Megacraspedus dolosellus (Zeller, 1839)

Huemer, Peter & Karsholt, Ole, 2018, Revision of the genus Megacraspedus Zeller, 1839, a challenging taxonomic tightrope of species delimitation (Lepidoptera, Gelechiidae), ZooKeys 800, pp. 1-278 : 43-51

Megacraspedus dolosellus (Zeller, 1839)

Ypsolophus (Megacraspedus) dolosellus Zeller, 1839: 190.

Ypsolophus separatellus Fischer von Röslerstamm, 1843: 300, 302, pl. 100, figs 1 a–d [on plate as’separatella‘], syn. n.

Megacraspedus incertellus Rebel, 1930: (14), syn. n.

Examined material.

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).

Redescription.

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.

Diagnosis.

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.

Molecular data.

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).

Distribution.

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.

Biology.

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 10th 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.

Remarks.

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).