Corresponding author: Peter Huemer (
Academic editor: E. van Nieukerken
The taxonomy of the Palearctic genus
DNA barcode sequences of the COI barcode fragment with > 500 bp were obtained from 264 specimens representing 62 species or about three-quarters of the species. Species delimitation is particularly difficult in a few widely distributed species with high and allegedly intraspecific DNA barcode divergence of nearly 14%, and with up to 23 BINs in a single species. Deep intraspecific or geographical splits in DNA barcode are frequently not supported by morphology, thus indicating a complex phylogeographic history or other unresolved molecular problems.
The following 44 new species (22 of them from Europe) are described:
Huemer P, Karsholt O (2018) Revision of the genus
The
The necessity of a monographic review became increasingly relevant when the authors of this study, in co-operation with Jan Šumpich, decided to publish an additional, third volume of the series “Microlepidoptera of Europe” (see
Our study is based on a large amount of material of
Lepidopteran DNA barcode sequences are based on a 658 base-pair long segment of the 5’ terminus of the mitochondrial COI gene (cytochrome c oxidase 1). DNA samples (dried legs) were prepared according to the prescribed standards. Legs from 418 specimens of
Photographs of the adults were taken with an Olympus SZX 10 binocular microscope and an Olympus E-3 digital camera over a dark grey background and developed using the software Helicon Focus 4.3 and Adobe Photoshop CS4 and Lightroom 2.3. Genitalia photographs were taken with an Olympus E1 Digital Camera from an Olympus BH2 microscope. Multiple photos of different focal planes were assembled into single deep-focus images both for adult specimens and genitalia. Photographs from external sources are used for few specimens that we were unable to study ourselves. Photographs of wing slides were taken with a Canon 750D Digital camera and lens Canon MP-E 65 mm. Multiple photos of different focal planes were stacked into single deep-focus images, assembled with Helicon Focus 6 and Adobe Photoshop CC 2017. Adults, as well as male and female genitalia are usually figured to the same scale.
The descriptive terminology of the genitalia structures generally follows
Some unpublished taxonomic changes made by Linda Pitkin and Klaus Sattler (both at
Gender agreement between specific and generic names as prescribed by article 31.2 of the
The systematics of the
Type species:
Type species:
Type species:
Type species:
Type species:
Type species:
Type species:
Labial palpus of
Adult head with pecten on antennal scape of
Wing venation of
Terminology of important characters in male genitalia of
Terminology of important characters in female genitalia of
The early stages of most
The genus
With 25 species, Spain has the most diverse fauna of
The present revision results in many changes and new country records, e.g., the most recent checklist of the
Brachyptery in
Female wing reduction in
COI sequences in
Species | Mean Intra-Sp | Max Intra-Sp | Nearest Neighbour | Nearest Species | Distance to NN |
---|---|---|---|---|---|
|
0.93 | 1.39 | LEAST584-17 |
|
5.09 |
|
4.45 | 4.45 | PHLAH795-12 |
|
11.22 |
|
1.19 | 2.24 | LEEUA729-12 |
|
7.56 |
|
2.08 | 4.19 | LEEUA685-12 |
|
6.77 |
|
0 | 0 | LEASS140-16 |
|
6.28 |
|
0 | 0 | LEATI093-15 |
|
10.66 |
|
0.15 | 0.15 | PHLAF012-11 |
|
11.99 |
|
N/A | 0 | PHLAH794-12 |
|
3.14 |
|
N/A | 0 | LEATJ1347-16 |
|
14.91 |
|
2.19 | 3.81 | LEASS112-16 |
|
8.97 |
|
3.74 | 5.97 | LEASS032-16 |
|
11.25 |
|
N/A | 0 | PHLAI412-13 |
|
7.55 |
|
N/A | 0 | LEALT202-16 |
|
6.64 |
|
N/A | 0 | LEASS012-16 |
|
3.14 |
|
2.16 | 2.16 | LEATI093-15 |
|
8.97 |
|
7.49 | 13.76 | PHLAF014-11 |
|
9.85 |
|
0.15 | 0.15 | PHLAI458-13 |
|
8.46 |
|
0.1 | 0.16 | LEAST573-17 |
|
5.56 |
|
1.04 | 1.57 | LEASS022-16 |
|
11.88 |
|
0.15 | 0.15 | PHLAH788-12 |
|
7.77 |
|
0.7 | 1.13 | LEASS026-16 |
|
6.24 |
|
N/A | 0 | PHLAG308-12 |
|
10.77 |
|
0.1 | 0.15 | PHLAJ385-14 |
|
8.78 |
|
N/A | 0 | LEASS012-16 |
|
10.01 |
|
0.49 | 0.77 | LECRT129-16 |
|
7.74 |
|
N/A | 0 | PHLAD606-11 |
|
9.1 |
|
N/A | 0 | LEAST388-17 |
|
7.46 |
|
3.38 | 4.6 | LEATJ1354-16 |
|
9.15 |
|
N/A | 0 | LECRT125-16 |
|
9.35 |
|
N/A | 0 | LEEUA949-12 |
|
5.09 |
|
1.2 | 2.82 | LEAST575-17 |
|
7.56 |
|
0.2 | 0.31 | LEASS618-17 |
|
10.13 |
|
0.15 | 0.15 | LEAST583-17 |
|
7.77 |
|
7.61 | 12.51 | LEASS101-16 |
|
9.85 |
|
0.03 | 0.19 | LEAST594-17 |
|
6.57 |
|
N/A | 0 | LEAST572-17 |
|
4.78 |
|
0.93 | 0.93 | LEATJ1383-16 |
|
2.33 |
|
N/A | 0 | PHLAD606-11 |
|
9.17 |
|
N/A | 0 | LEEUA846-12 |
|
8.13 |
|
0.7 | 1.4 | PHLAI416-13 |
|
4.78 |
|
N/A | 0 | LEFIJ2706-15 |
|
8.97 |
|
N/A | 0 | LEEUA779-12 |
|
8.13 |
|
0.56 | 0.93 | LEATJ1383-16 |
|
9.45 |
|
N/A | 0 | LEASS136-16 |
|
6.28 |
|
0 | 0 | LEASS005-16 |
|
9.14 |
|
N/A | 0 | LEASS120-16 |
|
8.43 |
|
0.97 | 1.55 | PHLAG675-12 |
|
9.69 |
|
0.15 | 0.15 | LEFIJ2706-15 |
|
6.24 |
|
0.79 | 1.19 | PHLAJ385-14 |
|
6.44 |
|
N/A | 0 | PHLSA052-11 |
|
7.97 |
|
0.32 | 0.64 | LECRT099-16 |
|
2.33 |
|
0.8 | 2.83 | LASTS762-15 |
|
7.97 |
|
0.45 | 0.95 | PHLAI458-13 |
|
9.3 |
|
1.71 | 2.5 | PHLAG308-12 |
|
11.41 |
|
0.62 | 0.62 | LEASS007-16 |
|
9.82 |
|
N/A | 0 | PHLAH508-12 |
|
7.74 |
|
2.99 | 7.45 | PHLAI454-13 |
|
9.69 |
|
N/A | 0 | PHLAH794-12 |
|
9.97 |
|
N/A | 0 | LEASS012-16 |
|
7.76 |
|
N/A | 0 | LECRT129-16 |
|
11.98 |
|
0 | 0 | LEASS108-16 |
|
6.44 |
|
N/A | 0 | LEASS005-16 |
|
7.46 |
279 specimens of
Maximum likelihood tree (built with MEGA7) of cytochrome
=
=
=
=
=
=
=
=
=
=
=
=
=
=
Numerous species from various regions have been described in
1 | Gnathos hook straight, massive and bulky, with longitudinal grooves (Figs |
|
– | Gnathos hook usually curved or bent, of various shape, without longitudinal grooves |
|
2 | Uncus apically rod-like, maximum half width of valva (Figs |
|
– | Uncus without rod-like apical part, usually with rounded apex, two-thirds to about twice width of valva |
|
3 | Sub-basal part of valva with long and curved thorn-like process (Fig. |
|
– | Sub-basal part of valva without process |
|
4 | Valva very broad, about width of uncus, with sclerotised subapical ridge (Figs |
|
– | Valva slender, rarely broad but without sclerotised subapical ridge |
|
5 | Valva distinctly exceeding apex of uncus; phallus long and slender, without distinct sclerotisations (Figs |
|
– | Valva at most length of uncus; phallus of different shape, partially with sclerotisations |
|
6 | Uncus basally constricted, short, suboval |
|
– | Uncus basally not constricted, sub-rectangular to sub-triangular |
|
7 | Sacculus usually present; phallus distally narrowing with sclerotised, denticulate ridge (Figs |
|
– | Sacculus absent; phallus distally with broadly sinusoid dorsal margin, without denticulate ridge (Figs |
|
8 | Phallus with long and slender thorn-like sclerite from about middle nearly to apex, parallel to phallus axis (Fig. |
|
– | Phallus of different shape, without long and slender thorn |
|
9 | Phallus bent distad of bulbous of coecum, with long and slender distal part, 1–2 strong medial or sub-apical teeth |
|
– | Phallus of different shape, without sclerotisation or with groups of smaller teeth or differently shaped sclerotisation |
|
10 | Sacculus well developed, digitate; phallus very long (Figs |
|
– | Sacculus absent, phallus shorter (Figs |
|
11 | Uncus sub-quadrate, large, nearly width of posterior part of tegumen; ductus ejaculatorius with double-twisted interior sclerotisation (Figs |
|
– | Uncus smaller, oblong; ductus ejaculatorius with or without interior sclerotisation of different shape |
|
12 | Sacculus well developed, digitate |
|
– | Sacculus absent |
|
13 | Uncus oblong 1.7 times longer than maximum width; gnathos hook weakly curved; ductus ejaculatorius with broad and short interior sclerotisation (Figs |
|
– | Uncus rounded, about same maximum length as width; ductus ejaculatorius with very long and 4–5× contorted interior sclerotisation (Figs |
|
14 | Saccus large, oblong, apically usually broadly rounded, rarely pointed, with longitudinal sclerotised ridge extending almost to apex |
|
– | Saccus shorter, sub-triangular, smoothly sclerotised without or with short ridge of various shape |
|
15 | Uncus small, about width of valva; gnathos hook straight; phallus with stout and broad distal part (Figs |
|
– | Uncus large, distinctly broader than valva; gnathos hook curved to strongly bent; phallus with oblong and usually slender distal part (Figs |
|
16 | Valva short and broad; saccus broadly suboval, with rounded apex (Figs |
|
– | Valva long, slender to moderately broad; saccus sub-triangular, with pointed apex |
|
17 | Saccus long and slender, ratio maximum width to length 0.6; phallus with small sub-apical spine and larger sub-apical tooth (Fig. |
|
– | Saccus short to moderately long, ratio maximum width to length minimum 0.8; phallus of different shape, exceptionally subapical tooth on right side and two small spines on left side |
|
18 | Gnathos hook long and slender, extending nearly to anterior margin of tegumen (Fig. |
|
– | Gnathos hook shorter, not extending to anterior margin of tegumen |
|
19 | Gnathos hook short, stout, curved; valva apically distinctly inflated, phallus with sclerotised, usually dentated ridge (Figs |
|
– | Gnathos hook moderately short to long, slender; valva apically narrowing or contorted and rounded; phallus without sclerotised, dentated ridge |
|
20 | Saccus with short, furcated posterior ridges (Figs |
|
– | Saccus without or with short, unfurcated ridge |
|
21 | Phallus with S-curvature (Figs |
|
– | Phallus without S-curvature |
|
22 | Gnathos hook long, bent at right angles at about two fifth (Fig. |
|
– | Gnathos hook shorter, nearly straight to weakly curved |
|
23 | Phallus with broadly lobed dorsal and rod-like ventral sclerotisation (Figs |
|
– | Phallus of different shape, without broadly lobed dorsal sclerotisation |
|
24 | Valva broad, with longitudinal ridge (Fig. |
|
– | Valva slender, without ridge (Figs |
The
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.
Adult.
Due to its variation
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
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
Southern parts of Europe, from the Balkan Peninsula to Italy, France, and Spain, also known from Germany (northwards to Rheinland-Pfalz (
Early stages are undescribed. According to Lhomme (1946: 538 as
Adult.
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
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
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Morocco. 2 ♂, Prov. Al Haouz, Oukaïmeden 1.5 km SE, 2660 m
Adult.
BIN BOLD:ADF2257 (n = 1). The distance to the nearest neighbour
Morocco (Middle Atlas, High Atlas).
Host plant and early stages are unknown. The type-series of
The
External morphology. Segment 2 of labial palpus with scale brush shorter or as long as segment 3; segment 3 about length of segment 2. Antennal scape without or with one or a few hairs. Wingspan (males) 10–16 mm. Forewing with white costa, and in some species with white veins, but no black dots. Known females distinctly to strongly brachypterous.
Genitalia morphology. Male genitalia. Uncus moderately slender, sub-triangular; gnathos hook long, slender, slightly longer than uncus; anterior margin of tegumen deeply emarginated; valva straight, moderately slender; saccular area setose, without separated sacculus; saccus nearly V-shaped, short, ratio maximum width to length approximately 0.8, posterior margin with distinct projection, distally abruptly tapered, without or with short, partially furcated medial ridge; phallus moderately stout, with globular coecum, distal portion weakly to distinctly S-curved, with broad dorsal and long and slender ventral lobes, apically tapered.
Female genitalia. Papilla analis small; apophysis posterior is very long; segment VIII long and slender, largely membranous; subgenital plate with sub-triangular subostial sclerotisation, anteromedially with short projection; apophysis anterior is rod-like, from posterior margin of segment VIII; colliculum short; signum small, rounded spiny plate.
Taxa of the
Adult. Male (Figure
Female (Figure
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia (Figure
BIN BOLD:ACA8694 (n = 1). The distance to the nearest neighbour
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.
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
BIN BOLD:ADB7272 (n = 1). The distance to the nearest neighbour
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.
Ukraine. 2 ♂, Crimea, Aj-Petri, 5–6.vii.2002, leg, Yu. Budashkin (ZMKU).
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
BIN BOLD:ACS7353 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour
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
Adult.
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
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
Adult.
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
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.
Adult.
BIN BOLD:ACM1095 (n = 1). The distance to the nearest congeneric neighbour
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.
Adult.
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
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
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
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.
The
External morphology. Segment 2 of labial palpus with scale brush shorter than segment 3; segment 3 as long as or longer than segment 2. Antennal scape without or with one fine hair. Wingspan (males) 11–14 mm. Forewing with three black dots, but without white costa and veins. Known females are brachypterous.
Genitalia morphology. Male genitalia. Uncus moderately small, about width of valva, sub-rectangular; gnathos hook moderately short, slender, straight, slightly longer than uncus; anterior margin of tegumen moderately emarginated; valva straight, stout; saccular area setose, with longitudinal ridge, without separated sacculus; saccus V-shaped, oblong, posterior margin with distinct projection, distally strongly tapered, with long medial ridge almost extended from posterior margin to apex; phallus weakly curved, with weakly inflated coecum and stout distal part, sclerotised dorsal ridge, ventrally wrinkled; ductus ejaculatorius with linear internal sclerotisation.
Female genitalia. Papilla analis small; apophysis posterior very long; segment VIII long and slender, largely membranous; subgenital plate with sub-triangular subostial sclerotisation, anteromedially with almost tubular projection; apophysis anterior rod-like, from posterior margin of segment VIII; colliculum short; signum small, rounded spiny plate.
Adult.
The male genitalia are similar overall to
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
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.).
Adult.
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
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.
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia unknown.
Adult.
BIN BOLD:ADI8272 (n = 1). The distance to the nearest neighbour
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.
The
External morphology. Segment 2of labial palpus with scale brush shorter to longer than segment 3; segment 3 as long as or longer than segment 2. Antennal scape without pecten. Wingspan (males) 10–14 mm. Forewing in some species with 1–2 indistinct black dots, but without white costa and veins. Females unknown.
Genitalia morphology. Male genitalia. Uncus small; gnathos hook stout, distally narrowing, curved and apically pointed, about length of uncus; anterior margin of tegumen with broadly rounded emargination; valva straight, stout, basal part distinctly wider than distal part, distal part weakly contorted; saccular area setose, without separated sacculus; saccus massive, larger than tegumen, semi-oval, with weakly pointed apex, posterior margin arched, with pointed mediolateral projections, medial part with strongly sclerotised longitudinal ridge, lateral sclerites short; phallus moderately slender, medially bent, orbicular coecum, distal two-thirds slender, sclerotised ridges with 1–2 strong teeth; ductus ejucalatorius with contorted linear interior sclerotisation.
Female genitalia unknown.
Adult.
BIN BOLD:ACC5029 (n = 1). The distance to the nearest neighbour
Southern France (
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.
Adult.
BIN BOLD: ADA0203 (n = 1). The distance to the nearest neighbour
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
Adult.
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
Adult.
BIN BOLD:ADF0469 (n = 1). The distance to the nearest neighbour
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
The
External morphology. Segment 2 of labial palpus with scale brush about same length as segment 3; segment 3 shorter or as long as segment 2. Antennal scape without pecten. Wingspan (males) 9–20 mm. Forewing with 0–3 black spots, but without white costa (except in
Genitalia morphology. Male genitalia. Uncus long and moderately to very slender; gnathos hook bulky, strongly sclerotised, particularly at apex, with longitudinal grooves, straight;
anterior margin of tegumen with strongly sclerotised edge, reversed V-shaped; valva straight, apically distorted, clubbed; saccular area densely covered with setae, without or with short sacculus; saccus broadly sub-triangular, posterior margin projected with medial indentation, sclerotised medial ridge of various length; phallus stout, almost straight, with moderately inflated coecum, distal two-thirds gradually tapered, predominantly with spiny dorsal zone, exceptionally with large spine, ductus ejucalatorius twirled, with contorted linear interior sclerotisation.
Female genitalia. Papilla analis small; apophysis posterior very long; segment VIII long and slender, largely membranous; subgenital plate with strongly sclerotised sub-triangular subostial sclerotisation, anteromedially with broadly convex projection; apophysis anterior rod-like, from posterior margin of segment VIII; colliculum strongly sclerotised; signum small, rounded spiny plate.
Adult.
BIN BOLD:ADA0606 (n = 1). The distance to the nearest congeneric neighbour
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.
Adult.
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.
Adult.
BIN BOLD:ACZ8007 (n = 1). The distance to the nearest neighbour
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.
Adult.
BIN BOLD:ACT1624 (n = 1). The distance to the nearest neighbour
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
Adult. Male (Figure
Female (Figure
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
Female genitalia (Figure
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
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
Adult. Male (Figure
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
Female genitalia. Unknown.
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
Adult. Male (Figure
Female. Unknown.
Variation. The yellowish streak in the fold can be more or less distinct.
Male genitalia (Figure
Female genitalia. Unknown.
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.
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
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
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.
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
BIN BOLD:ADF1915 (n = 1). The distance to the nearest neighbour
France (Pyrénées), Northern Spain.
Host plant and early stages are unknown. The adults have been collected from the middle of June to early July at altitudes from 560 m to 1800 m.
The species name is a compound word derived from the Latin words
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
The systematic position of the
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Not available, no suitable specimen was available for barcoding.
Malta and South Spain.
Host plant and early stages are unknown. The few examined specimens were collected in March to early April at unreported altitudes.
The type locality, Salina in northern Malter is now a nature reserve, and there is thus a possibility that
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
The systematic position of the
Adult.
Not available, no suitable specimen was available for barcoding.
Southern Spain (prov. Granada).
Host plant and early stages are unknown. The type series was collected at the end of April at unreported altitudes.
The
External morphology. Segment 2 of labial palpus with scale brush as long as or longer than segment 3; segment 3 as long as segment 2. Antennal scape without pecten. Wingspan (males) 13–19 mm. Forewing with two black dots, but without white costa and white veins. Known females brachypterous.
Genitalia morphology. Male genitalia. Uncus large, suboval to mitre-shaped; gnathos hook moderately slender, weakly curved; valva stout, digitate, basally with small hump, without separate sacculus; saccus moderately small, broadly V-shaped, with usually well developed forked sclerotised ridge from posterior margin; phallus with globular coecum, stout, distal part weakly to distinctly S-curved, without dentation.
Female genitalia. Papilla analis small; apophysis posterior very long; segment VIII moderately long and slender, membranous; subgenital plate with sub-triangular subostial sclerotisation, posteriorly with pointed sclerites of variable length, anterior edge medially with distinct sub-triangular projection; apophysis anterior rod-like, from posterior margin of segment VIII, posteriorly with sclerotised widening; colliculum short; signum small, rounded spiny plate.
Adult.
BIN BOLD:ADB7270 (n = 2). Genetically variable species. The intraspecific divergence of the barcode region is considerable with 2.2%. The distance to the nearest neighbour
Bulgaria, Romania.
Host plant and early stages are unknown. The adults have been collected from late April to late May from sea level to unreported altitudes in Dobrogea mountains.
The species name refers to the Latin noun
Adult.
BIN BOLD:ACM09062 (n = 5), BIN BOLD:ACW5732 (n = 1), BOLD:ADI7972 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 3 BINs with an average distance of 1% and a maximum divergence of 2% in BIN BOLD:ACM09062. The mean intraspecific divergence is 2.2%, with a maximum divergence of 3.8% indicating possible cryptic diversity. The minimum distance to the nearest neighbour
Austria, Czech Republic, Germany (northwards to Rheinland-Pfalz and Hessen) (
Early stages are unknown.
Adult.
BIN BOLD:ABA3165 (n = 3), BIN BOLD:ACZ2904 (n = 1), BIN BOLD:ACZ3665 (n = 1), BIN BOLD:ADJ3544 (n = 2). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 4 BINs with no divergence in BIN BOLD:ABA3165 but 0.9% variation in two specimens collected together and clustered in BIN BOLD:ADJ3544. The mean intraspecific divergence is 3.7%, with a maximum divergence of 6% indicating possible cryptic diversity. The minimum distance to the nearest neighbour
Albania, Greece, Macedonia, Montenegro.
Host plant and early stages are unknown. The adults have been collected from late May to early August at altitudes between 1550 and 2400 m.
The species name refers to the brachypterous female sex and is an adjective.
Adult.
. BIN BOLD:ABA2916 (n = 3). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour
Macedonia (Korab, Sar Planina).
Host plant and early stages are unknown. The few adults known to date have been collected in late July and early August at altitudes between 2400 and ca. 2700 m.
The name refers to the valuable method of DNA barcoding in identification of this and several other species of
The
External morphology. Segment 2 of labial palpus with scale brush shorter to longer than segment 3; segment 3 shorter than segment 2. Antennal scape with a single pecten. Wingspan (males) 12–13 mm. Forewing light with two black dots or brown with three white streaks. Females unknown.
Genitalia morphology. Male genitalia. Uncus moderately small, gradually tapered with rounded apical edge; gnathos hook massive, strongly sclerotised, weakly curved at apex; tegumen with short and slightly converging sclerotised ridges in anteriomedial third; valva moderately long and slender, straight, digitate, apex rounded; saccular area moderately setose, without separated sacculus; saccus elongated sub-triangular, posterior margin shallowly arched, with weak medial emargination, medial part smooth, without sclerotised ridge, lateral sclerites short; phallus straight, with bulbous coecum, distal two-thirds slender, rod-like, with sclerotised longitudinal ridge.
The three species in this species group are overall similar in the male genitalia. However, they strongly differ in phenotypic appearance and in DNA barcode divergence and thus the assignment to one species group is tentative. Studies of the female as well as molecular analysis of additional markers will be necessary to resolve this problem satisfactorily.
Adult.
BIN BOLD:ABU7227 (n = 1). The distance to the nearest congeneric neighbour
Northern Italy (Prov. Bergamo).
Host plant and early stages are unknown. The adults have been collected in the last third of July, flying freely around low vegetation in the early morning, just after sunrise. The habitat includes mainly grazed, south-exposed slopes with rich vegetation on siliceous soil at altitudes between about 1800 to 1900 m (
Adult.
BIN BOLD:ADG6163 (n = 1), BIN BOLD:ACS0692 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 2 BINs with 4.3% divergence, based on a single specimen of each cluster. The minimum distance to the nearest congeneric neighbour
France (Dep. Alpes Maritimes).
Host plant and early stages are unknown. The few type specimens were collected in July and August at altitudes between 2250 and 2450 m.
Adult.
BIN BOLD:ACM1096 (n = 5). The intraspecific divergence of the barcode region is considerable with mean 1.7% and maximum divergence of 2.5%. The distance to the nearest neighbour
Southern Spain.
Host plant and early stages are unknown. The adults have been collected from late April to early August at altitudes from sea level to 1300 m, indicating possible bivoltinism. Some specimens were collected in salt marshes.
The species name (a noun in the genitive case) is dedicated to Jan Šumpich, Czech Republic, who collected part of the type series and numerous other
The
External morphology. Segment 2 of labial palpus long to very long, porrect; segment 3 reduced. Antennal scape with pecten consisting of one to several hairs. Wingspan (males) 13–26 mm. Forewing with longitudinal white or grey streaks, and in some species with white costa, but without black spots. Known females (wingspan 10–30 mm) vary from fully winged to brachypterous.
Genitalia morphology. Male genitalia. Uncus longer than broad, with parallel outer margin, apex rounded; gnathos hook stout, massive, rarely moderately slender, curved, apically distinctly pointed; tegumen with broadly emarginated anterior margin, anteromedially with sclerotised ridge; valva extending to base of uncus, digitate, basally inflated, distal part tapered, apically weakly inflated, rounded, exceptionally narrowing; sacculus absent; saccus sub-triangular, longer than broad, posterior margin sinusoid, medial part smooth, without sclerotised ridge; phallus with inflated coecum, distal part with long sclerotised distodorsal ridge, with variable extent of dentation, ductus ejaculatorius without specialised sclerotisations.
Female genitalia. Papilla analis moderately small to large, apically rounded, rarely pointed; apophysis posterior of modest length, stout to moderately slender, sometimes with widening; segment VIII frequently with sclerotised posteriolateral part; subgenital plate with sub-triangular subostial sclerotisation, posteriorly with pointed sclerites of variable length, anterior edge medially with rounded to sub-triangular projection; apophysis anterior rod-like, extending from sclerotised widening at posterior margin of segment VIII, or reduced to free apical part; short colliculum or longer antrum present; signum of variable size, suboval spiny plate.
The majority of species of the
Adult.
BIN BOLD:ADF1917 (n = 2). The intraspecific divergence of the barcode region is low with maximum divergence of 0.6%. The distance to the nearest congeneric neighbour
North-western Morocco.
Host plant and early stages are unknown. The small type series was collected in middle of May at an altitude of 1125 m.
The species name (a noun in the genitive case) is dedicated to Jukka Tabell, Finland, who collected the type series of this species and significantly contributed to our work with most valuable material.
Adult.
BIN BOLD:ACF7111 (n = 5). The intraspecific divergence of the barcode region is moderate with mean 0.7% and maximum divergence of 1.1%. The distance to the nearest neighbour
Southern France, Spain.
Host plant and early stages are unknown. The adults have been collected from late April to late June at light at altitudes up to 1400 m.
This species is named after one of its places of occurrence: the Latin
Adult.
BIN BOLD:ACZ9288 (n = 2). The intraspecific divergence of the barcode region is low with maximum divergence of 0.2%. The distance to the nearest neighbour
Southern Spain.
Host plant and early stages are unknown. The adults have been collected from late May to late June at light at altitudes up to 1650 m.
Adult.
Not available, barcoding failed.
Libya, Morocco (Middle Atlas).
Host plant and early stages are unknown. The adults have been collected from late March to the end of April at light, at higher altitudes in late June.
This species is named after its place of occurrence, Libya. The name is an adjective.
Females from Morocco fully agree with material from Libya in the genitalia, whereas one male is slightly different in having a shorter saccus.
Adult.
BIN BOLD:ADA0605 (n = 1). The distance to the nearest neighbour
Libya, Morocco, Spain (Canary Islands: Fuerteventura), Tunisia.
Host plant and early stages are unknown. The few specimens known to date were collected from middle of February to early May April at altitudes from 10 to 1125 m.
Adult.
BIN BOLD:ACE2688 (n = 3). The intraspecific divergence of the barcode region is moderate with mean 0.9% and maximum divergence of 1.4%. The distance to the nearest neighbour
Czech Republic, Russia (S Ural).
Host plant and early stages are unknown. The type series was collected in June. The adults fly at night and come to artificial light. The habitat is grassy steppe (
Adult.
BIN BOLD:ACM1349 (n = 1). The distance to the nearest neighbour
Russia (S Ural), Ukraine (Bidzilya et al. 2011: 65).
Host plant and early stages are unknown. The type series were collected by artificial light from early June to early August at grassy lowland steppes (
Kazakhstan. 2 ♂, 20 km E Chelkar settl., Bolshoe Barsuk sands, 185 m, 16.v.2012, leg. K. Nupponen; 1 ♂, near Zhababulak vill., Kumzhargan sands by Emba river, 17.v.2012, leg. K. Nupponen, genitalia slide 17/1490 ♂ Huemer; 8 ♂, 17 km NE Emba vill., 300 m, 18.v.2012, leg. K. Nupponen; 1 ♂, same data, but 20.v.2012; 4 ♂, 45 km NE Zhana-Uzen town, Bostankum sands, 160 m, 26.v.2011, leg. K. Nupponen; 1 ♂, Sengirkum sands, Terekurpa well, 70 m, 27.v.2011, leg. K. Nupponen (
Adult.
BIN BOLD:ACB3210 (n = 4). The intraspecific divergence of the barcode region is moderate with mean 0.7% and maximum divergence of 1.4%. The distance to the nearest neighbour
Kazakhstan, Russia (S Ural).
Host plant and early stages are unknown. The adults have been collected in June at low altitudes.
Adult.
Not available, no suitable specimen was available for barcoding.
Iran, Israel/Palaestine.
Host plant and early stages are unknown. The few adults known to date have been collected in early April and early June at unreported altitudes.
Adult.
BIN BOLD:ACB0437 (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
Hungary, Kazahkstan, Russia (S Ural, Volgograd oblast), Turkey. According to
Early stages are unknown. The larva is stated to feed in stems of
Croatia. 1 ♂, Konjevrate, 6.vi.2005, leg. Z. Tokár; 1 ♂, 5 km SE Pirovac, 24.vi.2006, leg. Z. Tokár (all
Adult.
BIN BOLD:ADB9039 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour
Croatia, France, Italy, Spain.
Host plant and early stages are unknown. The adults have been collected from late May to late June in halophytic habitats (with a single specimen stated to have been collected at an altitude of 1000 m).
Austria. 2 ♂, Burgenland, Weiden am See S, Zitzmannsdorfer Wiesen, 24.vi.1961, leg. F. Kasy (
Adult.
BIN BOLD:ADB8683 (n = 1). The distance to the nearest neighbour
Austria, Hungary, Romania, Russia (S Ural, Altai mts), Ukraine.
Host plant and early stages are unknown. The adults have been collected from early June to early July at low altitudes.
Adult.
BIN BOLD:ADJ1190 (n = 1). The distance to the nearest neighbour
South-western Kazakhstan.
Host plant and early stages are unknown. The holotype was collected in late May in a sandy area at a low altitude of 70 m.
This species is named after its place of occurrence: the republic of Kazakhstan in Central Asia. The name is a masculine adjective.
Adult.
Not available, barcoding failed.
Spain (Canary Islands: Gran Canaria).
Host plant and early stages are unknown. The holotype was collected in the second half of July at an unreported altitude, and the paratype was collected in June in an altitude of 700 m.
The name (a noun in the genitive case) is dedicated to Knud Larsen, Denmark, who collected the holotype of this species and other valuable material for our study.
The systematic position of
The
External morphology. Segment 2 of labial palpus with scale brush longer than segment 3; segment 3 shorter than segment 2. Antennal scape with several long hairs. Wingspan (males) 17–21 mm. Forewing grey-brown with three black spots, or yellowish grey with white veins and no black spots. Females unknown.
Genitalia morphology. Male genitalia. Uncus broadly sub-rectangular; gnathos hook stout, longer than uncus, bent at about one-quarter, apex pointed; anterior margin of tegumen curved, sclerotised ridges joined medially in anterior part; valva digitate, weakly curved, distally slightly narrowing, apex rounded; saccular area without separated sacculus; saccus sub-triangular, basally broad, distal half tapered, with pointed apex, posterior margin arched, sinusoid, without medial emargination, medial part smooth, without sclerotised ridge; phallus straight, coecum inflated, distal two-thirds slender, with sclerotised dorsal and ventral zones, medially with small sclerotised patch, ductus ejaculatorius with double-twisted interior sclerotisation.
Adult.
Not available, no specimen was available for barcoding.
Kyrgyzstan (Alai Mountains).
Host plant and early stages are unknown. No specific collecting data of the type-series were published.
Adult.
Not available, barcoding failed.
South-eastern Kazahkstan.
Host plant and early stages are unknown. The holotype was collected at light in July on a steppe slope with
The species name is a compound word derived from the Latin words
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
The systematic position of the
Adult.
Not available, barcoding failed.
Morocco (High Atlas).
Host plant and early stages are unknown. The holotype was collected in mid August at an altitude of ca. 2000 m.
The species name is a compound word derived from the Latin words
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Adult.
BIN BOLD:ADF1285 (n = 1). The distance to the nearest neighbour, an unidentified
Morocco (High Atlas).
Host plant and early stages are unknown. The adults have been collected from the first half of June until late July at altitudes from ca. 1700 m to ca. 2800 m.
The species name is derived from combining of the Latin words
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Adult.
BIN BOLD:ABW9450 (n = 2), BOLD:ADB7271 (n = 1), BOLD:ACB3182 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 3 BINs with maximum divergence of 4.6% within all clusters. The minimum distance to the nearest neighbour
South-eastern Europe, with confirmed records from Austria, Bulgaria, Greece, Hungary, Russia (S. Ural, Caucasus), Romania, and Slovakia. A record from France (
Host plant and early stages are unknown. The type series of
The
External morphology. Segment 2 of labial palpus with scale brush longer than segment 3; segment 3 half as long to same length as segment 2. Antennal scape with pecten consisting of one to 10 hairs. Wingspan (males) 10–18 mm. Forewing with 2–3 black spots, in some species with longitudinal white or grey stripes. Known females vary from fully winged to brachypterous.
Genitalia morphology. Male genitalia. Uncus moderately large, sub-square to sub-rectangular; gnathos hook slender, curved to strongly bent in basal half; valva stout, digitate, without separate sacculus; saccus massive, U- to V-shaped, with longitudinal sclerotised ridge from posterior margin almost to apex, lateral sclerites of saccus apically often strongly bulged; phallus with distinctly inflated coecum, distal part long and slender, variably dentated, ductus ejaculatorius without specialised sclerotisation.
Female genitalia. Papilla analis medium-sized to large, apically rounded, predominantly membranous; apophysis posterior slender rod-like, short to moderate length, with small sclerotised posterior zone; segment VIII short, large sclerotised dorso- and ventrolateral zone, medially largely membranous, with or without microtrichia; subgenital plate with sub-triangular subostial sclerotisation, posteriorly extended into shortly pointed sub-medial sclerites, or subostial region membranous; apophysis anterior moderately stout, about length of segment VIII, posteriorly broadly connected with segment VIII by sclerotised band; colliculum short, sclerotised; signum small, weak spiny plate, or reduced.
Adult.
BIN BOLD:ACM0852 (n = 1) The distance to the nearest neighbour
Russia (S. Ural, Tuva rep.).
Host plant and early stages are unknown. The adults have been collected in early June at altitudes of ca. 700 m.
Adult.
BIN BOLD:ACB0748 (n = 1). The distance to the nearest neighbour
Russia (Buryatia rep.).
Host plant and early stages are unknown. The adults have been collected from late May to early June in forest steppe and sandy habitats at altitudes from ca. 550 to 700 m.
The species name (a noun in the genitive case) is dedicated to Kari Nupponen, Finland, who collected the type series of this species and significantly contributed to our work with extremely valuable material from Russia.
BIN BOLD:ACA8764 (n = 1). The distance to the nearest neighbour
Southern Bulgaria.
Host plant and early stages are unknown. The type-series was collected in late July at an altitude between 1600 m and 1700 m.
Adult.
BIN BOLD:ACE2700 (n = 3), BOLD:ACB0770 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is low in one cluster, with mean 0.1% and maximum divergence of 0.2% and unknown in the second cluster. The maximum distance between both clusters is 4.2%, indicating possible cryptic diversity, whereas the distance to the nearest neighbour
South-western Russia.
Host plant and early stages are unknown. The adults have been collected during July and August in steppe grassland at unreported altitudes (
Adult.
BIN BOLD:ADG5879 (n = 1). The distance to the nearest neighbour
Kyrgyzstan (Alai mountains).
Host plant and early stages are unknown. The few adults known to date were collected from the middle of July to the end of July at altitudes from ca. 2700 to 2900 m.
Adult.
BIN BOLD:ADI8386 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour
Kyrgyzstan (Alai mts).
Host plant and early stages are unknown. The type-series was collected in the last third of July at high altitudes of ca. 3600 to nearly 3700 m.
The species (a noun in the genitive case) name is dedicated to Alexander Pototski, Estonia, who made valuable material of
Adult.
BIN BOLD:ACB3260 (n = 10). The intraspecific divergence of the barcode region is low with mean 0.03% and maximum divergence of 0.2%. The distance to the nearest neighbour
Asian part of Russia, Mongolia.
Host plant and early stages are unknown.
Adult.
Not available, no specimen was available for barcoding.
Russia (S. Ural).
Host plant and early stages are unknown. The adults have been collected from June to early August by sweeping just before sunset on chalk steppe at low altitudes from 170 to 230 m (
The
External morphology. Labial palpus long, segment 2 with long scale brush; segment 3 short and narrow. Antennal scape without or with a single, soft hair. Wingspan (males) 17–26 mm. Forewing with whitish veins, and in some species with white costa, but without black spots. Known females (wingspan 21–25 mm) with slightly narrower wings, otherwise similar to males.
Genitalia morphology. Male genitalia. Uncus sub-triangular, short, apical part slender, partially rod-like, sub-basally with or without lateral flaps; gnathos hook straight, large to medium sized, strongly sclerotised; anterior margin of tegumen broadly emarginated with small medial excavation; valva broad, without separate sacculus, apex with distinct spine; saccus sub-triangular, posterior margin strongly arched, medial part without sclerotised ridge; phallus straight, with weakly bulbous coecum, distal three-quarters slender, rod-like, with small dorsoapical tooth.
Female genitalia. Papilla analis laterally compressed, extruding from tip of abdomen, strongly sclerotised, with or without fine longitudinal lines, apically pointed; apophysis posterior and anterior moderately short; segment VIII strongly sclerotised, subgenital plate without specialised structures, except for lateral ride delimiting ostium bursae; signum small, rounded plate.
Hungary. 1 ♂, Ofen [Budapest], 1868, leg. Anker, genitalia slide Mus. Vind. 16.656; 1 ♀, same data, but 1874, genitalia slide NM 16657 (all
Adult.
BIN BOLD:ACB0458 (n = 2). The intraspecific divergence of the barcode region is low with maximum divergence of 0.2%. The distance to the nearest neighbour
Hungary, Kazakhstan, Russia (S. Ural and Volgograd oblast).
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 600 m.
Adult.
Not available, no specimen was available for barcoding.
Northern China. Recently recorded from Korea (
Host plant and early stages are unknown. The adults have been collected from May to August at unreported elevation.
Adult.
BIN BOLD:ADJ1364 (n = 2). The intraspecific divergence of the barcode region is low with mean and maximum divergence of 0.2%. The distance to the nearest neighbour
Kazakhstan.
Host plant and early stages are unknown. The adults have been collected at the beginning of June at an altitude of ca. 500 m.
This species, being known only from females, is named in honour of the feminine gender. The name is a noun in apposition.
Adult.
It resembles
BIN BOLD:AAW5664 (n = 6), BOLD:ACB3361 (n = 2). Genetically variable species. The intraspecific divergence of the barcode region is low in both clusters, with mean 0.25% and maximum divergence 0.5 in BIN BOLD:AAW5664 and maximum divergence 0.31% in the second BIN. The maximum distance between both BINs is 2.8%, indicating possible cryptic diversity. The minimum distance to the nearest neighbour
Afghanistan, Armenia, Kazakhstan, Kyrgyzstan.
Host plant and early stages are unknown. The adults have been collected from early June to the second half of July at altitudes from ca. 500 to 2900 m. In Armenia the biotope is sandy steppe.
This species is named after one of the countries of occurrence: the Kyrgyz Republic (also known as Kirgizia or Kyrgyzstan). The name is a masculine adjective.
Adult.
BIN BOLD:ACB3166 (n = 5). The intraspecific divergence of the barcode region is moderate with mean 1.2% and maximum divergence 2.2%. The distance to the nearest neighbour
Armenia, Iran, Kyrgyzstan, Romania, Russia (S. Ural), Turkey. Also recorded from China (Xinjiang: Kuldja) (
Host plant and early stages are unknown. The adults have been collected from the second half of June to early August at altitudes ranging from lowland in Europe to 2550 m in Turkey.
This species has a distinct geographical variation: Specimens from southern areas (Armenia, Turkey and Iran) are larger on average and have lighter, slightly broader forewings with no brown area in the middle of the forewing compared with specimens from the north (Romania, Russia and Kyrgyzstan). The male genitalia from Iran are similar to those of a specimen from Ukraine (apart from a differently shaped saccus). Unfortunately we have no DNA barcode data of specimens from the south-eastern part of the distribution area, and therefore we leave it open for future studies as to whether one or two species or subspecies are involved.
The
External morphology. Segment 2 of labial palpus with short or long scale brush; segment 3 as long as or longer than segment 2. Antennal scape without or with a single, soft hair. Wingspan (males) 12–23 mm. Forewing with or without white costa and whitish veins and 2–3 black spots. Known female brachypterous.
Genitalia morphology. Male genitalia. Uncus large, sub-quadrate, apical corners rounded; gnathos hook moderately slender, about length of uncus, bent at right angles at one-quarter, apicaly pointed; anterior margin of tegumen with moderately shallow semi-oval emargination; pedunculi weakly demarcated, suboval; valva straight, stout, broadly digitate, extending to base of uncus, distal area covered with setae, without separated sacculus; posterior margin of vinculum with rounded lateral humps; saccus broadly suboval, with rounded apex, ratio maximum width to length nearly 1, posterior margin arched, with shallow medial emargination, medial part without sclerotised ridge, lateral sclerites moderately short; phallus moderately slender, stout, almost straight, suboval coecum weakly defined, digitate distal part with small dorsomedial tooth.
Female genitalia. Undescribed (see below).
The two species are externally very different and also differ in several characters in the male genitalia. Therefore, and in absence of comparative females, the common assignment to a species group is tentative, though supported by DNA barcodes.
Adult.
BIN BOLD:ACZ7298 (n = 1). The distance to the nearest neighbour
North Portugal and South Spain.
Host plant and early stages are unknown. The adults have been collected in the last third of September at altitudes between 560 and 1350 m.
This species is named after its place of occurrence: the Iberian Peninsula, including Portugal and Spain. The name is a masculine adjective.
Adult.
BIN BOLD:ADI7610 (n = 1). The distance to the nearest neighbour
Libya, Tunisia.
The larva and pupa are described in details by
A single male specimen from Sbeidla, Tunisia, coll. C. S. Larsen (
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Adult.
BIN BOLD:ACM0982 (n = 6). The intraspecific divergence of the barcode region is low with mean 0.5% and maximum divergence of 1%. The distance to the nearest neighbour
Spain, Tunisia.
The larva feeds within a stem of
The ‘allotype’ of
The
External morphology. Segment 2 of labial palpus with short or long scale brush; segment 3 as long as or longer than segment 2. Antennal scape without or with one to several hairs. Wingspan (males) 10–19 mm. Forewing with 0–4 black spots, in a few species with whitish or blackish veins, but without white costa. Known female slightly to moderately brachypterous.
Genitalia morphology. Male genitalia. Uncus moderately small, basally constricted, suboval; gnathos hook stout, short, medially widened, with ventromedial ridge, weakly curved; anterior margin with broad and shallow emargination, additional small emargination medially; valva long, nearly extending to apex of uncus; sacculus usually well developed, short and digitate, rarely reduced; saccus broadly V- to U-shaped, with or without short medial ridge; phallus stout, weakly separated coecum, distal part short and tapered, with sclerotised dorsal ridge, ductus ejaculatorius with long interior sclerotisation.
Female genitalia. Papilla analis large, apex broadly rounded; apophysis posterior rod-like, short, approximately 1.5 times length of papilla analis; segment VIII short, posteriolaterally sclerotised, medially membranous; subgenital plate with sub-triangular, subostial sclerotisation, broad and shallow projection anteriorly, sub-medial sclerites shortly pointed; apophysis anterior rod-like, about length of segment VIII; colliculum short, weakly sclerotised; signum small to medium-sized, spiny plate.
Adult.
Not available, barcoding failed.
France (Dep. Alpes-de-Haute-Provence).
Host plant and early stages are unknown. The type-series was collected in the middle of August at an altitude of ca. 900 m; a further specimen dates from early August.
Adult.
Not available, barcoding failed.
Morocco (High Atlas).
Host plant and early stages are unknown. The holotype was collected in early August at an altitude of ca. 2800 m.
The species name (a noun in the genitive case) is dedicated to Axel Steiner, Germany, who collected the holotype.
. Despite the limited diagnostic genitalia characters and the absence of molecular data, the separate species status is supported by the colour and pattern of the forewing.
Adult.
Not available, barcoding failed.
Algeria, Tunisia.
Host plant and early stages are unknown. The few specimens known were collected in October. The locality of the holotype is an oasis at the northern border of the Sahara.
The species (a noun in the genitive case) name is dedicated to Christian Gibeaux, France, in recognition of his invaluable help with dissection and photographs of type material from
The two paratypes from Algeria were found in the collection of P. Chrétien in the MHNP. They are mounted on the same polyporus strip and bear additional labels “?
Adult.
Not available, barcoding failed.
Central Turkey.
Host plant and early stages are unknown. The small type-series was collected in early September at an altitude of 1550 m.
The species name indicates the characteristic wing pattern and is derived from a combination of the Latin words
Adult.
BIN BOLD:ABW5890 (n = 6), BOLD:ADF1918 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 2 BINs with 5.4% divergence, based on a single specimen from Greece compared with a larger cluster from Croatia to Italy. Within the latter cluster average divergence is considerable with mean 1.6% and maximum divergence of 2.4%. The minimum distance to the nearest neighbour
Croatia, Greece, Italy, Slovenia.
Host plant and early stages are unknown. The adults have been collected from early August to early September at altitudes from sea level to ca. 1300 m.
This species is named after the most northern place of its occurrence: present day South Tyrol (Italy). The name is an adjective.
The considerable barcode split in two clusters is not reflected by the morphology. In particular the length of the sacculus shows some variation, e.g., dissected specimens of both clusters have a long sacculus, whereas it is short in one specimen from Greece. The separation of
Adult.
BIN BOLD:ABA2917 (n = 3). The intraspecific divergence of the barcode region is low with mean 0.2% and maximum divergence of 0.3%. The distance to the nearest neighbour
Macedonia.
Host plant and early stages are unknown. The few adults known to date were collected from the end of July to early August in alpine grassland at an altitude from 1500 to 2100 m.
This species is named after its place of occurrence: the Korab mountain range on the border between Albania and Macedonia. The name is a masculine adjective.
Adult.
BIN BOLD:AAU1830 (n = 5). The intraspecific divergence of the barcode region is moderate with mean 1% and maximum divergence of 1.6% (n = 5). The distance to the nearest neighbour
France, Spain.
Host plant and early stages are unknown. The adults have been collected from early August (rarely from the middle of July) to the end of September from the coast to altitudes of ca. 1450 m.
Adult.
BIN BOLD:AAJ3176 (n = 2). The intraspecific divergence of the barcode region is low with mean and maximum divergence of 0.2%. The distance to the nearest neighbour
Italy (Orobian Alps to Monte Baldo).
Host plant and early stages are unknown. Adults have been collected during the daytime by sweeping the vegetation with a net. Males are also attracted to light. The flight period ranges from the middle of May to the middle of August, depending on altitude. The habitat is characterised by scree and rock formations with alpine grassland vegetation at altitudes of between 1200 and 2100 m. A single specimen from the lowland locality Garda (
The
External morphology. Labial palpus long, porrect; segment 3 reduced. Antennal scape without pecten. Wingspan (males) 10–17 mm. Forewing white with grey or blackish scales forming longitudinal streaks especially in apical part and with 1–2 black dots; base of costa white. Female unknown.
Genitalia morphology. Male genitalia. Uncus moderately large, apically rounded; gnathos hook strongly sclerotised, stout, weakly curved; valva digitate, extraordinarily long, extending far beyond apex of uncus, apical quarter weakly curved, separated saccular area weakly bulged but without clearly separated sacculus; saccus sub-triangular, with pointed apex, moderately long posterior margin weakly arched, medial part smooth; phallus slender, straight, coecum weakly defined, distal three-quarters digitate, without specialised structures.
Female genitalia. Papilla analis large, apex broadly rounded; apophysis posterior short, about length of papilla analis; segment VIII short, posteriolaterally sclerotised, medially membranous; subgenital plate reduced to proximally convex subostial sclerotisation, laterally connected with base of apophysis anterior by sclerotised edge, sub-medial sclerites short, anteromedial edge of segment VIII membranous; apophysis anterior short, about length of segment VIII; colliculum short, sclerotised; signum small, irregularly shaped plate with two strong lateral teeth.
Adult.
BIN BOLD:ACM0982 (n = 5). The intraspecific divergence of the barcode region is low with mean 0.3% and maximum divergence of 0.6%. The distance to the nearest neighbour
Spain.
Host plant and early stages are unknown. The adults have been collected from the second half of June to early August at altitudes from lowland to 2130 m.
The species name (a noun in the genitive case) is dedicated to Bjarne Skule, Denmark, who collected part of the type-series of this species and numerous other
The species status of the closely allied and allopatric
Adult.
BIN BOLD:ADF1825 (n = 2). The intraspecific divergence of the barcode region is moderate with mean 0.9%. The distance to the nearest neighbour
Morocco (Middle Atlas and High Atlas).
Host plant and early stages are unknown. The type material was collected in the middle of April and from June to early July at altitudes ranging from ca. 1400 to 2400 m.
The species name is derived from a combination of the Latin words
DNA barcode divergence and diagnostic characters of adults including the male genitalia support a separate specific status of
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Adult.
BIN BOLD:ACC5030 (n = 4). The intraspecific divergence of the barcode region is moderate with mean 0.6% and maximum divergence of 0.9%. The distance to the nearest congeneric neighbour
Algeria, Spain.
The larva feeds within the lower part (above the earth surface) of a stem of
Although Le Cerf (op. cit.: 21) stated that his observations on the behavior of the larvae of
The
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
The systematic position of the
Adult.
Not available, barcoding failed.
Southern France (Pyrénées Orientales), northern Spain.
Host plant and early stages are unknown. The small type-series was collected from the middle of September to mid-October at altitudes of between 250 and 880 m.
The species name (a noun in the genitive case) is dedicated to Serge Peslier, France, who collected most specimens.
The
The
External morphology. Segment 2 of labial palpus with scale brush as long as or longer than segment 3; segment 3 as long as or shorter than segment 2. Antennal scape with a single hair. Wingspan (males) 15–16 mm. Forewing with 2–3 black spots, but without whitish veins and costa. Known female brachypterous.
Genitalia morphology. Male genitalia. Uncus moderately sub-rectangular, with broadly curved or indented apex; gnathos hook stout, sub-basally bent, with pointed apex; valva massive, short and broad, with proximal hump basally, distally tapered, with distinct flap-shaped ridge in distal half; saccular area densely covered with setae, without separated sacculus; saccus massive, V-shaped; phallus stout, straight, with weakly inflated coecum, distal two-thirds moderately broad, with two rod-like, weakly dentated sclerotisations or with prominent, weakly dentated thorn sub-basally.
Female genitalia. Papilla analis small, apically rounded, largely membranous; apophysis posterior long and slender; segment VIII weakly sclerotised to membranous; subgenital plate with large subostial sclerotisation, sub-triangular process from base of apophysis anterior pointed anteromedially, and with posteromedially pointed process, demarcating lateral part of ostium bursae by deeply sclerotised wall; apophysis anterior rod-like, posterior part with longitudinal sclerotised zone along entire segment VIII; colliculum short; signum a small, transverse, spiny plate.
Adult.
Not available, available specimens too old.
Western China (Pamir Mountains).
Host plant and early stages are unknown. The typeseries was collected on 8th July at an altitude of 4300 m (
Adult.
Not available, available specimens too old.
Central Afghanistan (prov. Bamyan).
Host plant and early stages are unknown. The adults have been collected at the end of July at high altitudes from 3000 to 3600 m.
The species name is derived from the Latin word
Adult.
Female genitalia (Figure
Not available, available specimens too old.
South-eastern Afghanistan.
Host plant and early stages are unknown. The small type series was collected in the last third of June at altitudes of ca. 2700 m.
The species name is derived from a combination of the Latin words
The phylogeny of numerous
The correct identification of
Large intraspecific barcode divergence as observed in
Similar to other
We have examined a vast number of
We are most grateful to Paul Hebert and the entire team at the Canadian Centre for DNA Barcoding (Guelph, Canada), whose sequencing work was enabled by funding from the Government of Canada to Genome Canada through the Ontario Genomics Institute. We also thank the Ontario Ministry of Research and Innovation and NSERC for their support of the BOLD informatics platform. PH is particularly indebted to the Promotion of Educational Policies, University and Research Department of the Autonomous Province of Bolzano - South Tyrol for helping to fund the projects “Genetische Artabgrenzung ausgewählter arktoalpiner und boreomontaner Tiere Südtirols” and “Erstellung einer DNA-Barcode-Bibliothek der Schmetterlinge des zentralen Alpenraumes (Süd-, Nord- und Osttirol)”. The authors thank the Department of Innovation, Research and University of the Autonomous Province of Bozen/Bolzano for covering the Open Access publication costs. Several Park authorities supported our field work and we thank the Ministry of the Environment and David Morichon, director of the Natural Reserve of Conat (France), for supporting an inventory of nocturnal
Stefan Heim (
Distribution catalogue
Data type: occurrence