, C.J. Borkent (1 ♀, LEM); Skagit Co., 32 km SE Marblemount Summit, North Cascades NP, MT, 18- 20.vi.1987, B. V. Brown (2 ♀, CNC); South Bend, 23.v.1917, ALM (1 ♀, USNM); Whatcom Co., Glacier, 25.v.1965, F. Schmid (1 ♂, CNC); Yakima Co., 5 mi SE Tieton RS nr Rimrock Ck, 23–24.vi.1974, W.J. Turner (1 ♀, WSU). Wyoming: Centennial, Snowy Ra, 10000', 9.viii.1950, R. R.D. & R.K. Schwab (1 ♀, CNC); Dunraven Pass, 26.vii.1956, ALM (1 ♀, USNM); Sierra Madre Ra, Battle L. Rd., 8500', 18.vii.1961, BHP (1 ♀, CNC); Teton Co., Togwotee Pass, 17.vii.1961, BHP (1 ♀, CNC).

Recognition. This species is distinguished by shiny spots on the anepisternum and katepisternum, biserial acrostichal setulae, elongate and apically expanded epandrium and coiled phallus.

Re-description. Wing length 2.6–4.2 mm. Male. Head subtriangular, black, with black setation; occiput greyish brown pollinose. Holoptic, with upper ommatidia enlarged. Frons represented by small triangular space just above antennae, bare, brownish pollinose. Ocellar triangle very prominent, with 2 pairs of proclinate setae; anterior ocellars moderately long, posterior ocellars minute. Postvertical and postocular setae moderately long, thin; additionally, occiput covered with numerous similar setae longer in lower part. Antenna brown; scape short, subequal to globular pedicel, both with short setae; postpedicel nearly 4.0 times longer than basal width, rather narrow basally, smoothly tapered; stylus very short, bristle-tipped, segment 9 shorter than wide, apical bristle somewhat longer than segment 9. Proboscis long, projected obliquely; labium about half as long as head is high; palpus projected parallel to labrum, nearly as long as labrum.

Thorax black, with brown to black setation; smoothly arched, prescutellar depression very slightly prominent; scutum viewed dorsally dull black brown (often rather subshining), viewed anteriorly velvety brown, with 2 indistinct paler vitae between acr and dc setae; katepisternum and anepisternum with large shining patch in middle; otherwise finely brownish grey pollinose (sometimes entirely finely pollinose; e.g. in some specimens taken from Khabarovsk Territory of Russia). Proepisternum with several brownish short bristly setae in lower and upper parts. Postpronotal lobe with 1–2 (usually 1) long and several short setae. Mesonotal setae prominent; acr short, arranged in 2 irregular, close rows (sometimes 3–4-serial anteriorly), broadly separated from dc, lacking on prescutellar depression; dc multiserial anteriorly, becoming uniserial towards scutellum, mostly subequal in length to acr, 2–3 prescutellar pairs longer; 1–2 presut spal (sometimes with additional setulae), 3–5 npl (usually with additional thinner setae of different lengths), 2–3 psut spal, 1 long pal and 5–9 (more often 6, but sometimes with additional short setae) pairs of sctl.

Legs almost entirely brownish, knees usually brownish yellow to yellow. Coxae and trochanters with numerous long bristly hairs. Fore femur with short, thin posteroventral setae longer subapically and with similar setae on dorsal and posterior faces. Mid femur with similar pattern of setation but posteroventral setae strong, spine-like on about apical half. Hind femur with moderately long, thin anteroventral and dorsal setae. Tibiae lacking prominent setae. Tarsomere 5 on all legs slightly flattened; pulvilli broad, shorter than tarsal claw.

Wing distinctly brownish infuscate, with brown veins; no bristle at wing base; stigma distinct brownish, elliptical, overlapping apex of vein R 1; anal lobe very prominent, acute. R 2+3 straight in apical part; radial fork with base proximal to apex of R 2+3 (R 4 sometimes absent); R 5 and M 1 divergent before wing-apex; cell d broad, longer than basal cells, hardly produced apically, truncate; m-m crossvein long, M branches widely separated; dm-cu crossvein slightly concave; M 4 somewhat shorter than dm-cu crossvein. Apex of cell cua slightly rounded, CuP+CuA long, ending just short of wing margin. Halter brown.

Abdomen brown, shining, covered with short, black, thin setae. Terminalia concolorous with abdomen, elongate. Hypandrium rounded, upcurved apically and extending into long, broad prolongation, with apical grooves supporting postgonites; gonocoxal apodeme broad and longer than ejaculatory apodeme. Postgonite posterior to phallus, extended well beyond hypandrium, parallel with phallus; apex sickle-shaped, roundly curved, mostly membranous. Epandrium (see Cumming et al. 1995, fig. 8) not greatly inflated laterally; dorsal bridge very narrow; elongate produced horizontally, apical half somewhat expanded, with apical margin broadly rounded; inner apical face expanded medially into ear-like surstylus; dorsal margin of inner apex with dark, knob-like projection obliquely ( Fig. 8B). Phallus long and slender, arched well beyond epandrium, with membranous subdivision at mid-length; apical half rounded, sickle-shaped arching beyond apex of epandrium; ejaculatory apodeme plate-like, rounded, slightly shorter than gonocoxal apodeme; dorsal margin of base of phallus produced posteriorly into slender lobe, strongly arched subapically forming slender trifid apex. Cercus long and slender, longer than half length of epandrium, projecting well beyond dorsal margin of epandrium; parallel-sided in lateral view with rounded apex, similar to lobes of hypoproct (see Cumming et al. 1995, fig. 8); hypoproct produced into pair of long, slender divergent lobes, half as long as length of cercus.

Female. Dichoptic, with equally small ommatidia. Frons very broad, widened toward ocellar tubercle, usually entirely shining (ocellar triangle pollinose), with short marginal setulae. Occiput with somewhat stronger postvertical and postocular setae and shorter bristly hairs in lower part. Scutum viewed dorsally subshining, finely brownish grey pollinose; presutural supra-alar and notopleural faces sometimes with shining patch. Scutellum with 3–5 pairs of setae. Legs greatly vary in colour; sometimes as in male or considerably paler, largely yellow (including coxae), or only extreme apex of femora yellow with tibiae and tarsi brownish to brownish yellow, sometimes tibiae and tarsi also yellowish basally. Fore and mid femora with short, thin posteroventral setae slightly longer subapically. Wing usually paler than in male. Abdomen brownish, viewed dorsally shining or subshining, with short, thin, brownish setae. Cercus elongate, concolorous with abdomen.

Distribution. This species is distributed across North America, extending as far south as New Mexico in the west and Smoky Mountains National Park in the east ( Fig. 10). In the Palearctic and Oriental regions, I. nitidula extends from Scandinavia to the Russian Far East and as far south as Bulgaria and Taiwan ( Fig. 9).

Remarks. In the Banff National Park region of Alberta ( Canada), I. nitidula was collected on Labrador tea ( Rhododendron groeniodicum (Oeder) Kron & Judd ), an evergreen shrub growing in moist to wet conditions. Melander (1946) reported I. nitidula was collected also on mountain ash. This species was also reported from the brown coal basin in Northwest Bohemia (Czeck Republic), considered the most anthropogenically damaged region in central Europe ( Chvála & Barták 2000).