The examined material cannot be unequivocally attached to barcode clusters due to lack of barcode sequences from several places, or partially sympatric clusters, and is thus listed in a standardized format in the alphabetic order of countries.

Syntypes [?], 2 ♂, Ypsolophus lanceolellus, [Italy] “Livorno Mn” “Gen. Präp. Mus. Vind. 16.653 ♂” (NHMW). Lectotype ♂, here designated, [Spain] Megacraspedus subdolellus, “Lecto-type” “20/8” “Granada m.” “Origin.” “Lectotype ♂ Megacraspedus subdolellus Stdgr teste K. Sattler, 1986” “ex coll. Staudinger” “GU 15/1404 ♂ P.Huemer” (ZMHU). Paralectotypes, 5 ♂, Megacraspedus subdolellus, all labelled “Para-lecto-type” “Origin.” “ex coll. Staudinger” (ZMHU). Holotype ♂, Megacraspedus tutti, [France] “Holo-type” “La Grave 5000 ft Dauphiné FRANCE VIII.1896 Tutt. 8157” “Type” “Walsingham Collection, 1910–427.” “Megacraspedus tutti ♂ Wlsm. Ent. Rec. IX.140. (1897) TYPE ♂ (BMNH). Paralectotypes, 2 ♂, Megacraspedus grossisquamпellus, [Spain] “TYPE” “grossisquamellus” “S. Ildef. 24.6.02” “25.6.02” “GENITALIA ♂ P. VIETTE PRÉP. NO. 3568” (MNHN) [photographs examined]; 2 ♂ (on same polyporus), “S.[an] Ildef.[onso] 7.02” “GU 15/1451 ♂ P.Huemer” (MNHN). Non-type material. Andorra. 1 ♂, Sant Julia, Fontaneda, 1300 m, 6.viii.2003, leg Viehmann (RCWS); 3 ♂, 1 ♀, Port de Cabús, 2290 m, 16.vii.2012, leg. P. Huemer, genitalia slides GEL 1192 ♂ Huemer, GEL 1257 ♀ Huemer (TLMF). Croatia. 1 ♀, Krk, Misučaynica, 5.viii.1967, leg. G. Baldizzone (ZMUC); 1 ♀, Krk, str. Krk-Vrbnik, 20.vii.1998, leg. G. Baldizzone (TLMF); 1 ♂, Krk isl., dint. di Poljica, 10.vii.2012; 4 ♂, Krk isl., Mt. Hiam, Branušine, 20–22.vi.2013, 180 m, leg. G. Baldizzone (RCGB; ZMUC); 2 ♂, Gorne Bilišane, 6.vii.2004, leg. Z. Tokár; 1 ♂, Bilišane, 300 m, 23.vi.2006, leg. Z. Tokár (all RCZT); 2 ♂, Pag, 10.vi.2015, leg. J. Junnilainen (RCJJ). Italy. 1 ♂, prov. Alessandria, Parco NR Cappane di Marcarolo, sent. Lago Badana, 800 m, 27.vi.2002, leg. G. Baldizzone; 1 ♂, same data, but Monto Peggio, 900 m, 12.vii.2002 (all RCGB); 4 ♂, prov. Pordenone, Vivaro, Magredi di Vivaro, 110 m, 6.vi.2008, leg. P. Huemer, genitalia slide GEL 1220 Huemer; 1 ♂, same data, but 7 km SSW Vivaro, 5.vi.2005; 1 ♂, prov. Pordenone, San Quirino, Magredi di San Quirino, 100 m, 11.vi.2001, leg. P. Huemer (all TLMF); 3 ♂, Livorno, 1872, leg. J. Mann (NHMW); 1 ♂, prov. Livorno, Antignana, 10 m, 26.vi.2004, leg. J. Liška (NMPC); 1 ♂, prov. Roma, Bracciano, 20–22.vi.2001, leg. A. Cox (ZMUC); 3 ♂, prov. Roma, Monti di Tolfa, dint. Manziana, 23–28.vi.1989, leg. G. Baldizzone (TLMF, RCGB); 1 ♂, prov. Latina, Aurunci, 4 km NW Castelforte, 400 m, 1.vii.1969, leg. R. Johansson, genitalia slide 3155 Karsholt; 1 ♂, prov. Latina, Maranola, 400 m, 19.vi.1969, leg. R. Johansson; 1 ♂, prov. Torino, Torre del Colle, Villa Dora (To), 500 m, 5.vii.1991, leg. U. Parenti; 1 ♀, same data, but 9.vii.1991; 2 ♂, prov. Puglia, 5 km SW Manfredònia, 50 m, 26.v.2005, leg. P. Skou (all ZMUC); 1 ♂, prov. Albenga, Salea, 200 m, 4.vii.2012, leg. J. Skyva; 2 ♂, same data, but 8.vii.1996, leg. J. Liška (all NMPC); 4 ♂, prov. Calábria, Monte Pollino, Timpa del Demonio, 1200 m, 11.vii.1991, leg. G. Baldizzone (RCGB); 4 ♂, same data, but 3 km N Civita, 800 m, 31.v.–1.vi.2005, leg. P. Skou (TLMF, ZMUC); 1 ♂, same data, but 2 km NNW Civita, 775 m, 26.vii.2011, leg. P. Skou & B. Skule (ZMUC). France. 1 ♂, Dep. Alpes Maritimes, Caussols, Mgne du Cheiron, 1150 m, 12.v.2007, leg. J. Skyva (NMPC); 1 ♂, Dep. Alpes Maritimes, Caussols, 1100 m, 22.vi.2002, leg. J. Nel; 6 ♂, same data, but 14.viii.1971, leg. F. Dujardin; 4 ♂, Dep. Alpes Maritimes, St. Barnabé, 950 m, 14.viii.1953, leg. F. Dujardin; 1 ♂, Dep. Alpes Maritimes, La Turbie, 250 m, 30.v.–2.vi.1959, leg. K. Burmann; 1 ♂, Dep. Alpes Maritimes, Coursegoules, Nougueiret, 945 m, 6.vii.1995, genitalia slide 3988 Nel; 1 ♂, Dep. Alpes-de-Haute-Provence, Col d´Allos, W, 2300 m, 25.vii.1999, leg. J. Nel, gen. slide 9390 Nel; 1 ♂, same data, but 9.vii.2005, gen. slide 19514 Nel; 1 ♂, Dep. Alpes-de-Haute-Provence, Allos, 1500 m, 30.vii.1972, leg. F. Dujardin; 1 ♂, Dep. Alpes-de-Haute-Provence, St. Etienne, St. Sebastiene, 12.v.2001, leg. J. Nel; 1 ♂, Dep. Alpes-de-Haute-Provence, Dabisse, 3.vii.2005, leg. J. Nel; 1 ♀, Dep. Alpes-de-Haute-Provence, Digne, 600 m, 2–5.vi.1959, leg. K. Burmann, genitalia prep. 3769 Z. Tokàr (in glycerin); 2 ♂, Dep. Hautes Alpes, Les Vigneaux, 1200 m, 25.vii.1990, leg. P. Huemer & G. Tarmann (all TLMF); 1 ♀, Dep. Hautes Alpes, Les Vigneaux, 1000 m, 12.vii.2002, leg. J. Junnilainen; 1 ♂, same data, but Les Vigneaux 5 km W, 1250 m, 5–6.vi.2003; 1 ♂, 1 ♀, Dep. Hautes Alpes, Embrun, 850 m, 6–7.vii.2002, leg. J. Junnilainen, genitalia slide GU 16/1472 ♀ Huemer; 1 ♂, Dep. Hautes Alpes, Col de la Cayolle, Estene, 1800 m, 25.vi.2006, leg. J. Junnilainen (all RCJJ); 1 ♂, Dep. Hautes Alpes, La Bessèe, 1100 m, 21.vii.1961, leg. K. Burmann; 1 ♂, Dep. Hautes Alpes, Ristolas, Le Brasc1640 m, 13.vii.1990, leg. R. Robineau, genitalia slide 2078 Nel; 1 ♂, Dep. Hautes Alpes, Col de Montgenèvre, 1800 m, 1.viii.1998, leg. J. Nel, genitalia slide 8536 Nel; 1 ♂, Dep. Hautes Alpes, Pelvoux, 2300 m, 29.vii.1994, leg. J. Nel, genitalia slide 2276 Nel; 1 ♂, Dep. Hautes Alpes, Villard, St. Crepin, 4.vi.2005, leg. J. Nel (all TLMF); 3 ♂, Dep. Hautes Alpes, Céreste, 8–17.vi.2010 (RCHW, ZMUC); 1 ♂, Dep. Ardèche, Lablachere, 2.vii.2004, leg. J. Procházka (NMPC); 1 ♂, Dep. Vaucluse, Mt. Ventoux, 18–19.v.1990, leg. G. Luquet, genitalia slide 4660 Nel; 2 ♂, same data, but 30–31.v.1990, genitalia slides 4660 Nel, 4654 Nel; 1 ♂, Dep. Vaucluse, Méthamis, 1.viii.1995, leg. G. Brusseaux; 1 ♂, same data, but 30.vii.1995, genitalia slide 44165 Nel; 2 ♀, Dep. Var, Val de Purian, Rougiers, 15.vi.1994, leg. J. Nel, gen. slide 2116 Nel; 1 ♂, Dep. Var, Camillier, Montmeyan, 550 m, 13.vi.1994, leg. T. Varenne, genitalia slide 2551 Nel; 1 ♂, Dep. Var, Caramy, 15.vi.1997, leg. J. Nel, genitalia slide 6048 Nel; 1 ♂, Dep. Var, Sommet Mt. Caume, 1.vii.1995, leg. J. Nel, genitalia slide 3517 Nel; 1 ♂, Dep. Var, Pourrières, riv. de l´Arc, 27.v.2000, leg. J. Nel, genitalia slide 11372 Nel; 1 ♂, Dep. Var, Mtgne. Lachens, 1600 m, 7.vii.1994, leg. J. Nel, genitalia slide 2225 Nel; 1 ♂, same data, but 11.vii.1995; 1 ♂, Dep. Var, Plan d´Aups, Bertagne, 850 m, 24.v.1999, leg. J. Nel, genitalia slide 10708 Nel; 1 ♂, Dep. Var, Puits de Rians, 20.vi.2992, leg. J. Nel, genitalia slide GEL 1222 Huemer; 1 ♂, Dep. Var, Signes, le Camp, 24.v.1998, leg. J. Nel (all TLMF); 1 ♂, Dep. Var, Montmeyan, 10.vi.1981, leg. Hahn (RCEA); 1 ♂, Dep. Bouches-du-Rhône, La Ciotat, Roumagoua, 30.v.1995, leg. J. Nel, genitalia slide 3449 Nel (TLMF); 1 ♂, Provence, 3 km W Cereste, 3–4.vii.1989, leg. B. Å. Bengtsson, genitalia slide Bengtsson 3266 (RCBB); 1 ♂, Rhône-Alpes, Montelimar, 27.vi.1980, leg. K. Schnack; 1 ♂, Alpes Maritimes, Col de Vence, 11–12.vi.1981, leg. Hahn (ZMUC); 1 ♂, Dep. Ardèche, St. Laurent-du-Pape, 400 m, 20–21.vi.1999, leg. P. Skou; 1 ♀, Dep. Hautes-Alpes, Châteauvieux, 750 m, 26.vi.1996, leg. A. Cox; 1 ♂, same data, but 3.vii.1998; 2 ♂, Dep. Hautes-Alpes, Réotier, 1000 m, 23–30.vii.1990, genitalia slide 4516, 4517 Hendriksen; 1 ♂, Dep. Hautes-Alpes, 1.2 km SSW Prelles, 1200 m, 2.vii.2017, leg. P.Skou (all ZMUC); 1 ♂, Dep. Var, Pont l’Arbuty, 800 m, 5.vii.1994, leg. K. Larsen (ZMUC); 1 ♂, Dep. Vaucluse, Vaison la Romaine, 200 m, 17.vii.1991, leg. A. Cox, genitalia slide 4518 Hendriksen (ZMUC). Montenegro. 3 ♂, Fundina, 15.vi.2011, leg. I. Richter. genitalia slide 5315 Karsholt (RCIR, ZMUC). Spain. 1 ♂, prov. Alicante, Sierra de Crevillente, 5 km NE Albatera 450 m, 23.v.2004, leg. P. Huemer, genitalia slide GEL 1238 Huemer (TLMF); 1 ♂, prov. Alicante, 3.8 km NW Torremendo, 5.v.2008, leg. J. Tabell (ZMUC); 2 ♂, prov. Almería, Sierra de los Filabres, Calar Alto, 2130 m, 5.vii.2015, leg. J. Tabell, genitalia slide GU 16/1430 Huemer (TLMF, ZMUC); 2 ♂, same data, but 2000 m, 1–2.viii.2010, leg. J. Šumpich, genitalia slide GU 16/1436 Huemer; 1 ♂, same data, but route Purchena - Senes, 1600 m, 16.vi.2007, leg. J. Šumpich (all NMPC); 2 ♂, same data, but Calar Alto, 1900–2022 m, 17–18.vi.2007, leg. J. Šumpich (NMPC); 1 ♂, prov. Almería, Tabernas, 380 m, 6–8.vii. 2007, leg. G. Jeppesen (ZMUC); 1 ♂, prov. Almería, Maria, 1200 m, 18–25.vi.2008, leg. M. Delnoye (ZMUC); 1 ♂, prov. Avila, Sierra de Villafrance, 1 km W La Herguijuela, 1650 m, 20.vii.2003, leg. B. Skule (ZMUC); 1 ♂, prov. Avila, Sierra de Gredos, below Platforme de Gredos, 1700 m, 21.vii.2003, leg. B. Skule; 1 ♂, prov. Burgos, Espinosa de Cervera, 24.vii.1988, leg. M. Hull (ZMUC); 1 ♂, same data, but 1650 m, 25.vi.1986, leg. P. Skou (ZMUC); 2 ♂, prov. Castellón, 20 km SE Morella, 15.vi.1989, leg. B. Å. Bengtsson, genitalia slide Bengtsson 3264 (RCBB, ZMUC); 1 ♂, prov. Castellon, 5 km E Cuevas de Vinroma, 200 m, 13.vii.1992, leg. M. Fibiger; 1 ♂, prov. Castellon, 25 km NW Castellon, La Banderetta pass, 800 m, 17.vii.1992, leg. M. Fibiger (all ZMUC); 1 ♂, prov. Girona, Port Bou, 0–600 m, 18.vi.–1.vii.1963, leg. M. & W. Glaser (SMNK); 2 ♂, same data, but 0–300 m, 9–24.vi.1964 (SMNK); 3 ♂, 1 ♀, prov. Girona, Cerdanya, Colle del Moixeró, 1972 m, 18.vii.2012, leg. J. Dantart, genitalia slide GU 16/1412 ♂ Huemer (RCJD, TLMF); 4 ♂, prov. Lleida, Puente de Montanana, 15.vi.2012, leg. P. Huemer & A. Mayr (TLMF); 3 ♂, prov. Zaragoza, Gelsa 8 km NE, 240 m, 19.v.2016, leg. J. Tabell (ZMUC); 5 ♂, prov. Zaragoza, Caspe 7 km N, 18.v.2004 (RCJJ); 6 ♂, prov. Huesca, Los Monegros, Castejón de Monegros, 570 m, 10.vi.2007, leg. J. Šumpich; 3 ♂, same data, but 20.vi. 2007; 1 ♂, same data, but 26–27.iv.2003 (all NMPC); 1 ♂, prov. Huesca, 8 km S Candasnos, Barranco de Valcuerna, 175 m, 5.vii.2002, leg. B. Skule (ZMUC); 2 ♂, same data, but 250 m, 28.vi.2005, leg. D. Feierabend; 1 ♂, same data, but 10 km S, 30.v.2015, leg. J. Viehmann; 1 ♂, same data, but 3.vii.2016 (all RCWS); 1 ♂, prov. Soria, El Burgo de Osma, 895 m, 14–15.vi.1995, leg. A. Cox, genitalia slide 6104 Wolf (RCHW); 2 ♂, prov. Soria, 30 km SW Soria, El Temerosa, 1080 m, 18.vii.2012, leg. T. Nupponen, genitalia slide 5015 Tabell (ZMUC); 1 ♂, prov. Tarragona, 2 km S Miami Hospital, 0 m, 12.vii.1992, leg. M. Fibiger (ZMUC); 2 ♂, prov. Teruel, Albarracin, 1100 m, 19.vi.2007, leg. J. Šumpich; 1 ♂, same data, but 3.v.2003; 5 ♂, 1 ♀, same data, but 28.vii.2010; 1 ♂, same data, but7.viii.2010; 1 ♂, same data, but 13.vii.2012, leg. M. Dvorak; 2 ♂, same data, but 14.vii.2012, leg. M. Dvorak (all NMPC); 26 ♂, 1 ♀, prov. Teruel, Sierra de Cucalon, Baguena 5 km E, 15.vi.2004, leg. J. Junnilainen (RCJJ); 1 ♂, prov. Teruel, Cosa, 2–13.viii.1989, leg. C. Gielis (RCHW); 1 ♂, prov. Teruel, Albarracin, 26.vi.1982, leg. C. Gielis; 1 ♂, same data, but 1000–1200 m, 4–8.viii.1989, leg. C. Gielis; 2 ♂, Teruel, 1 km E Albarracin, Valdevacar, 12.vii.2001, leg. C. Gielis (all RMNH); 6 ♂, prov. Teruel, Albarracin, 4.5 km NE, 1110 m, 7.vii.2016, leg. J. Tabell (TLMF, ZMUC); 3 ♂, prov. Teruel, Albarracin, Val de Vacar, 1200 m, 16.vii.1992, leg. M. Fibiger; 3 ♂, same data, but 1250 m, 17–18.vii.1988, leg. M. Fibiger, genitalia slide 4544 van der Wolf (ZMUC); 1 ♂, same data, but 5.vi.1993, leg. J. Wolschrijn (ZMUC); 1 ♂, same data, but 1100 m, 22.v.1998, leg. P. Skou; 1 ♂, same data, but 10.vi.1999; 4 ♂, same data, but 12.vi.1999 (ZMUC); 2 ♂, same data, but 15.vi.2004, leg. Viehmann, genitalia slide 9217 van der Wolf (RCWS); 1 ♂, same data, but 1200 m, 4.viii.2007, leg. B. Skule & P. Skou (ZMUC); 1 ♀, prov. Teruel, Albarracin, 1100 m, 14.vi.1986, leg. P. Skou; 3 ♂, same data, but 1000 m, 7.viii.1988 (ZMUC); 1 ♂, same data, but, 1170 m, 8–10.vi.1994, leg. A. Cox, genitalia slide 4514 Hendriksen (ZMUC); 1 ♂, same data, but, 1170 m, 18–20.vi.1996, leg. A. Cox, genitalia slide 4513 Hendriksen (ZMUC); 1 ♂, same data, but 2–4.vi.1999, leg. A. Cox (ZMUC); 2 ♂, same data, but 1200 m, 11.vi.1999, leg. P. Skou (ZMUC); 1 ♂, same data, but 3 km W, 1200 m, 1.vii.2005, leg. D. Feierabend (RCWS); 1 ♂, same data, but 3.5 km SWW, 1200 m, 13.vi.2010, leg. J. Tabel, genitalia in tube (ZMUC); 1 ♂, same data, but 2 km NW, 1100 m, 4.vii.2010, leg. Z. Tokár (RCZT); 7 ♂, same data, but 4.5 km N, 1110 m, 7.vii.2016, leg. J. Tabel (ZMUC); 1 ♂, prov. Teruel, Pozondon, 1500 m, 21.vii.1984, leg. W. O. De Prins (ZMUC); 2 ♂, prov. Teruel, 5 km NW Montalban, 950 m, 17.vii.2003, leg. B. Skule (TLMF, ZMUC); 2 ♂, same data, but 5.viii.2007, genitalia slide 6491 Hendriksen (ZMUC);1 ♂, prov. Teruel, 1 km E Tramacastillo, 1250 m, 3.ix.2001, leg. B. Skule & BC. Hviid (ZMUC); 13 ♂, prov. Teruel, Moscardon, 1500 m, 3.vii.2016, leg. J. Viehmann (RCWS); 1 ♂, 1 ♀ prov. Granada, Sierra de Alfacar, 1500 m, 27.vi.1968, leg. K. Sattler & D.J. Carter, gen. slides 33657 ♂, 33658 ♀ (BMNH); 4 ♂, prov. Granada, Sierra Nevada, El Pardor Nat, 2500 m, 21.vii.1980, leg. E. Traugot-Olsen; 1 ♂, same data, but 23.vii.1983, genitalia slide 5777 Traugott-Olsen; 1 ♂, prov. Granada, Sierra Nevada, Camino de Valeta, 2300 m, 23.vii.1983, leg. G. Baldizzone & P. Triberti; 1 ♂, same data, but 2250 m, 2.viii.1984, leg. E. Traugott-Olsen; 1 ♂, same data, but 17.viii.1984; 1 ♂, same data, but 2300 m, 19.viii.1984; 1 ♂, same data, but 2050 m, 6.viii.1986; 3 ♂, same data, but 2250 m, 21.vii.1985, leg. G. Baldizzone & E. Traugott-Olsen, genitalia slide 6534 Hendriksen; 1 ♂, same data, but 23.vii.1985 (all ZMUC); 1 ♂, same data, but 1.viii.1984, leg. E. Traugott-Olsen, genitalia prep. 4128 Z. Tokár (in glycerin) (NHMW); 1 ♂, Granada, Puerto de la Ragua, 1800 m, 23.vii.2003, leg. P. Skou; 1 ♂, Granada, Puerto de la Ragua, 2200 m, 3.viii.1982, leg. W.O. De Prins (all ZMUC); 1 ♂, Granada, Cam. Baza-Benamaurel, 15 km de Baza, 16.vii.1987, leg. G. Baldizzone & E. Traugott-Olsen; 3 ♂, same data, but 19.vii.1987; 2 ♂, same data, but 20.vii.1987 (all TLMF); 1 ♂, Segovia, San Ildefonso 26.vii.1884, coll Staudinger (ZMHU); 2 ♂, prov. Zaragoza, Tosos, 600 m, 16.vi.1999, leg. P. Skou (ZMUC); 1 ♂, prov. Zaragoza, Nuévalos, 1200 m, 9.vi.2003, leg. H. van der Wolf (RCHW); 5 ♂, prov. Zaragoza, 6 km W Bujaraloz, 300 m, 29.v.2015, leg. J. Viehmann (RCWS, ZMUC). Without locality: 2 ♂, [San Ildefonso?] 26.vi.[year unknown], coll. Staudinger; 1 ♂, same data, but 13.vii.; 1 ♂, same data, but 26.vii. (all ZMHU).

Adult. Male (Figs 7–8, 10–16). Wingspan 12–17 mm. Segment 2 of labial palpus with long scale brush, brownish at outer and inner surface, white at lower and upper surface; segment 3 about half length of segment 2, white with black tip. Antennal scape without pecten; flagellum ringed black and whitish brown. Head, thorax and tegula whitish brown. Forewing light yellow, mottled with brown-tipped scales, forming a darker area in costal part of wing; costa white; veins indistinctly white; a small black dot at end of cell; a dark streak towards apex; fringes whitish grey. Hindwing light grey with cream-white fringes.

Female (Figure 9). Wingspan 12–17 mm. Forewing ellipsoid with elongate apex; same colour as male, but apex with more black scales. Hindwing distinctly reduced in width; apex produced. Otherwise similar to male.

Variation. Megacraspedus lanceolellus shows both individual and also a tendency to geographical variation. The nominotypical form has light yellowish forewings with a dark shadow or streak towards the apex, but such individuals also occur elsewhere, e.g., in Spain. Specimens from south-western France and northern and central Spain (described as M. tutti and M. grossisquammellus, respectively) have a more or less pronounced white costa and can moreover often be recognized by the light yellow forewings with darker veins. Specimens from the Sierra Nevada in southern Spain (described as M. subdolellus) are larger on average (wingspan 14–20 mm) and are characterised by their more clear yellow forewings that almost lack black dots, and the whitish costa. Specimens from Montenegro have a black dot at 2/5 in the fold.

Male genitalia (Figs 153–156). Uncus moderately slender, nearly twice as long as maximum basal width, sub-triangular, gradually tapered to weakly rounded apex; gnathos hook stout, slightly longer than uncus, with curved and pointed apex; anterior margin of tegumen with broad, medially deep V-shaped emargination; pedunculi small; valva straight, slender, extending to about base of gnathos, apex weakly contorted, rounded, saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with moderately deep medial emargination, broadly rounded lateral humps, vincular sclerites broadly sub-rectangular, tapered towards valva, with sclerotised posterior edge; saccus long and moderately broad, almost V-shaped, with abruptly tapered distal part and pointed apex, ratio maximum width to length 0.75, posterior margin strongly bulged, with rounded mediolateral projections, separated by moderately shallow incision, medial part with elongated ridge from posterior edge to middle, lateral sclerites of variable length, slightly shorter to longer than maximum width of saccus; phallus about length of tegumen, stout, straight, with bulbous coecum, sclerotised band in anterior part of coecum, distal two-thirds stout, with rod-like ventral and lobed dorsal sclerotisations, ductus ejaculatorius with two linear interior sclerotisations.

Female genitalia (Figs 267–268). Papilla analis small, apically rounded; apophysis posterior slender rod-like, about 4 mm long, with short, bifurcate posterior end, bordered by minute sclerotised field, apex widened and rounded; segment VIII approximately 1.5 mm long, membranous; subgenital plate with sub-triangular subostial sclerotisation, posteriorly extended into shortly pointed sclerites, delimiting trapezoidal ostium bursae, anterior margin with rod-like edge connected with apophysis anterior, medially with long sinusoid projection; apophysis anterior slender, rod-like, at most one-third length of segment VIII, posteriorly becoming rod-like venula of segment VIII, extending to posterior margin; colliculum short, sclerotised; ductus bursae short, moderately broad; corpus bursae, moderately short and broad, weakly delimited from ductus bursae, entire length of ductus and corpus bursae nearly 2 mm; signum small, rounded spiny plate.

Due to its variation M. lanceolellus is difficult to characterise. It is a medium-sized species with light yellowish forewings, mottled with brown and black-tipped scales, a lighter costa, and sometimes one or two black dots in the middle of the wing and/or a black streak towards apex. See also M. bengtssoni sp. n. (p 37) and M. andreneli (p 96). The male genitalia are characterised by several structures such as the sub-triangular shape of the uncus, the slender and apically weakly contorted valva, the shape of the saccus with a long medial ridge, and particularly the massive distal part of the phallus with a broadly lobed dorsal and a slender ventral sclerotisation, as well as the two linear sclerotisations of the ductus ejaculatorius. They are somewhat similar to M. bengtssoni sp. n. but differ e.g., by the longer uncus, the medially strongly emarginated anterior margin of the tegumen, the longer and more slender valva, and the broader phallus.

The female genitalia are characterised by the long sinusoid anteriomedial projection of segment VIII, but in the absence of females of closely related species such as M. bengtssoni sp. n., these features may not be specifically diagnostic.

The extraordinary DNA barcode divergence is reflected by 19 BINs! The mean and maximum intraspecific divergence of the barcode region in this species is 7.6% and 12.5% respectively, largely reflecting a geographic pattern with several distinct clusters and a remarkable divergence of 4.3 to 8.1% to the nearest cluster. However, even within these clusters intraspecific variation of the barcode region is considerable with e.g., mean divergence of 2.5% in south-western alpine populations. Four specimens collected in the same microhabitat at Sierra de los Filabres (prov. Almería, Spain) form two distinct clusters with a variation of maximum 0.2% within clusters but minimum 8.6% between these clusters. Similarly diverging sympatric clusters are reported from prov. Teruel, and from Prov. Lleida.

The following 19 clusters are defined (based on sequenced material):

BIN lanc01 (Italy: Pordenone, Calabria; Croatia): BOLD:AAU2834 (n = 4).

BIN lanc02 (Italy: Calabria): BOLD:ACZ3381 (n = 1).

BIN lanc03 (Croatia): BOLD:ACZ2933 (n = 1).

BIN lanc04 (Italy: Livorno): BOLD:ACZ2657 (n = 1).

BIN lanc05 (Spain: Almeria): BOLD:ACZ9024 (n = 2).

BIN lanc06 (Andorra): BOLD:ACA9759 (n = 2)

BIN lanc07 (Spain: Barcelona): BOLD:ACZ8656 (n = 2).

BIN lanc08 (France: Alpes Maritimes): BOLD:ABA3648 (n = 1).

BIN lanc09 (Italy: Savona): BOLD:ACZ2656 (n = 1).

BIN lanc10 (France: Alpes Maritimes, Hautes Alpes, Var): BOLD:ABA3649 (n = 2).

BIN lanc11 (Spain: Almeria): BOLD:ACZ2607 (n = 2).

BIN lanc12 (Spain: Teruel, Valencia): BOLD:AAU1829 (n = 5).

BIN lanc13 (Spain: Soria): BOLD:ACZ8269 (n = 1).

BIN lanc14 (Spain: Teruel): BOLD:ADF2256 (n = 2).

BIN lanc15 (Spain: Teruel): BOLD:ACM1006 (n = 1).

BIN lanc16 (Spain: Zaragoza): BOLD:ADB7571 (n = 1).

BIN lanc17 (Spain: Zaragoza): BOLD:ADF2263 (n = 3).

BIN lanc18 (Spain: Lleida): BOLD:ADF2194 (n = 1).

BIN lanc19 (Spain: Lleida): BOLD:ADF1916 (n = 1).

The minimum distance to the nearest congeneric neighbour M. dolosellus is 9.9% (p-dist).

Southern parts of Europe, from the Balkan Peninsula to Italy, France, and Spain, also known from Germany (northwards to Rheinland-Pfalz (Hausenblas 2006: 10)). According to Mariani (1943: 174) also in Sicily and Corsica.

Early stages are undescribed. According to Lhomme (1946: 538 as M. tutti) the larva feeds within the stem of Festuca L. The adults have been collected from late April until the middle of August, from low altitudes to 2300 m in the Alps and 2500 m in the Sierra Nevada. There is probably only one generation each year, and the flight period seems to at least partly depend on the altitude of the collecting site.

Ypsolophus lanceolellus was described from an unstated number of males collected by J. Mann at Ardenza and Salviano, Italy (Zeller 1850).

Megacraspedus hessleriellus was described from two males collected at Biebrich and Mombach in Nassau, S Germany (Rössler 1866: 348). It was synonymised with M. lanceolellus by Heinemann (1870: 349).

Megacraspedus subdolellus was described from an unspecified number of both sexes collected in August at high altitudes (ca. 2700 m) in the Sierra Nevada, Spain (Staudinger 1859). Despite the slightly differing phenotypic appearance (see above) we found no sign of diagnostic characters in genitalia structures and thus consider M. subdolellus to be conspecific with M. lanceolellus (syn. n.). DNA barcodes are not yet known from the type locality. A lectotype, already labelled as such by K Sattler, is here designated in order to fix the identity of the species and conserve stability of nomenclature.

Megacraspedus tutti was described from a single male collected in August at La Grave, SE France by JW Tutt (Walsingham 1897). Despite of the slightly differing phenotypic appearance compared to nominotypic M. lanceolellus (see above) we found no sign of diagnostic characters in genitalia structures and thus consider M. tutti to be conspecific with M. lanceolellus (syn. n.). Megacraspedus grossisquammellus was described from an unstated number of specimens collected in Spain, Segovia, San Ildefonso, in June and July 1902. A lectotype was designated by Agenjo (1962: 161, pl. 2–3) and figured together with its genitalia. Two paralectotypes and numerous additional material from Spain examined by us show a considerable individual variation but cannot be reliably separated from M. lanceolellus in other areas, particularly in genitalia characters. We therefore consider M. grossisquamпellus to be conspecific with M. lanceolellus (syn. n.).

Megacraspedus lanceolellus shows a remarkable morphological and genetic variation. Barcode data clearly support several putative taxa, but the morphology gives a different and less straightforward picture. Though the phenotypic appearance partly depends on geography it is frequently impossible to reliably assign specimens to a barcode cluster. Male genitalia morphology is comparatively uniform and alleged diagnostic characters show intraspecific variation. Considering the intrapopulational barcode variation which exceeds 8% on one occasion we conclude that M. lanceolellus is a widespread, mainly western-Mediterranean species. The exceptional DNA barcode divergence may be explained by weak dispersal ability, particularly of the slightly brachypterous females, leading to several genetically isolated populations which are mainly restricted to various mountain systems. However, it seems unlikely that this is the sole cause for the observed intraspecific variation (see Discussion).

Holotype ♂, “SPAIN [prov.] Aragon Gelsa 8 km NE, 240 m N41.46004 W0.37642 19.5.2016 J. Tabell leg.” “DNA Barcode TLMF Lep 21279” (ZMUC). Paratypes. Spain. 1 ♂, prov. Alicante, Parcent, 450 m, 27.vi.2015, leg. H. Rietz; 1 ♂, prov. Alicante, Sierra de Crevillente, 400 m, 5.vi.2016, leg. H. Roweck & N. Savenkov (all ECKU); 1 ♂, prov. Castellon, 5 km E Cuevas de Vinroma, 200 m, 13.vii.1992, leg. M. Fibiger; 1 ♂, prov. Cuenca, Uña, 1150 m, 8–9.vii.2002, leg. B. Skule; 1 ♂, prov. Huesca, 6 km S Candasnos, Barranco de Valcuerna, 250 m, 19.vi.1998, leg. P. Skou, genitalia slide 3354 Hendriksen (all ZMUC); 10 ♂, prov. Huesca, 6 km SW Candasnos, Barranco de Valcuerna, 300 m, 15.vi.1999, leg. P. Skou, genitalia slide GU 15/1393 Huemer (TLMF, ZMUC); 1 ♂, prov. Cuenca, Cuenca, 29.vi.1982, leg. C. Gielis (RMNH); 1 ♂, prov. Cuenca, Uña, 1300 m, 12.ix.2007, leg. J. Viehmann; 1 ♂, prov. Huesca, Ontiñena, 300 m, 28.v.2015, leg. J. Viehmann; 1 ♂, prov. Huesca, 10 km SW Candasnos, 31.v.2016, leg. J. Viehmann (all RCWS); 5 ♂, same data, but 30.v.2015, leg. J. Viehmann (RCWS, ZMUC); 2 ♂, prov. Huesca, 10 km S Benabarre, Esteña, 800 m, 18.vi.1999, leg. P. Skou; 1 ♂, prov. Lleida, Artensa de Segre, Rubió, 14.vi.1989, leg. B. Å. Bengtsson, genitalia slide 3265 Bengtsson (RCBB); 1 ♂, prov. Lleida, 30 km NW Fraga, Ontinema, 250 m, 11.vii.1992, leg. M. Fibiger; 1 ♂, prov. Tarragona, Falset, 600 m, 19.vii.1984, leg. W. De Prins; 1 ♂, prov. Tarragona, 5 km S St. Carles de Ràpita, 20 m, 14.vi.1999, leg. P. Skou; 1 ♂, prov. Teruel, Cosa, 10.vii.1986, leg. C. Gielis; 1 ♂, prov. Teruel, 5 km NW Montalban, 950 m, 17.vii.2003, leg. B. Skule, genitalia slide GU 15/1394 Huemer (all ZMUC); 3 ♂, prov. Teruel, Sierra de Cucalon, 5 km E Baguena, 15.vi.2004, leg. J. Junnilainen, genitalia prep. (in glycerin); 2 ♂, prov. Zaragoza, Caspe, 6.v.2005, leg. J. Junnilainen (all RCJJ); 1 ♂, prov. Zaragoza, Tosos, 400m, 28.vi.1992, leg. P. Skou & B. Skule; 1 ♂, prov. Zaragoza, 4 km N Tosos, 400m, 28.vi.1997, leg. P. Skou (ZMUC).

Adult. Male (Figs 17–18). Wingspan 9–12 mm. Segment 2 of labial palpus with scale brush of about same length as segment 3, dark brown on outer surface, white mottled with brown on inner surface, white on upper surface; segment 3 about length of segment 2, white at base, becoming black towards tip. Antennal scape without pecten; flagellum ringed black and light grey-brown. Head cream-white; thorax and tegula light brown, the latter with whitish tip. Forewing light grey-brown with scattered black scales, especially in fold and apical part; costa white from 1/3; a black dot at end of cell; some black scales along termen; fringes light grey. Hindwing light grey with concolorous fringes.

Female. Unknown.

Variation. There is slight variation in the colour of the forewing (from light greyish towards light grey-brown) and in the amount of blackish scales. Some specimens have an indistinct black dot in the fold.

Male genitalia (Figs 157–158). Uncus moderately slender, 1.5 times as long as maximum basal width, sub-ovate, gradually tapered to weakly rounded apex; gnathos hook stout, slightly longer than uncus, straight, with pointed apex; anterior margin of tegumen with broadly rounded emargination, short ridges from anterior edge converging towards middle; pedunculi small, with additional sclerite; valva, straight, moderately stout, extending to about middle of uncus, apex weakly contorted, rounded, saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with shallow emargination, indistinct lateral humps, vincular sclerites broadly sub-rectangular, tapered towards valva, with sclerotised posterior edge; saccus moderately short and broad, almost U-shaped, with abruptly tapered and pointed apical fifth, ratio maximum width to length about 1, posterior margin strongly bulged, with rounded mediolateral projections, separated by moderately shallow incision, medial part with sclerotised ridge from posterior edge to first third, lateral sclerites about two-thirds to full length of maximum width of saccus; phallus slightly shorter than tegumen, weakly curved medially, with bulbous coecum, sclerotised band in anterior part of coecum, distal two-thirds slender, with weakly rod-like sclerotisation ventrally, subapical area with small sclerite, with or without minute tooth, ductus ejaculatorius with linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus bengtssoni sp. n. is characterised by its relatively small size and by its light greyish brown forewings with a single black dot at the end of the cell. It is similar to M. lanceolellus, but that species is larger, it has shorter segment 3 of the labial palps and the costa is white from base. See also M. dejectella (p 85). The male genitalia are somewhat similar to M. lanceolellus (Figs 153–156) but differ in several characters such as the shorter uncus, the medially less emarginated anterior margin of tegumen, the shorter valva, and the broader saccus with a shorter medial ridge.

BIN: BOLD:ACM1097 (n = 2). The intraspecific divergence of the barcode region has a maximum divergence of 0.2%. The distance to the nearest congeneric neighbour M. lanceolellus is 12% (p-dist).

Spain.

Host plant and early stages are unknown. The adults have been collected from the middle of May to the middle of July at altitudes up to 1150 m.

The species name, a noun in the genitive case, is dedicated to Bengt Å. Bengtsson, Sweden, who collected part of the type series of this species and other valuable Megacraspedus specimens used for our study.

Morocco. 2 ♂, Prov. Al Haouz, Oukaïmeden 1.5 km SE, 2660 m N31.19340 W7.85311, 2.vii.2016, leg. J. Tabell, DNA Barcode TLMF Lep 21271, genitalia slide GU 16/1453 Huemer (ZMUC); 7 ♂, same data, but 2400 m, 7–17.vi.1965, leg. Y. de Lajonquière (SMNK, ZMUC); 9 ♂, Middle Atlas, Ifrane, 5.–10.vii.1972, genitalia slide GU 5264 Stübner, leg. F. Hahn (SMNK, TLMF, ZSM).

Adult. Male (Figs 19–20). Wingspan 18–22 mm. Segment 2 of labial palpus with scale brush, light brown, on outer surface mottled with black-tipped scales, white on upper surface; segment 3 thin, longer than scale brush of segment 2, whitish brown mottled with black towards tip. Antennal scape without pecten; flagellum ringed with light brown and black. Head whitish brown; thorax and tegula light brown. Forewing light yellow-brown, mottled with scattered black scales, especially at base of costa and in apical area; veins indistinctly lighter; fringes light grey. Hindwing grey with grey fringes.

Female (based on original description). Wingspan 11–16.5 mm. Forewing lanceolate, shorter than the body, hindwing narrow; both pairs almost entirely reddish at the base.

Variation. A rather variable species as regards the amount of blackish brown scales on the forewings. Specimens with many such scales look greyish yellow. Material from Middle Atlas is rather reddish-brown than yellow-brown.

Male genitalia (Figure 159). Uncus large, approximately 1.3 times longer than wide, elongated suboval with evenly rounded apex; gnathos hook moderately slender, slightly longer than uncus, hardly curved with pointed apex; anterior margin of tegumen with deep V-shaped emargination, sclerotised edge merged mediad in sub-anterior part of tegumen; pedunculi distinct, suboval; valva massive, stout, thumb-shaped, extending slightly beyond base of uncus, apical part weakly divergent, rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with deep U-shaped emargination, weakly rounded lateral humps, suboval vincular sclerite with strongly sclerotised posterior edge; saccus long, sub-triangular, evenly tapered to pointed apex, ratio maximum width to length approximately 0.6, posterior margin with rounded mediolateral projections, separated by moderately broad and shallow incision, medial part with strongly sclerotised longitudinal ridge, forked in medial part of saccus and extending to anteriolateral edge, lateral sclerites about length of maximum width of saccus; phallus with weakly bulbous coecum, distal two-thirds stout, straight, medial ridge with small sub-apical spine, ventral edge with larger sub-apical tooth, both pointing ventrad, apex rounded, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Undescribed.

Megacraspedus homochroa is characterised by its rather large size and its light yellow-brown to reddish-brown forewings without spots. See also M. tenuignathos sp. n. (p 126). The male genitalia are unmistakable due to the combined shape of the large uncus and saccus, and the peculiar structures of the phallus.

BIN BOLD:ADF2257 (n = 1). The distance to the nearest neighbour M. golestanicus sp. n. is 9.1% (p-dist).

Morocco (Middle Atlas, High Atlas).

Host plant and early stages are unknown. The type-series of M. homochroa was collected in the Middle Atlas in August at an altitude of 2400 m, that of M. culminicola at high altitude of 3200 m in the middle of July. Additional samples date from about mid-June to early July and at altitudes from ca. 2400 to 2660 m.

Megacraspedus homochroa was described from two males collected in Morocco, Middle Atlas (Le Cerf 1932). The type-material could not be traced at MNHN, but the description of the adults leaves no doubt as to the identity of this species.

Megacraspedus culminicola was described from two males and two females taken in copula in Morocco, Middle Atlas (Le Cerf 1932). Like M. homochroa, the type-material could not be traced at MNHN, but descriptions of the males of both species agree in almost every detail and the type-localities are nearby. We therefore consider M. culminicola to be a junior synonym of M. homochroa (syn. n.).

Lectotype ♂, here designated, “Asia min. Penther ’02 Ilgin.” “Megacraspedus monolorellus Type Rbl.” “BC TLMF Lep 06704” “NM 16.649 ♂” (NHMW). Paralectotypes. 1 ♂, 1 ♀, [in copula] “Ilgün ´02 Penther” “Megacrasp Monolorellus Rbl Type” (NHMW). Turkey. 1 ♂, Silifke, Mut, 400 m, 31.v.1996, leg. F. Schepler, genitalia slide 5338 Karsholt (ZMUC); 1 ♂, 1 ♀, 30 km NE Konya, 27.v.1969, leg. E. Arenberger (RCEA); 16 ♂, prov. Konya, 25 km E Konya, 23.v.1997, leg. K. Nupponen & J. Junnilainen (RCKN).

Adult. Male (Figure 21). Wingspan 14–16 mm. Segment 2 of labial palpus with scale brush of same length as segment 3, brown on outer surface, whitish brown on inner surface, otherwise white; segment 3 about same length as segment 2, white mottled with light brown. Antennal scape with pecten of one hair; flagellum black, indistinctly ringed with grey. Head, thorax and tegula whitish brown. Forewing brown, slightly mottled with white scales; a distinct white stripe along costa, broadest in middle; base of costa blackish; fringes light grey. Hindwing grey with light grey fringes.

Female (Figure 22). Wingspan 11 mm. Labial palpus cream-white, segment 2 finely mottled with brown. Antennal scape with pecten of one hair; flagellum blackish brown, ringed with white. Forewing broadest in middle, apical half tapering into lanceolate, blackish tip. Hindwing very short (about one-fifth length of forewing), slender with rounded apex, whitish. Abdomen as long as forewing, whitish brown, becoming white at apical segments. Ovipositor long, protruding. Otherwise similar to male.

Variation. The examined specimens show only slight variation.

Male genitalia (Figure 160). Uncus moderately broad, approximately 1.6 times longer than maximum basal width, weakly tapered from base to distal third, apically constricted with rounded apex; gnathos hook moderately stout, about length of uncus, weakly curved, narrowing towards pointed apex; anterior margin of tegumen with moderately deep and broadly V-shaped emargination, medial ridge from anterior edge to middle, anteriolateral edge with small peg-like sclerite; pedunculi distinct, irregular shape; valva evenly broad, short, stout, about maximum width of uncus, apex rounded, extending slightly beyond middle of uncus; saccular area covered with numerous setae, without separated sacculus; posterior margin of vinculum with moderately deep medial emargination, lateral humps weakly developed, vincular sclerite oblong, with broadly sclerotised posteriomedial edges; saccus nearly V-shaped, apical half strongly narrowing, with pointed apex, ratio maximum width to length approximately 0.8, posterior margin arched, with small medial incision, medial part with short, furcated ridge from posterior edge almost extended to medial part, lateral sclerites nearly length of maximum width of saccus; phallus moderately stout, with globular coecum, distal portion nearly straight, with broad dorsal and slender ventral lobes, apically tapered.

Female genitalia (Figure 269). Papilla analis small, apically rounded; apophysis posterior slender rod-like, approximately 3.6 mm long, with short, bifurcate posterior end, bordered by minute sclerotised field, apex weakly widened and rounded; segment VIII approximately 1.8 mm long, membranous; subgenital plate with sub-triangular subostial sclerotisation, posteriorly extended into moderately long, pointed sub-medial sclerites, with weak medial flaps delimiting oblong ostium bursae, anterior margin with rod-like edge connected with apophysis anterior, medially membranous, without anterior projection; apophysis anterior slender, rod-like, 2 mm length, posteriorly becoming rod-like venula of segment VIII, extending to posterior margin of segment VIII; colliculum short, sclerotised; ductus bursae and corpus bursae weakly delimited, moderately short with entire length of approximately 3.1 mm; signum moderately large, suboval spiny plate.

Megacraspedus monolorellus is characterised by its brown forewings with white costa and without further markings. It is similar to M. uzunsyrtus (p 45) and M. similellus sp. n. (p 46). The male genitalia are very similar to M. junnilaineni sp. n. (Figure 161) but differ particularly by the broader uncus and gnathos hook, and the shorter and broader valva. From other related species such as M. uzunsyrtus (Figure 162) and M. similellus sp. n. (Figs 163–164) they can be separated e.g., by the gradually tapered uncus. The female genitalia differ from M. similellus sp. n. (Figure 270) and all other species of Megacraspedus with known females in particular by the extraordinarily long apophysis anterior.

BIN BOLD:ACA8694 (n = 1). The distance to the nearest neighbour M. golestanicus sp. n. is 9.2% (p-dist).

Turkey.

Host plant and early stages are unknown. The type series was collected in the first third of May, and also the few additional specimens examined were found in May.

Megacraspedus monolorellus was described from several specimens of both sexes collected on 10.v.1902 near Ilgin (“Ilgün”), central Turkey (Rebel 1905: 213). An especially large male mentioned by Rebel is here designated as the lectotype, in order to stabilize nomenclature, and the above description is in first hand based on that well preserved specimen.

Holotype ♂, “Turkey [prov. Nevşehir] Urgup [Ürgüp] 7 km E 18.5.2005 J. Junnilainen leg.” “DNA Barcode TLMF Lep 19975” “GU 16/1448 ♂ P. Huemer” (RCJJ). Paratypes. Turkey. 1 ♂, same data as holotype (TLMF); 1 ♂, Nevşehir, 10 km V Ürgüp, Göreme, ‘Love Valley’, 1300 m, 2.vii.1997, leg. M. Fibiger, genitalia slide 5325 Karsholt (ZMUC); 1 ♂, prov. Istanbul, 5 km NE Aksaray, 19.v.2005, leg. J. Junnilainen (TLMF).

Adult. Male (Figure 23). Wingspan 12–13 mm. Segment 2 of labial palpus with scale brush almost as long as segment 3, brown on outer surface, whitish brown on inner surface, otherwise white; segment 3 about same length as segment 2, white, darker on lower surface and towards tip. Antennal scape with pecten of one hair; flagellum black, indistinctly ringed with grey. Head, thorax and tegula whitish grey-brown. Forewing light greyish brown from brownish white and black-tipped scales; base of costa blackish brown, otherwise whitish; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. The examined specimens show only slight variation.

Male genitalia (Figure 161). Uncus basally very broad, slightly longer than maximum basal width, gradually and weakly tapered from base to broadly rounded apex; gnathos hook slender, about one-third longer than uncus, weakly curved, narrowing towards pointed apex; anterior margin of tegumen with moderately deep and broad emargination, medial ridge from anterior edge to middle, anteriolateral edge with small peg-like sclerite; pedunculi distinct, irregular shape; valva moderately slender, long, maximum two-thirds width of uncus, gradually tapered to slightly pointed apex, extending slightly beyond middle of uncus; saccular area covered with numerous setae, without separated sacculus; posterior margin of vinculum with shallow medial emargination, lateral humps weakly developed, vincular sclerite sub-rectangular, with broadly sclerotised posteriomedial edges; saccus nearly V-shaped, apical third strongly narrowing, with pointed apex, ratio maximum width to length nearly 1, posterior margin arched, with small medial incision, medial part with short, furcated ridge from posterior edge almost extended to medial part, lateral sclerites about three-quarters length of maximum width of saccus; phallus moderately stout, with globular coecum, distal portion S-curved, with broad dorsal and long and slender ventral lobes, apically tapered.

Female genitalia. Unknown.

Megacraspedus junnilaineni sp. n. is characterised by its light greyish brown forewings without markings. Externally it is hardly separable from M. golestanicus sp. n. (p 48). The male genitalia are very similar to M. monolorellus (Figure 160) but differ in particular by the more slender uncus and gnathos hook, and the longer and comparatively slender valva. From other related species such as M. uzunsyrtus (Figure 162) and M. similellus sp. n. (Figs 163–164) they can be separated e.g., by the gradually tapered uncus.

BIN BOLD:ADB7272 (n = 1). The distance to the nearest neighbour M. similellus sp. n. is 9.4% (p-dist).

Turkey.

Host plant and early stages are unknown. The adults have been collected from the middle of May to the middle of August at altitudes of between 1200 and 1300 m.

The species name (a noun in the genitive case) is dedicated to Jari Junnilainen, Finland, who collected part of the type series of this species and numerous other valuable specimens used for our study.

Paratypes. Ukraine. 2 ♂, Crimea, Koktebel vic., SW slope of Uzun-Syrt, evening collecting, 17.v.2014, leg. Yu. Budashkin, genitalia slide GU 16/1463 Huemer (ZMKU).

Ukraine. 2 ♂, Crimea, Aj-Petri, 5–6.vii.2002, leg, Yu. Budashkin (ZMKU).

Adult. Male (Figure 24). Wingspan 12–14 mm. Segment 2 of labial palpus with moderately long scale brush, brown on outer surface, whitish brown on inner surface, otherwise white; segment 3 white, darker on lower surface and towards tip. Antennal scape with pecten of one hair; flagellum light grey-brown ringed with black. Head, thorax and tegula as forewing. Forewing light greyish brown from white brownish- and black-tipped scales, darkest towards apex; a distinct white stripe along costa; base of costa blackish; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. The examined specimens show only slight variation.

Male genitalia (Figure 162). Uncus moderately slender, slightly more than twice as long as broad, parallel-sided, apical 1/3 weakly narrowed, apex rounded; gnathos hook stout, about length of uncus, bent near middle, apex curved, pointed; anterior margin of tegumen with moderately deep and broadly rounded emargination, long medial ridge from anterior edge to posterior third, anteriolateral edge with small peg-like sclerite; pedunculi distinct, suboval; valva broad, about one-third width of uncus, straight, of equal width, apex rounded, extending to middle of uncus, sub-basally with rounded hump; saccular area covered with setae, without separated sacculus; posterior margin of vinculum with shallow medial emargination, lateral humps weakly developed, basally broad vincular sclerite tapered posteriorly, with broadly sclerotised posteriomedial edges; saccus nearly V-shaped, with pointed, rod-like apical third, ratio maximum width to length approximately 0.8, posterior margin weakly arched, with small medial incision, medial part with short furcated ridge from posterior edge, lateral sclerites approximately 0.8 times length of maximum width of saccus; phallus with globular coecum, distal portion weakly S-curved, few minute teeth on dorsal edge, tapered apically.

Female genitalia. Unknown.

Megacraspedus uzunsyrtus is characterised by its light greyish brown forewings with a white costa and without further markings. It is very similar to M. monolorellus (Figure 21) which has more brownish forewings. M. similellus sp. n. (p 46). The male genitalia are particularly similar to M. similellus sp. n. (Figs 163–164) but differ by the abruptly bent gnathos hook, and the broader valva without distal constriction. From other related species they differ by several subtle characters, particularly the shape of the uncus with parallel-sided outer edges.

BIN BOLD:ACS7353 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour M. similellus sp. n. is 6.4% (p-dist).

Ukraine (Crimea).

Host plant and early stages are unknown. Adults have been collected in the middle of May in the evening before sunset on xerothermic slopes on Jurinea stoechadifolia (M. Bieb.) DC (Asteraceae) which is considered to be the larval host plant (Bidzilya and Budashkin 2015). Additional specimens were found in early July.

M. uzunsyrtus was described from four males collected in Crimea on 17.v.2014 (Bidzilya and Budashkin 2015: 222). Although these authors state that “Jurinea stoechadifolia … is undoubtedly the larval host plant” this needs to be confirmed by breeding as other host plant records of Megacraspedus species are from Poaceae. Bidzilya and Budashkin (op. cit.) stated that M. uzunsyrtus can be separated from M. monolorellus by having an antennal pecten of one hair, which should not be present in the latter species. However, examination of the lectotype of M. monolorellus shows that a pecten is present at the base of the left antenna.

Holotype ♂, “BULGARIA [Dobrich region] Tuzlata 3.6.2014 J.Junnilainen leg.” “GU 16/1449 ♂ P. Huemer” “DNA Barcode TLMF Lep 19950” (RCJJ). Paratypes. Bulgaria. 1 ♀, same data as holotype, but genitalia slide GU 16/1471 Huemer; 1 ♂, same data, but 28.v.2002, leg. J. Junnilainen; 1 ♂, same data, but 5.vi.2002, leg. J. Junnilainen (all RCJJ); 1 ♂, Dobrich reg., Kaverna, Chirakman, 21.v.2010, leg. B. S. Larsen (ZMUC). Romania. 1 ♂, Dobrogea, Jurilovca Capul, Dolosman, 16.vi.2012, leg. S. & Z. Kovacs, genitalia slide GU 16/1470 Huemer (RCKO). Turkey, 2 ♂, prov. Nigde, Bolkar Dağlari, northern slope, Maden, 1700 m, 29.vii.1997, leg. K. Larsen, genitalia slide 5326, 5345 Karsholt (ZMUC); 1 ♂, prov. Isparta, 35 km SW Askehir, Çetince, 1200 m, 14.viii.1999, leg. J. Junnilainen, genitalia slide GU 16/1462 Huemer (RCJJ); 1 ♂, 25 km E Konya, 23.v.1997, leg. K. Nupponen & J. Junnilainen; 1 ♂, 5 km N Urgüp, 25.v.1997, leg. K. Nupponen & J. Junnilainen; 1 ♂, 30 km SW Kayseri, Erciyes Dagi, 24.v.1997, leg. K. Nupponen & J. Junnilainen (all RCJJ).

Adult. Male (Figure 25). Wingspan 13–14 mm. Segment 2 of labial palpus with scale brush as long as segment 3, brown on outer surface, whitish brown on inner surface, otherwise white; segment 3 about same length as segment 2, dark brown. Antennal scape with pecten of one hair; flagellum black ringed with light brown. Head whitish grey; thorax as forewing; neck and tegula white. Forewing light grey brown from white brownish- and black-tipped scales, darkest towards apex; costa white; fringes light grey. Hindwing grey with light grey fringes.

Female (Figure 26). Wingspan 11 mm. Segment 3 of labial palpus white mottled with dark brown. Forewing broadest in middle, apical half tapering into lanceolate, blackish tip. Hindwing very short (about one-fifth length of forewing), slender with rounded apex, whitish. Otherwise similar to male.

Variation. There is some variation in the colour of the forewings: from lighter to darker brown.

Male genitalia (Figure 163–164). Uncus moderately slender, slightly more than twice as long as broad, parallel-sided, apical 1/3 weakly tapered, apex rounded; gnathos hook stout, about length of uncus, weakly and evenly curved, apex pointed; anterior margin of tegumen with moderately deep and broadly rounded emargination, long medial ridge from anterior edge to posterior third, anteriolateral edge with small peg-like sclerite; pedunculi distinct, suboval; valva moderately broad, basally slightly exceeding width of uncus, distal part narrowing, straight, apex rounded, extending to middle of uncus, sub-basally with rounded hump; saccular area covered with setae, without separated sacculus; posterior margin of vinculum with shallow medial emargination, lateral humps weakly developed, vincular sclerite broadly sub-rectangular, with broadly sclerotised posteriomedial edges; saccus nearly V-shaped, apical third evenly pointed, ratio maximum width to length approximately 0.8, posterior margin weakly arched, with small medial incision, medial part with short, furcated ridge from posterior edge, lateral sclerites approximately 0.8 times length of maximum width of saccus; phallus moderately stout, with globular coecum, distal portion weakly S-curved, tapered apically.

Female genitalia (Figure 270). Papilla analis small, apically rounded; apophysis posterior slender rod-like, about 4 mm long, with short, bifurcate posterior end, bordered by minute sclerotised field, apex weakly widened and rounded; segment VIII approximately 1.4 mm long, membranous; subgenital plate with sub-triangular subostial sclerotisation, posteriorly extended into long, pointed sub-medial sclerites, with medial flaps delimiting oblong ostium bursae, anterior margin with rod-like edge connected with apophysis anterior, straight, without anterior projection; apophysis anterior slender, rod-like, posteriorly becoming rod-like venula of segment VIII, extending to posterior margin of segment VIII; colliculum short, sclerotised; ductus bursae and corpus bursae weakly delimited, moderately short with entire length of approximately 2.5 mm; signum moderately large, suboval spiny plate.

Megacraspedus similellus sp. n. is characterised by its light greyish brown forewings with white costa and without further markings. Externally it is hardly separable from M. uzunsyrtus (Figure 24). It is also similar to M. monolorellus (Figs 21–22) which has more brownish forewings. The female is hardly separable from that of M. monolorellus. The male genitalia are very similar to M. uzunsyrtus (Figure 162) but differ by the evenly curved gnathos hook, and particularly the more slender and distally narrowing valva. From other related species they differ in several subtle characters, particularly the shape of the uncus with parallel-sided outer edges. The female genitalia differ from M. monolorellus (Figure 269) particularly by the distinctly shorter apophysis anterior, but females of several related species are unknown.

BIN BOLD:ADB8685 (n = 3). The intraspecific divergence of the barcode region is moderate with mean 0.8% and maximum divergence of 1.2%. The distance to the nearest neighbour M. uzunsyrtus is 6.4% (p-dist).

Bulgaria, Romania, Turkey.

Host plant and early stages are unknown. The adults have been collected from the middle of May to late July from sea level in the Balkans to 1700 m in Turkey.

The species name refers to the similarity to related taxa and is derived from the Latin word similis (meaning like), and the diminutive suffix –ellus. The name is a compound noun.

Holotype ♂, “Iran, [prov.] Khorassan NP [National Park] Golestan, Almeh 1770 m 2.5.2001 37°20,75'N, 56°07,02'E leg. P. Huemer” “BC TLMF Lep 03782” “P. Huemer GEL 1241 ♂” (TLMF). Paratypes. Iran. 4 ♂, same data as holotype, but BC TLMF Lep 03780, 03781 (TLMF).

Adult. Male (Figure 27). Wingspan 10–11 mm. Segment 2 of labial palpus with moderately long scale brush, brown on outer surface, whitish brown on inner surface, otherwise white; segment 3 white mottled with light grey-brown. Antennal scape with pecten of one hair; flagellum light brown, indistinctly ringed with black. Head, thorax and tegula as forewing. Forewing light greyish brown from white brownish- and black-tipped scales; base of costa blackish brown; margin of costa whitish; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. The few examined specimens show only slight variation. The margin of the forewing costa can be greyish or whitish. In one specimen there is no hair at the base of the antennal scape.

Male genitalia (Figure 165). Uncus approximately 1.5 times longer than broad, parallel-sided, apex rounded; gnathos slightly longer than uncus, moderately slender, curved, apex pointed; tegumen weakly constricted, anterior margin deeply excavated, pedunculi small, rounded; valva short, straight, of equal width, apex slightly distorted, rounded, extending to base of uncus; posterior margin of vinculum with broad lobes, anteromedial emargination broad, shallow; saccus broad at base, with long and narrow lateral projections, anterior edge with sinus-shaped projections, incised in-between, medial part with short sclerotised ridge, distal portion abruptly tapered, rod-like; phallus about length of tegumen without uncus, globular coecum, distal two-thirds weakly S-curved, tapered apically.

Female genitalia. Unknown.

Megacraspedus golestanicus sp. n. is characterised by its relatively small wingspan, and its light greyish brown forewings without markings. Externally it is hardly separable from M. junnilaineni sp. n. (Figure 23). The short medial ridge of the saccus and the abruptly tapered rod-like distal part are characteristic features of M. golestanicus sp. n.

BIN BOLD:AAU3338 (n = 3). The intraspecific divergence of the barcode region is low with mean 0.1% and maximum divergence of 0.2%. The distance to the nearest neighbour M. uzunsyrtus is 8.8% (p-dist).

Iran (Golestan).

Host plant and early stages are unknown. The habitat is dominated by steppe meadows. The adults have been collected in the last third of May at light at an altitude of 1770 m.

This species is named after its place of occurrence: the province of Golestan in north-eastern Iran. The name is a masculine adjective.

Holotype ♂, “CROATIA [Dalmatia region] Konjevrate 200 m 25.06.2006 leg. Z. Tokár” “Megacraspedus sp. det. Zdenko Tokár” “DNA Barcode TLMF Lep 16630” (RCZT). Paratypes. Croatia. 2 ♂, same data as holotype; 1 ♂, same data, but 25.vi.2003; 8 ♂, same data, but 28.vi.2003; 2 ♂, same data, but 6.vi.2005; 1 ♂, Gorne Bilišane, 6.vii.2004, leg. Z. Tokár; 1 ♂, Bilišane, 23.vi.2006, leg. Z. Tokár; 1 ♂, same data, but 15.ix.2007, genitalia slide 14/1384 Huemer (all RCZT); 2 ♂, Krk isl. Str. Krk-Vrbnik, 20.vii.1988, leg. G. Baldizzone (TLMF); 1 ♂, Krk isl., Punat-Stara Bavka, Trstenova, 18.vii.2013, leg. G. Baldizzone (RCGB); 2 ♂, Krk isl., Mt. Hiam, Branušine, 22.vi.2013, 180 m, leg. G. Baldizzone (RCGB; ZMUC).

Adult. Male (Figure 28). Wingspan 10 mm. Segment 2 of labial palpus with scale brush shorter than segment 3, brown on outer surface, whitish brown on inner surface, otherwise white; segment 3 about same length as segment 2, white. Antennal scape with pecten of a few hairs; flagellum black, indistinctly ringed with light brown. Head cream-coloured; thorax and tegula cream-coloured mottled with brownish. Forewing brown mottled with some yellow-white, especially along dorsum; a distinct white stripe along costa; fold yellowish; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. The examined specimens show no variation, but worn specimens become light greyish.

Male genitalia (Figure 166). Uncus moderately broad, 1.5 times as long as maximum basal width, sub-trapezoidal, weakly tapered to rounded apex; gnathos hook slender, straight, about one-quarter longer than uncus, with curved and pointed apex; anterior margin of tegumen with moderate, broadly rounded emargination, medially with longitudinal ridge, extending from anterior edge beyond middle of tegumen; pedunculi small, rounded, with sclerotised ridge; valva straight, slender, basally wider, apical part contorted, rounded, extending to about base of gnathos, saccular area covered with setae, with hardly separated sacculus, fused with valva; posterior margin of vinculum with shallow medial emargination, weakly rounded lateral humps, vincular sclerite suboval, posteriomedial edge strongly sclerotised; saccus moderately large, broadly V-shaped, with rod-like apical fifth, ratio maximum width to length approximately 0.9, posterior margin with broadly rounded projections, separated by V-shaped emargination, medial part of saccus with short sclerotised ridge, furcated at approximately one-third length of saccus, lateral sclerites approximately 0.8 times length of maximum width of saccus; phallus about length of tegumen, moderately stout, with bulbous coecum, distal two-thirds weakly curved, tapered apically, with rod-like ventral sclerotisation.

Female genitalia. Unknown.

Megacraspedus tokari sp. n. is characterised by cream-coloured head and its brown forewings with a white costa and without further markings. Its small size separates it from similar looking species. The male genitalia differ from other species of the M. dolosellus species group particularly by the furcated ridge of the saccus and the weakly curved phallus.

BIN BOLD:ACM1095 (n = 1). The distance to the nearest congeneric neighbour M. cuencellus is 10%, the distance to the nearest BIN in BOLD, Monochroa scutatella (Müller-Rutz, 1920), is 9.2% (p-dist).

Croatia.

Host plant and early stages are unknown. The adults have been collected from late June to the middle of September at low altitudes.

The species name (a noun in the genitive case) is dedicated to Zdenko Tokár, Slovakia, who collected most of the type series of this species and numerous other valuable specimens used for our study.

Lectotype ♂, Ypsolophus dolosellus, designated by Huemer & Karsholt (2001), “dolosella FR Wien” “Ypsolophus dolosellus Zeller Isis p 190 (1839) TYPE ♂” “Zeller Coll. Wlsm. Coll. B.M. 1910-427. ” “Lectotypus ♂ Ypsolophus dolosellus Select. K. Sattler, 1961” “B.M. Genitalia slide No. 7102 ♂“ ´Type” “LECTO-TYPE” (BMNH). Lectotype ♂, Megacraspedus incertellus, here designated, “LECTO-TYPE” “Alibotuschgebir Al K. Drenowski“ “Megacraspedus incertellus Rbl Type ♂” “LECTOTYPE ♂ Megacraspedus incertellus Rebel det. L.M. Pitkin, 1987” “Mus.Vind. Gen.Präp. 16.518 ♂” “BC TLMF Lep 06703” (NHMW). Albania. 2 ♂, Pashtrik, 7–15.viii.1918; 2 ♂, Gjalica Ljums, 17–26.vi.1918, genitalia slide Mus. Vind. 15.348 (NHMW). Austria. 1 ♀, without locality, leg. J. Mann (NHMW). Austria. 1 ♀ [form dolosellus], Wien, Prater, 1856 (NHMW); 2 ♂, Wien (ZSM); 1 ♂, Niederösterreich, Hainburg, 16.v.1909, leg. Zerny (NHMW). Bosnia and Herzegovina. 1 ♂, Trebovic, 5.vii.1898, genitalia slide Mus.Vind 15.343 ♂ (NHMW); 12 ♂, Dolovi, 10.vi.2011, leg. I. Richter; 1 ♂, same data, but 21.vi.2011 (RCIR, ZMUC); 1 ♂, Dolovi, 10.vi.2011, leg. I Richter (all NMPC). Bulgaria. 1 ♂, Jakoruda, 23.vi.2000, leg. J. Junnilainen, leg. J. Junnilainen, genitalia prep. (in glycerin); 1 ♂, Belashitsa, 2000 m, 27.vii.2013, leg. J. Junnilainen, genitalia slide 16/1465 Huemer [BOLD:ADB8789] (RCJJ); 1 ♂, 1 ♀ [form dolosellus], Rilo, 1600 m, 28.7.1902, genitalia slides Mus.Vind 16.523 ♂, Mus.Vind. 16.527 ♀; 2 ♂, Slivno, vi.1896, leg. H. Rebel, genitalia slide Mus.Vind 16.526 ♂ (all NHMW); 1 ♂, 5 km NNE Pastra, 31.vii.2013, leg. B. Å. Bengtsson; 1 ♂, Pirin mts, Oreliak, 1850–1950 m, 9.viii.2013, leg. O. Karsholt & B. Zlatkov (all ZMUC); 5 ♂, same data, but 1800–1950 m, 24.vi.2014, leg. Z. Tokár, genitalia slide GEL 1216 Huemer (RCZT, TLMF); 1 ♂, but 2000 m, 4.vii.2014, leg. J. Junnilainen, genitalia in gylcerin (RCJJ); 3 ♂, Pirin mts, Vichren-Kazana, 2000 m, 26–30.vii.1986, leg. J. Liška (NMPC); 12 ♂, Pirin mts, Popovi, Livadi, 2000 m, 22.vi.2001, leg. J. Junnilainen, genitalia slide 16/1468 Huemer; 1 ♂, 10 km E Ilindentci, 1200 m, 10.vii.2014, leg. J. Junnilainen (all RCJJ). Croatia. 3 ♂, 1 ♀ [form dolosellus], Petrinjski kras, Petrinja, 3.vi.2005, leg. Z. Tokár (RCZT); 1 ♂, Pag, 10.vi.2015, leg. J. Junnilainen [BOLD:ADB8686] (RCJJ); 6 ♂, N Velebit, Pandore, 800 m, 21.vi.2006, leg. Z. Tokár; 6 ♂, same data, but 21.vi.2006, genitalia prep. (in glycerin); 6 ♂, Biokovo mts, Vošac, 1370 m, 27.vi.2006, leg. Z. Tokár, genitalia prep. (in glycerin); 1 ♀, Malovan-Gracac, 10.vii.2004, leg. Z. Tokár (all RCZT). France. 1 ♀ [form dolosellus], Dep. Pyrénées-Orientales, Vernet, 1.vii.1895, leg. P. Chrétien; 1 ♂, 1 ♀ [form dolosellus], Dep. Paris, Seine, leg. P. Chrétien; 1 ♂, Paris, leg. P. Chrétien (all ZSM); 2 ♂, Dep. Cantal, Joursac, 4.vii.1996, leg. J. Nel, genitalia slides 5005 Nel, 5363 Nel; 1 ♂, Dep. Marne, Reims, Université, 4.vii.1997, leg. J. Nel, genitalia slide 5945 Nel (all TLMF). Greece. 1 ♂, 1 ♀ [form separatellus], Olympos, Kataphygion, 2000 m, 27–30.vi.1962, leg. J. Klimesch; 2 ♀ [form dolosellus], same data, but 2–-7.vii.1962, leg. J. Klimesch (all ZSM); 2 ♂, same data, but 2100 m, 12–18.vii.1962 leg. F. Kasy (NHMW, ZSM); 1 ♂, 1 ♀ [form separatellus], Chelmos, 2200–2300 m, 24.vi. 1958, leg. J. Klimesch; 1 ♂, same data, but 24.vi.1968, leg. J. Klimesch (all ZSM); 1 ♂, same data, but 30.v.2009, leg. J. Junnilainen [BOLD:AAG0031] (RCJJ); 4 ♂, Epirus, above Monodendri, 1300 m, 16.vi.2010, leg. P. Skou; 3 ♂, Florina, above Germanos, 2000 m, 27.vii.2013, leg. B. Skule & C. Hviid; 1 ♂, Fokida, Mt. Parnassos, road to southern ski resort, 1650 m, 10.vi.2010, leg. P. Skou; 1 ♂, same data, but 2.6 km N Kellaria Ski Center, 1600 m, 4.vi.2013; 1 ♂, Makedonia, Olympos, 700–2100 m, 21–26.v.1990, leg. O. Karsholt; 1 ♂, Makedonia, Kavala, Pangeo, 1700 m, 24.viii.1989, leg. K. Larsen; 1 ♂, Larissa, Olympos mts, below Skolio, 11 km NE Kalvia, 1850 m, 20.vii.1998, leg. B. Skule & D. Nilsson; 1 ♂, same data, but 3.vii.2004, leg. B. Skule; 1 ♀ [form dolosellus], Makedonia, 1 km NE Agios, Germanus, 1060 m, 14.vi.2013, leg. P. Skou; 7 ♂, Makedonia, 2.4 km SE Pisoderi, Vigia, 1550 m, 18.vi.2013, leg. P. Skou; 1 ♂, Makedonia, Mt. Voras, 4 km N Zervi, road to Voras Ski Resort, 1850 m, 5.vii.2016, leg. P. Skou (all ZMUC); 1 ♂, Makedonia, Kavala, Ofryno, Strymon Delta, 24.v.2009, leg. W. Schmitz; 2 ♂, Askion Oros, Vlasti, 1400 m, 7.vi.2014, leg. W. Schmitz (all RCWS); 4 ♂, Makedonia, 15 km W Kozani, Xerolimni, 21–23.v.2003, leg. J. Junnilainen; 1 ♂, same data, but 11.vi.2010, leg. J. Junnilainen; 7 ♂, Makedonia, Kozani NE, 23–24.v.2003, leg. J. Junnilainen; 26 ♂, Makedonia, Askio Vellia, 1450 m, 23.v.2003, leg. J. Junnilainen; 1 ♂, 1 ♀ [form separatellus], Makedonia, 15 km W Olympos, Leptokaria, 750 m, 27.v.2001, leg. J. Junnilainen, genitalia slides GU 16/1446 ♂ Huemer, GU 16/1467 ♀ Huemer (all RCJJ); 1 ♂, Epirus, Parga, 0 m, 1.vi.2002, leg. W. Schmitz (ZMUC); 1 ♂, Fokida, Parnassos, Itea/Desfina, 550 m, 21.v.2009, leg. W. Schmitz (RCWS).; 15 ♂, Drama, Phalakro mts, above Volas, 1700 m, 20.vii.1987, leg. M. Fibiger; 7 ♂, same data, but 15.vii.1998, leg. B. Skule & D. Nilsson (all ZMUC); 6 ♂, Peloponnes, Arkadien, Mari env. 620 m, 17.v.2009, leg. T. Mayr (TLMF, RCTM); 1 ♂, Peloponnes, Chelmos, 1600 m, 12.vi.2008, leg. J. Skyva (NMPC); 27 ♂, 1 ♀, Ipiros, Katara Pass, 15–1700 m, 24–27.v.1994, leg. O. Karsholt, genitalia slide GU 00/888 Huemer; 4 ♂, Ioánina, 8 km above Monodendri, Mt. Timfi, 1380 m, 22.vi.2004, leg. B. Skule; 1 ♂, Mt. Phalakron, above Volas, 6.vii.1986, leg. M. Fibiger, genitalia slide 6518 H. Hendriksen; 1 ♂, Makedonia/Thessalia, Olympos mts east, 900–1500 m, 17–18.v.1994, leg. O. Karsholt; 2 ♂, same data, but 700–2100 m, 21–26.v.1994; 1 ♂, Peloponnesos, Taygetos mts, 1000 m, 11.vi.1980, leg. G. Christensen; 2 ♂, same data, but 1700 m, 28–29.vi.1982, leg. B. Skule & S. Langemark; 2 ♂, 1 ♀, same data, but, above Trapenzandi, 5.vii.1984, 1800 m, leg. B. Skule, genitalia slide 5327 Karsholt; 2 ♂, Peloponnesos, Chelmos mts, above Kalavrita, 1700 m, 17–19.vi.1982; leg. B. Skule & S. Langemark; 1 ♂, Peloponnesos, Parnon Oros, 1700 m, 8.vii.1982; leg. B. Skule & S. Langemark; 2 ♂, same data, but western slope, 1000–1500 m, 3.vi.1994, leg. O. Karsholt; 1 ♂, Peloponnesos, Mt. Kyllini, above Trikala, 1650 m, 13.vi.2010, leg. P. Skou; 1 ♂, Thessaly, 3.1 km NE Metsovo, Katara Pass, 1580 m, 19.vi.2013, leg. P. Skou; 3 ♂, Epirus, 0.9 km NNE Kapesovo, 1160 m, 21.vi.2013, leg. P. Skou, genitalia slide GU 16/1441 ♂ P. Huemer (all ZMUC); 1 ♂, Crete, Mt. Ida, Südhang, Rouwawald, 1300 m, 15–31.vii.1938, leg. Dürck, genitalia slide in vial (ZSM). Hungary. 1 ♂, Szentggotthárd, 4.vi.1910, leg. Schmidt; 5 ♂, Leanyfalu, 5.–10.vii.1997, leg. B. S. Larsen; 2 ♂, 2 km N Börgönd, 24–25.vi.1998, leg. B. S. Larsen (all ZMUC); 1 ♂, Budafok, 20.vi.1916, leg. Uhrik; 1 ♂, 2 km SW Agárd, 20.vi.1998, leg. B. S. Larsen; 2 ♂, 3 km E Öksü, 29.vi.1998, leg. B. S. Larsen (all ZMUC); 1 ♂, Hortobágy, 18.v.1912; 1 ♀ [form dolosellus], same data, but 2.vi.1912 (all NHMW); 1 ♂, Csákberény, Bucka-hegy, 21.v.2005, leg. Z. Tokár; 1 ♂, Bélmegyer env., salt marshes, 9.v.2014, leg. Z. Tokár [BOLD:AAG0031]. Italy. 1 ♂, prov. Cuneo, Briga Alta, Monte Tanarello, 2000 m, 20.vii.1993, leg. G. Delmastro (TLMF); 2 ♂, prov. L´Aquila, NP Gran Sasso, Campo Imperatore, E Observatorio, 2010 m, 13.vii.2010, leg. P. Huemer, genitalia slide GEL 1198, 1200 Huemer, BC TLMF Lep 01493 (TLMF); 1 ♂, prov. Rieti, Monte Terminillo, 1730–1780 m, 12.vii.2010, leg. P. Huemer, genitalia slide GEL 1239 Huemer, BC TLMF Lep 04316 (TLMF); 2 ♂, prov. Roma, Monte Lepini, 4 km SE Carpineto Romano, 700 m, 20.v.2014, leg. J. Tabell, genitalia slide 16/1443 Huemer (ZMUC). Macedonia. 1 ♂, Treska Schlucht, 4–10.v.1963, leg. J. Klimesch; 1 ♂, Ohrid, Petrina Planina, 17–26.vi.1959, leg. J. Klimesch; 1 ♂, Shar Planina, Ljuboten, 1500–2000 m, 22.vi.1955, leg. J. Klimesch (all ZSM); 5 ♂, Galicica Planina, Stara Planina, Monte Magaro N, 2240 m, 26.vi.2009, leg. G. Tarmann; 9 ♂, NP Mavrovo, Korab, eastern ridge, 2325–2400 m, 28.vii.–1.viii.2011, leg. P. Huemer & G. Tarmann; 42 ♂, 2 ♀ [form separatellus], NP Mavrovo, Korab, Korabska jezero, Kobilino pole, 2080–2180 m, 28.vii.–1.viii.2011, leg. P. Huemer & G. Tarmann; 7 ♂, 1 ♀ [form separatellus], same data, but leg. P. Huemer & G. Tarmann [BOLD:ABA2915] (all TLMF); 5 ♂, same data, but 20–21.vii.2015, 2300 m, leg. I. Richter (RCIR); 1 ♂, Galicica, 12–13.vi. 2014, leg. I. Richter; 1 ♂, same data, but NP Galicica, 12.vi.2016; 1 ♂, same data, but NP Galicica, 15–16.vi.2017; 2 ♂, Bjelovodica, Mermerno jezero, 13–14.vi.2017, leg. I. Richter (all NMPC); 2 ♂, Ohrid, Galicica, 1400 m, 28.vi.2014, leg. J. Junnilainen [BOLD:ADB8790] (RCJJ); 1 ♂, NP Pelister, Golemo Jezero, 1.viii.2015, leg. I. Richter [BOLD:ABA2915] (RCIR); 2 ♂, Galicica NP, Galicica Saddle, 1380 m, 15–16.vi.2013, leg. P. Skou (ZMUC); 1 ♂, Prilep, Treskavac, 6.vi.2014, leg. I. Richter; 1 ♂, same data, but 10.vi.2016; 1 ♂, NP Mavrovo, 10 km N. Nalicnik, 20.vi.2016, leg. I. Richter; 4 ♂, Pivska pl. Pešče, 21.vi.2012, leg. I Richter (all NMPC). Montenegro. 1 ♂, Durmitor NP, 8.7 km NW Zabljak, Mali Stouc, 1880 m, 20–21.vii.2014, leg. C. Hviid & O. Karsholt [BOLD:ACS7352]; 1 ♂, Durmitor NP, 9 km E Zabljak, 1280 m, 24.vi.2013, leg. C. Hviid & O. Karsholt [BOLD:ACS7982]; 15 ♂, Durmitor National Park, 4 km S Žabljak, Virak, 1550 m, 22–28.vi.2013, leg. C. Hviid, O. Karsholt & K. Larsen; 1 ♂, Durmitor National Park, 6.8 km NW Žabljak, 1740 m, 23.vii.2014, leg. C. Hviid & O. Karsholt; 3 ♂, Durmitor National Park, 4 km S Žabljak, Virak, 1550 m, 19–24.vii.2014, leg. C. Hviid & O. Karsholt; 4 ♂, Durmitor National Park, 13 km SW Žabljak, Sedlo Pass, 1900 m, 19–24.vii.2014, leg. C. Hviid & O. Karsholt; 1 ♂, Durmitor National Park, 8 km S Žabljak, Virak, 1600 m, 19–24.vii.2014, leg. C. Hviid & O. Karsholt; 1 ♂, Durmitor National Park, 8.7 km NW Žabljak, Mali Stouc, 1880 m, 20–21.vii.2014, leg. C. Hviid & O. Karsholt (all ZMUC). Nepal. 1 ♂, Binap-Nager, 6.x.1996, leg. V. Cikolovecs, genitalia slide 17/1494 Huemer (RCZT). Romania. 1 ♂, Carpatii orientali, Mni. Baraolt, Ariusd, 600 m, 2.vi.1996, leg. S. & Z. Kovács; 1 ♂, Carpatii orientali, Muntii Ciucului, Racu, Cseretetö, 650 m, 13.vii.2004, leg. S. & Z. Kovács; 1 ♂, same data, but 14.vii.2005 (all RCKO); 6 ♂, 1 ♀ [form dolosellus], Rimetea area, 600 m, 29–31.v.2009, leg. O. Karsholt (ZMUC); 1 ♀ [form dolosellus], same data, leg. O. Karsholt [BOLD:AAG0031]; 1 ♂, same data, but 30.v.2009, leg. J. Junnilainen [BOLD:AAG0031] (RCJJ). Russia. 1 ♂, N Caucasus, Kabardino-Balkaria, river Gundelen, 1400 m, 10–20.vii.2012, leg. L. Srnka, genitalia prep. (in glycerin) (RCZT); 6 ♂, 3 ♀ [form separatellus], S Ural, Cheliabinsk distr., Arkaim reserve, near Akurskii vill., 16.vi.1996, leg. K. Nupponen et al.; 1 ♂, S Ural, Orenburg distr., 20 km S Pokrovka vill., Schibendy valley, 3–7.vi.1998, leg. J. Junnilainen (all RCJJ); 1 ♀ [form separatellus], same data, but 22.vi.1999, leg. T. & K. Nupponnen; 2 ♂, same data, but 22.vi.1999 (all ZMUC); 1 ♂, S-Ural, Moskovo env., 500 m, 18.vi.2009, leg. J. Šumpich; 2 ♂, same data, but 575 m, 19.vi.2009, leg. J. Šumpich; 2 ♂, same data, but 15–18.vii.2011, leg. J. Šumpich; 4 ♂, S-Ural, Moskovo, village, 6–7.vii.2013, leg. L. Srnka (all NNMP); 1 ♂, S Ural, Cheliabinsk distr., Kizilskoye village, 1.vii.2017, leg. H. Roweck & N. Savenkov (ECKU); 7 ♂, SW Altai, Katun valley, 10 km W Katanda, 1200 m, 22.vi.–8.vii.1983, leg. K. Mikkola, H. Hippa & J. Jalava, genitalia slide Hendriksen 2979 (MZH); 2 ♂, Altai mts, 10 km SE Aktash, 1500 m, 13.vii.1997, leg. C. Gielis (RCHW); 4 ♂, 3 ♀, Altai mts, Kuraisky hrebet, 2300 m, 10.vii.2001, leg. K. Nupponen (ZMUC); 2 ♂, Rep. Altai, Aktash vill., 1400 m, 21.vi.2015, lg. J. Šumpich; 2 ♂, Rep. Altai, 3 km SE Aktash, 1300 m, 17.vi.2012, leg. B. Schacht, genitalia slide GEL 1246 Huemer (all TLMF). Slovakia, 1 ♂, Plešivec, 12.vi.1987, leg. B. Å Bengtsson (ZMUC); 2 ♂, Komárno, 1.vi.1991, leg. G. Pastorális; 2 ♂, 3 ♀ [form dolosellus], Šal’a, Váh, 1.v.2007, leg. S. Tokár (NMPC, ZMUC); 1 ♂, Plášt’ovce, 2.vi.1986, leg. J. Patočka (ZMUC); 2 ♀ [form dolosellus], Šal’a, okolie Váhu, 2.v.2009, leg. Z. Tokár; 1 ♂, same data, leg. Z. Tokár [BOLD:ACS9604]; 1 ♀ [form dolosellus], same data, leg. Z. Tokár [BOLD:AAG0031]; 1 ♂, same data, but 11.v.2009, leg. Z. Tokár (all RCZT); 1 ♂, Slovenský Kras, Zádiel, 27.v.2001 (ZMUC). Slovenia. 3 ♂, Karst, Presnica, 15.v.2002, leg. H. Deutsch, genitalia slide GEL 1190 Huemer (TLMF); 3 ♂, Senezece, 500 m, 30.v.2008, leg. J. Junnilainen (RCJJ). Turkey. 3 ♂, Prov. Mersin, Çamliyayla, 5.vi.1984, leg. G. F. Curletti, genitalia slide Hendriksen 3339 (ZMUC). Ukraine. 1 ♂, Odessa, Borodino, Tarutinskaya steppe, 7.vi.2011, leg. A. Bidzilya, genitalia prep. (in glycerin) [BOLD:ADA0140] (RCAB); 1 ♂, Crimea, Karadag, 5.vii.1987, leg. R. Puplesis (ZMUC); 1 ♂, same data but 3.vi.2004, leg. Yu. Budashkin, genitalia slide GU 16/1464 P. Huemer; 1 ♂, same data, but 6.vi.2004; 1 ♂, same data, but 10.vi.2004 (all RCAB). Locality illegible. 1 ♀ [form separatellus], 22.vi.1902 (ZMUC).

Adult. Male (Figs 29, 31, 33–38). Wingspan 10–15 mm. Segment 2 of labial palpus with scale brush about as long as segment 3, brown on outer and inner surface, white on lower and upper surface; segment 3 white. Antennal scape without pecten, flagellum ringed blackish and dirty white to nearly unicolorous black. Head, thorax and tegula cream coloured, mottled with brown. Forewing light brown (lightest towards dorsum) to light clay-brown; veins and costa clear white to finely dirty white; fringes grey. Hindwing grey with light grey fringes.

Female (Figs 30, 32). Wingspan 11–12 mm. Forewing broadest in middle, apical part pointed; fringes light grey. Hindwing lanceolate, about two-thirds length of forewing, whitish, with a few dark scales in apex; only scattered whitish fringes (form dolosellus) or hindwing slender with pointed tip, about one-fifth the length of forewing, whitish (form separatellus), hindwing about one-third length of forewings in several females from Altai mts Otherwise similar to male.

Variation. There is some variation in the forewing colour (more or less yellowish to greyish or brownish), and in the distinctness of the whitish veins. Worn specimens become lighter. Specimens from Turkey are larger (wingspan 14–18 mm) than European ones, but otherwise similar.

Male genitalia (Figs 167–172). Uncus moderately broad, 1.5 times as long as maximum basal width, sub-trapezoidal, weakly tapered to rounded apex; gnathos hook slender, straight, about length of uncus, with hardly curved and pointed apex; anterior margin of tegumen with moderate, broadly rounded emargination, sclerotised ridges extending from anterior margin and merged in medial zone; pedunculi small, rounded, with scleotized ridge; valva straight, slender, basally wider, apical part contorted, rounded, extending to about base of gnathos, saccular area covered with setae, with hardly separated sacculus, fused with valva; posterior margin of vinculum with curved medial emargination, broadly rounded lateral humps, vincular sclerite oblong, posterior edge strongly sclerotised; saccus moderately large, broadly V-shaped, with broad and short rod-like apex, ratio maximum width to length about 0.85, posterior margin with broadly rounded projections, separated by V-shaped emargination, medial smooth with at most very short sclerotised ridge, lateral sclerites broad, approximately 0.85 times length of maximum width of saccus; phallus about length of tegumen, stout, with bulbous coecum, distal two-thirds strongly S-curved, tapered apically, with rod-like ventral sclerotisation.

Female genitalia (Figs 271–274). Papilla analis small, apically rounded; apophysis posterior slender rod-like, approximately2.2–2.6 mm long, with short, weakly bifurcate posterior end; segment VIII 0.9–1.0 mm long, membranous; subgenital plate with sub-triangular subostial sclerotisation, posteriorly extended into shortly pointed sclerites, large medial flaps delimiting ostium bursae, anterior margin with rod-like edge connected with apophysis anterior, medially with broadly sinusoid projection; apophysis anterior 1.4–1.8 mm, slender, rod-like, posteriorly becoming rod-like venula of segment VIII, extending to posterior margin of segment VIII; ductus bursae short, moderately broad, about same length as corpus bursae; corpus bursae oblong, weakly delimited from ductus bursae, entire length of ductus and corpus bursae approximately 2.5 mm; signum small, variably shaped spiny plate.

Males are variable from light brown, with clear white veins and costa to light clay-brown and with veins and costa finely dirty white, ocassionally white without distinct markings. Females of form dolosellus have slender lanceolate hindwings, whereas females of form separatellus have very short ellipsoid hindwings. In a series of 5 females collected at the same place and date in Russia (Altai mts) 4 females have lanceolate hindwings but shorter than in the European form dolosellus, whereas a single female comes close to form separatellus. The male genitalia are very similar to the closely related M. tokari sp. n. (Figure 166) but can be separated in particular by the distinctly curved phallus. The female genitalia are somewhat similar to those of M. lanceolellus (Figs 267–268) but differ by the much shorter apophysis posterior and the shorter sinusoid anteriomedial projection of segment VIII. From the more closely related M. sumpichi sp. n. (Figure 279) they differ in the much longer apophysis anterior and the large medial flaps delimiting the ostium bursae. However, in the absence of females in other closely related species such as M. tokari sp. n. the diagnostic value of these characters for species delimitation remains uncertain.

The extraordinary DNA barcode divergence is reflected by 23 BINs! The intraspecific divergence of the barcode region in this species is mean 7.5% and maximum 13.8%, largely reflecting a geographic pattern. However, on several occasions large intrapopulational divergence was detected, e.g., in specimens from Italy (Gran Sasso), Bulgaria (Pirin Mts Blagoewgrad), Montenegro (Durmitor), and Slovakia with two sympatric clusters in each region. Individual variation of specimens collected at the same time in the same microhabitat may be high in these cases as e.g., in clusters dolo03 and dolo04 with two specimens diverging 6.9% in DNA barcode.

The following 23 clusters are defined (based on sequenced material):

BIN dolo01 (Greece: Ioannina): BOLD:ACS7817 (n = 1).

BIN dolo02 (Bulgaria: Alibotusch, Pirin mts): BOLD:ACA9065 (n = 3).

BIN dolo03 (Italy: l`Aquila): BOLD:AA03318 (n = 1).

BIN dolo04 (Italy: l`Aquila): BOLD:AA03319 (n = 1).

BIN dolo05 (Italy: Rieti): BOLD:AAX3311 (n = 1).

BIN dolo06 (Italy: Lazio): BOLD:ACZ7902 (n = 1).

BIN dolo07 (Greece: Peloponnes): BOLD:ACQ6924 (n = 2).

BIN dolo08 (Bulgaria: Kyustendil): BOLD:ACZ9025 (n = 1).

BIN dolo09 (Bulgaria: Pirin mts): BOLD:ACR2396 (n = 5).

BIN dolo10 (Bulgaria: Blagoewgrad): BOLD:ADB8684 (n = 1).

BIN dolo11 (Montenegro: Durmitor): BOLD:ACS7982 (n = 1)

BIN dolo12 (Macedonia: Galicica): BOLD:ADB8790 (n = 2).

BIN dolo13 (Macedonia: Korab, Pelister): BIN BOLD:ABA2915 (n = 10).

BIN dolo14 (Montenegro: Durmitor): BIN BOLD:ACS7352 (n = 1).

BIN dolo15 (Croatia: Pag): BIN BOLD:ADB8686 (n = 1).

BIN dolo16 (Ukraine: Odessa): BOLD:ADA0140 (n = 1).

BIN dolo17 (Bulgaria: Blagoewgrad): BOLD:ADB8789 (n = 1).

BIN dolo18 (Slovenia): BOLD:AAV7561 (n = 3).

BIN dolo19 (Russia: Orenburg obl.): BOLD:ACZ3281 (n = 3).

BIN dolo20 (Russia: Altai mts): BOLD:ACB3319 (n = 1).

BIN dolo21 (Russia: Altai mts): BOLD:ACZ3530 (n = 4).

BIN dolo22 (Hungary, Romania, Slovakia): BOLD:AAG0031 (n = 4).

BIN dolo23 (Austria, Slovakia): BOLD:ACS9604 (n = 3).

The minimum distance to the nearest neighbour M. lanceolellus is 9.9% (p-dist).

Widely distributed from western Europe (northwards to ca. 49°N latitude) to Central Asia (Altai Mts). A specimen from the Himalaya Mountains needs verification. According to Mariani (1943: 174) also in Sicily. Absent from the Iberian Peninsula and the northern part of Central Europe as well as northern Europe.

The larva was described in great detail (570 words, but no figures!) by Joannis (1923: 156–158). It lives from March to May underground in the rhizome of a wild grass (“Graminées sauvages”) (Joannis op. cit., Trouvelot 1923: 158–160). Although the larva makes long galleries within a rhizome, the plant does not seem to suffer from its presence because the roots beyond the attacked point are sufficient to ensure its nourishment. If wheat (Triticum aestivum L.) is sown in an abandoned field where M. dolosellus occurs, the larvae will attack the young wheat plants. The young larva occupies a similar situation, attacking one of the buried internodes of the roots, but this is serious because it causes the complete death of the affected plant (Trouvelot op. cit.) since wheat does not have rhizomes. The pupation takes place at the end of May, and Trouvelot obtained the first adults on 10^th June; they do not fly, but stay hidden or run at ground level (Trouvelot, op. cit.). Trouvelot did not state in which species of wild grass he found the larvae of M. dolosellus. According to Joannis (1923: 156) it was probably Elymus repens (L.) Gould (“chiendent?”). Lhomme (1946: 539) referred to “chiendent” as Cynodon dactylon Rich. [recte (L.) Pers.].

Chrétien (1924: 66–67) bred M. dolosellus from eggs. He placed some females in a container where he had planted Poa annua L., Lolium perenne L., and Trifolium repens L. The females placed their eggs in the leaf sheaths of the grasses or between stipules of the leaves and the stems of T. repens. The larvae hatched within 10–12 days, and the young larva descends to the base of the stems of the grasses. It bores into the stem and occupies a small space, depositing its frass inside the stem. Chrétien (op. cit.) did not explicitly write that larvae of M. dolosellus feed within stems or roots of Trifolium, and that plant is unlikely as a host for a Megacraspedus species, but he also recorded that he bred M. dolosellus from roots of Poa trivialis L.

The adults have been collected from early May to late August at altitudes from lowland localities to ca. 2400 m. The single record from Nepal dates from October. However, the identity of this specimen remains doubtful.

Ypsolophus (Megacraspedus) dolosellus was described from four males collected in June and July in the surroundings of Vienna, Austria (Zeller 1839: 190). Later it was redescribed in detail and both sexes figured in colour by Fischer von Röslerstamm (1843: 301, pl. 99, fig. 3).

Ypsolophus separatellus was described from an unstated number of males and two females collected in the surroundings of Vienna (Austria) by J. Mann (Fischer von Röslerstamm 1843). The correct identification of males of M. dolosellus and M. separatellus was always considered to be an extremely difficult task as both taxa were considered as hardly separable in the male sex (Kasy 1987: 13). In contrast the two female morphotypes with longer lanceolate (form dolosellus) or short and pointed (form separatellus) hindwings were widely accepted as diagnostic for species identification by Kasy (op. cit.) and other authors. However, additional females from Russia (Altai mts) show intermediate characters with a tendency to individual infrasubspecific variation. We therefore cross-checked genetic divergence in both female morphotypes. Sequences of three morphologically identified females of M. dolosellus exhibit a considerable intraspecific divergence of mean 1.2% and maximum 1.9% with a barcode gap of 4.9% to M. separatellus. The intraspecific barcode divergence in two sequenced females of M. separatellus is even more pronounced at 2.3%. Alleged differences in forewing colour and pattern (light brown, with clear white veins and costa in M. dolosellus, light clay-brown and with veins and costa finely dirty white in M. separatellus) similarly underly infrasubspecific individual and geographic variation. Joannis (1923: 156) had already observed, based on seven specimens (apparently all males) bred from larvae, that fresh specimens were referable to M. dolosellus, whereas a worn specimen looked more like M. separatellus. He moreover noted that the colour of living specimens differed from dead and set specimens: J’ajoute que, après étalage, les lignes blanches étaient moins visibles et la couleur semblait plus foncée. De plus l’aspect luisant que signalent les descriptions pour les ailes était insensible chez l’animal vivant, très sensible au contraire chez l’insecte étalé. [I would add that, after mounting, the white lines were less visible and the colour looked darker. Moreover, the glossy aspect of the descriptions of the wings was indiscernible in the living animal, on the contrary very discernible in the spread insect”] (Joannis op. cit.). Finally, both the male and female genitalia of M. dolosellus are inseparable from those of M. separatellus, and alleged specific differences figured by Elsner et al. (1999: pl. 2, figs 7–8) proved infrasubspecific. We therefore consider M. separatellus to be a junior synonym of M. dolosellus (syn. n.).

Megacraspedus incertellus was described from an unstated number of males collected by AK Drenowski in the Alibotusch Mountains, S Bulgaria from 22–23.vii.1929 (Rebel 1930). A lectotype is here designated in order to fix the identity of the species and conserve stability of nomenclature. Specimens from South Bulgaria and North Greece are highly variable in DNA barcode with at least five strongly divergent clusters, either separated geographically or occurring sympatrically, e.g., in Pirin Mts. They differ from nominotypical M. dolosellus in the darker antennae, and the light grey-brown forewings with a white costa and reduced markings. However, such specimens are inseparable from nominotypical M. dolosellus in the male genitalia (female unknown). We therefore consider M. incertellus as a local form at most and a synonym of M. dolosellus (syn. n.), the latter of which shows an exceptional genetic and to some extent morphological variation. Barcode data clearly support several putative taxa, but the morphology is less straightforward. Phenotypic appearance is only partially congruent with geography and it is frequently impossible to reliably assign specimens to a barcode cluster. Male genitalia morphology is uniform and alleged subtle diagnostic characters underlie intraspecific variation. Considering the observed intrapopulational barcode variation of nearly 8% we conclude that M. dolosellus is as a widespread species, ranging from Eastern Europe to Central Asia. The exceptional DNA barcode divergence may be explained by weak dispersal ability, particularly of the strongly brachypterous females, leading to several genetically isolated populations. However, it seems unlikely that this is the sole cause for the observed intraspecific variation (see Discussion).

Holotype ♂, [France] “NEL Jacques [leg.] [Dep. Vaucluse, Montange de] Lure. Ou. 1700 m. 12.VI.1999” “P. Huemer GEL 1218 ♂” (TLMF). Paratypes. Italy. 1 ♂, prov. Torino, Valsusa, Mompantero, Monte Rocciamelone, 2200 m, 3.vii.1993, leg. G. Bassi, genitalia prep. (in glycerin) (ZMUC); 1 ♂, same data, but loc. Riposa, 2200 m, 16.vii.1993, leg. G. B. Delmastro (TLMF).

Adult. Male (Figure 39). Wingspan 11 mm. Segment 2 of labial palpus with scale brush shorter than segment 3, blackish brown on outer surface, white mottled with brown on inner surface, white on lower and upper surface; segment 3 longer than segment 2, cream-white with darker tip. Antennal scape with a single fine pecten; flagellum blackish brown ringed with grey. Head, thorax and tegula cream-white. Forewing cream coloured mottled with light brown and some black-tipped scales; base of costa dark grey; indistinct black dots in fold and at 3/5 in middle of wing and at end of cell; some black-tipped scales along termen; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. Unknown.

Male genitalia (Figure 173). Uncus moderately small, sub-rectangular, apical corners rounded, apical edge nearly straight; gnathos hook moderately slender, straight, apically pointed, 1.3 times length of uncus; anterior margin of tegumen with suboval emargination, sclerotised ridges from anterior edge converged in medial part of tegumen; pedunculi small, suboval, with irregular ridge; valva about width of uncus, stout, extending to about base of uncus, digitate distal part, apex rounded; saccular area densely covered with setae, with longitudinal ridge, without separated sacculus; posterior margin of vinculum distinctly emarginated, with shallow lateral humps, vincular sclerite elongated sub-ovate, with sclerotised posterior edge; saccus broadly V-shaped, ratio maximum width to length approximately 0.85, posterior margin with distinct projections, separated by broad incision, medial part with long sclerotised ridge from posterior margin to apical third, lateral sclerites 0.7 times length of maximum width of saccus; phallus weakly curved at about one-quarter, with weakly inflated coecum, 1.5 times wider than distal part, distal part 3 times length of coecum, sclerotised dorsal ridge, ventrally wrinkled, apex broadly rounded; ductus ejaculatorius with linear internal sclerotisation.

Female genitalia. Unknown.

Megacraspedus neli sp. n. is a rather uncharacteristic species of Megacraspedus, which may be recognized by its small size with a single pecten on the antennal scape. The holotype is somewhat worn and specimens in better condition may reveal additional diagnostic characters. It resembles M. pentheres (Figure 130), but that species has more distinct black dots in the forewing.

The male genitalia are similar overall to M. faunierensis sp. n. (Figure 174) but differ in several characters such as the larger uncus, the more slender valva, and in particular the much broader and shorter saccus.

Not available, barcoding failed.

France (Dep. Vaucluse), Italy (prov. Torino).

Host plant and early stages are unknown. The species was collected from the middle of June to mid-July at altitudes from ca. 1700 to 2200 m.

The species name (a noun in the genitive case) is dedicated to Jacques Nel, France, who collected the holotype of this species and numerous other Megacraspedus specimens used for our study.

The holotype was collected in the western part of Montagne de Lure, at the edge of the road at 1700 m altitude, between “the Refuge the Lure” and the “Signal de Lure” (J Nel in litt.).

Holotype ♂, “Italien, Prov. Cuneo Alpi Cozie, Demonte NW Colle Valcavera NE, 2420 m 7°6'23"E, 44°23'04"N 2.8.2010, leg. Huemer TLMF 2011-010” (TLMF). Paratypes. Italy. 26 ♂, 9 ♀, same data as holotype, but genitalia slides GEL 1219 ♂ Huemer, GEL 1235 ♀ Huemer (TLMF, ZMUC); 12 ♂, Prov. Cuneo, Demonte NW, Colle Fauniera Umgebung, 2480–2500 m, 3.viii.2008, leg. P. Huemer; 3 ♂, same data, but Colle Valcavera Umgebung, 2400–2500 m, 5.viii.2008, leg. P. Huemer; 1 ♂, same data, but Colle Valcavera NE, 2420 m, 4.viii.2008, leg. P. Huemer; 2 ♂, same data, but 23.vii.2009, leg. P. Huemer; 21 ♂, 3 ♀, same data, but 28.vii.2009, leg. P. Huemer (all TLMF); 11 ♂, 1 ♀, same data, but 2420 m, 27.vii.2009, leg. T. Mayr; 1 ♂, same data, but 17.vii.2012, leg. T. Mayr; 2 ♂, same data, but 18.vii.2012, leg. T. Mayr; 1 ♀, same data, but 16.viii.2013, leg. T. Mayr; 10 ♂, same data, but 17.viii.2013, leg. T. Mayr; 4 ♂, same data, but 2450 m, 22.vii.2018, leg. T. Mayr; 1 ♂, Prov. Cuneo, Gias Valcavera, 2050 m, 23.vii.2009, leg. T. Mayr; 14 ♂, 1 ♀, same data, but 22.vii.2018, leg. T. Mayr (all RCTM); 2 ♂, Prov. Cuneo, Valdieri N, RN Juniperus phoenicea, 900–1000 m, 29.vi.2008. leg. P. Huemer; 1 ♂, same data, but 8.vi.2009 (all TLMF).

Adult. Male (Figure 40). Wingspan 13–14 mm. Segment 2 of labial palpus with moderately long scale brush, brown on outer surface, white mottled with brown on inner surface, white on lower and upper surface; segment 3 cream-white. Antennal scape without pecten; flagellum dark brown ringed with white. Head, thorax and tegula cream-white. Forewing cream coloured mottled with some brown-tipped scales, especially in apical part; base of costa dark grey; a black dot in fold at 2/5 and two black dots at 3/5 in middle of wing and at end of cell; some black-tipped scales along termen; fringes light grey. Hindwing light grey with concolorous fringes. Female (Figure 41). Wingspan 11 mm. Forewing with apical part pointed, light yellowish brown, darker towards apex, fringes whitish. Hindwing lanceolate, whitish grey. Otherwise similar to male.

Variation. The amount of black scales on the forewings is variable. Rarely the black dot before the apex is absent. There is a slight variation in the colour of the forewings; worn specimens look more greyish.

Male genitalia (Figure 174). Uncus small, nearly sub-rectangular, basally weakly widened, apical corners rounded, apical edge straight with weak medial emargination; gnathos hook moderately slender, straight, apically pointed, approximately 1.2 times length of uncus; anterior margin of tegumen with suboval emargination, sclerotised ridges from anterior edge converge in medial part of tegumen; pedunculi small, suboval, with small transverse ridge; valva slightly broader than uncus, stout, extending slightly beyond base of uncus, digitate distal part, apex broadly rounded; saccular area densely covered with setae, with longitudinal ridge, without separated sacculus; posterior margin of vinculum weakly emarginated, without distinct lateral humps, vincular sclerite elongated sub-ovate, with sclerotised posterior edge; saccus prominent, slightly longer than valva, slender V-shaped, ratio maximum width to length 0.6, posterior margin with distinct projections, separated by broad incision, medial part with long sclerotised ridge from posterior margin to apex, lateral sclerites approximately 0.8 times length of maximum width of saccus; phallus weakly curved at about one-quarter, with weakly inflated coecum, 1.5 times wider than distal part, distal part 3 times length of coecum, sclerotised dorsal ridge, ventrally wrinkled, apex broadly rounded; ductus ejaculatorius with linear internal sclerotisation.

Female genitalia (Figure 275). Papilla analis small, apically rounded; apophysis posterior slender rod-like, 2.5 mm long, with short, bifurcate posterior end, bordered by minute sclerotised field; segment VIII 0.8 mm long, membranous; subgenital plate with sub-triangular subostial sclerotisation, posteriorly extended into long and pointed sclerites delimiting small ostium bursae, anterior margin with rod-like edge connected with apophysis anterior, medially with moderately short nearly tubular projection; apophysis anterior slender, rod-like, about half length of segment VIII, posteriorly becoming rod-like venula of segment VIII, extending to posterior margin of segment VIII; ductus bursae short, moderately broad; corpus bursae, moderately short and broad, weakly delimited from ductus bursae, entire length of ductus and corpus bursae nearly 2 mm; signum small, rounded spiny plate.

Megacraspedus faunierensis sp. n. is characterised by its cream coloured forewings with three distinct black spots. It resembles M. tristictus (Figure 48) which has darker forewings, but M. faunierensis sp. n. has the head white and the antennae darker. It is also similar to M. pentheres (Figure 130), which shares the yellowish brown colour of the forewings with M. tristictus, but has the dot at the end of the cell largely reduced, and there are several small dark spots in the terminal area. The male genitalia differ from the probably closest species M. neli sp. n. (Figure 173) particularly in the smaller uncus, broader valva, and the shape of the saccus. They somewhat resemble those of M. leuca (Figure 237) in this character, although the saccus is V-shaped and not U-shaped, but differ in many other structures. The female genitalia are similar to several species such as M. sumpichi sp. n. (Figure 279) and differ in particular by the tubular rather than sinusoid projection of the anterior edge of segment VIII. However, this character is unknown for related species and maybe of limited diagnostic value in the delimitation of species.

BIN BOLD:AAJ3164 (n = 3). The intraspecific divergence of the barcode region is considerable with mean 1% and maximum divergence of 1.6%. However, this divergence is exclusively based on a divergent specimen from a lowland locality (Prov. Cuneo, Valdieri N, RN Juniperus phoenicea, 900–1000 m, 29.vi.2008. leg. P. Huemer). The distance to the nearest congeneric neighbour M. sumpichi sp. n. is 11.9%, the distance to the nearest BIN in BOLD, an unnamed Limacodidae, is 9.1% (p-dist).

North-western Italy.

Host plant and early stages are unknown. The adults have been collected from early June to mid-August, depending on the altitude. In alpine habitats they were found flying freely in the early morning about sunrise. At this time several specimens could be collected in copula, mainly sitting on low vegetation, particularly on blades of grass. The altitudes range from ca. 900 to 2500 m.

This species is named after the place of occurrence of part of the type series: the Colle di Fauniera pass in northern Italy. The name is a noun in apposition.

Holotype ♂, “Hispania, [prov. Avila] 19.7.1980 Sierra de Gredos Navacepeda, 1500 m M. u. E. Arenberger” “GU 16/1416 ♂ P. Huemer“ (RCEA). Paratypes. Spain. 1 ♂, same data as holotype (TLMF); 1 ♂, prov. Avila, Sierra de Villafranca, 1 km W La Herguijulea, 1650 m, 20.vii.2003, leg. B. Skule, genitalia slide 5314 Karsholt (ZMUC); 1 ♂, same data, but 15.vi.2012, leg. T. Nupponen, genitalia slide 17/1489 Huemer (RCKN).

Adult. Male (Figure 42). Wingspan 15 mm. Segment 2 of labial palpus with scale brush about as long as segment 3, dark brown on outer surface, whitish brown on inner surface, white on lower and upper surface; segment 3 white mottled with some black. Antennal scape without pecten; flagellum blackish brown, indistinctly ringed with dirty white. Head, thorax and tegula cream coloured, mottled with brown. Forewing yellow-brown, mottled with grey towards costa; costa basally grey-brown, otherwise pure white; veins indistinctly white; fringes light grey. Hindwing dark grey with grey fringes.

Female. Unknown.

Variation. Unknown.

Male genitalia (Figure 175–176). Uncus moderately slender, approximately 2.5 times as long as maximum basal width, with parallel outer margin, distal part with weakly rounded apex; gnathos hook stout, about length of uncus, bent near base and slightly curved at apex, distal part club-shaped, apically pointed; anterior margin of tegumen with broadly rounded emargination; pedunculi moderately small, with additional sclerotisation; valva moderately slender, basal part distinctly wider than distal part, nearly straight with weakly curved and inflated apical part, extending to about base of uncus; sacculus long, slender digitate; posterior margin of vinculum with distinct medial emargination, broadly rounded lateral humps, vincular sclerites slender, with sclerotised proximal edge; saccus irregularly V-shaped, medially bulged, apical fifth abruptly tapered, ratio maximum width to length approximately 0.65, posterior margin with sub-triangular mediolateral projections, separated by deep incision, media part with long ridge, forked in middle of saccus, lateral sclerites stout, about length of maximum width of saccus; phallus with bulbous coecum, distal three-quarters slender, basally curved, a short sclerotised ridge with a single tooth subapically on ventral surface.

Female genitalia. Unknown.

Megacraspedus gredosensis sp. n. is characterised by its yellow-brown forewings with a clear white costa and no black dots, and the dark grey hindwigs. It may resemble species of the M. dolosellus-complex, but these have more pure whitish veins, but not a pure white costa and lighter hindwings. The male genitalia are somewhat similar to M. glaberipalpus sp. n. (Figs 226–227) but are unmistakable due to the characteristic shape of the phallus.

BIN BOLD:ADI8272 (n = 1). The distance to the nearest neighbour M. bidentatus sp. n. is 10.01% (p-dist).

Central Spain (prov. Avila).

Host plant and early stages are unknown. The few adults known to date have been collected in the middle of July at altitudes between 1500 and 1650 m.

This species is named after its place of occurrence: the mountain range of Sierra de Gredos in Central Spain. The name is an adjective.

Holotype ♂, “Cuenca V Korb” “HOLOTYPE Megacraspedus cuencellus Car. ♂ ROMANIA” “CIS-Korea Microlep. 4216 Megacraspedus cuencellus Car. – Cuena [sic], Spain K. T. Park” (MGAB) [photographs examined]. Non type-material. France. 1 ♂, Alpes Maritimes, Caussols, 1100 m, 22.v.2002, leg. J. Nel, genitalia slide 14295 Nel (TLMF). Spain. 1 ♂, prov. Teruel, Carretera Moscardón, 1600 m, 5.vii.2010, leg. Z. Tokár, genitalia slide GU 15/1401 Huemer (RCZT).

Adult. Male (Figure 43). Wingspan 10–14 mm. Segment 2 of labial palpus with large scale brush, dark brown on outer surface, white mottled with brown on inner and lower surface, white upper surface; segment 3 white with black tip. Antennal scape without pecten; flagellum black, indistinctly lighter ringed. Head, thorax, and tegula light grey. Forewing dark grey from blackish, light grey-based scales; fringes grey. Hindwing dark grey with grey fringes.

Female. Unknown.

Variation. The forewing colour varies slightly from brownish grey to darker grey. The holotype (examined from a photograph) differs in having black streaks in the fold and towards apex.

Male genitalia (Figure 177). Uncus moderately small and slender, about two times longer than wide, sub-rectangular with rounded apex; gnathos hook strong, about length of uncus, distally moderately curved with pointed apex; anterior margin of tegumen with deep, U-shaped emargination, pedunculi distinct, suboval; valva nearly straight, stout, extending almost to apex of uncus, basal part distinctly wider than distal part, distal part weakly curved dorsad, apically rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with moderate medial emargination, weakly rounded lateral humps, broadly sub-rectangular vincular sclerites extending from sclerotised posterior edge of saccus to sub-basal part of valva; saccus massive, semi-oval, with weakly pointed apex, ratio maximum width to length about 1, posterior margin arched, with pointed mediolateral projections, separated by shallow medial emargination, medial part with strongly sclerotised longitudinal ridge extending almost to middle of saccus, lateral sclerites short, nearly about half length of maximum width of saccus; phallus moderately slender, medially bent, orbicular coecum, distal two-thirds slender, sclerotised medial ridge with strong tooth, ventral edge with strongly sclerotised subapical tooth, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus cuencellus is characterised by its almost plain dark grey forewings and dark hindwings but largely agrees with M. bidentatus sp. n. (p 65), M. fuscus sp. n. (p 67) and M. skoui sp. n. (p 74). The male genitalia are unmistakable due to the unique structures of the phallus. M. cuencellus differs from other species particularly in the thorns of the phallus having a different position.

BIN BOLD:ACC5029 (n = 1). The distance to the nearest neighbour M. bidentatus sp. n. is 3.1% (p-dist).

Southern France (Nel and Varenne 1997), Spain.

Host plant and early stages are unknown. The few adults have been collected from late May to early July at altitudes of between 1500 and 1650 m.

Megacraspedus cuencellus was described from one male in good condition from the province of Cuenca in Central Spain (Caradja 1920: 117). The genitalia of the holotype were figured by Park (1996: 75, figs 50–52). A male specimen from France corresponds with M. cuencellus from Spain in the genitalia and in the short COI sequence of 407 bp.

Holotype ♂, “SPAIN [prov. Barcelona] Gironella 680 m N42.03975 E2.02987 9.5.2013 J. Tabell leg.” “GU 16/1433 ♂ P. Huemer” “DNA Barcode TLMF Lep 19855” (ZMUC).

Paratype. Spain. 1 ♂, same data as holotype (ZMUC).

Adult. Male (Figure 44). Wingspan 13–14 mm. Segment 2 of labial palpus with scale brush of about length of segment 3, dark brown on outer surface, dark brown mottled with white on inner surface, white on lower and upper surface; segment 3 cream-white with brown tip. Antennal scape without pecten; flagellum blackish brown. Head cream-white mottled with light brown; neck white; thorax as forewing; tegula whitish brown. Forewing greyish brown from light black- and brown-tipped scales, costal third more blackish; a narrow yellow line in fold ending in a small black dot; a small black dot at end of cell; termen with some black scales; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. One of the two examined specimens has the rather denuded forewings light greyish.

Male genitalia (Figure 178). Uncus moderately small, 1.7 times longer than maximum basal width, sub-triangular, evenly tapered to rounded apex; gnathos hook strong, about length of uncus, distally curved with pointed apex; anterior margin of tegumen with deep, U-shaped emargination, pedunculi distinct, suboval; valva nearly straight, stout, extending almost to apex of uncus, basal part distinctly wider than distal part, distal part weakly curved dorsad, apically rounded; saccular area covered with setae, without separated sacculus; posterior margin of vinculum with shallow medial emargination, weakly rounded lateral humps, broad vincular sclerites extending from sclerotised posterior edge of saccus to sub-basal part of valva; saccus massive, semi-oval, with pointed rod-like apex, ratio maximum width to length approaching 1, posterior margin arched, with pointed mediolateral projections, separated by shallow medial emargination, medial part with strongly sclerotised longitudinal ridge extending almost to middle of saccus, lateral sclerites short, about half length of maximum width of saccus; phallus moderately slender, medially bent, orbicular coecum, distal two-thirds slender, sclerotised medial ridge with subapical tooth, ventral edge with subapical tooth, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus bidentatus sp. n. is characterised by its greyish brown forewings, which are darker in the costal third and have a distinct black dot at end of the fold and at the cell. It is similar to M. alfacarellus (Figs 49–50) which is sligtly larger and has more yellow in the fold and more indistinct black dots on the forewings. It is also similar M. pusillus (Figure 51), but in that species the forewings are not darker in the costal third. M. fuscus sp. n. (Figure 45) differs in being almost unicolorous blackish grey. See also M. cuencellus (p 64). The male genitalia of M. bidentatus sp. n. are almost unmistakable due to unique structures, particularly the phallus, somewhat similar to M. fuscus sp. n. (Figure 179) from which it differs in several characters such the shape of the uncus, valva and the saccus. From the similar M. cuencellus (Figure 177) it differs in the shape of the uncus and the larger thorns of the phallus having a different position.

BIN BOLD: ADA0203 (n = 1). The distance to the nearest neighbour M. cuencellus is 3.1% (p-dist).

North-eastern Spain (prov. Barcelona).

Host plant and early stages are unknown. The two type specimens were collected in early May at an altitude of 680 m.

The species name refers to the sclerotisations of the phallus and is derived from the Latin word bidentatus (having two teeth). The name is a masculine adjective.

Holotype ♂, “Spain, Extremadura [prov. Cáceres] 39°34'11"N, 06°06'34"W 11 km SE Monroy 4. iv. 2009, 375 m leg. B. Skule” “DNA Barcode TLMF Lep 21299” “GU 17/1478 ♂ P. Huemer” (ZMUC). Paratype. Spain. 1 ♂, same data as holotype (ZMUC); 1 ♂, prov. Salamanca, Belena, 8.v.1979, leg. C. Gielis, genitalia slide 5344 Karsholt (RMNH).

Adult. Male (Figure 45). Wingspan 13 mm. Segment 2 of labial palpus with scale brush shorter than segment 3, blackish brown, mottled with white on inner surface; segment 3 longer than segment 2, black mottled with white on inner surface. Antennal scape without pecten; flagellum black. Head, thorax and tegula blackish grey. Forewing blackish grey; fold slightly lighter; an indistinct black dot in fold and at end of cell; fringes dark grey. Hindwing dark grey with grey fringes. Abdomen black.

Female. Unknown.

Variation. Unknown.

Male genitalia (Figure 179). Uncus moderately small, approximately 1.5 times longer than maximum basal width, sub-rectangular, apical edge weakly rounded; gnathos hook strong, about one-third longer than uncus, distally weakly curved with pointed apex; anterior margin of tegumen with broad and deep U-shaped emargination, pedunculi distinct, suboval; valva nearly straight, stout, extending almost to apex of uncus, basal part distinctly wider than distal part, distal part weakly curved dorsad, apically rounded, sub-basal edge with long and sword-shaped sclerite, pointed disto-posteriad; saccular area covered with setae, without separated sacculus; posterior margin of vinculum with shallow medial emargination, weakly rounded lateral humps, broadly suboval vinucular sclerites extending from sclerotised posterior edge of saccus to sub-basal part of valva; saccus nearly V-shaped, with rounded apex, ratio maximum width to length approximately 0.6, posteriolateral margin medially projected, without incision, medial part with broad and strongly sclerotised longitudinal ridge extending almost to apex of saccus, lateral sclerites slightly longer than maximum width of saccus; phallus moderately slender, nearly straight, orbicular coecum, distal two-thirds slender, sclerotised medial ridge with medial and ventral tooth, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus fuscus sp. n. is characterised by its almost plain blackish grey forewings and dark hindwings. Externally it mostly resembles M. grisea from Asia, but that species has ringed antennae and some lighter scales along the termen. It is also similar to, but darker than, M. cuencellus (Figure 43), M. pusillus (Figure 51) and M. skoui sp. n. (Figure 52). The male genitalia are unmistakable, due to unique structures, such as the sword-shaped sclerites of the valva, the peculiar saccus with a broad longitudinal ridge, and the dentation of the phallus situated mediad compared to M. bidentatus (Figure 178).

Not available, barcoding failed.

Western Spain (prov. Cáceres, Salamanca).

Host plant and early stages are unknown. The type specimens were collected in early April and early May at altitudes from 375 m to ca. 900 m.

The name of this species is derived from the Latin word fuscus (meaning dark) after the colour of the forewings. The name is a masculine adjective.

Megacraspedus fuscus sp. n. is tentatively attributed to the M. cuencellus species group due to the overall similarity in the male genitalia. Molecular data will be necessary to confirm this classification.

Holotype ♂, “Portugal BB [= prov. Beira Baixa] Serra de Estréla Torre, 1600–1950 m 10.-11.vii.1986 O. Karsholt” “Gen. Præparat N^o 3355 ♂ H. Hendriksen” (ZMUC). Paratypes. Portugal. 12 ♂, same data as holotype, but genitalia slide GEL 1206 Huemer (TLMF, ZMUC); 1 ♂, Braçais, Castello de Vide, Alto Alentejo, 9.v.1999, leg. M. Corley, genitalia slide 2003 Corley (RCMC); 3 ♂, Poco do Inferno, 1100 m, 6.vii.2014, leg. M.F.V. Corley, genitalia prep. (in glycerin) (RCMC). Spain. 1 ♂, Castilia, viii.1886, leg. O. Staudinger, genitalia slide Mus. Vind. 16.655 (NHMW); 1 ♂, prov. Segovia, San Ildefonso, 23.vi. [without year], leg. Staudinger, genitalia prep. (in glycerin); 1 ♂, same data, but 21.vi., leg. O. Staudinger; 1 ♂, same data, but 10.vii, leg. O. Staudinger (ZMHU). No collecting data. 1 ♂, 25.vi., coll. O. Staudinger (ZMHU). Excluded from the type-series (see Remarks): Spain. 1 ♂, prov. Avila, 5 km NW Guisando, 1150–1400 m, 21.vi.2012, leg. T. Nupponen, genitalia slide 5014 Tabell (ZMUC).

Adult. Male (Figure 46). Wingspan 12–13 mm. Segment 2 of labial palpus with moderately long scale brush, brownish on outer surface, white mottled with brown on inner surface, otherwise white; segment 3 white, mottled with black on outer surface and with black tip. Antennal scape without pecten; flagellum blackish brown ringed with light grey. Head cream-white, thorax, and tegula as forewing. Forewing bone white, mottled with light brown in costal half; a weak black dot at end of cell; costa blackish brown at base, otherwise white; fringes light grey. Hindwing greyish with concolorous fringes.

Female. Unknown.

Variation. A rather variable species. The colour of the forewing varies from almost plain bone white to light brownish with a white costa. A rather indistinct black dot is sometimes present at the end of the cell. Sometimes there are a few black scales at the end of the cell and/or near apex.

Male genitalia (Figs 180–181). Uncus moderately slender, about two times longer than wide, sub-rectangular with broadly rounded apex; gnathos hook strong, about length of uncus, weakly curved with pointed apex; anterior margin of tegumen with deep V-shaped emargination, pedunculi small, suboval; valva straight, moderately slender, basal part wider than distal part, extending to about apex of uncus, apically rounded; short, digitate sacculus at medial part of valva, weakly separated; posterior margin of vinculum with moderate medial emargination, weakly rounded lateral humps, broad semi-oval sclerites extending from sclerotised posterior edge of saccus to sub-basal part of valva; saccus massive, suboval, with abruptly tapered apex, ratio maximum width to length approaching 1, posterior margin arched, with shallow medial emargination, medial part with strongly sclerotised longitudinal ridge extending to anterior third of saccus, lateral sclerites short, about half length of maximum width of saccus; phallus slender, medially bent, orbicular coecum, distal three-quarters slender, rod-like, distal half with strongly sclerotised ridge and a single postmedial tooth, subapical area with few small thorns, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus trineae sp. n. is a rather uncharacteristic Megacraspedus which can be recognized by lacking black dots (apart from an indistinct one at the end of the cell) and the whitish coloured veins on the forewings. It is very similar to M. occidentellus sp. n. (p 79). The male genitalia differ from other species of the M. cuencellus species group in the characteristic postmedial tooth and the presence of a short sacculus.

BIN BOLD:ADF0469 (n = 1). The distance to the nearest neighbour Megacraspedus bidentatus sp. n. is 7.8% (p-dist).

Portugal and Spain.

Host plant and early stages are unknown. The adults have been collected in early May and in the first half of July at altitudes from 1100 to ca. 1900 m.

The species name (a noun in the genitive case) is dedicated to Trine Karsholt, Denmark, who assisted OK when collecting the holotype and a part of the type series of this species.

A single male from Central Spain (Figure 47) differs in the broader phallus and sclerotised ridge (Figure 182), whereas other characters, e.g., the presence of a short sacculus, cannot be determined with certainty due to the traditional preparation technique employed. As well as the obvious differences in the genitalia morphology the considerable DNA barcode distance (BIN BOLD:ACZ8654) of 7.2% to M. trineae sp. n. indicates a, presumably, different species. However, in the absence of sufficient material we postpone a description.

Holotype ♂, [France] “Holo-type” “Cannes S FRANCE 22.V.1892 Wlsm. 81169” “Type” “Walsingham Collection, 1910–427.” “Megacraspedus tristictus ♂ Wlsm. Ent. Mo. Mag. 46.231-2 sp. 3009.1 (1910) TYPE ♂ (f) deser 81169” (BMNH). Non-type material. France. 2 ♂, Dep. Alpes Maritimes, S of Tende, Saorge, 482 m, 3–4.vii.2008, leg. O. Karsholt (ZMUC); 1 ♂, same data, but 6.vi.2009, leg. P. Huemer; 1 ♂, same data, but 7.vi.2009, leg. P. Huemer, genitalia prep. (in glycerin); 1 ♂, Alpes Maritimes, Les Ferres, 3.vii.1971, leg. F. Dujardin; 1 ♂, Dep. Alpes Maritimes, St. Barnabé, 7.vi.1970, leg. F. Dujardin; 1 ♂, Dep. Alpes Maritimes, Èze, Fort de la Revere, 700 m, 23.vi.1995, leg. J. Nel, genitalia slide 4105 Nel (all TLMF); 1 ♂, Dep. Alpes Maritimes, Col de Vence 11–12.vi.1981, leg. Hahn (RCEA); 1 ♂, Dep. Alpes Maritimes, Gourdon, late vi.1968, leg. F. Zürnbauer, genital slide GEL 1185 Huemer (TLMF); 1 ♂, same data, but 13.v.1994, leg. Petru (NMPC); 1 ♂, Dep. Drôme, Col de l’Homme Mort, 1212 m, 17.vi.1982, leg. C. Gibeaux & Lovenu (ZMUC); 2 ♂, Dep. Var, Puits de Rians, 27.v.2000, leg. J. Nel, genitalia slide 11295 Nel; 2 ♂, Dep. Var, Puits de Rians, la Gardiole, 3.vi.2000, leg. J. Nel; 1 ♂, same data, but 19.vi.2001, leg. J. Nel; 1 ♂, same data, but 4.vi.1995, leg. J. Nel, genitalia slide 3819 Nel; 1 ♂, same data, but 30.v.1999, leg. J. Nel; 1 ♂, Dep. Var, Puits de Rians, la Plarée, 31.v.2004, leg. J. Nel; 1 ♂, Dep. Var, Mt. Caume, l´ubac, 750 m, 5.vi.2000, leg. J. Nel; 1 ♂, Dep. Var, Mt. Caume, ouest, 740 m, 29.v.1994, leg. J. Nel, genitalia slide 2064 Nel; 1 ♂, Dep. Var, Draguignan, 6.vi.1993, leg. J. Nel, genitalia slide 1170 Nel; 1 ♂, same data, but 31.v.1992, leg. J. Nel, genitalia slide 2543 Nel (all TLMF); 1 ♂, Dep. Var, Frejus, 20.v.1998, leg. H. K. Jensen (ZMUC). Italy. 1 ♂, prov. Savona, Conna, 300 m, 28.vi.1976, leg. G. Baldizzone (ZMUC).

Adult. Male (Figure 48). Wingspan 10–13 mm. Segment 2 of labial palpus with moderately long scale brush, light brown on outer surface, white mottled with brown on inner surface, white on lower and upper surface; segment 3 white with black tip. Antennal scape without pecten; flagellum light brown ringed with black. Head very light brown; thorax and tegula yellowish brown. Forewing light brown, from dorsum to fold yellowish brown, costa whitish brown apart from dark base; a black dot in fold at 2/5 and two black dots at 3/5 in middle of wing and at end of cell; a few blackish scales along termen; fringes light grey. Hindwing light grey with concolorous fringes.

Female. Unknown.

Variation. There is some variation in size, including among specimens from the same locality.

Male genitalia (Figure 183). Uncus moderately slender, about one-third times longer than maximum basal width, weakly tapered, apical edge evenly rounded; gnathos hook bulky, strongly sclerotised, particularly at apex, with longitudinal grooves, straight, approximately 1.5 times length of uncus, apically pointed; anterior margin of tegumen with deep V-shaped emargination, suboval pedunculi small; valva straight, stout, extending to about middle of uncus, distorted apical part rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with deep U-shaped medial emargination, broadly rounded lateral humps, suboval vincular sclerites with sclerotised posterior edge; saccus broadly sub-triangular, abruptly tapered at about two-thirds, ratio maximum width to length about 1, posterior margin with pointed mediolateral projections, separated by moderately deep incision, medial part with strongly sclerotised longitudinal ridge extending nearly to apex of saccus, lateral sclerites slightly shorter than maximum width of saccus; phallus slightly shorter than tegumen, almost straight, with moderately inflated coecum, distal two-thirds gradually tapered, ventrally with sclerotised ridge, dorsomedial area broadly sclerotised, ductus ejucalatorius twirled, with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus tristictus is characterised by its light yellowish brown forewings with three distinct black dots. It resembles M. pentheres (Figure 130). M. faunierensis sp. n. (Figure 40) differs in having the head white and the antennae darker. The male genitalia are similar overall to other species of the M. pusillus species group but differ e.g., by the distinctly shorter uncus.

BIN BOLD:ADA0606 (n = 1). The distance to the nearest congeneric neighbour M. tenuiuncus sp. n. is 12% , the distance to the nearest BIN in BOLD, Emarginea combusta (Noctuidae), is 10.3% (p-dist).

South-eastern France, north-western Italy.

Host plant and early stages are unknown. The adults have been collected from the middle of May to early July at altitudes from 300 to 1200 m.

Megacraspedus tristictus was described from one male collected by Walsingham at Cannes, SE France, 22.v.1892 (Walsingham 1910).

Holotype ♂, “Type” [Spain, prov. Granada] “Sierra Alfacar 18/19 VI 25 Dr. E. Wehrli, Basel” “alfacarellus Sa. Alfacar ♂ + 18.VI.25. W. ” “GU 16/1414 ♂ P. Huemer” (NHMB). Non-type material. Spain. 1 ♂, prov. Granada, Sierra de Alfacar, 24.vi.1880, coll. Staudinger; 1 ♂, Granada, coll. Staudinger (all ZMHU).

Adult. Male (Figs 49–50). Wingspan 16 mm. Segment 2 of labial palpus with long scale brush, dark brown on outer surface, white mottled with brown on inner surface, white on lower and upper surface; segment 3 whitish brown. Antennal scape without pecten; flagellum black. Head whitish brown with white neck; thorax and tegula as forewing. Forewing dark brown; a yellow line in fold ending in an elongate black dot; a small black dot in fold and at end of cell; fringes grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. The limited material available so far shows no variation.

Male genitalia (Figs 184–185). Uncus slender, about two times longer than maximum basal width, evenly tapered towards weakly pointed apex; gnathos hook bulky, with longitudinal grooves, straight, slightly longer than uncus, apically strongly sclerotised, pointed; anterior margin of tegumen with deep V-shaped emargination, suboval pedunculi distinct; valva straight, stout, extending to about middle of uncus, distorted apical part rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with deep U-shaped medial emargination, broadly rounded lateral humps, suboval vincular sclerites with sclerotised posterior edge; saccus sub-triangular, apically abruptly tapered, rod-like, ratio maximum width to length approximately 0.6, posterior margin with pointed mediolateral projections, separated by moderately deep incision, medial part with strongly sclerotised longitudinal ridge extending to anterior part of saccus, with or without forked anterior end, lateral sclerites about length of maximum width of saccus; phallus slightly shorter than tegumen, almost straight, with moderately inflated coecum, distal two-thirds gradually tapered, ventral margin with sclerotised ridge, dorsomedial area with long row of spines, ductus ejucalatorius twirled, with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus alfacarellus is characterised by its black antennae, and by the dark brown forewings with two weak black dots. It is similar to some forms of M. spinophallus sp. n. (Figs 53–54) and M. bidentatus sp. n. (Figure 44). The male genitalia are very similar to other species of the M. pusillus species group and mainly differ in the distally tapered phallus and the row of dorsomedial spines; they are only weakly separated from M. skoui sp. n. (Figure 52) by the larger uncus and the less pointed apex of the saccus.

Not available, no suitable specimen was available for barcoding.

Southern Spain (prov. Granada).

Host plant and early stages are unknown. The few adults known to date have been collected in the second half of June at unreported altitudes.

Megacraspedus alfacarellus was described from one male collected between 18–19.vi.1925 in Sierra Alfacar, Andalusia, Spain (Wehrli 1926).

Paratype ♂, “Sierra Nevada Granada ; SPAIN 3 VI 1901 Wlsm 86728” “Paratype” “Megacraspedus ♂ pusillus WLSM 2/3” “Walsingham Collection, 1910-427.”, genitalia slide 33661 (BMNH). Non-type material. Spain. 1 ♂, prov. Granada, Sierra Nevada, El Parador NP, 2500 m, 21.vii.1980, leg. E. Traugott-Olsen, genitalia slide 5621 Traugott-Olsen; 1 ♂, prov. Granada, Sierra Nevada, Camino de Valeta, 1600 m, 13.vi.1986, leg. E. Traugott-Olsen; 1 ♂, prov. Granada, Sierra Nevada, 3.4 km SSE Güejar Sierra, 1830 m, 4.vii.2015, leg. J. Tabell, genitalia slide 16/1432 Huemer; 1 ♂, prov. Almería, Sierra de los Filabres, Calar Alto, 2130 m, 5.vii.2015, leg. J. Tabell (all ZMUC).

Adult. Male (Figure 51). Wingspan 12–13 mm. Segment 2 of labial palpus with scale brush of about length of segment 3, dark brown on outer and inner surface, white on lower and upper surface; segment 3 cream-white with brown tip. Antennal scape without pecten; flagellum dark brown. Head whitish brown with white neck; thorax and tegula as forewing. Forewing light grey-brown, darkest at base of costa; a narrow yellow line in fold with an indistinct elongate black dot; a distinct black dot at end of cell; fringes light grey. Hindwing grey with light grey fringes.

Female. Unknown.

Variation. The black dot in the fold is absent in some specimens.

Male genitalia (Figs 186–187). Uncus moderately slender, approximately 1.7 times longer than maximum basal width, evenly tapered towards rounded apex; gnathos hook bulky, with longitudinal grooves, straight, slightly longer than uncus, apically strongly sclerotised, pointed; anterior margin of tegumen with deep V-shaped emargination, rounded pedunculi distinct; valva straight, moderately broad, extending slightly beyond base of gnathos, distorted apical part rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with deep medial emargination, broadly rounded lateral humps, suboval vincular sclerites with sclerotised posterior edge; saccus sub-triangular, with rod-like apex, ratio maximum width to length approximately 0.6, posterior margin with pointed mediolateral projections, separated by shallow incision, medial part with strongly sclerotised longitudinal ridge extending to middle of saccus, with weakly forked anterior end, lateral sclerites slightly longer than maximum width of saccus; phallus about length of tegumen, weakly curved, with weakly inflated coecum, distal two-thirds stout, with rod-like ventral sclerite extending to apex, broader and shorter dorsal sclerotisation with few minute teeth, ductus ejaculatorius with long and slender band-like interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus pusillus is characterised by its light grey-brown forewings with a distinct black dot at end of the cell. It is particularly similar to M. skoui sp. n. (Figure 52), but also to M. bidentatus sp. n. (Figure 44) and M. fuscus sp. n. (Figure 45). The male genitalia differ from those of M. skoui sp. n. (Figure 188) and M. spinophallus sp. n. (Figs 189–190) in the shallow medial incision of the posterior margin of the saccus and the shorter and less forked medial ridge, and the absence of a distinct row of dorsal and ventral teeth of the phallus.

BIN BOLD:ACZ8007 (n = 1). The distance to the nearest neighbour M. spinophallus sp. n. is 8.4% (p-dist).

Southern Spain (prov. Granada and Almería).

Host plant and early stages are unknown. The few adults known to date have been collected from the middle of June to the second half of July, at altitudes from between 1600 and 2500 m.

Megacraspedus pusillus was described from four males collected in the Sierra Nevada on 2.vi.1901 (Walsingham 1903: 266).

Holotype ♂, “Spain, [prov.] Granada, Sierra Nevada, Camino del Veleta 1650 m, 25.VI.1986, leg. Peder Skou” “Gen. prep. 5318 ♂ O. Karsholt” (ZMUC). Paratypes. Spain. 3 ♂, prov. Granada, Puerto de la Mora, 1300 m, 6.vi.1974, leg. W. Glaser, genitalia slide 5319 Karsholt (SMNK, ZMUC); 1 ♂, prov. Granada, 2 km W Diezma, 1400 m, 9–10.vii.2010, leg. Z. Tokár, genitalia slide GU 16/1405 ♂ Huemer (RCZT).

Adult. Male (Figure 52). Wingspan 11–13 mm. Segment 2 of labial palpus with scale brush almost as long as segment 3, brown on outer surface, brown mottle with white on inner surface, white on upper and lower surface; segment 3 longer than segment 2, black with white tip. Antennal scape without pecten; flagellum black, indistinctly ringed with light brown. Head, thorax and tegula cream-white mottled with light grey-brown. Forewing light grey-brown; base of costa and apex darker; fringes grey. Hindwing grey with grey fringes.

Female. Unknown.

Variation. One specimen has an indistinct black dot in the fold of the forewings.

Male genitalia (Figure 188). Uncus moderately slender, 1.7 times longer than maximum basal width, evenly tapered towards rounded apex; gnathos hook bulky, with longitudinal grooves, straight, slightly longer than uncus, apically strongly sclerotised, pointed; anterior margin of tegumen with deep V-shaped emargination, rounded pedunculi distinct; valva straight, moderately broad, extending slightly beyond base of gnathos, distorted apical part rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with deep medial emargination, broadly rounded lateral humps, suboval vincular sclerites with sclerotised posterior edge; saccus sub-triangular, with rod-like apex, ratio maximum width to length 0.7, posterior margin with pointed mediolateral projections, separated by moderately deep V-shaped incision, medial part with strongly sclerotised ridge strongly forked in medial part of saccus, lateral sclerites slightly longer than maximum width of saccus; phallus about length of tegumen, weakly curved, with weakly inflated coecum, distal two-thirds stout, ventral margin with sclerotised ridge extending to apex, weakly dentated near base, broader and shorter dorsal sclerotisation with row of minute teeth extending over entire length, ductus ejaculatorius with long and slender band-like interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus skoui sp. n. is characterised by its almost unicolorous light grey forewings. It resembles several smaller, unicolorous Megacraspedus species, especially M. pusillus, which has a distinct black dot at end of the cell (Figs 186–187). The male genitalia are very similar overall to other species of the M. pusillus species group and differ in particular in the characteristic dentations of the phallus. In this character the species resembles M. alfacarellus (Figs 184–185) but differs in the smaller uncus and some minute details such as the longer rod-like apex of the saccus.

BIN BOLD:ACT1624 (n = 1). The distance to the nearest neighbour M. spinophallus sp. n. is 7.97% (p-dist).

Spain (prov. Granada).

Host plant and early stages are unknown. Adults have been collected in June and July at altitudes between 1300 and 1650 m.

The species name (a noun in the genitive case) is dedicated to Peder Skou, Denmark, who collected the holotype of this species and numerous other Megacraspedus specimens used for our study.

Holotype ♂, “Spanien, [prov.] Alicante Sierra de Crevillente 5 km NE Albatera, 450 m, 38°15,22'N, 00°54,86'W 26.5.2004 leg. Huemer TLMF 2005-04” “BC TLMF Lep 03227” “P. Huemer GEL 1208 ♂” (TLMF). Paratypes. Spain. 8 ♂, same data as holotype (TLMF); 2 ♂, prov. Alicante, Rebate, 26.vi.1989, leg. B. Å Bengtsson, genitalia slide Bengtsson 3268 (RCBB, ZMUC); 2 ♂, prov. Alicante, 4 km E Aspe, Rio Vialopo, 300 m, 24.v.1998, leg. P. Skou (ZMUC); 2 ♂, prov. Alicante, 8 km N Albatera, 300 m, leg. J. Šumpich (NMPC); 1 ♂, prov. Alicante, Sierra Alta, Aitana, 1200 m, 18.vi.2011, leg. H. Rietz; 1 ♂, prov. Alicante, Parcent, 8.vi.2014, leg. H. Rietz; 1 ♂, prov. Alicante, Beniarbeig, 16.vi.2015, leg. H. Rietz; 1 ♂, prov. Alicante, Pto de la Carresqueta, 1050 m, 24.vi.2015, leg. H. Rietz (all ECKU); 1 ♂, prov. Alicante, Alcoj, Font Roja, 27.vi.2010, leg. A. Stübner (TLMF); 1 ♂, prov. Almería, 5 km SW Tabernas, Rambla de Tabernas, 200 m, 28.v.1998, leg. P. Skou; 1 ♂, prov. Almería, 10 km E Bedar, El Pinar, 325 m, 19–27.iv.2001, leg. P. Skou & B. Skule; 3 ♂, prov. Almería, El Pozo del Esparto, 20 m, 22–26.iv.2001, leg. C. Hviid, P. Skou & B. Skule (all ZMUC); 2 ♂, 3 km SW Pulpi, 200 m, 2.vi.2002, leg. W. Schmitz (RCWS); 1 ♂, prov. Almería, Tabernas, 380 m, 6–8.vii.2007, leg. G. Jeppesen, genitalia slide 6490 Hendriksen) (ZMUC); 1 ♂, prov. Almería, Tabernas env., Aghuilla Salada, 550 m, 7.vii.2010, leg. Z. Tokár; 1 ♂, prov. Almería, Sierra de Alhamilla, Huebro, 700–800 m, 29.iv.2008, leg. Z. Tokár (all RCZT); 6 ♂, same data, but leg. J. Šumpich; Sierra de Alhamilla, vicinity of Huebro, 700–800 m, 29.iv.2008, leg. J. Šumpich; 1 ♂, prov. Almería, Sierra Cabrera, Mojácar, 50 m, 4.v.2008, leg. J. Sumpich; 25 ♂, prov. Almería, Sierra de Alhamilla, Turrillas env., route Colativi, 1000 m, 15–19.vi.2007, leg. J. Šumpich; 8 ♂, Sierra de Alhamilla, vicinity of Níjar, 560 m, 29.iv.2008, leg. J. Šumpich (all NMPC); 1 ♂, prov. Almeria, Nijar, Huebro, 10.v.2014, leg. A. Stübner (ZSM); 1 ♂, prov. Castellon, 5 km E Cuevas de Vinroma, 200 m, 13.vii.1992, leg. M. Fibiger; 16 ♂, prov. Castellon, 25 km NW Castellon, La Banderetta pass, 800 m, 17.vii.1992, leg. M. Fibiger, genitalia slide 6521 Hendriksen, 15/1403 Huemer (ZMUC); 1 ♂, 1 ♀, prov. Granada, valley of Rio Guadelfo, South side, E of Orgiva, 1000 m, 5.vii.1969, leg. K. Sattler & D.J. Carter, gen. slides 33659 ♂, 33660 ♀ (BMNH); 1 ♂, prov. Huesca, 6 km SW Candasnos, Barranco de Valcuerna, 300 m, 15.vi.1999, leg. P. Skou; 1 ♂, same data, but 8 km S Candasnos, 175 m, 5.vi.2002, leg. B. Skule (all ZMUC); 2 ♂, prov. Huesca, 6 km W Ontiñena, 7.vi.2002, leg. W. Schmitz; 1 ♂, prov. Huesca, 10 km S Candasnos, 300 m, 13.vi.2004, leg. W. Schmitz; 4 ♂, prov. Huesca, Ontiñena, 300 m, 28.v.2015, leg. J. Viehmann (all RCWS); 1 ♂, prov. Huesca, 10 km SW Candasnos, 30.v.2015, leg. J. Viehmann (ZMUC); 5 ♂, prov. Huesca, Los Monegros, La Zaida, 19.v.2004, leg. J. Junnilainen; 2 ♂, same data, but 10 km NW Gelsa, 21.v.2004, leg. J. Junnilainen (all RCJJ); 2 ♂, prov. Lleida, 30 km NW Fraga, Ontinema, 250 m, 11.vii.1992, leg. M. Fibiger; 1 ♂, prov. Málaga, Camino de Ojen, 150 m, 21.vi.1980, leg. E. Traugott-Olsen; 1 ♂, same data, but 8.vi.1986; 1 ♂, prov. Málaga, Sierra de Marbella, 14.vii.1980, leg. E. Traugott-Olsen; 1 ♂, prov. Valencia, Porta Coeli, 13.vii.1988, leg. J. Baixeras & M. Domingues (all ZMUC); 2 ♂, prov. Valencia, Villargordo del Cabriel, 20.vi.2010, leg. A. Stübner; 3 ♂, same data, but 19.vi.2010; 1 ♂, same data, but 21.vi.2010 (all ZSM); 1 ♂, 1 ♀, prov. Zaragoza, Belchite, 350 m, 3.vii.2004, leg. J. Procházka (NMPC).

Adult. Male (Figure 53). Wingspan 11–20 mm. Segment 2 of labial palpus with long scale brush, dark brown on outer surface, white mottled with brown on inner surface, white on lower and upper surface; segment 3 white with some black towards tip. Antennal scape without pecten; flagellum black. Head whitish brown with white neck; thorax and tegula as forewing. Costal half and apical part of forewing dark brown; dorsal part light brown, darker towards dorsum; elongate black dots in fold at 1/3 and at end of cell; a few black scales at tornus; fringes light brownish grey. Hindwing grey with light grey fringes.

Female (Figure 54). Wingspan 19 mm. Forewing light yellow-brown, darker towards costa and especially at base of costa. An indistinct black dot at end of cell. Hindwing more slender than in male. Otherwise similar to male.

Variation. Highly variable in size, with specimens from southern Spain being generally larger. The colour of the head varies from almost cream coloured to grey-brown, and thorax and tegula vary accordingly. In some specimens the forewing (apart from the fold) is almost plain brown. One specimen has an indistinct black streak in the middle of the forewing. One of the two female specimens has no black dot in the fold.

Male genitalia (Figs 189–190). Uncus slender, about two times longer than maximum basal width, evenly tapered towards rounded apex; gnathos hook bulky, with longitudinal grooves, straight, slightly longer than uncus, apically strongly sclerotised, pointed; anterior margin of tegumen with deep V-shaped emargination, suboval pedunculi distinct; valva straight, stout, extending to about middle of uncus, distorted apical part rounded; saccular area densely covered with setae, without separated sacculus; posterior margin of vinculum with deep U-shaped medial emargination, broadly rounded lateral humps, suboval vincular sclerites with sclerotised posterior edge; saccus sub-triangular, apically abruptly tapered, rod-like, ratio maximum width to length 0.6, posterior margin with pointed mediolateral projections, separated by moderately deep incision, medial part with strongly sclerotised longitudinal ridge extending to anterior part of saccus, with distinctly forked anterior end, lateral sclerites about length of maximum width of saccus; phallus slightly shorter than tegumen, almost straight, with moderately inflated coecum, distal two-thirds gradually tapered, ventral margin with sclerotised ridge, dorsomedial area with large group of spines, ductus ejucalatorius twirled, with contorted linear interior sclerotisation.

Female genitalia (Figure 276). Papilla analis small, apically rounded; apophysis posterior slender rod-like, long, with short, bifurcate posterior end, bordered by sclerotised field; segment VIII approximately 1.5 mm long, membranous; subgenital plate with sub-triangular subostial sclerotisation, without pointed sclerites posteriorly, posteromedially broadly merged with medial flaps delimiting rounded ostium bursae, anterior margin with rod-like edge connected with apophysis anterior, medially with rounded projection; apophysis anterior slender, rod-like, about length of segment VIII, posteriorly becoming rod-like venula of segment VIII, extending to posterior margin; colliculum short, strongly sclerotised; ductus bursae short, broad; corpus bursae moderately short and broad, weakly delimited from ductus bursae; signum small, rounded spiny plate.

Megacraspedus spinophallus sp. n. is characterised by its black antennae, and by the dark brown forewings having two elongate black dots and being yellowish in dorsal half. It is very similar to M. alfacarellus (Figs 49–50), but that species has more plain dark brown forewings (apart from the yellow fold) with two small black dots. The male genitalia are very similar to other species of the M. pusillus species group but differ in the characteristic long and furcated medial ridge of the saccus and particularly the large field of spines of the phallus. The female genitalia are similar overall to several taxa in other species groups but differ in the subgenital plate without pointed sclerites. However, it remains unknown if this character is shared with other species in the M. pusillus species group.

BIN BOLD:AAU1828 (n = 7), BIN BOLD:ACT2894 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 2 BINs with 2.8% maximum divergence, based on a single specimen compared with a larger cluster. Within the latter average divergence is only 0.2% and maximum divergence is 0.6% (n = 2). The minimum distance to the nearest neighbour M. skoui sp. n. is 8% (p-dist).

Spain.

Host plant and early stages are unknown. The adults have been collected from late April to the middle of July at altitudes from sea level to 1200 m.

The species name is a compound word derived from the Latin adjective spinosus (meaning spiny) and noun phallus, referring to the spiny phallus. The name is a noun in apposition.

Holotype ♂, “Portugal, Ext. [=prov. Estremadura] Ericeira, 30 m 4.-8.vii.1986 O. Karsholt” “GU 13/1352 ♂ Huemer” (ZMUC). Paratypes. Portugal. 3 ♂, same data as holotype, but genitalia slide 6520 Hendriksen (TLMF, ZMUC); 1 ♂, Estremadura, 3 km E Capo da Roca, 200 m, 8.vii.1986, leg. O. Karsholt (ZMUC).

Adult. Male (Figure 55). Wingspan 9–12 mm. Segment 2 of labial palpus with moderately long scale brush, brownish on outer surface and on apical half of inner surface, otherwise white; segment 3 white mottled with blackish brown on lower and outer surface and with black tip. Antennal scape without pecten; flagellum dark brown, indistinctly ringed with black. Head, thorax and tegula as forewing. Dorsal half of forewing light yellow, costal half light brown, extending into a dark brown apical streak; costa whitish; fringes light grey. Hindwing grey with light grey fringes. Female. Unknown.

Variation. The few examined specimens show a slight variation in the forewing colour which, however, may be due to a more or less fresh condition.

Male genitalia (Figure 191). Uncus slender digitate, about 3 times longer than maximum basal width, apex rounded; gnathos hook bulky, with longitudinal grooves, slightly longer than uncus, straight, apically strongly sclerotised, pointed; anterior margin of tegumen with deep V-shaped emargination, pedunculi small, drop-shaped; valva straight, stout, extending slightly beyond middle of uncus, distorted apical part rounded; saccular area densely covered with setae, with weakly separated sacculus; posterior margin of vinculum with U-shaped emargination, broadly rounded lateral humps, suboval vincular sclerites with sclerotised posterior edge; saccus semi-oval, apically tapered, about same length and width, posterior edge weakly arched, with shallow medial incision, medial part with strongly sclerotised furcated longitudinal ridge, lateral sclerites shorter than maximum width of saccus; phallus stout, inflated coecum about twice width and almost length of distal part, sclerotised ridge in distal part without group of spines medially, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus occidentellus sp. n. is a rather uncharacteristic Megacraspedus which can be recognized by lacking black dots and whitish coloured veins on the forewing. It is similar, e.g., to the slightly larger M. trineae sp. n. (Figure 46), but can be recognized by its more uniformly coloured dark antennae, and by having the forewing colour separated by a lighter dorsal part and a darker costal part, the latter extending into a dark brown apical streak. The male genitalia differ from other species of the M. pusillus species group in having a slender uncus and in particular the posteriorly deeply incised saccus and the stout phallus with a comparatively large coecum.

Not available, barcoding failed.

Western Portugal.

Host plant and early stages are unknown. The type series was collected in early July at low altitudes.

This species is named after its occurrence at the most western part of continental Europe. The species name is derived from a combination of the Latin occidentalis (meaning western) and the diminutive suffix –ellus. The name is a compound noun.

Holotype ♂, “Hispania. [prov. Granada] Andalucia. Sierra Nevada. Cam.d.Valeta 1600 m 14.6.1986 E. Traugott-Olsen” “GU 16/1458 ♂ P. Huemer” (ZMUC). Paratypes. Spain. 1 ♂, prov. Granada, Sierra Nevada, Camino de Valeta, 1600 m, 12.vi.1986, leg. E. Traugott-Olsen; 2 ♂, same data, but 13.vi.1986; 4 ♂, same data, but 14.vi.1986, genitalia slide 6535 Hendriksen (ZMUC).

Adult. Male (Figure 56). Wingspan 13–16 mm. Segment 2 of labial palpus with scale brush about length of segment 3, brown on outer surface, brown mottled with white on inner surface, white on lower and upper surface; segment 3 about same length as segment 2, white mottled with black especially on lower surface and at tip. Antennal scape without pecten; flagellum indistinctly ringed black and brownish. Head light brown mottled with white; thorax cream coloured slightly mottled with brown; tegula brown at base, whitish towards apex. Forewing light grey-brown from whitish brown- and black-tipped scales, lighter in dorsal part below fold; costa blackish brown at base, otherwise greyish white; fold indistinctly light yellowish; some black scales along termen; fringes light grey. Hindwing dark grey with concolorous fringes.

Female. Unknown.

Variation. The yellowish streak in the fold can be more or less distinct.

Male genitalia (Figure 192). Uncus evenly slender, digitate, about four times longer than broad, basally slightly widened, apically converged; gnathos hook massive, with longitudinal grooves, slightly longer than uncus, basally about two-third width, evenly tapered with pointed apex; anterior edge of tegumen with deep suboval emargination; pedunculi suboval; valva extending to about base of uncus, broad, apical sixth abruptly tapered, distorted, with rounded apex; saccular area densely covered with setae, without clearly separated sacculus; posterior margin of vinculum medially emarginated, laterally with hump, suboval vincular sclerites with sclerotised posterior and proximal edge; saccus sub-triangular, approximately 1.4 times longer than broad, posterior edge weakly sinusoid, medial part with strongly sclerotised ridge extending from posterior edge to apex, lateral sclerites slightly shorter than maximum width of saccus; phallus with weakly inflated coecum, about one-quarter length of broadly digiate and weakly curved distal part, medial part with large group of small sclerotised knobs, apex rounded, ductus ejucalatorius with two contorted linear interior sclerotisations.

Female genitalia. Unknown.

Megacraspedus granadensis sp. n. is characterised in having the forewings covered with whitish brown- and black-tipped scales, in having the costa blackish brown at base and whitish grey in middle, and having no black dots. Also the dark grey hindwings are characteristic. The male genitalia differ from the particularly similar M. spinophallus sp. n. (Figs 189–190) in the knob-like sclerites of the phallus which is furthermore of a different shape.

Not available, barcoding failed.

Spain (prov. Granada).

Host plant and early stages are unknown. The adults have been collected in the middle of June at an altitude of 1600 m.

This species is named after its place of occurrence: the province of Granada in southern Spain. The name is a masculine adjective.

Holotype ♂, “ESPANA, Prov. Cantabria PN Picos de Europa Fuente De, El Cable Bergst. 4°48,53'W, 43°09,55'N 1870 m, 11.7.2012 leg. Huemer TLMF 2012-011” “BC TLMF Lep 08326” (TLMF). Paratypes. Spain. 1 ♂, same data as holotype; 7 ♂, same data, but TLMF 2013-011 (all TLMF); 30 ♂, same data, but leg. T. Mayr (RCTM); 1 ♂, Prov. Cantabria, PN Picos de Europa, above Fuente Dé, 1780 m, 9.vii.1999, leg. R. J. Heckford, genitalia slide 949 Heckford (RCRH); 21 ♂, prov. Cantabria, PN Picos de Europa, Espinama, Portillas des Poqueion, 1340 m, 11.vii.2012, leg. P. Huemer; 2 ♂, prov. Leon, Picos de Europa, Portilla de la Reina env., 1230 m, 9.vii.2012, leg. P. Huemer, genitalia slide GEL 1205 Huemer (RCTM, TLMF, ZMUC).

Adult. Male (Figure 57). Wingspan 12–14 mm. Segment 2 of labial palpus with scale brush as long as segment 3, grey-brown on outer surface, brown mottled with white on inner surface, white on lower and upper surface; segment 3 as long as segment 2, white with black tip. Antennal scape without pecten; flagellum black, indistinctly ringed with grey-brown. Head, thorax and tegula cream-white. Forewing light greyish brown from light brown dark-tipped scales; costa blackish brown at base, otherwise whitish; fringes light grey. Hindwing grey with grey fringes.

Female. Unknown.

Variation. The examined specimens show only slight variation.

Male genitalia (Figure 193). Uncus evenly slender, digitate, about four times longer than broad, apex rounded; gnathos hook massive, with longitudinal grooves, slightly longer than uncus, basally same width, evenly tapered to pointed apex; anterior edge of tegumen with deep suboval emargination; pedunculi sub-triangular, posteriorly with sclerotised ridge; valva extending to about middle of uncus, broad, apical sixth abruptly tapered, distorted, with rounded apex; saccular area densely covered with setae, without clearly separated sacculus; posterior margin of vinculum medially emarginated, laterally with hump, suboval vincular sclerites with sclerotised posterior edge; saccus sub-triangular, approximately 1.5 times longer than broad, posterior edge weakly sinusoid, medial part with strongly sclerotised ridge extending from posterior edge to apex, lateral sclerites slightly shorter than maximum width of saccus; phallus with inflated coecum, about twice width and one-third length of distal part, strongly sclerotised spine of about ¾ length of coecum emerging from distodorsal part, directed towards apex of phallus in moderate curvation, distal part of phallus slightly curved, apex rounded.

Female genitalia. Unknown.

Megacraspedus heckfordi sp. n. is characterised by its plain light greyish brown forewings without markings. It can be separated from other species without markings on the forewings by its cream-white head. The male genitalia are unmistakable within Megacraspedus and can be easily identified by the characteristic large spine of the phallus.

BIN BOLD:ACA9758 (n = 4). The intraspecific divergence of the barcode region is low with mean 0.5% and maximum divergence of 0.8%. The distance to the nearest neighbour M. tenuiuncus sp. n. is 7.8% (p-dist).

Spain (prov. Cantabria).

Host plant and early stages are unknown. The adults have been collected in the middle of July at altitudes from 1230 to 1870 m.

The species name (a noun in the genitive case) is dedicated to Robert (Bob) Heckford, U.K., who collected the first specimen and provided his valuable material available to our study.

Holotype ♂, “Spain [prov.] Lerida Roni near Sort 7.vii.1993, 1000 m P. Skou” “GU 15/1402 ♂ P. Huemer” (ZMUC). Paratypes. France. 1 ♂, Dep. Pyrénées-Orientales, Les Queres, Angoustrine, 1410 m, 19.v.2004, leg. T. Varenne, genitalia slide 17979 Nel; 3 ♂, same data, but Les Fontêtes, 1500 m, 13.vii.2005, genitalia slide 19284 Nel (all RCTV). Spain. 1 ♂, same data as holotype, but 8.vii.1993, leg. P. Skou (ZMUC); 2 ♂, prov. Barcelona, Castellfollit de Riubregós, Bosc de Roters, 560 m, 14.vi.2012, leg. F. Vallhonrat & C. Coll, genitalia slide 2152 Requena and 26231 Nel (RCCR, RCJN); 1 ♂, prov. Barcelona, Castellfollit de Riubregós, Torrent del Magrà, 716 m, 15.vi.2012, leg. F. Vallhonrat & C. Coll (RCCR); 1 ♂, prov. Girona, La Molina, 1800 m, 23–26.vi.2004, leg. A. Blumberg, genitalia slide 5343 Karsholt (RCWS).

Adult. Male (Figure 58). Wingspan 13 mm. Segment 2 of labial palpus with scale brush about same length as segment 3, brown on outer surface, white mottled with brown on inner surface, white on upper surface; segment 3 rather short, white. Antennal scape without pecten; flagellum black. Head cream-white; thorax and tegula light brown, the latter with whitish tip. Forewing light greyish; costa white from base; fold and veins in apical part of wing whitish; an indistinct black dot in fold and a black dot at end of cell; termen with some blackish scales; fringes grey. Hindwing grey with concolourus fringes.

Female. Unknown.

Variation. The examined specimens show only slight variation.

Male genitalia (Figure 194). Uncus extraordinarily long and slender, about five times longer than broad, basally weakly widened, apex slighty pointed; gnathos hook bulky, with longitudinal grooves, straight, slightly longer and broader than uncus, apically strongly sclerotised, curved tip; anterior margin of tegumen with deep, U-shaped emargination, teguminal wall with two longitudinal ridges anteriorly, converging mediad; suboval pedunculi with small supplementing hump; valva straight, stout, extending to about middle of uncus, distorted apical part rounded; saccular area densely covered with setae, distally with short digitate sacculus; posterior margin of vinculum with deep U-shaped medial emargination, broadly rounded lateral humps, vincular sclerites elongated, posteriomedial edge strongly sclerotised; saccus sub-triangular, apically abruptly tapered, short, ratio maximum width to length approximately 1.1, posterior margin with pointed mediolateral projections, separated by moderately deep V-shaped incision, medial part with strongly sclerotised longitudinal ridge extending to anterior part of saccus, with weakly forked anterior end, lateral sclerites about three-quarters length of maximum width of saccus; phallus with weakly inflated coecum, about twice width of digitate distal four-fifths, distomedial part curved, with small spines ventrally, ductus ejucalatorius with contorted linear interior sclerotisation.

Female genitalia. Unknown.

Megacraspedus tenuiuncus sp. n. is characterised by its black antennae, and by its light greyish forewings with two black dots. It is similar to M. lanceolellus (Figs 7–16), but that species has ringed antennae, and less distinct black dots on the forewing. The male genitalia of M. tenuiuncus sp. n. are unmistakable and in particular the extremely long and slender uncus is unique in Megacraspedus. It furthermore differs from most other species of the M. pusillus species group by the presence of a short sacculus.