Pteremis fenestralis ( Fallén, 1820 )

Roháček, Jindřich, 2024, The genus Pteremis (Diptera: Sphaeroceridae) in the West Palaearctic area, with description of four new species, Acta Entomologica Musei Nationalis Pragae 64 (1), pp. 153-206 : 158-168

Pteremis fenestralis ( Fallén, 1820)

( Figs 1–4 , 7–39 )

Copromyza fenestralis Fallén, 1820: 8 (both sexes). Type locality (by neotype designation): Sweden, Huddinge, Gömmaren lake res. Neotype (designated by ICZN 2014: Opinion 2341, see also RoHÁĆFK 2012a: 102): J (NHRS).

Limosina fenestralis : ZFTTFRsTFDT (1847): 2504 (generic combination).

Limosina (Stenhammaria) fenestralis : DUDA (1918): 88 –90 (subgeneric combination, redescription, illustr.); DUDA (1938): 90 –91 (redescription, illustr.).

Leptocera (Stenhammaria) fenestralis : DUDA (1925): 73 (generic combination).

Leptocera (Pteremis) fenestralis : RICHARDs (1930): 845 (subgeneric combination); PAPP (1973b): 66 (key, illustr.); OKFLY (1974): 49 (puparium, illustr.).

Pteremis fenestralis : HACKMAN (1963): 59 (generic combination); PAPP (1984): 87 (Palaearctic catalog); PITKIN (1988): 39, 103, 111, 123, 135, 149, 167 (key, genitalia, illustr.); RoHÁĆFK et al. (2001): 211 (world catalog); MARsHALL et al.(2011):265 (world catalog update); RoHÁĆFK (2012b): 540 (wing polymorphism).

Borborus nivalis Haliday, 1833: 178 (both sexes?).Type locality: Ireland, Downshire , Holywood. Lectotype (designated here): ♀ (NMID).

Borborus nivalis : CoLLIN (1956): 178 (synonymy).

Limosina nivalis : HALIDAY (1836): 330 (generic combination).

Pteremis nivalis : RoNDANI (1856): 124 (generic combination).

Limosina (Pteremis) nivalis : DUDA (1918): 90 –92 (subgeneric combination, redescription); DUDA (1938): 89 –90 (redescription, illustr.).

Leptocera (Pteremis) nivalis : DUDA (1925): 73 (generic combination).

Stenhammaria nivalis : CoLLIN (1956): 178 (generic combination).

Limosina erratica Haliday, 1836: 330 (both sexes?). Type locality: not given. Syntypes (not found in NMID, cf. CoLLIN 1914: 244, probably lost).

Limosina erratica : CoLLIN (1914): 244 (synonymy).

Limosina paradoxa Stenhammar, 1855: 399 (female). Type locality: Sweden, ‘ Ostrogothia paroecia Wärna’. Holotype: ♀ (not located in UZIU, cf. KIM 1972: 205–207, probably lost).

Limosina paradoxa : DUDA (1918): 90 (synonymy).

Pteremis subapterus : FRFY (1941): 26 (nomen nudum).

Pteremis subapterus Frey, 1947: 68 (both sexes, illustr.).Type locality (by lectotype designation): Finland, Suomussalmi. Lectotype (designated here): J (MZHF)

Pteremis subapterus : HACKMAN (1964): 75, 84 (synonymy).

Type material examined. Copromyza fenestralis Fallén : NFoTYPF (designated by ICZN 2014: Opinion 2341, see also RoHÁĆFK 2012a: 102): J ( NHRS), labelled: ‘ SWEDEN: Huddinge, Gömmaren lake res., 59°15′15″N, 17°55′40″E, 58 m, J. Roháček leg.’, ‘ 7.7.2011, peat-bog, sifting Sphagnum , moss and grass’, ‘ NEOTYPUS J, Copromyza fenestralis Fallén, 1820 , J. Roháček des. 2011’ (red label) and ‘ Pteremis fenestralis (Fallén) J, J. Roháček det. 2011’ (see Figs 8, 9 ). The specimen is intact, glued onto pinned triangular card ( Fig. 7 ).

Borborus nivalis Haliday : LFCToTYPF (designated here): ♀ ( NMID), labelled: ‘Holywood’ (green label), ‘Haliday 20. 2. ’82’ (registration number of museum), ‘Named by J. E. Collin, Limosina nivalis’ (pencil handwritten), ‘ LECTOTYPUS ♀ Borborus nivalis Haliday, 1833 , J. Roháček des. 2022’ (red label), ‘ Pteremis fenestralis (Fallén, 1920) ♀, J. Roháček det. 2022’ ( Fig. 12 ). The specimen is intact ( Fig. 10 ), mounted in Haliday’s style, glued on pinned quadrangular card ( Fig. 12 ).

Pteremis subapterus Frey : LFCToTYPF (designated here): J ( MZHF), labelled: ‘Suomussalmi’, ‘Hellén’, ‘553’, ‘Mus. Zool. H:fors, Spec. typ No. 8182, Pteremis subapterus Frey’ (faded green label, partly handwritten), ‘ Pteremis J fenestralis (Fall.) f. subapterus’, ‘J. ROHÁČEK det. 1985’ (partly handwritten), ‘Mus. Zool. Helsinki, Loan No. DIP 2010–17’ (yellow label) and ‘ LECTOTYPUS J Pteremis subapterus Frey, 1947 , J. Roháček des. 2023’ (red label) (see Fig. 15 ).The specimen is intact ( Figs 13, 14 ), minutia pinned and mounted on pinned polyporus bricket. PARALFCToTYPF: ♀, labelled as lectotype, except for sex symbols and ‘516’, ‘Spec. typ No. 8182’ and ‘ PARALECTOTYPUS ♀ ’ ( MZHF). Note. These two specimens are listed as ‘Typen Nr. 8181–8182’ in FRFY (1947: 68). However, FRFY (1947) listed a total of 1 J 4 ♀♀, i.e. also 3 additional females from localities N. Finnland: Om. Brahestad (G. Wuorentaus leg.), Ok. Kajana, 29.v.1917 (W. Hellén leg.) and Ok. Suomussalmi, 26.vi.1917, gesiebt (W. Hellén leg.). All these specimens are preserved in MZHF (J. Kahanpaa, pers. communication, 2023) but only the last one has been examined. According to article 72.4.6 of the Code ( ICZN 1999) these three females do not belong to the type series and, consequently, cannot be designated as paralectotypes.

Additional material examined. AUSTRIA: Ebensee (am Traunsee), ~ 50 km E Salzburg, grass roots nr. River, 26.viii.1986, 2JJ 1♀ f. brach., S. A Marshall leg. ( DEBU, 1 ♀ genit. prep.); Tirol: Igls, 900 m, 12. vii.1953, 1 J, J. R. Vockeroth leg. ( HNHM); Obergurgl Mt., 1950 m, 4.vii.1975, 1 J, 2.x.1975, 1 J, H. Troger leg.; Badgastein, 16.vi.1977, 2 JJ (f. brach.), H. Stockner leg. ( IZIU). BULGARIA: Geleznitza nr. Sofia, 13.vii.1974, 1 J, 24.vii.1977, 1 J, P. Snejanka leg.; Zlatni piasatzi, 23.viii.1974, 1 J; Drangovo-Petritsch, 12.iv.1973, 1 J 1 ♀; Papaz Tchair, 16.vi.1975, 1 ♀; Pirdop, 12.v.1973, 1 ♀, 1.vi.1977, 1 ♀; Rila-hut Musala, 2200 m, 2.x.1973, 1 ♀; Sandanski, 11.iv.1973, 1 ♀; Bistritza, vodopada, 24.ix.1972, 1 ♀, all V. Beschovski leg. ( IZS). CZECH REPUBLIC (more than 1000 specimens, deposited in JRO, MBP, MMBC, NMPC, SMOC, many genit. prep.), examples of localities: BOHEMIA: W Bohemia: Doupovské hory Mts-Lochotín, Jelení nr. Karlovy Vary (Škaloudová leg.), Čistá nr. Sokolov (Dlabola leg.), Bílina-Chloumek, Bílina Štěpánov, Duchcov env. (M. Barták leg.); S Bohemia: Lipno nad Vltavou, Šumava Mts-Zhůřské slatě, Šumava Mts-Horská Kvilda, Šumava Mts-Pěkná (J. Roháček leg.), Šumava Mts-Rakouská louka, Šumava-Horská Kvilda, Vlkov nad Lužnicí (M. Barták leg.), České Budě- jovice-Švábův Hrádek (J. Olejníček leg.), Hluboká (I. Kovář leg.), Filipov, Žíšov-Doubí (J. Máca leg.), Palupín nr. Strmilov, Dačice-Malý Pěčín (J. Roháček leg.); C Bohemia: Srbsko (J. Zuska leg.), Břežany, Lešany nr. Benešov (I. Kovář leg.), Ondřejov (M. Chvála leg.), Praha-Holešovice, Praha-Bohnice, Praha-Trója, Praha-Šárka, Třebotov, Úvaly nr. Praha, Srbsko-Koda, Jíloviště, Kunice,Kunice-Hůra,Velké Popovice, Struhařov, Sokoleč, Veltrusy-deer park (M. Barták leg.); N Bohemia: Holany 2.1 km NW-Dolské údolí valley, Hradčany 1.5– 2 km W-Ploučnice valley, Mimoň 6 km E-Novodvorský rybník 5, Staré Splavy 1.2 km NE nr Máchovo jezero, Doksy 4.3 km NE-Břehyňský rybník res., Obora nr. Doksy 0.6 km SE-Poselský pond, Jestřebí 1.7 km SE-Jestřebské slatiny res., Jizerské hory Mts.-Jizerka (all J. Roháček leg.), Krkonoše Mts-Vítkovice (Zuska leg.); E Bohemia: Jilemnice (Dlabola leg.), Bohdaneč, Černá u Bohdanče (B. Mocek leg.). MORAVIA: W Moravia: Třešť- Kaz, Třešť- Jelenice, Třešť- Zákotský rybník (pond), Třešť- Otov, Třešť- Louč- ky, Řásná nr. Telč- Pařezitý rybník (pond) (J. Roháček leg.); S Moravia: Havraníky-Havranické vřesoviště, Lednice, Pouzdřany-Kolby (B. Mocek leg.), Lednice-park, Pálava Mts, Nejdek nr. Lednice, Sedlec u Mikulova-Skalky, Sedlec-Slanisko nr. Nesyt pond, Vranovská přehrada-Cornštějn, Bílé Karpaty-Radějov (J. Roháček leg.), Moravský Krumlov (M. Barták leg.), Havraníky, Hnanice, Horní Břečkov, Čížovský rybník (pond), Liščí skála nr. Podmolí, Zadní Hamry, Braitava, Široká pole, Hardegg vyhlídka, Fládnická chata (M. Barták et al. leg.); C Moravia: Nedvědice-Chlébské, Moravský kras-Líšeň, Moravský kras-Mariánské údolí, Brno-Lesná, Brno-Obřany, Dolní Loučky nr. Tišnov (B. Mocek leg.), Ohrozim (M. Vála leg.), Moravičany nr. Mohelnice, Hostýnské vrchy Mts-Rajnochovice (J. Roháček leg.); N Moravia:Rychlebské hory Mts-Nýznerov, Vidnavské louky res. nr. Vidnava, Supkovice 1.1 km NE-sand-pit, Kolnovice 1 km SW-sand-pit, Bohušov nr. Osoblaha 0.7 km S-sand-pit, Hrubý Jeseník Mts-Kouty n. Desnou, Hrubý Jeseník Mts-Skřítek peat-bog, Hrubý Jeseník Mts-Keprník-Vozka peat-bog, Hrubý Jeseník Mts-Rejvíz peat-bog, Bělá pod Pradědem, Hrubý Jeseník Mts-Vleká kotlina, Nízký Jeseník Mts-Slunečná Mt., Spálené-Sokolí důl, Horní Benešov, Hanušovická vrchovina-Jeřáb Mt., Karlova Pláň- Volárenský potok, Karlova Pláň- Karlovec, Klokočov nr. Vítkov, Úvalenské louky res. nr. Krnov, Hradec nad Moravicí-Půlles, Vršovice nr. Opava, Chvalíkovice nr. Opava, Opava, Hněvošice, Bělá nr. Chuchelná, Závada 1.8 km SE-sand-pit, Moravskoslezské Beskydy Mts-Tanečnica Mt. (J. Roháček leg.), Moravskoslezské Beskydy Mts-Muřinkový vrch Mt., Třinec-Tyra (M. Barták leg.). DENMARK: Lyngbymoor, 15.ix.1912, 1 J, no collector ( ZMHB); NEJ: Frederikshavn, 15.vi.1919, 1 J, T. Mortensen leg.; LFM: Radsted, 17.vii.1964, 1 ♀, N. M. Anderssen leg. (both ZMUC). FINLAND: Helsinki, 11.iv.1952, 1 ♀, L. Tiensuu leg.; 1 J 3 ♀♀ (f. nivalis ), 1 ♀, R. Frey leg.; Helsinge, 111, 1J (genit. prep.), R. Frey leg.; Kyrkslätt, 5341, 2 ♀♀, R. Frey leg.; Muonio, 2 JJ 1 ♀ (f. subapterus ), J. Sahlberg leg.; 2 ♀♀ (f. subapterus ), Palmén leg.; Suomussalmi, 538, 1 ♀ (f. subapterus , genit. prep.), Hellén leg.; Kajana, 29.v, 1 ♀ (f. subapterus ), Hellén leg.; Esbo, Westend, Sorkhål, falla nr 4, 10.viii.1957, 1 ♀, W. Hackman leg. (all MZHF); Ab. Kaarina Järvelä, 67145:32457, pitfall, 17.v.–3.vi.2017, 1 ♀ (genit. prep.). 3.vi.–10. vii.2017, 1 ♀♀, V.-M. Mukkala leg. ( SMOC). FRANCE: Arc, 30. viii.1986, 1 J (f. brach.); 9 km S Chambery, 1000 m, grass roots, 31. viii.1986, 1 J 1 ♀, all S. A Marshall leg. ( DEBU, 1 J genit. prep.); Hte Savoie, Le Môle, mousses, 1800 m, 17.vii.1979, 1 J, Besuchet leg. ( HNHM). GERMANY: Westfalen:Herten, 18.iii.1916, 1♀, 20.iii.1916, 1 ♀; S. Harz: Ilfeld, 18.iv.1915, 1 ♀, 25.iv.1915, 1 ♀, 28.iv.1915, 1 J, all O. Duda leg. ( ZMHB); Frankfurt an Oder, 30.x.1927, 1 J; Frankfurt an Oder-F See, 10.iv.1935, 1 J; Frankfurt an Oder-Eichwald, horse excrement, 21.iii.1935, 1 ♀, all M. P. Riedel leg. ( ZMHB). GREAT BRITAIN: ENGLAND: Oxon: Bagley Wood nr, Oxford, nest of Evotomys , 9.xii.1925, 1 ♀ f. brach., O. W. Richards leg. ( ZMHB); Oxford, University Park env., sifting grass and Carex tufts, 10.viii.1998, 2 ♀♀ (1 ♀ genit.prep.), J. Roháček leg.( SMOC); Suffolk:Lakenheath warren, breck grass & heather heath, suction sampler, 13.vii.2003, 2 JJ 3 ♀♀ (1 J 3 ♀♀ f. brach.); Suffolk: Maidscross Hill, acid grassland, 13.vii.2003, 2 JJ 2 ♀♀, all P. Gatt leg. ( SMOC). GREECE: Pieria: Olympos Mts, Karyá 5 km E, 39°59′N, 22°26′E, 750 m, sweeping undergrowth of mixed forest, 3.vi.2007, 2 JJ 1 ♀ (1 J genit. prep.); Karyá 3 km E, 39°59′N, 22°25′E, 800 m, under tufts of grass in damp meadow, 3.vi.2007, 6 ♀♀; NW Peloponnese:Ano Vlasia 2.9 km S, 37°58′32″N, 21°54′10″E, 1020 m, sifting leaf litter under Platanus , 29.v.2015, 1 J (genit. prep.), all J. Roháček leg. ( SMOC); Viotia, Parnassos, Ski Centre, 1900 m, 6.vi.1982, 1 J, R. Danielsson leg.( HNHM). HUNGARY: Kiskunsági N.P., Fülöpháza, homokbuckás, 26.vii.1978, 1 J; Kiskunsági N.P., Ágasegyháza, homokbuckás, 27.vii.1978, 1 J; Hortobágy N.P., Egyek, Ohati erdö, 17.vii.1978, 1 J, all L. Papp leg. ( HNHM); Bátorliget, talajcsapda, iii.1990, 1 J, I. Loksa leg. ( HNHM); Csik Szépvir, 9.v.1917, Fodor leg. ( ZMHB). ICELAND: S. Iceland: Seljalandsfoss, Water falls, stream, 7.v.2013, 1 ex (sex unknown, abdomen damaged), P. Gatt leg. ( SMOC). ITALY: N. Italy: Bibione env., seashore wrack, 16.vi.1994, 1 ♀ (genit. prep.), J. Roháček leg. ( JRO); Volano, Strada spiaggia, aeroplancton, 5.x.1983, 17:30, 2♀♀, no collector;Volano,oRe 12, Tamarix , 12.xi.1972, 1 ♀, no collector; Arta Fornaci, prato umido, 22.iii.1964, 3 ♀♀, no collector; Vatle[s] Bur[gusio], 29.iv.1967, 1 J, no collector; Padova: Luvigliano, 22.vi.1988, 1 ♀, Scarpa leg.; Venezia: Mestre, S. Giuliano Terriccio, 20.ii.1988, 1♀; Monti Lessini: NE di Cortiolo, 897 m, stagno, 28.vi.1989, 1 J, L. Munari leg. (all MCNV); Abruzzo: ( AQ), Monte Velino, Fontecchia, 1250 m, 17.v.2006, G. Lo Giudice leg. ( SMOC); Puglia: Gargano, S Giov. Rotondo, Coppa di Mezzo, 850 m, Fagus and Quercus grazed forest, 4.vii.2005, 1J (genit. prep.), P. Gatt leg.( SMOC); Calabria: Serre Calabresi Mts, Mongiana 2.4 km N, 38°32′05″N, 16°19′06″E, 1000 m, in tufts of Juncus in alder forest, 25.v.2018, 4 JJ 4 ♀♀ (1 J 1 ♀ genit prep.); same, sweeping in alder forest and over meadow nr. brook, 25.v.2018, 1J; Mongiana 2.5 km NNE, 38°32′09″N, 16°19′33″E, 1035 m, sweeping vegetation along brook in alder forest, 22.v.2018, 1 ♀, all J. Roháček leg. ( SMOC). SARDINIA: Pattada 2.6 km SE, 40°34′12″N, 9°08′17″E, 575 m, aspirated from tufts of grass, 10.v.2014, 1 ♀ (genit. prep.), J. Roháček leg. ( SMOC). SICILY: Messina: Nebrodi, Monte Sorro, 1700 m, Fagus , Quercus , 7.vi.1999, 1 ♀ (genit. prep.),P.Gatt leg.( SMOC); Parco di Nebrodi,Lago Pisciotto,37°5848″N, 14°50′49″E, 1240 m, sweeping swampy shores of lake, 21.iv.2016, 2JJ (1 J genit. prep.), J. Roháček leg. ( SMOC). POLAND: W. Poland: Gleiwitz, 21.iii.1934, 1 ♀; Wustung bei Habelschwerdt, 6.iv.1925 – 11. iv.1929, ca 40 specimens, all O. Duda leg. ( ZMHB); E. Poland: Bondary, Narew river shore, under tufts of grass, 26.vi.2005, 1 J 2 ♀♀ (1 ♀ genit. prep.); Białowieża-Budy 2 km NE, under tufts of grass in damp meadow, 24.vi.2005, 1 ♀; Białowieża-Budy 4 km S, Czerlon, on carrion of Bison bonasus , 25.vi.2005, 2 ♀♀, all J. Roháček leg. ( SMOC). PORTUGAL: Algarve: Tavira 3 km N, 37°08′52″N, 7°39′12″W, 24 m, sweeping over boggy meadow, 1.iv.2009, 1 J (genit. prep.), J. Roháček leg. ( SMOC); Manteigas, Serra de Estrela, 40°19′23″N, 7°36′09″W, 1860 m, grazed bog, steppe, stream, 29.ix.2014, 1J 1♀; Seia,Aldeia da Serra, 40°25′03″N, 7°40′38″W, 893 m, grasses & scrub, 28.ix.2014, 1♀, P. Gatt leg.( SMOC). ROMANIA: Banat: Sfânta Elena 2.5 km NE, 44°41′44″N, 21°43′10″E, 420 m, sweeping over meadow, 1.vi.2008, 1♀ (genit.prep.), J. Roháček leg. ( SMOC). SLOVAKIA (about 400 specimens, deposited in JRO, MBP, SMOC, many genit. prep.), examples of localities: S Slovakia: Patince nr. Komárno, Búč nr. Komárno, Kamenín nr. Štúrovo, Nová Vieska nr. Štúrovo (J. Roháček leg.), Hegyfarok, Štúrovo, Bieľ, Pribylina (M. Barták leg.); C Slovakia: Kremnické pohorie Mts-Turček, Detva-Horná Chrapková, Hrochoťská dolina-Hrochoť 3 km E,, Kyslinky-Dolná Zálomská, Kyslinky- Hrochoťská dolina, Kyslinky-Majerová, Predná Poľana Mt.-Žliebky, Sihla env., Sihla-Kamenistý potok, Zákľuky Mt., Stankovany nr. Kraľovany, Chočské vrchy Mts-Škútova dolina, Muránska Huta-Šiance res., Muránska planina NP-Šiance res.top plateau, Pohronská Polhora 5.3 km SE-Rosiarka res., Tisovec 3.9 km NW-Trstie res., Jestice-Hradisko Mt. (all J. Roháček leg.); N Slovakia: Tatranská Lomnica (M. Barták leg.), Vysoké Tatry Mts-Batizovská dolina, Vysoké Tatry Mts-Velické pleso, Vysoké Tatry Mts-Štrbské pleso, Belianske Tatry Mts-Tatranská Kotlina, Tatranská Kotlina-Šarpanec res., Regetovka res. peat-bog (J. Roháček leg.); E Slovakia: Levočské pohorie Mts- -Branisko Mt., Boľ nr. Král. Chlmec, Boťany nr. Latorica, Stakčín, Vihorlat Mts-Sninský kameň Mt., Hostovice-Hostovické lúky res., Nová Sedlica-Stužica res. (J. Roháček leg.). SPAIN: S. Spain: Sierra Nevada Mts, Llano Prado, 2 200 m, on decayed grass, 14.v.1979, 2 JJ 3 ♀♀ (1 J 1 ♀ genit. prep.), in runs of Pitymys and Arvicola sp. , 14.v.1979, 1 J 1 ♀; Grazalema nr. Ronda, 1000 m, 16.–17.v.1979, sifting moss, 1 J (genit. prep.), sweeping over meadow, 1 ♀, sweeping by stream, 1 ♀, all J. Roháček leg. ( JRO); Sierra Nevada, Hwy, 1860 m, edge of small, cold stream, 7.viii.1986, 1 J 1 ♀; Sierra Nevada, Hwy, 2100 m, along small stream, 9.viii.1986, 1 J 3 ♀♀ (1♀ genit. prep.); Sierra Nevada, Hwy, 2 270 m, grass of dry stream, 11.viii.1986, 2 JJ 2 ♀♀ (1 J 1 ♀ genit. prep.), J. R. Vockeroth leg. ( DEBU). SWEDEN: Huddinge, Gömmaren lake res., 59°15′15″N, 17°55′40″E, 58 m, peat-bog, sifting Sphagnum , moss and grass, 7.vii.2011, 3 JJ 1 ♀ (2 JJ 1 ♀ genit. prep.), all collected together with the male neotype, J. Roháček leg. ( SMOC). SWITZERLAND: Delémont, 500 m, conifer slash (fresh), 28.viii.1986, 1 J 3 ♀♀ (1 ♀ genit.prep.), S.A. Marshall leg. ( DEBU); ZH: Unterengstringen, 15.ix.1997, 1 J, G. Bächli leg. ( HNHM).

Redescription. Male ( Figs 2, 3 , 7 ). Total body length 1.38–1.98 mm; general colour brown to dark brown with greyish brown to grey microtomentum, subshining dorsally on thorax and abdomen), duller on head, thoracic pleuron and ventral abdominal sterna.

Head ( Figs 4 , 11, 14 ) about 1.4× higher than long, blackish posterodorsally, orange to yellow anteroventrally. Frons dark brown to blackish except for orange-brown anterior margin, distinctly microtomentose; occiput almost black, with dark grey microtomentum. Orbits, interfrontalia (very narrow) and frontal (including ocellar) triangle with silvery grey glittering microtomentum and all separated by dull blackish (often with some bluish tinge) stripes almost forming M-shaped mark; frontal triangle distinctly delimited and reaching to anterior third of frons; anterior margin of frons orange to orange-brown, wider laterally. Frontal lunule relatively long (although shorter than wide), yellow, whitish microtomentose, distinctly lighter than anterior margin of frons. Face normally yellow (darkened in some specimens, for detail see Variability below), whitish microtomentose including facial cavities below antennae; medial carina slightly developed, distinct only dorsally, below frontal lunule. Gena yellow to ochreous, paler anteriorly, all greyish microtomentose and sharply separated from dark occiput by dark and shining perpendicular stripe on postgena. Cephalic chaetotaxy ( Fig. 4 ): pvt small and weak but strongly convergent or crossed; a pair of minute convergent postocellar setulae between pvt and posterior ocelli also present; occe and occi subequal (or occi slightly longer) and about two-thirds of vti; vti and vte subequal, robust and longest of frontal bristles; oc distinctly shorter than vti; 2 ors, posterior as long as oc and about 1.4 times as long as anterior ors; 4 (more rarely only 3) ifr, two middle pairs usually longer and more robust, sometimes with 1 microseta in front of anterior ifr in addition; 2 or 3 very minute ads inside and below ors; g and 1 or 2 setae behind it weak, hardly longer than anterior peristomal setula; vi very long and robust, about as long as vti; peristomal setulae sparse (only 4 or 5) and about as long as 2 weak postgenal setae; postocular setulae shorter than peristomals, in single long row. Eye subcircular (19:17), of moderate size, with longest diameter about 4 times as long as smallest genal height. Antenna with scape ochreous to yellow, pedicel blackish and 1 st flagellomere dark brown, the latter relatively short (as long as pedicel), suboval to ovoid, with whitish ciliation on apex distinctly longer than cilia on arista. Arista relatively long, about 4.2 times as long as antenna, with short dense ciliation.

Thorax dark brown to brown (ventral part of pleuron palest) and dark grey microtomentose; mesonotum subshining, pleuron and scutellum duller. Sutures between pleural sclerites pale brown to ochreous. Scutellum large and relatively long, flat on disc, little transversely (3:2), rounded subtriangular. Thoracic chaetotaxy: mesonotal macrosetae relatively long and robust; 1 hu (as long as anterior npl) and 1 microseta on humeral callus; 2 relatively short npl (posterior shorter); 1 prs (= posthumeral) and 1 sa, both short; 2 pa, the outer very long, longer than dc, the inner small; only 1 robust (but shorter than laterobasal sc) dc in prescutellar position; dc microsetae (in front of dc) somewhat longer than ac microsetae); 8 rows of ac microsetae on suture but only 4 in front of scutellum; medial prescutellar ac pair small, hardly or only slightly longer than other ac microsetae; 2 very long and robust sc, laterobasal longer than scutellum, apical (longest thoracic seta) 1.5‒1.6 times as long as laterobasal; 2 stpl but only posterior long (as long as laterobasal sc but finer), anterior reduced to microseta.

Legs brown or dark brown (femora, coxae, t 3) to ochreous or dirty yellow (trochanters, tibiae, tarsi) but colouration is variable, generally darker in montane and boreal specimens and more yellow in brachypterous forms. f 1 with base and apex usually yellow; also f 2 and f 3 with apex (knee) often ochreous, and t 3 with both ends ochreous to yellow. On the contrary, the largely yellow t 1 and t 2 can be darkened about middle as is also hind tarsus or its basal segments. Pedal chaetotaxies: f 1 with 2 rows of setae: a short posterodorsal row with 3 or 4 setae in the middle of femur, and a longer row of posteroventral setae, 3 of which in distal half of femur are enlarged. f 2 anteriorly with 1 robust subapical seta and 1 short in front of it, and posteriorly with 1 shorter posteroapical seta. t 2 ventrally (see Fig. 23 ) with 3 setae, viz. 1 long robust va (directed in axis of tibia), 1 similarly long and strong vpa and 1 short av near middle; dorsally ( Fig. 22 ) there are two groups of setae: proximal group, with 1 longer anterodorsal seta + 1 shorter above it and 1 shorter posterodorsal seta in about proximal fourth; distal group with 1 shorter anterodorsal and 1 very long posterodorsal (rather subdorsal) seta in about distal fourth, each surmounted with a short seta (that subdorsal can be reduced); in addition there are 2 small posterior setae in proximal third and above distal fourth ( Fig. 22 ); apex of tibia provided with 3 short (1 or 2 longer) subapical setae anteriorly ( Fig. 23 ) and 2 posteriorly ( Fig. 24 ). f 3 with 1 short and weak anterior subapical seta. Other parts of femora and tibiae uniformly finely setulose. Mid basitarsus (mt 2) sometimes with 1 enlarged ventral (or anteroventral) setula (similarly as in Fig. 57 ). Ratio t

2

: mt

2

= 1.81–1.94.

Wing normal (macropterous form, Fig. 1 ) or reduced (brachypterous form, Figs 2, 3 ). Membrane distinctly brownish fumose, veins brown to dark brown. Macropterous form: wing ( Fig. 16 ) relatively broad; C distinctly produced beyond apex of R 4+5. Cs 1 with markedly longer setulae than rest of C. R 2+3 very slightly sinuate but apically distinctly upcurved to C; R 4+5 distinctly sinuate, distally upcurved to C and diverging from and ending about as far from apex of wing as venal fold of M. Discal cell (dm) of moderate length, relatively broad but distally tapered, with small process of M and longer one of CuA 1 beyond dm-cu; that of M continued by venal fold; both outer corners of cell dm normally obtuse-angled. A 1 relatively long but only small basal part fully developed, more distally formed by brownish coloured fold not reaching wing margin. Anal lobe large, well developed; alula relatively small and narrow. Wing measurements: length 1.19–1.71 mm, width 0.50–0.78 mm, C-index = 0.80–0.96, r-m\dm-cu: dm-cu = 1.64–2.09. Brachypterous form: wing size and venation variously reduced ( Figs 17–21 ). C shortly produced beyond apex of R 4+5. R 4+5 rather curved (slightly to distinctly) than sinuate. Discal cell (dm) very shortened beyond r-m, usually with cross-vein dm-cu absent (f. nivalis , Figs 18, 20 ), more rarely with it present and posterior outer corner of dm cell rounded ( Fig. 17 ); also R 2+3 can sometimes be more or less shortened (f. subapterus , Figs 19, 21 ) or apex of wing reduced and rounded ( Fig. 20 ). A 1 always with coloured distal venal fold shortened or discoloured and hardly visible. Wing measurements: length 0.75–1.03 mm, width 0.37–0.49 mm, C-index = 0.80–0.95. Haltere bicolorous, with ochreous-yellow stem and dark brown knob.

Abdomen blackish brown, dorsally (particularly medially) less microtomentose and more shining than ventrally. Basal preabdominal terga relatively broad (as wide as thorax) and sparsely dark grey microtomentose; T2–T5 sparsely and relatively shortly setose, with only setae in posterior corners long and robust (cf. Fig. 25 ). T1+2 largest tergum (about 1.4 times as long as T4 or T5), narrower anteriorly, widest at posterior margin; with pale brown base (on original T1) and the rest blackish brown. T3 widest tergum but clearly shorter than T4 (thus most transverse), the latter slightly narrowed posteriorly and about as long as T5; T5 markedly narrower than T4 and posteriorly more distinctly tapered. Preabdominal sterna: S1+2 (or S2) smallest, transversely trapezoidal (narrowed anteriorly), pale brown to pale ochreous (lightest anteriorly), contrasting with dark brown S3–S5 and sparsely shortly setose; S3 and, particularly, S4 large, broad, blackish brown and strongly sclerotized; S3 only slightly narrower anteriorly, shorter than S4, the latter largest sternum, transversely suboblong and distinctly longer than (equally broad) S5. S2–S4 with sparse, short setae, finer than those on adjacent terga; also setae in their posterior corners short and weak. S5 ( Figs 26, 27 ) shorter and, hence more transverse than S4, very slightly asymmetrical, with relatively large, (usually) pale-pigmented and finely micropubescent semicircular area in front of simple posteromedial comb of spines (this extended over about medial third of posterior margin of S5); lateral setae of S5 more robust and more or less separated from two groups of finer setae surrounding lateral margins of posteromedial semicircular area. S6+7 strongly asymmetrical, situated left ventro- to dorsolaterally ( Figs 25, 27 ) and dorsally fused with S8; the latter less asymmetrical and situated dorsally. S6+7 with transverse dark-pigmented ledge and S7 with irregular posteroventral projection bent inside postabdomen; right laterally, near right end of S6, there is an enlarged, annular, 6 th right spiracle ( Fig. 27 , rs). S6+7 with 2 pairs of setae ( Fig. 25 ) and S8 with only a pair of small dorsal setae near middle ( Figs 25, 27 ).

Genitalia. Epandrium ( Figs 28, 29 ) of medium length and width, almost symmetrical in caudal view ( Fig. 28 ), with rather uniform setosity, longest setae posteroventrally near margin of anal fissure. Anal fissure relatively high and narrow, elongately suboval. Cerci large but completely fused with epandrium and ventromedially separated by acute incision; each cercus ventrally terminated by a small subtriangular tubercle ( Fig. 28 ) and with 1 long sinuate seta and 2 small setae. Medandrium low (short) and broad, with long lateral arms, each connected with posterior part of gonostylus ( Fig. 28 ). Hypandrium genus-characteristic, short, laterally fused with epandrium, posteroventrally projecting on each side to connect with postgonites and medially with very short, strongly asymmetrical (right directed) flattened apodeme ( Fig. 31 ). Gonostylus ( Figs 28–30 ) of complex structure, composed of larger anterior (more lateral) lobe and of smaller (shorter and more medial) posterior lobe. Anterior lobe ( Fig. 30 ) with short pointed anterodorsal process, 3 robust curved setae anteroventrally, 1 robust darkened ventral, posteriorly directed projection (finely spinose subapically but with simple apex) and posterolaterally with oblique longitudinal spinulose ledge; a single short external seta on small process in front of proximal end of this ledge is also characteristic. Posterior lobe (dotted in Fig. 30 ) terminated by very robust spine and 2 subapical setae and posteriorly with subconical process having a robust, medially bent seta on apex (see Fig. 28 ). Aedeagal complex ( Fig. 32 ) of relatively simple construction. Phallapodeme rod-like, about as long as aedeagus, with moderate, pale-pigmented, dorsal keel. Aedeagus proximally with short but compact, laterally somewhat flattened, phallophore ( Fig. 32 ) having simply pointed but not acute (ventral) apex. Distiphallus relatively weakly sclerotized and pale-pigmented, basally narrow, widened laterally (but hardly so dorsoventrally) towards apex, proximoventrally and dorsally more sclerotized, and with a pair of slender band-like lateral sclerites, each with thinly pointed apex. Middle part of dorsal side of distiphallus finely haired and its dorsal sclerite subapically forked and bent laterally on side of distiphallus ( Fig. 32 ). Postgonite relatively large (as long as distiphallus), sinuate in lateral view, broader proximally, strongly tapered distally but with apex knob-like dilated, with 2–3 microsetae anteriorly in distal third, some minute sensilla on apex, and with minute sclerite (= remnant of pregonite) with 1 or 2 setulae, inserted in anterodorsal emargination of postgonite ( Fig. 32 ). Ejacapodeme small, having short distal digitiform process on a wider proximal part (see Fig. 32 ) attached to ejaculatory duct.

Female ( Fig. 1 ). Head, thorax, legs and wings very similar to those of male unless mentioned otherwise below. Total body length 1.43–2.10 mm. t 2 often with anterior subapical setae (or one of them) longer; mt 2 slightly longer on the average and with more frequent presence of 1 enlarged ventral setula in proximal third. Ratio t 2: mt 2 = 1.78–1.94. Wing measurements: macropterous form: length 1.38–1.79 mm, width 0.64–0.81 mm, C-index = 0.78–1.01, r-m\dm-cu: dm-cu = 1.67–2.00; brachypterous form: length 0.59–0.99 mm, width 0.29–0.50 mm, C-index = 0.84–1.08.Abdomen more ovoid in dorsal view, basally broadest. T1+2 largest and widest tergum; other preabdominal terga (T3–T5) distinctly shorter and more transverse than in male, becoming narrower posteriorly, subequal in length or T5 slightly longer. Preabdominal sterna: S1+2 formed and pale as in male but with only a few short setae; S3–S5 dark brown as in male but S3 shorter and also narrower than S4 (largest sternum, slightly trapezoidal, wider posteriorly) and S5 almost as long as S4 but distinctly narrower (as wide as S3), unmodified, transversely suboblong, with rounded corners.

Postabdomen ( Figs 34–36 ) relatively long when exposed, telescopic and retractible into preabdomen, basally (base of 6 th abdominal segment) 0.7–0.8× as wide as 5 th segment posteriorly, with subsequent segments becoming narrower (cf. Figs 35, 36 ). T6 transversely subtrapezoidal to suboblong, much narrower than T5 but wider and longer than S6, with posterior margin pale and sparse setae in posterior twofifths, those in posterior corners long and robust ( Fig. 35 ); T7 markedly shorter and narrower than T6 and bent farther laterally (see Fig. 34 ), with single row of sparse posterior setae on each side. T8 dorsally with tripartite pigmentation composed of a paler brown tongue-shaped medial part and larger darker lateral parts ( Fig. 35 ), micropubescent in only posterior half, with a few setae (one longer) laterally ( Figs 34, 35 ). T10 about as long as wide, roughly pentagonal, micropubescent only posteromedially and with a pair of relatively short setae in the middle ( Fig. 35 ). S6 transversely trapezoidal ( Fig. 36 ), narrower, and shorter but with more setae than T6. S7 simple, transversely suboblong, much narrower than S6 and also T7, with more setae than on T7, all in posterior half. S8 ( Figs 36, 38 ) relatively narrow, slightly wider than long, rounded pentagonal to subtriangular (largest extension view, see Fig. 38 ), rather convex, with 2 or only 1 lateral setae in posterior half, and posteromedially with a group of (usually) 4 short setae on elevated sockets, which can also be situated in membranous part of sclerite; disc of sclerite micropubescent in posterior two-thirds ( Fig. 38 ). S10 (in largest extension) only slightly shorter and as wide as S8, transversely subpentagonal, but anteriorly broadly emarginate, largely micropubescent (only anteriorly projecting corners and anterior margin bare) and relatively long setose in front of posterior margin ( Fig. 39 ). Spectacles-shaped sclerite (= sclerotization of female genital chamber) poorly developed as a pair of ovoid rings, almost invisible because unpigmented and membranous. Spermathecae 2+1 ( Figs 33, 37 ) blackish, each ovoid, with somewhat wrinkled surface and a group of spine-like tubercles on tapered base; terminal parts of ducts long, slender, well-sclerotized and dark-pigmented, those of paired spermathecae connected far from their bodies ( Fig. 37 ). Cerci ( Figs 34, 35 ) relatively broad but dorsoventrally flattened (see Fig. 34 ), micropubescent, each with 2 long sinuate setae (dorsopreapical and apical) and several short fine setae.

Variability. Body of brachypterous form is on average distinctly smaller, particularly so in specimens with wings most reduced (form subapterus ). In smallest specimens also ifr setae are weaker and in only 3 pairs. Male S5 is also somewhat variable depending on body size of specimens. Smallest flies (usually brachypterous form) have posterior comb with spines less numerous and, consequently, the comb is narrower than in large specimens. Female S8 sometimes (in largest specimens) with an additional pair of setae near middle. Two pairs of short posteromedial setae can often be situated in membrane just behind posterior margin of pigmented part of S8.

Comments. Pteremis fenestralis ( Fallén, 1820) is a wing polymorphic (for detail see RoHÁĆFK 2012b), externally seemingly variable species. Its brachypterous form, having besides the shortened wing also a narrowed thorax (caused by reduction of wing musculature) and, hence, relatively large head and generally smaller body, has long been considered a separate species, even placed in a different (sub)genus than was the fully winged form. Due to its striking appearance and variable wing venation (cf. Figs 17–21 ), the brachypterous form was described three times, as Borborus nivalis Haliday, 1833 , Limosina paradoxa Stenhammar, 1855 and Pteremis subapterus Frey, 1947 . However, the chaetotaxy and structures of the male S5, male terminalia and female postabdomen of the brachypterous form are practically identical to those of the more common macropterous form regardless of some size variability of setae. Although this fact was first recognized already by CoLLIN (1956) and HACKMAN (1964) terminalia of numerous specimens of both forms (including topotypic specimens) have been checked during this revision to confirm their conspecificity. Moreover, the barcoding region of COI of both forms from the Czech Republic have been compared with no difference recognized (see ID record DNMPC020-17 = f. brach. and DNMPC023-17 = f. macropt. in the BOLD database).

Pteremis fenestralis is readily distinguished from all similar relatives by the shape and armature of the male cercus (with only a small apical tubercle, Fig. 28 ), the gonostylus (with an apically simple ventral process on the anterior lobe and a robust terminal spine on the posterior lobe), the distiphallus (not dorsoventrally dilated in the middle) and the postgonite (with a knob-like apex, Fig. 32 ). In the female sex, it distinctly differs from all relatives by the spermathecae (ovoid rather than pyriform, with minute tubercle-like spines basally and sclerotized ducts connected far from bodies of paired spermathecae, Fig. 37 ) and S8 (of characteristic shape and posteromedially having only two pairs of small setae, Fig. 38 ). The armature and setosity of male S5 ( Fig. 26 ) and the somewhat dorsoventrally flattened female cerci (see Figs 34–36 ) are also different from those of its congeners. For other diagnostic characters of the species see the key below.

The relationships of P. fenestralis have not been unambiguously recognized. Judging from the chaetotaxy of the mid tibia (with both va and vpa setae developed), similar short anterodorsal of anterior lobe of distiphallus and chaetotaxy of male S 5 P. fenestralis seems to be more closely allied to P. pulliceps sp. nov. and (perhaps) also to P. tenebricus sp. nov. than to other species treated here. However, the presence of both va and vpa setae on t 2 seems to be a ground-plan character of the whole genus, not a synapomorphy of the above three species, and also the other two features can be plesiomorphic within the genus. On the other hand, the pyriform shape and ringed surface of spermathecae seems to be a synapomorphy of P. pulliceps sp. nov., P. canaria , P. ferreus sp. nov. and possibly also P. apterina sp. nov. If so, this assemablage can be a sister group to P. fenestralis + P. wirthi (see below). The female is unknown in P. tenebricus , and, consequently, its position in this group is uncertain.

The Nearctic, also wing-polymorphic, P. wirthi ( Marshall, 1984) obviously is currently the closest relative or even sister-species of P. fenestralis . Examination of several specimens (1 J and 1 ♀ dissected) from Canada (Ontario), identified by S. A. Marshall, resulted in finding that both species share several features, some probably synapomorphic, e.g. the very similar male S5, almost identical ventral process of anterior lobe of gonostylus, relatively short (not ventrally prolonged and flattened) male cercus with an apical tubercle (although being wider in P. wirthi ), similarly armed spermathecae (basally with small surface tubercles) and broad, dorsoventrally flattened female cerci. For differences between P. fenestralis and P. wirthi see comments under the latter species below.

Biology. The species is mainly associated with wet grassy and mossy sites, both in open (meadows, fens, peat-bogs, see DAHL 1909, RICHARDs 1930, DUDA 1938, BÄHRMANN 1984, RoHÁĆFK 1984, FLoRḖN 1989, NoWAKoWsKI 1989) but also in woodland (RoHÁĆFK 2012b) habitats. Adults live terricolously near the ground among tufts of graminoid plants or in moss, Sphagnum , leaf litter, and often also in runs and nests of various small mammals ( RICHARDs 1930; HACKMAN 1963, 1964, 1967; IsMAY 1978; RoHÁĆFK 1984, 2019a) and can be easily collected by means of soil traps (RoHÁĆFK 1980, 1984; PFRssoN 1983; BÄHRMANN 1987). Adults are attracted to various rotting matter, including wet excrement, particularly in runs of small mammals and the microsaprophagous larvae surely develop in these substrates. Pteremis fenestralis occurs regularly in various types of peat-bogs, often in Sphagnum , and, therefore, Ro- HÁĆFK (1984) and RoHÁĆFK & BARTÁK (1999) classified it as a tyrphophilous species. OKFLY (1974) reared both macropterous and brachypterous specimens from puparia obtained from bait (boiled grass cuttings) exposed in burrow tunnels of the rabbit.

RoHÁĆFK (2012b) found that wing polymorphic populations are only known from North and Central Europe (here usually at higher altitudes) while in southern areas only macropterous specimens occur. Thus the presence of brachypterous forms in populations seems to be geographically dependent, inasmuch as the percentage of short- -winged specimens increases in more northern latitudes and at higher altitudes (RoHÁĆFK 1975). Macropterous specimens, particularly gravid females, serve as the main agent for dispersal of the species because they are (more or exclusively) represented in samples obtained by sweeping, Malaise and light traps ( PITKIN 1986) or car-netting (MUNARI & SCARPA 1989). Adults were recorded in all months of the year (in winter in mainly subterranean microhabitats), most commonly in spring (combined published data and material examined).

Distribution. Widespread in Europe: Andorra, Austria, Belgium, Bulgaria, Czech Republic, Denmark, Estonia, Finland, France, Germany, Great Britain, Hungary, Ireland, Italy (incl. Sardinia, new, and Sicily), Latvia, Netherlands, Norway, Poland, Portugal, Russia (Central European territory), Slovakia, Spain, Sweden, Switzerland (records summarized by RoHÁĆFK et al. 2001, MARsHALL et al. 2011). Additional country records are as follows: Belarus (several records in BOLD System, e.g. https://www. boldsystems.org/index.php/Public_RecordView?processi- d=GMBMA074-17), Greece (new), Iceland (new), and Romania (new).

Note. The records of P. fenestralis from Madeira (RoHÁ- ĆFK 2007, MARsHALL et al. 2011) are erroneous because they are based on misidentification of P. pulliceps sp. nov. (described below). Also the record from the Far East of Russia, listed by RoHÁĆFK et al. (2001) and MARsHALL et al. (2011), is probably a mistake.

Figures

Figs 1–6. Pteremis fenestralis (Fallén, 1820) and its habitats. 1 – female, f. macropt., laterally (Czech Republic); 2 – male, f. brach., laterally (Czech Republic); 3 – male, f. brach., living specimen on grass stem (Czech Republic); 4 – male (topotypic specimen, Sweden), head dorsofrontally; 5 – peat-bog meadow, habitat in the (neo)type locality (Sweden: Huddinge, Gömmaren lake reserve); 6 – montane meadow with tufts of grasses, habitat near Jizerka in the Jizerské hory Mts (Czech Republic). Scales = 0.5 mm. Photo by M. Deml (1, 2) and J. Roháček (others).

Figs 7–15. Pteremis fenestralis (Fallén, 1820), type specimens. 7 – neotype male of Copromyza fenestralis Fallén, laterally; 8 – labels of the neotype; 9 – neotype mounted and pinned, with all labels; 10 – lectotype female of Borborus nivalis Haliday, 1833, dorsally; 11 – ditto, head dorsofrontally; 12 – lectotype mounted on card and all its labels; 13 – lectotype male of Pteremis subapterus Frey, 1947, laterally; 14 – ditto, head laterofrontally; 15 – all labels of this lectotype. Photo by A. O’Hanlon (10–12) and J. Roháček (others).

Figs 16–21. Pteremis fenestralis (Fallén, 1820), wings. 16 – f. macropt., male; 17–21 – f. brach.; 17 – female with dm-cu preserved; 18 – female (f. nivalis); 19 – male (f. subapterus); 20 – female (f. nivalis); 21 – male (f. subapterus). Figs 16–20 are based on specimens from Rejvíz, peat-bog meadow (Czech Republic); Fig. 21 on specimen from Muonio (Finland). Scale = 0.5 mm. Photo by J. Roháček.

Figs 22–27. Pteremis fenestralis (Fallén, 1820), topotypic male (Sweden: Huddinge). 22 – left t2 and mt2 dorsally; 23 – ditto, anteriorly; 24 – apex of left t posteriorly; 25 – male 5th abdominal segment and postabdomen, left laterally; 26 – male S5, ventrally; 27 – 5th abdominal segment and postabdomen 2 (genitalia omitted), ventrally. Scales = 0.2 mm (Figs 22–24), 0.1 mm (others). For abbreviations see Material and methods.

Figs 28–32. Pteremis fenestralis (Fallén, 1820), topotypic male (Sweden: Huddinge). 28 – external genitalia, caudally (aedeagal complex omitted); 29 – ditto, left laterally; 30 – gonostylus, laterally; 31 – epandrium and hypandrium (setosity omitted), anteriorly; 32 – aedeagal complex, laterally. Scales = 0.05 mm (Fig. 30), 0.1 mm (others). For abbreviations see Material and methods.

Figs 33–39.Pteremis fenestralis (Fallén, 1820), female.33 – spermatheca; 34 – postabdomen,laterally; 35 – ditto, dorsally; 36 – ditto, ventrally; 37 – paired spermathecae; 38 – S8, ventrally; 39 – S10, ventrally. Scales = 0.05 mm (Figs 33, 37–39), 0.1 mm (others). For abbreviations see Material and methods.

Figs 55–61. Pteremis pulliceps sp. nov., male, female, paratype. 55 – male left t2 and mt2 dorsally; 56 – ditto, posteriorly; 57 – ditto, anteriorly; 58 – spectacles-shaped sclerite, ventrally; 59 – S8, ventrally; 60 – S10, ventrally; 61 – spermathecae. Scales = 0.2 mm (Figs 55–57), 0.05 mm (others).