(Figs. 3, 4, 11 –13, 19– 22)
Nepticula headleyella Stainton 1854: 300. 2 Syntypes: [England, London]: Headley Lane, August, leg. Douglas [probably lost].
Nepticula argyrostigma Frey 1856: 261. Lectotype ♂ (designated by van Nieukerken & Johansson 1990: 262), Switzerland: Zürich, Frey coll., Genitalia slide 24089 (Natural History Museum London) (Synonymised by Frey 1880: 425) [examined].
Nepticula dubiella Hauder 1912: 273. Lectotype ♀ (designated by van Nieukerken & Johansson 1990: 262), Austria: Kirchdorf, 21.v. 1900, leg. Hauder, genitalia slide EJvN 2609 (see Fig. 22) (NMW) (Synonymised by Klimesch 1948: 76) [examined].
Trifurcula rodella Svensson 1982: 299. Holotype ♂, Sweden: Vg., Kinnekulle, 1–2.vii. 1966, I. Svensson, Genitalia slide IS 4550 (Lund) (Synonymised by van Nieukerken 1986 a: 5) [examined].
Fedalmia headleyella; Beirne 1945: 207 [new genus, new combination]; Emmet 1976: 208 [redescription].
Trifurcula (Fedalmia) headleyella; Johansson 1971: 245. [new combination].
Trifurcula (Glaucolepis) headleyella; van Nieukerken 1986 a: 15 [revised combination]; van Nieukerken & Johansson 1990: 261 [redescription]; Laštůvka & Laštůvka 1997: 125 [redescription]; Bengtsson et al. 2008: 234 [redescription].
Diagnosis. Trifurcula headleyella can be separated from other European Trifurcula, except T. lituanica (see there for differences), by the postmedial opposite metallic spots on the forewings in both sexes, and in the female (Fig. 4) in addition by the black head and strongly leaden shining basal third of the forewing. T. (Levarchama) eurema (Tutt, 1899), also has opposite spots, but never metallic, and often forming a fascia. Males of that species also possess a hairpencil on hindwing.
Descriptive notes. Male: Forewing length 1.8–2.5 mm, wingspan 4.1–5.7 mm, antenna with 35–41 segments (Bengtsson et al. 2008 give up to 45). Female: Forewing length 1.9–2.4 mm, wingspan 4.2–5.4 mm, antenna with 35–41 segments. T. headleyella is one of the few nepticulid species were the female has as many antennal segments as the male.
Male genitalia (Figs. 11–13): capsule 255–305 µm long, valva length 120–160 µm, aedeagus 220–275 µm long, cornutus 60–85 µm long.
Female genitalia (Figs. 19–22): total bursa length ca. 900 µm, signa 240–310 µm long, in some specimens very dissimilar, in others equally long. Anal papillae with ca. 9–10 setae, T 8 with some setae and scales. Ductus spermathecae with 2.5–3 convolutions.
Biology. Hostplants: Prunella vulgaris, P. grandiflora and P. laciniata (Lamiaceae). Egg on upper surface of a leaf. The larva usually makes a short gallery in the first leaf and mines down the petiole, into the stem and then via the petioles into a second and often third leaf. The mine is linear, with linear frass, later becoming wider and more irregular, the frass often not in the middle. By mining the petiole the larva often partly cuts off the sap stream, leading to purple discoloration of one or more leaves (often those without mines!) (illustrated by Huisman et al. 2004). Discoloured leaves may be an indication for finding mines. In larger plants the larva may just use one leaf.
Distribution. Widespread, but very localised in many parts of Europe: southern England (Emmet 1976; Edmunds 2011), Denmark, southern Sweden and southern Finland (Bengtsson et al. 2008), * Netherlands (Huisman et al. 2004), Germany (van Nieukerken et al. 2010), * Poland (Borkowski 1975), France (Nieukerken et al. 2006), * Spain (van Nieukerken et al. 2004), Switzerland (SwissLepTeam 2010), Austria, Italy, Czech Republic, Slovakia (Laštůvka & Laštůvka 1994; Tokár et al. 2002), Hungary, Croatia, Romania, Greece, * Russia: Karelia (Kutenkova 1989), Estonia, Latvia, * Lithuania (Ivinskis 2004), * Ukraine (general source: van Nieukerken 2011, other cited references contain detailed distribution information and/or maps). From countries with an asterisk just a single record or locality is known, for the country names in italics, detailed records are provided below.
Habitat. Considering how widespread and abundant the hostplant Prunella is, T. headleyella is remarkably local and rare. It is most frequently found in limestone grasslands, downland, dune slacks and alpine meadows. Where it occurs, it can be abundant, but still occurs very locally.
Remarks. Although the three synonyms and their types had been checked before (van Nieukerken & Johansson 1990), we re-examined the information on these types, including photographs of genitalia slides, in order to see whether one of these names could be the new species. Clearly all types show the characters of T. headleyella. The female lectotype of Nepticula dubiella (Fig. 22) can now be evaluated more accurately against the very different female genitalia of T. lituanica. In fact the complete type series of Nepticula dubiella and most other specimens under this name in the collections in Linz and Munich are after dissection shown to be T. headleyella, and only the four Klimesch specimens from Steyregg, Pfenningberg near Linz, cited above, belong to T. lituanica. Although Klimesch (Klimesch 1948) had synonymised dubiella with headleyella, he later changed his mind (Klimesch 1990), probably on the basis of the information that the single specimen dissected by Roland Johansson belonged to a different species. The confusion with N. dubiella lead EJvN (in Kasy 1985) to cite T. headleyella incorrectly as new for Austria.
Additional material examined. AUSTRIA: 13, Kärnthen, Wippach, 1.vi. 1909, Polica (Vienna); 13, Niederösterreich, Buchberg, Spitz a. D., 8.v. 1902, Preissecker (Vienna); 23, 1Ƥ, Niederösterreich, Gramatneusiedl: Fischawiesen (Fürbachwiesen), larvae 2.x. 1983 on Prunella grandiflora, emerged 25.iv– 10.v. 1984, E.J. van Nieukerken & J. Boomsma (Leiden); Oberösterreich: 13, Berg, 17.vii. 1912, Wolfschläger (Linz); Hinterstoder, 10.v. 1936, J. Klimesch (Munich); 23, Kirchdorf, 13.v.1893, 21.v. 1900, Hauder (2 PLT dubiella, Linz); 13, Linz a. D., Prebenau, 17.viii. 1910 (Linz); 13, Linz, Plesching, 25.v. 1967, J. Klimesch (Munich); 13, Pregarten (Prägarten), 31.v. 1909, Knitschke (PLT dubiella, Linz); 23, Prägarten, 26.v. 1910, Hauder & [Knitschke] (2 PLT dubiella, Linz); 23, Puchenau [Puchenauer Graben], 17.viii. 1910, [Frau Frazeni] (2 PLT dubiella, Linz); 23, Puchenau [Puchenauer Graben], 31.v. 1912, Hauder (labelled as type dubiella, but not cited in description, Linz, Vienna); 13, Niederösterreich, Buchberg, Spitz a. D., 8.v. 1902, Preissecker (Vienna); 13, Ruefling Linz, v. 1912 (Ende Mai 1912), Knitschke (PLT dubiella, Linz); 13, Steiermark, Altaussee, Seewiese, 1.vi. 1969, J. Klimesch (Munich); 13, Wien, Haschberg, 21.v. 1916 (Vienna). CROATIA: 23, 1Ƥ, Slavonija, Banicevac, 7 km NE Cernik, larvae 17.x. 1983 on Prunella laciniata, emerged 17.iv– 8.v. 1984, E.J. van Nieukerken & J. Boomsma (Leiden); 33, Dalmatia, Baska, Voda (Makarska), 18.v. 1979, J. Klimesch (Munich). GREECE: 13, Ipiros, Peristrei Mts. S. Metsovo, 12–1900 m, 27– 28.v. 1994, O. Karsholt (Copenhagen); 13, Trikala, Oxinia, 8.vi. 1999, A. & Z. Lastuvka (coll. A. Laštůvka). ITALY: 13, Cuneo, Pamparato, St. Gree, 25.vi. 1987, G. Bassi (coll. Bassi); 13, Torino, Almese, Falde del Mt. Musiné, 450 m., 26.v. 1983, U. Parenti (Leiden); 83, Verona, Monte Baldo, Rifuggio Novezzina, 1250–1350 m, 14.vii.1979, 24.vii.1984, 16.vii.1985, 16.vii. 1987, U. Parenti (Leiden, Torino); Roma: Monti d. Tolfa, dint. Manziana, 300 m, 23– 28.vi. 1989, G. Baldizzone (coll. Baldizzone); 5 Ƥ, Trento: Mte Maranza, NW slopes, 4 km E Trento, 900 m, larvae 10.x. 1983 on Prunella grandiflora, emerged 8–16.v. 1984, E.J. van Nieukerken (Leiden); 13, 1Ƥ, Veneto, Sappada, Passo Siera, 1600 m, 6.vii. 1991, B.Å. Bengtsson (coll. Bengtsson). ROMANIA: larvae, leafmines, Covazna: Ozunca- Bai S, 770 m, Prunella vulgaris, E.J. van Nieukerken (Leiden). UKRAINE: 13, Karpaty, Iv.-Frankovskaya obl. s. Yaremche, 600 m, 28.vi. 2003, A. Bidzilya (Kiev).
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