Monograph |
Corresponding author: Peter Huemer ( p.huemer@tiroler-landesmuseen.at ) Academic editor: Erik J. van Nieukerken
© 2018 Peter Huemer, Ole Karsholt.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Huemer P, Karsholt O (2018) Revision of the genus Megacraspedus Zeller, 1839, a challenging taxonomic tightrope of species delimitation (Lepidoptera, Gelechiidae). ZooKeys 800: 1-278. https://doi.org/10.3897/zookeys.800.26292
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The taxonomy of the Palearctic genus Megacraspedus Zeller, 1839 (Lepidoptera, Gelechiidae) is revised, based on external morphology, genitalia and DNA barcodes. An integrative taxonomic approach supports the existence of 85 species which are arranged in 24 species groups (disputed taxa from other faunal regions are discussed). Morphology of all species is described and figured in detail. For 35 species both sexes are described; for 46 species only the male sex is reported, in one species the male is unknown, whereas in three species the female adult and/or genitalia morphology could not be analysed due to lack of material.
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: Megacraspedus bengtssoni sp. n. (Spain), M. junnilaineni sp. n. (Turkey), M. similellus sp. n. (Bulgaria, Romania, Turkey), M. golestanicus sp. n. (Iran), M. tokari sp. n. (Croatia), M. neli sp. n. (France, Italy), M. faunierensis sp. n. (Italy), M. gredosensis sp. n. (Spain), M. bidentatus sp. n. (Spain), M. fuscus sp. n. (Spain), M. trineae sp. n. (Portugal, Spain), M. skoui sp. n. (Spain), M. spinophallus sp. n. (Spain), M. occidentellus sp. n. (Portugal), M. granadensis sp. n. (Spain), M. heckfordi sp. n. (Spain), M. tenuiuncus sp. n. (France, Spain), M. devorator sp. n. (Bulgaria, Romania), M. brachypteris sp. n. (Albania, Greece, Macedonia, Montenegro), M. barcodiellus sp. n. (Macedonia), M. sumpichi sp. n. (Spain), M. tabelli sp. n. (Morocco), M. gallicus sp. n. (France, Spain), M. libycus sp. n. (Libya, Morocco), M. latiuncus sp. n. (Kazahkstan), M. kazakhstanicus sp. n. (Kazahkstan), M. knudlarseni sp. n. (Spain), M. tenuignathos sp. n. (Morocco), M. glaberipalpus sp. n. (Morocco), M. nupponeni sp. n. (Russia), M. pototskii sp. n. (Kyrgyzstan), M. feminensis sp. n. (Kazakhstan), M. kirgizicus sp. n. (Afghanistan, Kazakhstan, Kyrgyzstan), M. ibericus sp. n. (Portugal, Spain), M. steineri sp. n. (Morocco), M. gibeauxi sp. n. (Algeria, Tunisia), M. multipunctellus sp. n. (Turkey), M. teriolensis sp. n. (Croatia, Greece, Italy, Slovenia), M. korabicus sp. n. (Macedonia), M. skulei sp. n. (Spain), M. longivalvellus sp. n. (Morocco), M. peslieri sp. n. (France, Spain), M. pacificus sp. n. (Afghanistan), and M. armatophallus sp. n. (Afghanistan). Nevadia Caradja, 1920, syn. n. (homonym), Cauloecista Dumont, 1928, syn. n., Reichardtiella Filipjev, 1931, syn. n., and Vadenia Caradja, 1933, syn. n. are treated as junior synonyms of Megacraspedus. Furthermore the following species are synonymised: M. subdolellus Staudinger, 1859, syn. n., M. tutti Walsingham, 1897, syn. n., and M. grossisquammellus Chrétien, 1925, syn. n. of M. lanceolellus (Zeller, 1850); M. culminicola Le Cerf, 1932, syn. n. of M. homochroa Le Cerf, 1932; M. separatellus (Fischer von Röslerstamm, 1843), syn. n. and M. incertellus Rebel, 1930, syn. n. of M. dolosellus (Zeller, 1839); M. mareotidellus Turati, 1924, syn. n. of M. numidellus (Chrétien, 1915); M. litovalvellus Junnilainen, 2010, syn. n. of M. imparellus (Fischer von Röslerstamm, 1843); M. kaszabianus Povolný, 1982, syn. n. of M. leuca (Filipjev, 1929); M. chretienella (Dumont, 1928), syn. n., M. halfella (Dumont, 1928), syn. n., and M. arnaldi (Turati & Krüger, 1936), syn. n. of M. violacellum (Chrétien, 1915); M. escalerellus Schmidt, 1941, syn. n. of M. squalida Meyrick, 1926. Megacraspedus ribbeella (Caradja, 1920), comb. n., M. numidellus (Chrétien, 1915), comb. n., M. albella (Amsel, 1935), comb. n., M. violacellum (Chrétien, 1915), comb. n., and M. grisea (Filipjev, 1931), comb. n. are newly combined in Megacraspedus.
Brachyptery, DNA barcoding, Gelechiidae , Lepidoptera , Megacraspedus , new species, Palearctic, taxonomy
The Gelechiidae are a mega-diverse family of Lepidoptera with approximately 4,700 described species (van
Megacraspedus Zeller, 1839 is one of these widely neglected genera of Palearctic Gelechiidae, unknown to the vast majority of lepidopterists. Among experts in Gelechiidae it is considered as one of the taxonomically most difficult genera and few have dealt with it in the last 100 years, usually with isolated descriptions of single species (e.g.,
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 Megacraspedus from various institutional and private collections (see below). Most of the material was traditionally pinned, set and dried or alternatively spread; few specimens are only pinned. Genitalia preparations followed standard techniques (
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 Megacraspedus were processed at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph) to obtain DNA barcodes using the standard high-throughput protocol described in
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
Abbreviations of private and institutional collections
ECKU Collection of Ecology-Centre, Kiel University, Germany
RCBB Research Collection Bengt Åke Bengtsson, Färjestaden, Sweden
RCAB Research Collection Aleksei Bidzilya, Kiev, Ukraine
RCAP Research Collection Alexander Pototski, Tallinn, Estonia
RCAW Research Collection Andreas Werno, Nunkirchen, Germany
RCCDL Research Collection Carmello De Lucca, Malta
RCEA Research Collection Ernst Arenberger, Vienna, Austria
RCER Research Collection Emili Requena Miret, Gurb, Spain
RCGB Research Collection Giorgio Baldizzone, Asti, Italy
RCGT Research Collection Giovanni Timossi, Oderzo, Italy
RCHW Research Collection Hugo van der Wolf, Nuenen, The Netherlands
RCJD Research Collection Jordi Dantart, Barcelona, Spain
RCJJ Research Collection Jari Junnilainen, Vantaa, Finland
RCJN Research Collection Jacques Nel, La Ciotat, France
RCJR Reseach Collection Jorge Rosete, Louriçal, Portugal
RCKO Research Collection Sándor Kovács & Zoltán Kovács, Miercurea Ciuc, Romania
RCKN Research Collection Kari Nupponen, Espoo, Finland
RCMC Research Collection Martin Corley, Faringdon, U.K.
RCPF Research Collection Per Falck, Nexø, Denmark
RCPT Research Collection Paolo Triberti, Verona, Italy
RCRH Research Collection Robert J. Heckford, Plympton, Plymouth, U.K.
RCSP Research Collection Serge Peslier, Perpignan, France
RCTM Research Collection Toni Mayr, Feldkirch, Austria
RCTV Research Collection Thierry Varenne, Nice, France
RCTZ Research Collection Tomasz Rynarzewski, Poznań, Poland
RCZT Research Collection Zdenko Tokár, Šal’a, Slovakia
RCWS Research Collection Willibald Schmitz, Bonn, Germany
The systematics of the Anomologinae on tribal and generic level is still imperfectly known, and it is outside the scope of the present study to determine the exact phylogenetic placement of Megacraspedus. However, there is an overall agreement about the systematic position of Megacraspedus.
Megacraspedus Zeller, 1839
Megacraspedus Zeller, 1839: 189.
Type species: Ypsolophus (Megacraspedus) dolosellus Zeller, 1839. Designated by
Chilopselaphus Mann, 1867: 850.
Type species: Chilopselaphus fallax Mann, 1867. By monotypy.
Chilopselaphalus Rebel, 1901: 161 (incorrect subsequent spelling).
Toxoceras Chrétien, 1915: 329, fig. 5 (homonym). By monotypy.
Type species: Toxoceras violacellum Chrétien, 1915.
Toxidoceras Chrétien, 1923: 168 (objective replacement name of Toxoceras Chrétien).
Type species: Toxoceras violacellum Chrétien, 1915.
Nevadia Caradja, 1920: 117 (homonym) syn. n. By monotypy.
Cauloecista Dumont, 1928: 33 syn. n.
Type species: Cauloecista chretienella Dumont, 1928. By original designation.
Reichardtiella Filipjev, 1931: 167 syn. n.
Type species: Reichardtiella grisea Filipjev, 1931. By monotypy.
Vadenia Caradja, 1933: 94 syn. n.
Type species: Nevadia ribbeella Caradja, 1920 (objective replacement name of Nevadia Caradja).
Megacraspedus was described as a subgenus of the genus Ypsolophus Fabricius, 1798 by Zeller and included two species: “striatellus S.V.” [= Isophrictis striatella ([Denis and Schiffermüller], 1775)] and “dolosellus FR.” [= Megacraspedus dolosellus (Zeller, 1839)]. The latter was subsequently designated as the type species of Megacraspedus (Walsingham 1909: 21).
Chilopselaphus was described by Mann for his new species C. fallax. Among the several details stated to be peculiar to Chilopselaphus were the long labial palps, whereas the venation of the hindwings was stated to be similar to that of Megacraspedus (
Toxoceras was described by Chrétien to include a single species T. violacellum from North Africa. It was compared with the genera Paltodora Meyrick, 1894 and Ypsolophus, but not with Megacraspedus. It was listed as a synonym of the latter by
Nevadia was described by Caradja to include a single species N. ribbeella. It was compared with Megacraspedus and Chilopselaphus, but stated to differ in the shape of the forewing and its cell. According to its genitalia it is nested within the M. fallax species group. Nevadia Caradja is a homonym of Nevadia Walcott, 1910 (Trilobita). The objective replacement name is Vadenia
Cauloecista was described by Dumont to include C. chretienella (genus type) and C. halfella from Tunisia. Our studies have shown that both are synonyms of M. violacellum (see above under Toxoceras). See also the section on Formal synonymisations.
Reichardtiella was described by Filipjev to include a single species R. grisea. It was compared with Megacraspedus and Pycnostola, but stated to differ in details of wing venation. Together with M. pacificus sp. n. and M. armatophallus sp. n., M. grisea forms our M. grisea species group. Meyrick (1926) described two species (M. neurophanes and M. niphorrhoa) in the genus Trichembola Meyrick, 1918. However, the type species of Trichembola has no uncus in the male genitalia (K Sattler in litt.) and is thus not a synonym of Megacraspedus.
Neda Chambers, 1874, with the type species Neda plutella Chambers, 1874 from North America, is a homonym of Neda Mulsant, 1850 (Coleoptera) which was later replaced by Autoneda Busck, 1903 (
Adults. Small to large gelechiids (wingspan 8–26 mm). Segment 2 of labial palpus with brush of long scales, these being most often light at base and becoming darker towards tip, especially on lower and outer surface; segment 3 as long as or shorter than segment 2, sometimes largely reduced, slender, either pointed upwards or porrect and more or less hidden in the scales of segment 2 (Figure
Wing venation (Figure
Wing venation of Megacraspedus. a M. lagopellus Herrich-Schäffer, 1860 – male, Armenia, (
Male pregenital segments. Sternite and tergite VIII without distinct modifications.
Male genitalia (Figure
Female pregenital segments. Sternite and tergite VII without distinct modifications, except for M. monolorellus with strongly elongated sternite and tergite VII.
Female genitalia (Figure
The early stages of most Megacraspedus species are unknown. Descriptions of larvae have only been published for three species (M. dolosellus, M. peyerimhoffi, and M. violacellum), although they are very detailed (and all dating back to the first part of the 20th century). There are host plant records for a few further species. All confirmed host plants belong in the Poaceae. The larvae live within the rhizomes or subterranean stem of ‘soft’ grasses like Elymus repens (L.) Gould, Poa annua L. or Lolium perenne L. (M. dolosellus), or in the lower stem of the more ‘stiff’ grass Macrochloa tenacissima (Loefl. ex L.) (M. peyerimhoffi and M. violacellum). The larva of M. dolosellus has been reported as harmful to wheat (Triticum aestivum L.) (
Megacraspedus species occur at altitudes ranging from sea level to high mountains (2500 m in the Alps, Sierra Nevada, and Turkey, 2700 m in the Balkans, 3200 m in the Atlas Mountains, and up to 4300 m in the Pamir Mountains of Central Asia (M. grisea). The habitats are different kind of grassland (e.g., halophytic meadows, grassy steppes, or alpine grassland). Adults have been collected in all months from February to November, but there is no evidence that any species have more than one annual generation. Most species are active during the night and are attracted to artificial lights. Several species seem to be active also in the early morning, where females are reported to run up and down grass stems, apparently calling for males, and they can occasionally been found in copula on the stems (Huemer pers. obs.,
The genus Megacraspedus is widely distributed throughout the south-western part of the Palearctic region (Supplementary material
With 25 species, Spain has the most diverse fauna of Megacraspedus, followed by Russia (14 species) and France (12 species). No Megacraspedus species were found from Switzerland, although several species occur in the adjacent parts of France and Italy, and the genus is not listed in the checklist of Swiss Lepidoptera (
The present revision results in many changes and new country records, e.g., the most recent checklist of the Lepidoptera of Spain (
Pecten. Several species of Megacraspedus have a pecten at the base of the antennal scape. This may consist of one, or from two to about ten hair-like scales. The number of such scales seems to be consistent within each species. In worn specimens (obtained e.g., by sweeping or from automatic light traps) one or rarely both pectens may be lost. A pecten usually consists of a row of long erect scales. It is found in most families of the Gelechioidea and may be a groundplan character of this superfamily. It is rare within the Gelechiidae where it is known from members of the Apatetrinae and Anomologinae. In the former subfamily the pecten consists of a row of long erect scales, and is present in both the Apatetrini and Pexicopini. In the Anomologinae a pecten is present in several genera, either in all species within a genus (e.g., Bryotropha Heinemann, 1870 where it is a single, long and strong, narrow scale) or only in single species of, e.g.,, Aristotelia Hübner, 1825 or Monochroa Heinemann, 1870 where it is a more deciduous hair-like scale (
Labial palpus. Two different types of labial palpi are present in Megacraspedus (Figure
Wing venation. The overall rather uniform wing venation within Gelechiidae was in former days used as an important character for classifying the Gelechiidae (e.g.,
Wing reduction. Females of many Megacraspedus species are brachypterous. According to
Brachyptery in Lepidoptera is not a uniform phenomenon but has various reasons which should be looked at on their own merit (
Female wing reduction in Lepidoptera is rarely polymorphic. One classical example is the aquatic moth Acentria ephemerella (Denis & Schiffermüller, 1775) (Crambidae), which has both winged and wingless females.
Female genitalia. Female genitalia are only known for 34 of 85 species, and even for several species groups they are currently undescribed. Therefore the description of female genitalia characters on a generic level is tentative.
COI sequences in Megacraspedus. Intraspecific mean K2P (Kimura 2 parameter) divergences, maximum pairwise distances, and distance to the nearest neighbour in percentage.
Species | Mean Intra-Sp | Max Intra-Sp | Nearest Neighbour | Nearest Species | Distance to NN |
---|---|---|---|---|---|
M. albovenata | 0.93 | 1.39 | LEAST584-17 | M. kazakhstanicus | 5.09 |
M. andreneli | 4.45 | 4.45 | PHLAH795-12 | M. tokari | 11.22 |
M. argyroneurellus | 1.19 | 2.24 | LEEUA729-12 | M. kirgizicus | 7.56 |
M. attritellus | 2.08 | 4.19 | LEEUA685-12 | M. leuca | 6.77 |
M. balneariellus | 0 | 0 | LEASS140-16 | M. podolicus | 6.28 |
M. barcodiellus | 0 | 0 | LEATI093-15 | M. binotella | 10.66 |
M. bengtssoni | 0.15 | 0.15 | PHLAF012-11 | M. lanceolellus | 11.99 |
M. bidentatus | N/A | 0 | PHLAH794-12 | M. cuencellus | 3.14 |
M. bilineatella | N/A | 0 | LEATJ1347-16 | M. fallax | 14.91 |
M. binotella | 2.19 | 3.81 | LEASS112-16 | M. devorator | 8.97 |
M. brachypteris | 3.74 | 5.97 | LEASS032-16 | M. dolosellus | 11.25 |
M. cerussatellus | N/A | 0 | PHLAI412-13 | M. attritellus | 7.55 |
M. consortiella | N/A | 0 | LEALT202-16 | M. leuca | 6.64 |
M. cuencellus | N/A | 0 | LEASS012-16 | M. bidentatus | 3.14 |
M. devorator | 2.16 | 2.16 | LEATI093-15 | M. binotella | 8.97 |
M. dolosellus | 7.49 | 13.76 | PHLAF014-11 | M. lanceolellus | 9.85 |
M. eburnellus | 0.15 | 0.15 | PHLAI458-13 | M. skulei | 8.46 |
M. fallax | 0.1 | 0.16 | LEAST573-17 | M. niphorrhoa | 5.56 |
M. faunierensis | 1.04 | 1.57 | LEASS022-16 | M. sumpichi | 11.88 |
M. feminensis | 0.15 | 0.15 | PHLAH788-12 | M. lagopellus | 7.77 |
M. gallicus | 0.7 | 1.13 | LEASS026-16 | M. ribbeella | 6.24 |
M. glaberipalpus | N/A | 0 | PHLAG308-12 | M. cerussatellus | 10.77 |
M. golestanicus | 0.1 | 0.15 | PHLAJ385-14 | M. uzunsyrtus | 8.78 |
M. gredosensis | N/A | 0 | LEASS012-16 | M. bidentatus | 10.01 |
M. heckfordi | 0.49 | 0.77 | LECRT129-16 | M. tenuiuncus | 7.74 |
M. homochroa | N/A | 0 | PHLAD606-11 | M. golestanicus | 9.1 |
M. ibericus | N/A | 0 | LEAST388-17 | M. violacellum | 7.46 |
M. imparellus | 3.38 | 4.6 | LEATJ1354-16 | M. attritellus | 9.15 |
M. junnilaineni | N/A | 0 | LECRT125-16 | M. similellus | 9.35 |
M. kazakhstanicus | N/A | 0 | LEEUA949-12 | M. albovenata | 5.09 |
M. kirgizicus | 1.2 | 2.82 | LEAST575-17 | M. argyroneurellus | 7.56 |
M. korabicus | 0.2 | 0.31 | LEASS618-17 | M. quadristictus | 10.13 |
M. lagopellus | 0.15 | 0.15 | LEAST583-17 | M. feminensis | 7.77 |
M. lanceolellus | 7.61 | 12.51 | LEASS101-16 | M. dolosellus | 9.85 |
M. leuca | 0.03 | 0.19 | LEAST594-17 | M. skulei | 6.57 |
M. longipalpella | N/A | 0 | LEAST572-17 | M. niphorrhoa | 4.78 |
M. longivalvellus | 0.93 | 0.93 | LEATJ1383-16 | M. skulei | 2.33 |
M. monolorellus | N/A | 0 | PHLAD606-11 | M. golestanicus | 9.17 |
M. multispinella | N/A | 0 | LEEUA846-12 | M. nupponeni | 8.13 |
M. niphorrhoa | 0.7 | 1.4 | PHLAI416-13 | M. longipalpella | 4.78 |
M. numidellus | N/A | 0 | LEFIJ2706-15 | M. gallicus | 8.97 |
M. nupponeni | N/A | 0 | LEEUA779-12 | M. multispinella | 8.13 |
M. peyerimhoffi | 0.56 | 0.93 | LEATJ1383-16 | M. skulei | 9.45 |
M. podolicus | N/A | 0 | LEASS136-16 | M. balneariellus | 6.28 |
M. pototskii | 0 | 0 | LEASS005-16 | M. ibericus | 9.14 |
M. pusillus | N/A | 0 | LEASS120-16 | M. spinophallus | 8.43 |
M. quadristictus | 0.97 | 1.55 | PHLAG675-12 | M. teriolensis | 9.69 |
M. ribbeella | 0.15 | 0.15 | LEFIJ2706-15 | M. gallicus | 6.24 |
M. similellus | 0.79 | 1.19 | PHLAJ385-14 | M. uzunsyrtus | 6.44 |
M. skoui | N/A | 0 | PHLSA052-11 | M. spinophallus | 7.97 |
M. skulei | 0.32 | 0.64 | LECRT099-16 | M. longivalvellus | 2.33 |
M. spinophallus | 0.8 | 2.83 | LASTS762-15 | M. skoui | 7.97 |
M. squalida | 0.45 | 0.95 | PHLAI458-13 | M. skulei | 9.3 |
M. sumpichi | 1.71 | 2.5 | PHLAG308-12 | M. cerussatellus | 11.41 |
M. tabelli | 0.62 | 0.62 | LEASS007-16 | M. numidellus | 9.82 |
M. tenuiuncus | N/A | 0 | PHLAH508-12 | M. heckfordi | 7.74 |
M. teriolensis | 2.99 | 7.45 | PHLAI454-13 | M. quadristictus | 9.69 |
M. tokari | N/A | 0 | PHLAH794-12 | M. cuencellus | 9.97 |
M. trineae | N/A | 0 | LEASS012-16 | M. bidentatus | 7.76 |
M. tristictus | N/A | 0 | LECRT129-16 | M. tenuiuncus | 11.98 |
M. uzunsyrtus | 0 | 0 | LEASS108-16 | M. similellus | 6.44 |
M. violacellum | N/A | 0 | LEASS005-16 | M. ibericus | 7.46 |
279 specimens of Megacraspedus were successfully sequenced, with 264 COI sequences longer than 500 bp. The sequenced 62 species group in different clusters (Figure
Maximum likelihood tree (built with MEGA7) of cytochrome c oxidase subunit I (COI) barcode fragments. Values on branches represent bootstrap values of ≥ 50 % inferred from 500 replicates, scale bar represents substitutions per site. Note: the scale bar only applies to internal branches between species. The width of the triangles represents the sample size, and the depth the relative genetic variation within the cluster (2× scale bar). Source: DNA Barcode data from BOLD (Barcode of Life Database, cf.
Species groups. Closely related species are treated within informal species groups. These are mainly based on male genitalia characters (females are frequently unknown), supported by molecular data of the DNA barcode (available for the majority of species), and supplemented by external features of the adults. Several of these groups, particularly those without molecular support, may not be monophyletic and similarities could occasionally reflect convergences only. Except for a few widely distributed species, the vast majority of species groups are geographically restricted, indicating a common evolutionary scenario of species.
Systematic arrangement of species. The sequence of species groups and species largely follows a maximum likelihood analysis of DNA barcode data, but also considers genitalia morphology which is partially but not fully concordant with molecular results. Species without molecular data are arranged according to morphological similarities.
Megacraspedus Zeller, 1839
Chilopselaphus Mann, 1867
Chilopselaphalus Rebel, 1901 (incorrect subsequent spelling)
Toxoceras Chrétien, 1915 (homonym)
Toxidoceras Chrétien, 1923
Nevadia Caradja, 1920 (homonym)
Cauloecista Dumont, 1928, syn. n.
Reichardtiella Filipjev, 1931, syn. n.
Vadenia Caradja, 1933, syn. n.
Megacraspedus lanceolellus species group
Megacraspedus lanceolellus (Zeller, 1850)
= M. hessleriellus Rössler, 1868
= M. subdolellus Staudinger, 1859, syn. n.
= M. tutti Walsingham, 1897, syn. n.
= M. grossisquammellus Chrétien, 1925, syn. n.
Megacraspedus bengtssoni sp. n.
Megacraspedus homochroa species group
Megacraspedus homochroa Le Cerf, 1932
= M. culminicola Le Cerf, 1932, syn. n.
Megacraspedus dolosellus species group
Megacraspedus monolorellus Rebel, 1905
Megacraspedus junnilaineni sp. n.
Megacraspedus uzunsyrtus Bidzilya & Budashkin, 2015
Megacraspedus similellus sp. n.
Megacraspedus golestanicus sp. n.
Megacraspedus tokari sp. n.
Megacraspedus dolosellus (Zeller, 1839)
= M. separatellus (Fischer von Röslerstamm, 1843), syn. n.
= M. incertellus Rebel, 1930, syn. n.
Megacraspedus faunierensis species group
Megacraspedus neli sp. n.
Megacraspedus faunierensis sp. n.
Megacraspedus gredosenis species group
Megacraspedus gredosensis sp. n.
Megacraspedus cuencellus species group
Megacraspedus cuencellus Caradja, 1920
Megacraspedus bidentatus sp. n.
Megacraspedus fuscus sp. n.
Megacraspedus trineae sp. n.
Megacraspedus pusillus species group
Megacraspedus tristictus Walsingham, 1910
Megacraspedus alfacarellus Wehrli, 1926
Megacraspedus pusillus Walsingham, 1903
Megacraspedus skoui sp. n.
Megacraspedus spinophallus sp. n.
Megacraspedus occidentellus sp. n.
Megacraspedus granadensis sp. n.
Megacraspedus heckfordi sp. n.
Megacraspedus tenuiuncus sp. n.
Megacraspedus lativalvellus species group
Megacraspedus lativalvellus Amsel, 1954
Megacraspedus dejectella species group
Megacraspedus dejectella (Staudinger, 1859)
Megacraspedus binotella species group
Megacraspedus devorator sp. n.
Megacraspedus binotella (Duponchel, 1843)
Megacraspedus brachypteris sp. n.
Megacraspedus barcodiellus sp. n.
Megacraspedus bilineatella species group
Megacraspedus bilineatella Huemer & Karsholt, 1996
Megacraspedus andreneli Varenne & Nel, 2014
Megacraspedus sumpichi sp. n.
Megacraspedus fallax species group
Megacraspedus tabelli sp. n.
Megacraspedus gallicus sp. n.
Megacraspedus ribbeella (Caradja, 1920), comb. n.
Megacraspedus libycus sp. n.
Megacraspedus numidellus (Chrétien, 1915), comb. n.
= M. mareotidellus Turati, 1924, syn. n.
Megacraspedus albovenata Junnilainen, 2010
Megacraspedus longipalpella Junnilainen, 2010
Megacraspedus niphorrhoa (Meyrick, 1926)
Megacraspedus albella (Amsel, 1935), comb. n.
Megacraspedus fallax (Mann, 1867)
Megacraspedus balneariellus (Chrétien, 1907)
Megacraspedus podolicus (Toll, 1942)
Megacraspedus kazakhstanicus sp. n.
Megacraspedus knudlarseni sp. n.
Megacraspedus majorella species group
Megacraspedus majorella Caradja, 1920
Megacraspedus latiuncus sp. n.
Megacraspedus tenuignathos species group
Megacraspedus tenuignathos sp. n.
Megacraspedus glaberipalpus species group
Megacraspedus glaberipalpus sp. n.
Megacraspedus imparellus species group
Megacraspedus imparellus (Fischer von Röslerstamm, 1843)
= M. litovalvellus Junnilainen, 2010, syn. n.
Megacraspedus consortiella species group
Megacraspedus multispinella Junnilainen & Nupponen, 2010
Megacraspedus nupponeni sp. n.
Megacraspedus cerussatellus Rebel, 1930
Megacraspedus attritellus Staudinger, 1871
Megacraspedus consortiella Caradja, 1920
Megacraspedus pototskii sp. n.
Megacraspedus leuca (Filipjev, 1929)
= M. kaszabianus Povolný, 1982, syn. n.
Megacraspedus orenburgensis Junnilainen & Nupponen, 2010
Megacraspedus lagopellus species group
Megacraspedus lagopellus Herrich-Schäffer, 1860
Megacraspedus coleophorodes (Li & Zheng, 1995)
Megacraspedus feminensis sp. n.
Megacraspedus kirgizicus sp. n.
Megacraspedus argyroneurellus Staudinger, 1871
Megacraspedus violacellum species group
Megacraspedus ibericus sp. n.
Megacraspedus violacellum (Chrétien, 1915), comb. n.
= M. chretienella (Dumont, 1928), syn. n.
= M. halfella (Dumont, 1928), syn. n.
= M. arnaldi (Turati & Krüger, 1936), syn. n.
Megacraspedus squalida species group
Megacraspedus squalida Meyrick, 1926
= M. escalerellus Schmidt, 1941, syn. n.
Megacraspedus pentheres species group
Megacraspedus pentheres Walsingham, 1920
Megacraspedus steineri sp. n.
Megacraspedus gibeauxi sp. n.
Megacraspedus multipunctellus sp. n.
Megacraspedus teriolensis sp. n.
Megacraspedus korabicus sp. n.
Megacraspedus quadristictus Lhomme, 1946
Megacraspedus eburnellus Huemer & Karsholt, 2001
Megacraspedus longivalvellus species group
Megacraspedus skulei sp. n.
Megacraspedus longivalvellus sp. n.
Megacraspedus peyerimhoffi species group
Megacraspedus peyerimhoffi Le Cerf, 1925
Megacraspedus peslieri species group
Megacraspedus peslieri sp. n.
Megacraspedus grisea species group
Megacraspedus grisea (Filipjev, 1931), comb. n.
Megacraspedus pacificus sp. n.
Megacraspedus armatophallus sp. n.
Numerous species from various regions have been described in Megacraspedus, but could not be confirmed to belong unequivocally to this genus. Although a revisionary work of these problems is outside the scope of our paper we present a short overview. Several species from the Afrotropical region were described in Megacraspedus or Chilopselaphus (Janse, 1958: 136), but none of these could be confirmed to belong in Megacraspedus. M. brachypogon Meyrick, 1937 and M. coniogramma Meyrick, 1921 are now placed in Ephysteris (Janse 1958: 136,
1 | Gnathos hook straight, massive and bulky, with longitudinal grooves (Figs |
M. pusillus species group |
– | Gnathos hook usually curved or bent, of various shape, without longitudinal grooves | 2 |
2 | Uncus apically rod-like, maximum half width of valva (Figs |
M. lagopellus species group |
– | Uncus without rod-like apical part, usually with rounded apex, two-thirds to about twice width of valva | 3 |
3 | Sub-basal part of valva with long and curved thorn-like process (Fig. |
M. squalida species group |
– | Sub-basal part of valva without process | 4 |
4 | Valva very broad, about width of uncus, with sclerotised subapical ridge (Figs |
M. grisea species group |
– | Valva slender, rarely broad but without sclerotised subapical ridge | 5 |
5 | Valva distinctly exceeding apex of uncus; phallus long and slender, without distinct sclerotisations (Figs |
M. longivalvellus species group |
– | Valva at most length of uncus; phallus of different shape, partially with sclerotisations | 6 |
6 | Uncus basally constricted, short, suboval | 7 |
– | Uncus basally not constricted, sub-rectangular to sub-triangular | 8 |
7 | Sacculus usually present; phallus distally narrowing with sclerotised, denticulate ridge (Figs |
M. pentheres species group |
– | Sacculus absent; phallus distally with broadly sinusoid dorsal margin, without denticulate ridge (Figs |
M. peyerimhoffi species group |
8 | Phallus with long and slender thorn-like sclerite from about middle nearly to apex, parallel to phallus axis (Fig. |
M. peslieri species group |
– | Phallus of different shape, without long and slender thorn | 9 |
9 | Phallus bent distad of bulbous of coecum, with long and slender distal part, 1–2 strong medial or sub-apical teeth | 10 |
– | Phallus of different shape, without sclerotisation or with groups of smaller teeth or differently shaped sclerotisation | 11 |
10 | Sacculus well developed, digitate; phallus very long (Figs |
M. gredosensis species group |
– | Sacculus absent, phallus shorter (Figs |
M. cuencellus species group |
11 | Uncus sub-quadrate, large, nearly width of posterior part of tegumen; ductus ejaculatorius with double-twisted interior sclerotisation (Figs |
M. majorella species group |
– | Uncus smaller, oblong; ductus ejaculatorius with or without interior sclerotisation of different shape | 12 |
12 | Sacculus well developed, digitate | 13 |
– | Sacculus absent | 14 |
13 | Uncus oblong 1.7 times longer than maximum width; gnathos hook weakly curved; ductus ejaculatorius with broad and short interior sclerotisation (Figs |
M. glaberipalpus species group |
– | Uncus rounded, about same maximum length as width; ductus ejaculatorius with very long and 4–5× contorted interior sclerotisation (Figs |
M. imparellus species group |
14 | Saccus large, oblong, apically usually broadly rounded, rarely pointed, with longitudinal sclerotised ridge extending almost to apex | 15 |
– | Saccus shorter, sub-triangular, smoothly sclerotised without or with short ridge of various shape | 16 |
15 | Uncus small, about width of valva; gnathos hook straight; phallus with stout and broad distal part (Figs |
M. faunierensis species group |
– | Uncus large, distinctly broader than valva; gnathos hook curved to strongly bent; phallus with oblong and usually slender distal part (Figs |
M. consortiella species group |
16 | Valva short and broad; saccus broadly suboval, with rounded apex (Figs |
M. violacellum species group |
– | Valva long, slender to moderately broad; saccus sub-triangular, with pointed apex | 17 |
17 | Saccus long and slender, ratio maximum width to length 0.6; phallus with small sub-apical spine and larger sub-apical tooth (Fig. |
M. homochroa species group |
– | 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 |
18 | Gnathos hook long and slender, extending nearly to anterior margin of tegumen (Fig. |
M. tenuignathos species group |
– | Gnathos hook shorter, not extending to anterior margin of tegumen | 19 |
19 | Gnathos hook short, stout, curved; valva apically distinctly inflated, phallus with sclerotised, usually dentated ridge (Figs |
M. fallax species group |
– | Gnathos hook moderately short to long, slender; valva apically narrowing or contorted and rounded; phallus without sclerotised, dentated ridge | 20 |
20 | Saccus with short, furcated posterior ridges (Figs |
M. binotella species group |
– | Saccus without or with short, unfurcated ridge | 21 |
21 | Phallus with S-curvature (Figs |
M. dolosellus species group |
– | Phallus without S-curvature | 22 |
22 | Gnathos hook long, bent at right angles at about two fifth (Fig. |
M. lativalvellus species group |
– | Gnathos hook shorter, nearly straight to weakly curved | 23 |
23 | Phallus with broadly lobed dorsal and rod-like ventral sclerotisation (Figs |
M. lanceolellus species group |
– | Phallus of different shape, without broadly lobed dorsal sclerotisation | 24 |
24 | Valva broad, with longitudinal ridge (Fig. |
M. dejectella species group |
– | Valva slender, without ridge (Figs |
M. bilineatella species group |
The M. lanceolellus species group includes two species: M. lanceolellus and M. bengtssoni sp. n.
External morphology. Segment 2 of labial palpus with long scale brush; segment 3 shorter or about length of segment 2. Antennal scape without pecten. Wingspan (males) 12–15 mm. Forewing light greyish or yellowish with indistinctly lighter veins and 1–2 black dots. Known females slightly brachypterous.
Genitalia morphology. Male genitalia. Uncus moderately slender, sub-triangular; gnathos hook long and stout, about length of uncus; anterior margin of tegumen deeply emarginated; valva straight, slender, with weakly contorted and rounded apical part; saccular area setose, without separated sacculus; saccus nearly V-shaped, broad, abruptly tapered distally; phallus with band-like sclerotisation of coecum, broadly lobed dorsal and rod-like ventral sclerotisation, and one to two short and broadly linear interior sclerotisation of ductus ejaculatorius. 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 long sinusoid projection; apophysis anterior rod-like, from posterior margin of segment VIII; colliculum short; signum small, rounded spiny plate.
Diagnostic remarks. The M. lanceolellus species group is defined by a unique combination of several structures in the male genitalia. The moderately large and apically narrowing uncus, the nearly straight gnathos hook, the slender and apically weakly contorted valva, the nearly V-shaped, broad saccus with a long medial ridge, and particularly the shape of the phallus with a broad dorsal lobe and a slender ventral sclerotisation as well as the linear sclerotisation of the ductus ejaculatorius are characteristic. The female genitalia overall largely agree with several taxa in other species groups.
Ypsolophus lanceolellus Zeller, 1850: 143.
Megacraspedus hessleriellus Rössler, 1868: 347.
Megacraspedus subdolellus Staudinger, 1859: 243, syn. n.
Megacraspedus tutti Walsingham, 1897: 140, syn. n.
Megacraspedus grossisquammellus Chrétien, 1925: 257, syn. n.
The examined material cannot be unequivocally attached to barcode clusters due to lack of barcode sequences from several places, or partially sympatric clusters, and is thus listed in a standardized format in the alphabetic order of countries.
Syntypes [?], 2 ♂, Ypsolophus lanceolellus, [Italy] “Livorno Mn” “Gen. Präp. Mus. Vind. 16.653 ♂” (
Adult. Male (Figs
Female (Figure
Variation. Megacraspedus lanceolellus shows both individual and also a tendency to geographical variation. The nominotypical form has light yellowish forewings with a dark shadow or streak towards the apex, but such individuals also occur elsewhere, e.g., in Spain. Specimens from south-western France and northern and central Spain (described as M. tutti and M. grossisquammellus, respectively) have a more or less pronounced white costa and can moreover often be recognized by the light yellow forewings with darker veins. Specimens from the Sierra Nevada in southern Spain (described as M. subdolellus) are larger on average (wingspan 14–20 mm) and are characterised by their more clear yellow forewings that almost lack black dots, and the whitish costa. Specimens from Montenegro have a black dot at 2/5 in the fold.
Male genitalia (Figs
Female genitalia (Figs
Due to its variation M. lanceolellus is difficult to characterise. It is a medium-sized species with light yellowish forewings, mottled with brown and black-tipped scales, a lighter costa, and sometimes one or two black dots in the middle of the wing and/or a black streak towards apex. See also M. bengtssoni sp. n. (p 37) and M. andreneli (p 96). The male genitalia are characterised by several structures such as the sub-triangular shape of the uncus, the slender and apically weakly contorted valva, the shape of the saccus with a long medial ridge, and particularly the massive distal part of the phallus with a broadly lobed dorsal and a slender ventral sclerotisation, as well as the two linear sclerotisations of the ductus ejaculatorius. They are somewhat similar to M. bengtssoni sp. n. but differ e.g., by the longer uncus, the medially strongly emarginated anterior margin of the tegumen, the longer and more slender valva, and the broader phallus.
The female genitalia are characterised by the long sinusoid anteriomedial projection of segment VIII, but in the absence of females of closely related species such as M. bengtssoni sp. n., these features may not be specifically diagnostic.
The extraordinary DNA barcode divergence is reflected by 19 BINs! The mean and maximum intraspecific divergence of the barcode region in this species is 7.6% and 12.5% respectively, largely reflecting a geographic pattern with several distinct clusters and a remarkable divergence of 4.3 to 8.1% to the nearest cluster. However, even within these clusters intraspecific variation of the barcode region is considerable with e.g., mean divergence of 2.5% in south-western alpine populations. Four specimens collected in the same microhabitat at Sierra de los Filabres (prov. Almería, Spain) form two distinct clusters with a variation of maximum 0.2% within clusters but minimum 8.6% between these clusters. Similarly diverging sympatric clusters are reported from prov. Teruel, and from Prov. Lleida.
The following 19 clusters are defined (based on sequenced material):
BIN lanc01 (Italy: Pordenone, Calabria; Croatia): BOLD:AAU2834 (n = 4).
BIN lanc02 (Italy: Calabria): BOLD:ACZ3381 (n = 1).
BIN lanc03 (Croatia): BOLD:ACZ2933 (n = 1).
BIN lanc04 (Italy: Livorno): BOLD:ACZ2657 (n = 1).
BIN lanc05 (Spain: Almeria): BOLD:ACZ9024 (n = 2).
BIN lanc06 (Andorra): BOLD:ACA9759 (n = 2)
BIN lanc07 (Spain: Barcelona): BOLD:ACZ8656 (n = 2).
BIN lanc08 (France: Alpes Maritimes): BOLD:ABA3648 (n = 1).
BIN lanc09 (Italy: Savona): BOLD:ACZ2656 (n = 1).
BIN lanc10 (France: Alpes Maritimes, Hautes Alpes, Var): BOLD:ABA3649 (n = 2).
BIN lanc11 (Spain: Almeria): BOLD:ACZ2607 (n = 2).
BIN lanc12 (Spain: Teruel, Valencia): BOLD:AAU1829 (n = 5).
BIN lanc13 (Spain: Soria): BOLD:ACZ8269 (n = 1).
BIN lanc14 (Spain: Teruel): BOLD:ADF2256 (n = 2).
BIN lanc15 (Spain: Teruel): BOLD:ACM1006 (n = 1).
BIN lanc16 (Spain: Zaragoza): BOLD:ADB7571 (n = 1).
BIN lanc17 (Spain: Zaragoza): BOLD:ADF2263 (n = 3).
BIN lanc18 (Spain: Lleida): BOLD:ADF2194 (n = 1).
BIN lanc19 (Spain: Lleida): BOLD:ADF1916 (n = 1).
The minimum distance to the nearest congeneric neighbour M. dolosellus is 9.9% (p-dist).
Southern parts of Europe, from the Balkan Peninsula to Italy, France, and Spain, also known from Germany (northwards to Rheinland-Pfalz (
Early stages are undescribed. According to Lhomme (1946: 538 as M. tutti) the larva feeds within the stem of Festuca L. The adults have been collected from late April until the middle of August, from low altitudes to 2300 m in the Alps and 2500 m in the Sierra Nevada. There is probably only one generation each year, and the flight period seems to at least partly depend on the altitude of the collecting site.
Ypsolophus lanceolellus was described from an unstated number of males collected by J. Mann at Ardenza and Salviano, Italy (Zeller 1850).
Megacraspedus hessleriellus was described from two males collected at Biebrich and Mombach in Nassau, S Germany (
Megacraspedus subdolellus was described from an unspecified number of both sexes collected in August at high altitudes (ca. 2700 m) in the Sierra Nevada, Spain (
Megacraspedus tutti was described from a single male collected in August at La Grave, SE France by JW Tutt (
Megacraspedus lanceolellus shows a remarkable morphological and genetic variation. Barcode data clearly support several putative taxa, but the morphology gives a different and less straightforward picture. Though the phenotypic appearance partly depends on geography it is frequently impossible to reliably assign specimens to a barcode cluster. Male genitalia morphology is comparatively uniform and alleged diagnostic characters show intraspecific variation. Considering the intrapopulational barcode variation which exceeds 8% on one occasion we conclude that M. lanceolellus is a widespread, mainly western-Mediterranean species. The exceptional DNA barcode divergence may be explained by weak dispersal ability, particularly of the slightly brachypterous females, leading to several genetically isolated populations which are mainly restricted to various mountain systems. However, it seems unlikely that this is the sole cause for the observed intraspecific variation (see Discussion).
Holotype ♂, “SPAIN [prov.] Aragon Gelsa 8 km NE, 240 m N41.46004 W0.37642 19.5.2016 J. Tabell leg.” “DNA Barcode
Adult. Male (Figs
Female. Unknown.
Variation. There is slight variation in the colour of the forewing (from light greyish towards light grey-brown) and in the amount of blackish scales. Some specimens have an indistinct black dot in the fold.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus bengtssoni sp. n. is characterised by its relatively small size and by its light greyish brown forewings with a single black dot at the end of the cell. It is similar to M. lanceolellus, but that species is larger, it has shorter segment 3 of the labial palps and the costa is white from base. See also M. dejectella (p 85). The male genitalia are somewhat similar to M. lanceolellus (Figs
BIN: BOLD:ACM1097 (n = 2). The intraspecific divergence of the barcode region has a maximum divergence of 0.2%. The distance to the nearest congeneric neighbour M. lanceolellus is 12% (p-dist).
Spain.
Host plant and early stages are unknown. The adults have been collected from the middle of May to the middle of July at altitudes up to 1150 m.
The species name, a noun in the genitive case, is dedicated to Bengt Å. Bengtsson, Sweden, who collected part of the type series of this species and other valuable Megacraspedus specimens used for our study.
The M. homochroa species group includes one species: M. homochroa.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Diagnostic remarks. The M. homochroa species group is defined by unique structures in the male genitalia. Particularly the long and slender saccus with a long and forked medial ridge and the sub-apical spine and larger sub-apical tooth of the phallus are diagnostic. Furthermore the broad valva is rarely present in other species groups.
Megacraspedus homochroa Le Cerf, 1932: 165.
Megacraspedus culminicola Le Cerf, 1932: 165, syn. n.
Morocco. 2 ♂, Prov. Al Haouz, Oukaïmeden 1.5 km SE, 2660 m N31.19340 W7.85311, 2.vii.2016, leg. J. Tabell, DNA Barcode
Adult. Male (Figs
Female (based on original description). Wingspan 11–16.5 mm. Forewing lanceolate, shorter than the body, hindwing narrow; both pairs almost entirely reddish at the base.
Variation. A rather variable species as regards the amount of blackish brown scales on the forewings. Specimens with many such scales look greyish yellow. Material from Middle Atlas is rather reddish-brown than yellow-brown.
Male genitalia (Figure
Female genitalia. Undescribed.
Megacraspedus homochroa is characterised by its rather large size and its light yellow-brown to reddish-brown forewings without spots. See also M. tenuignathos sp. n. (p 126). The male genitalia are unmistakable due to the combined shape of the large uncus and saccus, and the peculiar structures of the phallus.
BIN BOLD:ADF2257 (n = 1). The distance to the nearest neighbour M. golestanicus sp. n. is 9.1% (p-dist).
Morocco (Middle Atlas, High Atlas).
Host plant and early stages are unknown. The type-series of M. homochroa was collected in the Middle Atlas in August at an altitude of 2400 m, that of M. culminicola at high altitude of 3200 m in the middle of July. Additional samples date from about mid-June to early July and at altitudes from ca. 2400 to 2660 m.
Megacraspedus homochroa was described from two males collected in Morocco, Middle Atlas (
Megacraspedus culminicola was described from two males and two females taken in copula in Morocco, Middle Atlas (
The M. dolosellus species group includes seven species: M. monolorellus, M. junnilaineni sp. n., M. uzunsyrtus, M. similellus sp. n., M. golestanicus sp. n., M. tokari sp. n., and M. dolosellus.
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.
Diagnostic remarks. The M. dolosellus species group is particularly defined by the usually distinctly S-curved phallus in combination with several other characters of the male genitalia such as the long and slender gnathos hook, the moderately slender valva, and the broadly sub-triangular saccus with or without a short medial ridge.
Taxa of the M. dolosellus species group are difficult to separate by phenotypic appearance, with only subtle differences in the forewing colour and pattern. Similarly diagnostic characters of male genitalia are cryptic in some species with only subtle differences.
Megacraspedus monolorellus Rebel, 1905: 213.
Lectotype ♂, here designated, “Asia min. Penther ’02 Ilgin.” “Megacraspedus monolorellus Type Rbl.” “BC
Adult. Male (Figure
Female (Figure
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus monolorellus is characterised by its brown forewings with white costa and without further markings. It is similar to M. uzunsyrtus (p 45) and M. similellus sp. n. (p 46). The male genitalia are very similar to M. junnilaineni sp. n. (Figure
BIN BOLD:ACA8694 (n = 1). The distance to the nearest neighbour M. golestanicus sp. n. is 9.2% (p-dist).
Turkey.
Host plant and early stages are unknown. The type series was collected in the first third of May, and also the few additional specimens examined were found in May.
Megacraspedus monolorellus was described from several specimens of both sexes collected on 10.v.1902 near Ilgin (“Ilgün”), central Turkey (
Holotype ♂, “Turkey [prov. Nevşehir] Urgup [Ürgüp] 7 km E 18.5.2005 J. Junnilainen leg.” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus junnilaineni sp. n. is characterised by its light greyish brown forewings without markings. Externally it is hardly separable from M. golestanicus sp. n. (p 48). The male genitalia are very similar to M. monolorellus (Figure
BIN BOLD:ADB7272 (n = 1). The distance to the nearest neighbour M. similellus sp. n. is 9.4% (p-dist).
Turkey.
Host plant and early stages are unknown. The adults have been collected from the middle of May to the middle of August at altitudes of between 1200 and 1300 m.
The species name (a noun in the genitive case) is dedicated to Jari Junnilainen, Finland, who collected part of the type series of this species and numerous other valuable specimens used for our study.
Megacraspedus uzunsyrtus Bidzilya & Budashkin, 2015: 222, figs 19–22.
Paratypes. Ukraine. 2 ♂, Crimea, Koktebel vic., SW slope of Uzun-Syrt, evening collecting, 17.v.2014, leg. Yu. Budashkin, genitalia slide GU 16/1463 Huemer (ZMKU).
Ukraine. 2 ♂, Crimea, Aj-Petri, 5–6.vii.2002, leg, Yu. Budashkin (ZMKU).
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus uzunsyrtus is characterised by its light greyish brown forewings with a white costa and without further markings. It is very similar to M. monolorellus (Figure
BIN BOLD:ACS7353 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour M. similellus sp. n. is 6.4% (p-dist).
Ukraine (Crimea).
Host plant and early stages are unknown. Adults have been collected in the middle of May in the evening before sunset on xerothermic slopes on Jurinea stoechadifolia (M. Bieb.) DC (Asteraceae) which is considered to be the larval host plant (
M. uzunsyrtus was described from four males collected in Crimea on 17.v.2014 (
Holotype ♂, “BULGARIA [Dobrich region] Tuzlata 3.6.2014 J.Junnilainen leg.” “GU 16/1449 ♂ P. Huemer” “DNA Barcode
Adult. Male (Figure
Female (Figure
Variation. There is some variation in the colour of the forewings: from lighter to darker brown.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus similellus sp. n. is characterised by its light greyish brown forewings with white costa and without further markings. Externally it is hardly separable from M. uzunsyrtus (Figure
BIN BOLD:ADB8685 (n = 3). The intraspecific divergence of the barcode region is moderate with mean 0.8% and maximum divergence of 1.2%. The distance to the nearest neighbour M. uzunsyrtus is 6.4% (p-dist).
Bulgaria, Romania, Turkey.
Host plant and early stages are unknown. The adults have been collected from the middle of May to late July from sea level in the Balkans to 1700 m in Turkey.
The species name refers to the similarity to related taxa and is derived from the Latin word similis (meaning like), and the diminutive suffix –ellus. The name is a compound noun.
Holotype ♂, “Iran, [prov.] Khorassan NP [National Park] Golestan, Almeh 1770 m 2.5.2001 37°20,75'N, 56°07,02'E leg. P. Huemer” “BC
Adult. Male (Figure
Female. Unknown.
Variation. The few examined specimens show only slight variation. The margin of the forewing costa can be greyish or whitish. In one specimen there is no hair at the base of the antennal scape.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus golestanicus sp. n. is characterised by its relatively small wingspan, and its light greyish brown forewings without markings. Externally it is hardly separable from M. junnilaineni sp. n. (Figure
BIN BOLD:AAU3338 (n = 3). The intraspecific divergence of the barcode region is low with mean 0.1% and maximum divergence of 0.2%. The distance to the nearest neighbour M. uzunsyrtus is 8.8% (p-dist).
Iran (Golestan).
Host plant and early stages are unknown. The habitat is dominated by steppe meadows. The adults have been collected in the last third of May at light at an altitude of 1770 m.
This species is named after its place of occurrence: the province of Golestan in north-eastern Iran. The name is a masculine adjective.
Holotype ♂, “CROATIA [Dalmatia region] Konjevrate 200 m 25.06.2006 leg. Z. Tokár” “Megacraspedus sp. det. Zdenko Tokár” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show no variation, but worn specimens become light greyish.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus tokari sp. n. is characterised by cream-coloured head and its brown forewings with a white costa and without further markings. Its small size separates it from similar looking species. The male genitalia differ from other species of the M. dolosellus species group particularly by the furcated ridge of the saccus and the weakly curved phallus.
BIN BOLD:ACM1095 (n = 1). The distance to the nearest congeneric neighbour M. cuencellus is 10%, the distance to the nearest BIN in BOLD, Monochroa scutatella (Müller-Rutz, 1920), is 9.2% (p-dist).
Croatia.
Host plant and early stages are unknown. The adults have been collected from late June to the middle of September at low altitudes.
The species name (a noun in the genitive case) is dedicated to Zdenko Tokár, Slovakia, who collected most of the type series of this species and numerous other valuable specimens used for our study.
Ypsolophus (Megacraspedus) dolosellus Zeller, 1839: 190.
Ypsolophus separatellus Fischer von Röslerstamm, 1843: 300, 302, pl. 100, figs 1a–d [on plate as’separatella‘], syn. n.
Megacraspedus incertellus Rebel, 1930: (14), syn. n.
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” (
Adult. Male (Figs
Female (Figs
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
Female genitalia (Figs
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
The extraordinary DNA barcode divergence is reflected by 23 BINs! The intraspecific divergence of the barcode region in this species is mean 7.5% and maximum 13.8%, largely reflecting a geographic pattern. However, on several occasions large intrapopulational divergence was detected, e.g., in specimens from Italy (Gran Sasso), Bulgaria (Pirin Mts Blagoewgrad), Montenegro (Durmitor), and Slovakia with two sympatric clusters in each region. Individual variation of specimens collected at the same time in the same microhabitat may be high in these cases as e.g., in clusters dolo03 and dolo04 with two specimens diverging 6.9% in DNA barcode.
The following 23 clusters are defined (based on sequenced material):
BIN dolo01 (Greece: Ioannina): BOLD:ACS7817 (n = 1).
BIN dolo02 (Bulgaria: Alibotusch, Pirin mts): BOLD:ACA9065 (n = 3).
BIN dolo03 (Italy: l`Aquila): BOLD:AA03318 (n = 1).
BIN dolo04 (Italy: l`Aquila): BOLD:AA03319 (n = 1).
BIN dolo05 (Italy: Rieti): BOLD:AAX3311 (n = 1).
BIN dolo06 (Italy: Lazio): BOLD:ACZ7902 (n = 1).
BIN dolo07 (Greece: Peloponnes): BOLD:ACQ6924 (n = 2).
BIN dolo08 (Bulgaria: Kyustendil): BOLD:ACZ9025 (n = 1).
BIN dolo09 (Bulgaria: Pirin mts): BOLD:ACR2396 (n = 5).
BIN dolo10 (Bulgaria: Blagoewgrad): BOLD:ADB8684 (n = 1).
BIN dolo11 (Montenegro: Durmitor): BOLD:ACS7982 (n = 1)
BIN dolo12 (Macedonia: Galicica): BOLD:ADB8790 (n = 2).
BIN dolo13 (Macedonia: Korab, Pelister): BIN BOLD:ABA2915 (n = 10).
BIN dolo14 (Montenegro: Durmitor): BIN BOLD:ACS7352 (n = 1).
BIN dolo15 (Croatia: Pag): BIN BOLD:ADB8686 (n = 1).
BIN dolo16 (Ukraine: Odessa): BOLD:ADA0140 (n = 1).
BIN dolo17 (Bulgaria: Blagoewgrad): BOLD:ADB8789 (n = 1).
BIN dolo18 (Slovenia): BOLD:AAV7561 (n = 3).
BIN dolo19 (Russia: Orenburg obl.): BOLD:ACZ3281 (n = 3).
BIN dolo20 (Russia: Altai mts): BOLD:ACB3319 (n = 1).
BIN dolo21 (Russia: Altai mts): BOLD:ACZ3530 (n = 4).
BIN dolo22 (Hungary, Romania, Slovakia): BOLD:AAG0031 (n = 4).
BIN dolo23 (Austria, Slovakia): BOLD:ACS9604 (n = 3).
The minimum distance to the nearest neighbour M. lanceolellus is 9.9% (p-dist).
Widely distributed from western Europe (northwards to ca. 49°N latitude) to Central Asia (Altai Mts). A specimen from the Himalaya Mountains needs verification. According to Mariani (1943: 174) also in Sicily. Absent from the Iberian Peninsula and the northern part of Central Europe as well as northern Europe.
The larva was described in great detail (570 words, but no figures!) by
The adults have been collected from early May to late August at altitudes from lowland localities to ca. 2400 m. The single record from Nepal dates from October. However, the identity of this specimen remains doubtful.
Ypsolophus (Megacraspedus) dolosellus was described from four males collected in June and July in the surroundings of Vienna, Austria (
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 (
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 (
The M. faunierensis species group includes two species: M. neli sp. n. and M. faunierensis sp. n.
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.
Diagnostic remarks. The M. faunierensis species group is characterised by a combination of several diagnostic structures, particularly the small uncus, the straight and long gnathos hook, the longitudinal ridge of the saccular area, the oblong saccus with a longitudinal ridge and the shape of the phallus with wrinkles.
Holotype ♂, [France] “NEL Jacques [leg.] [Dep. Vaucluse, Montange de] Lure. Ou. 1700 m. 12.VI.1999” “P. Huemer GEL 1218 ♂” (
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus neli sp. n. is a rather uncharacteristic species of Megacraspedus, which may be recognized by its small size with a single pecten on the antennal scape. The holotype is somewhat worn and specimens in better condition may reveal additional diagnostic characters. It resembles M. pentheres (Figure
The male genitalia are similar overall to M. faunierensis sp. n. (Figure
Not available, barcoding failed.
France (Dep. Vaucluse), Italy (prov. Torino).
Host plant and early stages are unknown. The species was collected from the middle of June to mid-July at altitudes from ca. 1700 to 2200 m.
The species name (a noun in the genitive case) is dedicated to Jacques Nel, France, who collected the holotype of this species and numerous other Megacraspedus specimens used for our study.
The holotype was collected in the western part of Montagne de Lure, at the edge of the road at 1700 m altitude, between “the Refuge the Lure” and the “Signal de Lure” (J Nel in litt.).
Holotype ♂, “Italien, Prov. Cuneo Alpi Cozie, Demonte NW Colle Valcavera NE, 2420 m 7°6'23"E, 44°23'04"N 2.8.2010, leg. Huemer
Adult. Male (Figure
Variation. The amount of black scales on the forewings is variable. Rarely the black dot before the apex is absent. There is a slight variation in the colour of the forewings; worn specimens look more greyish.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus faunierensis sp. n. is characterised by its cream coloured forewings with three distinct black spots. It resembles M. tristictus (Figure
BIN BOLD:AAJ3164 (n = 3). The intraspecific divergence of the barcode region is considerable with mean 1% and maximum divergence of 1.6%. However, this divergence is exclusively based on a divergent specimen from a lowland locality (Prov. Cuneo, Valdieri N, RN Juniperus phoenicea, 900–1000 m, 29.vi.2008. leg. P. Huemer). The distance to the nearest congeneric neighbour M. sumpichi sp. n. is 11.9%, the distance to the nearest BIN in BOLD, an unnamed Limacodidae, is 9.1% (p-dist).
North-western Italy.
Host plant and early stages are unknown. The adults have been collected from early June to mid-August, depending on the altitude. In alpine habitats they were found flying freely in the early morning about sunrise. At this time several specimens could be collected in copula, mainly sitting on low vegetation, particularly on blades of grass. The altitudes range from ca. 900 to 2500 m.
This species is named after the place of occurrence of part of the type series: the Colle di Fauniera pass in northern Italy. The name is a noun in apposition.
The M. gredosensis species group includes one species, M. gredosensis sp. n.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia unknown.
Diagnostic remarks. The M. gredosensis species group is defined by unique structures in the male genitalia. In particular the distinctly digitate sacculus and the phallus with a small coecum, and a long and slender distal part with subapical tooth are characteristic structures.
Holotype ♂, “Hispania, [prov. Avila] 19.7.1980 Sierra de Gredos Navacepeda, 1500 m M. u. E. Arenberger” “GU 16/1416 ♂ P. Huemer“ (RCEA). Paratypes. Spain. 1 ♂, same data as holotype (
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus gredosensis sp. n. is characterised by its yellow-brown forewings with a clear white costa and no black dots, and the dark grey hindwigs. It may resemble species of the M. dolosellus-complex, but these have more pure whitish veins, but not a pure white costa and lighter hindwings. The male genitalia are somewhat similar to M. glaberipalpus sp. n. (Figs
BIN BOLD:ADI8272 (n = 1). The distance to the nearest neighbour M. bidentatus sp. n. is 10.01% (p-dist).
Central Spain (prov. Avila).
Host plant and early stages are unknown. The few adults known to date have been collected in the middle of July at altitudes between 1500 and 1650 m.
This species is named after its place of occurrence: the mountain range of Sierra de Gredos in Central Spain. The name is an adjective.
The M. cuencellus species group includes four species: M. cuencellus, M. bidentatus sp. n., M. fuscus sp. n., and M. trineae sp. n.
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.
Diagnostic remarks. The M. cuencellus species group is unmistakable due to several characters of the male genitalia such as the small uncus compared to the tegumen, the massive saccus, and the dentation of the phallus.
Megacraspedus cuencellus Caradja, 1920: 117.
Holotype ♂, “Cuenca V Korb” “HOLOTYPE Megacraspedus cuencellus Car. ♂ ROMANIA” “CIS-Korea Microlep. 4216 Megacraspedus cuencellus Car. – Cuena [sic], Spain K. T. Park” (
Adult. Male (Figure
Female. Unknown.
Variation. The forewing colour varies slightly from brownish grey to darker grey. The holotype (examined from a photograph) differs in having black streaks in the fold and towards apex.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus cuencellus is characterised by its almost plain dark grey forewings and dark hindwings but largely agrees with M. bidentatus sp. n. (p 65), M. fuscus sp. n. (p 67) and M. skoui sp. n. (p 74). The male genitalia are unmistakable due to the unique structures of the phallus. M. cuencellus differs from other species particularly in the thorns of the phallus having a different position.
BIN BOLD:ACC5029 (n = 1). The distance to the nearest neighbour M. bidentatus sp. n. is 3.1% (p-dist).
Southern France (
Host plant and early stages are unknown. The few adults have been collected from late May to early July at altitudes of between 1500 and 1650 m.
Megacraspedus cuencellus was described from one male in good condition from the province of Cuenca in Central Spain (
Holotype ♂, “SPAIN [prov. Barcelona] Gironella 680 m N42.03975 E2.02987 9.5.2013 J. Tabell leg.” “GU 16/1433 ♂ P. Huemer” “DNA Barcode
Paratype. Spain. 1 ♂, same data as holotype (
Adult. Male (Figure
Female. Unknown.
Variation. One of the two examined specimens has the rather denuded forewings light greyish.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus bidentatus sp. n. is characterised by its greyish brown forewings, which are darker in the costal third and have a distinct black dot at end of the fold and at the cell. It is similar to M. alfacarellus (Figs
BIN BOLD: ADA0203 (n = 1). The distance to the nearest neighbour M. cuencellus is 3.1% (p-dist).
North-eastern Spain (prov. Barcelona).
Host plant and early stages are unknown. The two type specimens were collected in early May at an altitude of 680 m.
The species name refers to the sclerotisations of the phallus and is derived from the Latin word bidentatus (having two teeth). The name is a masculine adjective.
Holotype ♂, “Spain, Extremadura [prov. Cáceres] 39°34'11"N, 06°06'34"W 11 km SE Monroy 4. iv. 2009, 375 m leg. B. Skule” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus fuscus sp. n. is characterised by its almost plain blackish grey forewings and dark hindwings. Externally it mostly resembles M. grisea from Asia, but that species has ringed antennae and some lighter scales along the termen. It is also similar to, but darker than, M. cuencellus (Figure
Not available, barcoding failed.
Western Spain (prov. Cáceres, Salamanca).
Host plant and early stages are unknown. The type specimens were collected in early April and early May at altitudes from 375 m to ca. 900 m.
The name of this species is derived from the Latin word fuscus (meaning dark) after the colour of the forewings. The name is a masculine adjective.
Megacraspedus fuscus sp. n. is tentatively attributed to the M. cuencellus species group due to the overall similarity in the male genitalia. Molecular data will be necessary to confirm this classification.
Holotype ♂, “Portugal BB [= prov. Beira Baixa] Serra de Estréla Torre, 1600–1950 m 10.-11.vii.1986 O. Karsholt” “Gen. Præparat No 3355 ♂ H. Hendriksen” (
Adult. Male (Figure
Female. Unknown.
Variation. A rather variable species. The colour of the forewing varies from almost plain bone white to light brownish with a white costa. A rather indistinct black dot is sometimes present at the end of the cell. Sometimes there are a few black scales at the end of the cell and/or near apex.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus trineae sp. n. is a rather uncharacteristic Megacraspedus which can be recognized by lacking black dots (apart from an indistinct one at the end of the cell) and the whitish coloured veins on the forewings. It is very similar to M. occidentellus sp. n. (p 79). The male genitalia differ from other species of the M. cuencellus species group in the characteristic postmedial tooth and the presence of a short sacculus.
BIN BOLD:ADF0469 (n = 1). The distance to the nearest neighbour Megacraspedus bidentatus sp. n. is 7.8% (p-dist).
Portugal and Spain.
Host plant and early stages are unknown. The adults have been collected in early May and in the first half of July at altitudes from 1100 to ca. 1900 m.
The species name (a noun in the genitive case) is dedicated to Trine Karsholt, Denmark, who assisted OK when collecting the holotype and a part of the type series of this species.
A single male from Central Spain (Figure
The M. pusillus species group includes nine species: M. tristictus, M. alfacarellus, M. pusillus, M. skoui sp. n., M. spinophallus sp. n., M. occidentellus sp. n., M. granadensis sp. n., M. heckfordi sp. n., and M. tenuiuncus sp. n.
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 M. tenuiuncus sp. n.), in some species with lighter dorsum. Female (only known for M. spinophallus sp. n.) moderately brachypterous.
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.
Diagnostic remarks. The M. pusillus species group is characterised by unique structures, in particular the massive and bulky gnathos hook with longitudinal grooves is highly diagnostic. Furthermore the combination of characters such as the distorted and clubbed apex of the valva, the strongly setose saccular area with a tendency for separation of the sacculus and the medial ridge of the saccus are important characters. Female genitalia differ from other species groups having an overall similarity by the absence of posteriorly pointed sclerites of the subgenital plate. However, females are unknown for the vast majority of species and the diagnostic value of this character is therefore unknown.
Megacraspedus tristictus Walsingham, 1910: 231.
Holotype ♂, [France] “Holo-type” “Cannes S FRANCE 22.V.1892 Wlsm. 81169” “Type” “Walsingham Collection, 1910–427.” “Megacraspedus tristictus ♂ Wlsm. Ent. Mo. Mag. 46.231-2 sp. 3009.1 (1910) TYPE ♂ (f) deser 81169” (
Adult. Male (Figure
Female. Unknown.
Variation. There is some variation in size, including among specimens from the same locality.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus tristictus is characterised by its light yellowish brown forewings with three distinct black dots. It resembles M. pentheres (Figure
BIN BOLD:ADA0606 (n = 1). The distance to the nearest congeneric neighbour M. tenuiuncus sp. n. is 12% , the distance to the nearest BIN in BOLD, Emarginea combusta (Noctuidae), is 10.3% (p-dist).
South-eastern France, north-western Italy.
Host plant and early stages are unknown. The adults have been collected from the middle of May to early July at altitudes from 300 to 1200 m.
Megacraspedus tristictus was described from one male collected by Walsingham at Cannes, SE France, 22.v.1892 (Walsingham 1910).
Megacraspedus alfacarellus Wehrli, 1926: 163.
Holotype ♂, “Type” [Spain, prov. Granada] “Sierra Alfacar 18/19 VI 25 Dr. E. Wehrli, Basel” “alfacarellus Sa. Alfacar ♂ + 18.VI.25. W. ” “GU 16/1414 ♂ P. Huemer” (
Adult. Male (Figs
Female. Unknown.
Variation. The limited material available so far shows no variation.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus alfacarellus is characterised by its black antennae, and by the dark brown forewings with two weak black dots. It is similar to some forms of M. spinophallus sp. n. (Figs
Not available, no suitable specimen was available for barcoding.
Southern Spain (prov. Granada).
Host plant and early stages are unknown. The few adults known to date have been collected in the second half of June at unreported altitudes.
Megacraspedus alfacarellus was described from one male collected between 18–19.vi.1925 in Sierra Alfacar, Andalusia, Spain (
Megacraspedus pusillus Walsingham, 1903: 266.
Paratype ♂, “Sierra Nevada Granada ; SPAIN 3 VI 1901 Wlsm 86728” “Paratype” “Megacraspedus ♂ pusillus WLSM 2/3” “Walsingham Collection, 1910-427.”, genitalia slide 33661 (
Adult. Male (Figure
Female. Unknown.
Variation. The black dot in the fold is absent in some specimens.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus pusillus is characterised by its light grey-brown forewings with a distinct black dot at end of the cell. It is particularly similar to M. skoui sp. n. (Figure
BIN BOLD:ACZ8007 (n = 1). The distance to the nearest neighbour M. spinophallus sp. n. is 8.4% (p-dist).
Southern Spain (prov. Granada and Almería).
Host plant and early stages are unknown. The few adults known to date have been collected from the middle of June to the second half of July, at altitudes from between 1600 and 2500 m.
Megacraspedus pusillus was described from four males collected in the Sierra Nevada on 2.vi.1901 (Walsingham 1903: 266).
Holotype ♂, “Spain, [prov.] Granada, Sierra Nevada, Camino del Veleta 1650 m, 25.VI.1986, leg. Peder Skou” “Gen. prep. 5318 ♂ O. Karsholt” (
Adult. Male (Figure
Female. Unknown.
Variation. One specimen has an indistinct black dot in the fold of the forewings.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus skoui sp. n. is characterised by its almost unicolorous light grey forewings. It resembles several smaller, unicolorous Megacraspedus species, especially M. pusillus, which has a distinct black dot at end of the cell (Figs
BIN BOLD:ACT1624 (n = 1). The distance to the nearest neighbour M. spinophallus sp. n. is 7.97% (p-dist).
Spain (prov. Granada).
Host plant and early stages are unknown. Adults have been collected in June and July at altitudes between 1300 and 1650 m.
The species name (a noun in the genitive case) is dedicated to Peder Skou, Denmark, who collected the holotype of this species and numerous other Megacraspedus specimens used for our study.
Holotype ♂, “Spanien, [prov.] Alicante Sierra de Crevillente 5 km NE Albatera, 450 m, 38°15,22'N, 00°54,86'W 26.5.2004 leg. Huemer
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
Megacraspedus spinophallus sp. n. is characterised by its black antennae, and by the dark brown forewings having two elongate black dots and being yellowish in dorsal half. It is very similar to M. alfacarellus (Figs
BIN BOLD:AAU1828 (n = 7), BIN BOLD:ACT2894 (n = 1). Genetically variable species. The intraspecific divergence of the barcode region is large and reflected by 2 BINs with 2.8% maximum divergence, based on a single specimen compared with a larger cluster. Within the latter average divergence is only 0.2% and maximum divergence is 0.6% (n = 2). The minimum distance to the nearest neighbour M. skoui sp. n. is 8% (p-dist).
Spain.
Host plant and early stages are unknown. The adults have been collected from late April to the middle of July at altitudes from sea level to 1200 m.
The species name is a compound word derived from the Latin adjective spinosus (meaning spiny) and noun phallus, referring to the spiny phallus. The name is a noun in apposition.
Holotype ♂, “Portugal, Ext. [=prov. Estremadura] Ericeira, 30 m 4.-8.vii.1986 O. Karsholt” “GU 13/1352 ♂ Huemer” (
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.
Megacraspedus occidentellus sp. n. is a rather uncharacteristic Megacraspedus which can be recognized by lacking black dots and whitish coloured veins on the forewing. It is similar, e.g., to the slightly larger M. trineae sp. n. (Figure
Not available, barcoding failed.
Western Portugal.
Host plant and early stages are unknown. The type series was collected in early July at low altitudes.
This species is named after its occurrence at the most western part of continental Europe. The species name is derived from a combination of the Latin occidentalis (meaning western) and the diminutive suffix –ellus. The name is a compound noun.
Holotype ♂, “Hispania. [prov. Granada] Andalucia. Sierra Nevada. Cam.d.Valeta 1600 m 14.6.1986 E. Traugott-Olsen” “GU 16/1458 ♂ P. Huemer” (
Adult. Male (Figure
Female. Unknown.
Variation. The yellowish streak in the fold can be more or less distinct.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus granadensis sp. n. is characterised in having the forewings covered with whitish brown- and black-tipped scales, in having the costa blackish brown at base and whitish grey in middle, and having no black dots. Also the dark grey hindwings are characteristic. The male genitalia differ from the particularly similar M. spinophallus sp. n. (Figs
Not available, barcoding failed.
Spain (prov. Granada).
Host plant and early stages are unknown. The adults have been collected in the middle of June at an altitude of 1600 m.
This species is named after its place of occurrence: the province of Granada in southern Spain. The name is a masculine adjective.
Holotype ♂, “ESPANA, Prov. Cantabria PN Picos de Europa Fuente De, El Cable Bergst. 4°48,53'W, 43°09,55'N 1870 m, 11.7.2012 leg. Huemer
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus heckfordi sp. n. is characterised by its plain light greyish brown forewings without markings. It can be separated from other species without markings on the forewings by its cream-white head. The male genitalia are unmistakable within Megacraspedus and can be easily identified by the characteristic large spine of the phallus.
BIN BOLD:ACA9758 (n = 4). The intraspecific divergence of the barcode region is low with mean 0.5% and maximum divergence of 0.8%. The distance to the nearest neighbour M. tenuiuncus sp. n. is 7.8% (p-dist).
Spain (prov. Cantabria).
Host plant and early stages are unknown. The adults have been collected in the middle of July at altitudes from 1230 to 1870 m.
The species name (a noun in the genitive case) is dedicated to Robert (Bob) Heckford, U.K., who collected the first specimen and provided his valuable material available to our study.
Holotype ♂, “Spain [prov.] Lerida Roni near Sort 7.vii.1993, 1000 m P. Skou” “GU 15/1402 ♂ P. Huemer” (
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus tenuiuncus sp. n. is characterised by its black antennae, and by its light greyish forewings with two black dots. It is similar to M. lanceolellus (Figs
BIN BOLD:ADF1915 (n = 1). The distance to the nearest neighbour M. heckfordi sp. n. is 7.7% (p-dist).
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 tenuis (meaning small) and uncus, referring to the slender uncus. The name is a noun in apposition.
The M. lativalvellus species group includes one species: M. lativalvellus.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Diagnostic remarks. The M. lativalvellus species group is defined by structures of the male genitalia, particularly the long gnathos hook, submedially bent at right angles, the short and smoothly sclerotised saccus without ridge, and the sclerotisation of the phallus with postmedial thorn are characteristic.
The systematic position of the M. lativalvellus species group is uncertain and tentative due to the preparation-related lack of a ductus ejaculatorius, and in the lack of females and supportive molecular data.
Megacraspedus lativalvellus Amsel, 1954: 54, fig. 4.
Holotype ♂, “MALTA SALINA 16.III.52. Coll. Delucca” “Type” “Gen. prep. Nr. 5092 ♂ O. Karsholt” (RCCDL). Paratype. 1 ♂, same data as holotype, but 31.iii.1952, genitalia slide 1850 Amsel (
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus lativalvellus is similar overall to M. dolosellus and related species. It can be separated from these by the longer segment 3 of the labial palpus, and by having a pecten of one or a few hairs on the antennal scape. The male genitalia are easily recognized by the characteristic gnathos hook, only shared by a few other species such as M. leuca (Figure
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.
Megacraspedus lativalvellus was described from three males collected in Malta in March 1952 by C. De Lucca (Amsel, 1954). The holotype and one paratype were kept in the collection of Carmello De Lucca. After his death in 1971 the collection stayed with the family and was not curated, with the result that it suffered much from attack by pest insects (P Sammut in litt.). With the help from Paul Sammut we were able to borrow these two type specimens, but only the pins and labels were left – apart from the tip of the abdomen of the holotype, from which an almost complete genitalia slide could be prepared, however, leaving the apex of the phallus broken. The second paratype in
The type locality, Salina in northern Malter is now a nature reserve, and there is thus a possibility that M. lativalvellus is still present there.
The M. dejectella species group includes one species: M. dejectella.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Diagnostic remarks. The M. dejectella species group is defined by structures of the male genitalia, particularly the small uncus, and the stout and apically curved gnathos hook, the distinct longitudinal ridge of the entire saccular area, and the posteriorly arched and smoothly sclerotised saccus are characteristic.
The systematic position of the M. dejectella species group is uncertain and tentative in the absence of females and supportive molecular data.
Gelechia dejectella Staudinger, 1859: 242.
Lectotype ♂, designated by
Adult. Male (Figure
Female. Unknown.
Variation. The above description is based on one rather worn specimen, having the labial palps totally denuded.
Male genitalia (Figure
Female genitalia. Undescribed.
Megacraspedus dejectella is characterised by its relatively small size, and by the light grey-brown forewings with a white costa and fold and being without black spots. It is similar to M. sumpichi sp. n. (Figs
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.
Megacraspedus dejectella was described from three specimens of both sexes collected in the surroundings of Granada, southern Spain (
The M. binotella species group includes four species: M. devorator sp. n., M. binotella, M. brachypteris sp. n., and M. barcodiellus sp. n.
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.
Diagnostic remarks. The M. binotella species group is defined by the combined structures of the male genitalia, particularly the nearly mitre-shaped uncus, the slender gnathos hook, the short and comparatively small saccus with long lateral sclerites, and particularly the S-curved phallus that is only shared wirth few other species of the M. dolosellus species group. The female genitalia largely agree overall with several taxa in other species groups.
Holotype ♂, “BULGARIA [Dobrich region] Nos Sabla [Shabla] 27.5.2002 J. Junnilainen leg.” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. There is considerable variation in wingspan.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus devorator sp. n. is characterised by its light brown forewings with a white costal edge and two elongate black spots. It is very similar to M. binotella (Figs
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 M. binotella is 9% (p-dist).
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 devorator (=food), and refers to eating (too) much after hunger during an expedition to Altai mts in search for Megacraspedus together with Christoph Wieser.
Palpula binotella Duponchel, 1843: 256, pl. 72, fig. 7.
Ypsolophus binotellus Fischer von Röslerstamm, 1843: 300, 301, pl. 99, figs 2a, 2b; homonym and synonym of Palpula binotella Duponchel, 1843.
Syntype ♂, “binotella FR” “TYPE” “Duponchel” “2169” [without abdomen] (
Adult. Male (Figs
Female (Figure
Variation. The forewing can be more or less mottled with brown-tipped scales, and rarely the veins are whitish. Some specimens have darker scales along the termen. Also the head can be light brown. The tip of segment 3 of the labial palps is sometimes black. M. binotella furthermore shows some variation in size (see also M. brachypteris sp.n.) with e.g., two specimens collected simultaneously ranging between 14 and 17 mm wingspan.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus binotella is characterised by the creamy forewings with two distinct black dots. It resembles well-marked specimens of M. brachypteris sp. n., a species which usually differs in the whitish veins and the short-winged female. See also M. devorator sp. n. (p 86). The male genitalia are very similar to M. barcodiellus sp. n. (see also Figure
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 M. devorator sp. n. is 9% (p-dist).
Austria, Czech Republic, Germany (northwards to Rheinland-Pfalz and Hessen) (
Early stages are unknown.
Palpula binotella was described from an unstated number of specimens which Duponchel had received through the insect dealer Parreyss under the name “binotella Fischer de Röslerstamm” (Duponchel 1843). Megacraspedus binotella was previously attributed to Fischer von Röslerstamm. However,
Holotype ♂, “Macedonia NP [National Park] Mavrovo Korab [mountain], eastern ridge ca. 2325–2400 m 20°34'46"E, 41°47'08"N 28.7.-1.8.2011 leg. Huemer & Tarmann” “BC
Adult. Male (Figure
Female (Figure
Variation. The variation in the forewing colour and labial palpi is probably due to faded compared with fresh specimens. Similarly the extent of the the two black dots of the forewing shows some variation, particularly in worn specimens they are largely reduced or denuded. Specimens from Albania (Pashtrik, Kula Ljums) have distinct black spots and are larger on average (17–19 mm) than those from other populations.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus brachypteris sp. n. is characterised by its light brown forewings with faintly whitish veins and with two black spots present or reduced. It is sometimes hardly separable from certain forms of M. binotella (Figs
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 M. dolosellus is 11.3% (p-dist).
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.
Megacraspedus brachypteris sp. n. shows considerable phenotypic variation both individually and to some extent also geographically. However, the large intraspecific barcode divergence neither shows a clear geographic structure nor a concordant pattern to phenotypic appearance. Similarly the genitalia do not support cryptic diversity and therefore we accept only one species.
Holotype ♂, “Macedonia NP [National Park] Mavrovo Korab [mountain], Korabska jezero, Kobilino pole, 2080–2180 m 20°34'55"E, 41°46'42"N 28.7.-1.8.2011 leg. Huemer & Tarmann” “BC
Adult. Male (Figure
Female. Unknown.
Variation. The spots of the forewing are sometimes reduced, but this may be due to worn specimens which furthermore become lighter and almost whitish.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus barcodiellus sp. n. is characterised by its plain cream-white forewings with two distinct black spots. It is very similar to M. andreneli (p 93). The male genitalia are similar to those of other species of the M. binotella species group but differ in particular in the strongly curved distal part of the phallus and the broad valva. These characters are similar to M. brachypteris sp. n. (Figs
. BIN BOLD:ABA2916 (n = 3). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour M. binotella is 10.7% (p-dist).
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 Megacraspedus. It is derived from the latinized vernacular barcode, and the diminutive suffix -ellus. The name is a compound noun in apposition.
The M. bilineatella species group includes three species: M. bilineatella, M. andreneli, and M. sumpichi sp. n.
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.
Diagnostic remarks. The M. bilineatella species group is defined by several structures of the male genitalia, particularly the small and distally narrowing uncus, the comparatively massive gnathos hook, the slender and digitate valva without sacculus, the smoothly sclerotised saccus, and the phallus without distinct sclerotisations are characteristic in this combination.
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.
Megacraspedus bilineatella Huemer & Karsholt, 1996: 251, figs 1–10.
Holotype ♂, “ITALIA sept. Prov. Bergamo Alpi Orobie W Ca. San Marco, 1800–1900 m, 22.7.1992 leg. Huemer & Tarmann” “Holotype Megacraspedus bilineatella Huemer & Karsholt, 1995” (
Adult. Male (Figure
Female. Unknown.
Variation. The few examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus bilineatella is unique within the genus in having the forewings brown with two distinct, longitudinal white streaks along costa and through the middle of the wing and no other markings. The male genitalia are somewhat similar to several other congeners and differ from the closest, M. andreneli (Figure
BIN BOLD:ABU7227 (n = 1). The distance to the nearest congeneric neighbour M. fallax is extraordinarily large with 14.9% (p-dist), whereas several other Gelechiidae in BOLD have smaller distances.
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 (
Megacraspedus bilineatella was described from nine males collected in the Orobian Alps in northern Italy (
Megacraspedus andreneli Varenne & Nel, 2014: 61, figs 7–9.
Holotype ♂, France, Alpes-Maritimes, Saint-Sauveur-sur-Tinée, Lac Nègre, Parc national du Mercantour, 10 août 2013, 2450 m Th. Varenne leg., genitalia slide JN n°27182 (RCTV) [photographs examined]. Paratype. France. 1 ♂, Alpes-Maritimes, Valdeblore, Col de Veillos, 2250 m, 20.vii.2010, leg. A. Nel, genitalia slide 24592 J. Nel (
Adult. Male (Figure
Female. Unknown.
Variation. The forewings of the paratype are darker than those of the holotype, probably because the latter is more worn.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus andreneli is a rather uncharacteristic species. It can be confused with several other species having two black dots in the forewing, e.g., M. binotella and M. barcodiellus sp. n. (flagellum distinctly ringed black and white), and M. lanceolellus (costa white). The male genitalia are similar to those of M. bilineatella (Figure
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 Megacraspedus tokari sp. n. is 11.2% (p-dist).
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.
Megacraspedus andreneli was described from two males collected at two localities in the Alpes Maritimes (
Holotype ♂, “HISP.Prov.d.Granada Sierra de Baza 1300 m 18.7.1987 leg.G. Baldizzone E.Traugott-Olsen” “P.Huemer GEL 1244 ♂” (
Adult. Male (Figure
Female (Figure
Variation. There is some variation in the amount of grey-brown scales in the forewing. In specimens with many such scales the white veins look more distinct. Worn specimens become whitish.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus sumpichi sp. n. is characterised by its relatively small size, by the white head and the light grey-brown forewings without black dots and with white costa. It is very similar to M. dejectella (p 85). The male genitalia differ from related species particularly in the V-shaped and smooth saccus. The female genitalia are similar to those of M. lanceolellus (Figs
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 M. cerussatellus is 11.4% (p-dist).
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 Megacraspedus specimens used for our study.
The M. fallax species group includes 14 species: M. tabelli sp. n., M. gallicus sp. n., M. ribbeella, M. libycus sp. n., M. numidellus, M. albovenata, M. longipalpella, M. niphorrhoa, M. albella, M. fallax, M. balneariellus, M. podolicus, M. kazakhstanicus sp. n., and M. knudlarseni sp. n.
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.
Diagnostic remarks. The M. fallax species group differs from the vast majority of Megcraspedus by the long and porrect labial palpus with reduced segment 3. It is furthermore defined by combined structures of the male genitalia, particularly the shape of the uncus with a parallel outer edge, the stout gnathos hook, the digitate valva with usually weakly inflated apex and the comparatively small, sub-triangular saccus without sclerotised ridges. The female genitalia vary in several characters and agree overall with some taxa in other species groups, but females are unknown for the majority of Megacraspedus.
The majority of species of the M. fallax species group are very similar in the male and, as far as known, female genitalia, usually differing at most by subtle characters. However, all these species clearly differ in phenotypic appearance, supported by DNA barcode divergence in several taxa.
Holotype ♂, “MOROCCO Settat Prov. Sidi Said Machou 2.5 km E 1125 m, N33.14137 W8.09181 18.5.2016 J. Tabell leg.” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. The two examined specimens are similar except for being in different conditions.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus tabelli sp. n. is characterised by its light yellow-brown forewings with white veins. It may resemble M. albovenata (Figs
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 M. numidellus sp. n. is 9.8% (p-dist).
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.
Holotype ♂, “SPAIN [prov.] Almería Sierra de Alhamilla 800–900 m route Huebro - Colotivi 1.5.2008 Zdenko Tokár leg.” “Gen. pr. Z. Tokár ♂ No 12180 [in glycerin]” “Megacraspedus fallax det. Zdenko Tokár” “DNA Barcode
Adult. Male (Figure
Female (Figure
Variation. The forewing colour of both males and females can be lighter or darker brownish.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus gallicus sp. n. is characterised by its relatively large size, its long labial palps and the light brownish forewings with white veins in costal half. It can be confused with M. fallax (Figure
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 M. ribbeella is 6.2% (p-dist).
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 Gallia which included present-day France and adjacent territories. The name is a masculine adjective.
Nevadia ribbeella Caradja, 1920: 118.
Holotype ♀, [Spain] “Nevadia ribbeella”, “HOLOTYPE Vadenia (=Nevadia) ribbeella Car. ♀ ROMÄNIA” “Sra Nevada” “Genit. praep. ♀ N 187797 Euparal 2003 Ponomarenko” (
Adult. Male (Figure
Female (Figure
Variation. Very variable in size. The dark parts of the forewings can be lighter or darker brown. One specimen has a broader white streak along the costa. The dark streak on the thorax is often obsolete because specimens tend to become greasy. Worn specimens become lighter.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus ribbeella is characterised by its large size, and by its contrasting brownish and white forewings. Small specimens resemble M. albovenata (Figs
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 Megacraspedus gallicus sp. n. is 6.2% (p-dist) (p-dist).
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.
Nevadia ribbeella was described from one specimen from Sierra Nevada, Spain (
Holotype ♂, “[northern] LIBYA, Gharian Wadi el Hira, 7.iv.1983 Uffe Seneca” “GU 16/1435 ♂ P. Huemer” (
Adult. Male (Figure
Female (Figure
Variation. The examined specimens exhibit only slight variation. The dark streak on the thorax is often obsolete because specimens tend to be greasy. Worn specimens become lighter. Specimens from Morocco are generally lighter both on the forewings and hindwings.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus libycus sp. n. is characterised by its large size, and by its contrasting brownish and white forewings. Externally it is hardly separable from M. ribbeella (Figs
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.
Chilopselaphus numidellus Chrétien, 1915: 333.
Megacraspedus mareotidellus Turati, 1924: 169, pl. 6, fig. 15, syn. n.
Lectotype ♂, Chilopselaphus numidellus, here designated, “TYPE” “Chilopselaphus numidellus” “Dj. Gafsa 28.4.09” genitalia slide PGCGN 8017 (
Adult. Male (Figs
Female. Unknown.
Variation. The few examined specimens from Northern Africa are rather worn with a tendency of white rather than yellow stripes. These stripes also show some variation in extension and the amount of blackish brown scales on the forewing is also variable. The patch between 3/5 and 4/5 in middle of the forewing wing is sometimes yellowish.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus numidellus is characterised by its moderately long labial palps with reduced segment 3, by the antennal scape with the pecten consisting of several hairs and by the forewings having three white streaks, but no black dots. The male genitalia are similar to M. albovenata (Figure
BIN BOLD:ADA0605 (n = 1). The distance to the nearest neighbour M. gallicus sp. n. is 9% (p-dist).
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.
Chilopselaphus numidellus was described from an unspecified number of specimens (probably more than one as the wingspan is stated to be “15–16 mm”) collected at Gafsa in Tunisia in May. It was compared with C. fallax Mann (
Megacraspedus albovenata Junnilainen, 2010, in Junnilainen & Nupponen, 2010: 1, figs 22, 28.
Paratypes. Russia. 3 ♂, S Ural, Cheliabinsk district, Arkaim reserve near Amurskii village, 16.vi.1996, leg. K. Nupponen, J.-P. Kaitila, J. Junnilainen & M. Ahola (RCKN,
Adult. Male (Figure
Female (Figure
Variation. The examined specimens show only slight variation, and specimens from central Europe are similar to those from the southern Urals.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus albovenata is characterised by its light yellow, brownish dusted forewings with clear white veins, giving it a striped look. It is similar to M. longipalpella (Figs
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 M. kazakhstanicus is 5.1% (p-dist).
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 (
Megacraspedus albovenata was described from numerous males collected in the southern Ural Mountains, Russia (
Megacraspedus
longipalpella
Junnilainen, 2010, in
Paratypes. Russia. 5 ♂, S Ural, Cheliabinsk district, Troizkii reserve near Berlin village, 30.vi.1997, leg. K. Nupponen & J. Junnilainen, genitalia slides no. 1/7.IV.2008 K. Nupponen, 02022402 J. Junnilainen (RCKN,
Adult. Male (Figure
Female (Figure
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus longipalpella is characterised by its light yellow, brownish dusted forewings with clear white veins. It is similar to M. albovenata (Figs
BIN BOLD:ACM1349 (n = 1). The distance to the nearest neighbour M. niphorrhoa is 4.8% (p-dist).
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 (
Megacraspedus longipalpella was described from numerous males collected in the southern Urals, Russia (
Trichembola niphorrhoa Meyrick, 1926: 272.
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 (RCKN,
Adult. Male (Figure
Female (Figure
Variation. The examined specimens show only slight variation.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus niphorrhoa is characterised by its light greyish forewings mottled with dark brown scales especially in the middle and apical part of the wing, and with three longitudinal yellow streaks. It is very similar to M. podolicus (Figure
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 M. longipalpella is 4.8% (p-dist).
Kazakhstan, Russia (S Ural).
Host plant and early stages are unknown. The adults have been collected in June at low altitudes.
Trichembola niphorrhoa was described from one male collected by Bartel at Lake Indersky, W. Kazakhstan (Meyrick 1926: 272).
Chilopselaphus albella Amsel, 1935: 302, pl. 10, fig. 57
Lectotype ♂, here designated, [Israel/Palaestine, West Bank]“Georgsklost. Wad. Kelt Lichtfang 1.4.1930“ “Palästina Expedition 18.2.-4.6.30 H. Amsel” “GU 699” “Typus leg. H. Amsel % Chilopselaphus albella” “coll.
Adult. Male (Figure
Female (Figure
Variation. Not observed because of very limited material.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus albella is characterised by its long labial palps with reduced segment 3, by the antennal scape with pecten consisting of several hairs, and by the light yellowish forewings with brown streaks/patches in medial part and by having white veins. It resembles M. numidellus (Figs
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.
Chilopsephalus albella Amsel, 1935 was described from few specimens (“selten”) collected by HG Amsel at “Georgskloster”, now Israel, 1 April 1930. A lectotype is here designated in order to fix the identity of the species and conserve stability of nomenclature. The lectotype is rather worn and bleached. The original male genitalia slide (without phallus) has been remounted. It fully agrees with specimens from Iran and we thus consider these samples conspecific.
Chilopselaphus fallax Mann, 1867: 850.
Trichembola neurophanes Meyrick, 1926: 271.
Holotype ♂, Chilopselaphus fallax, “Ofen 1866” “
Adult. Male (Figure
Female. Unknown.
Variation. The amount of dark brown scales on the forewings vary, and in some specimens the apical area (between veins) is almost brown. Specimens tend to become greasy and the whitish parts of the head and body thereby become brown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus fallax is characterised by the distinct, silvery white sub-costal streak on the forewings. M. gallicus sp. n. (p 101). The male genitalia are very similar to M. albella (Figure
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 M. niphorrhoa is 5.6% (p-dist).
Hungary, Kazahkstan, Russia (S Ural, Volgograd oblast), Turkey. According to
Early stages are unknown. The larva is stated to feed in stems of Stipa (
Chilopselaphus fallax was described from one specimen collected by Rudolf Anker at Ofen (now part of Budapest), Hungary (
Chilopselaphus balneariellus Chrétien, 1907: 179.
Croatia. 1 ♂, Konjevrate, 6.vi.2005, leg. Z. Tokár; 1 ♂, 5 km SE Pirovac, 24.vi.2006, leg. Z. Tokár (all RCZT); 3 ♂, 2 ♀, Pag, 10.vi.2015, leg. J. Junnilainen, genitalia slide GU 16/1457 Huemer (RCJJ); 1 ♂, 5 km SE Pirovac, 24.vi.2006, leg. Z. Tokár (RCZT). France. 1 ♂, Dep. Gard, Aigues-Mortes, 3.vii.1988, leg. K. Larsen (
Adult. Male (Figure
Female (Figure
Variation. The examined specimens exhibit only minor variation. One specimen has a wingspan of only 15 mm.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus balneariellus is characterised by its yellow forewings with greyish veins. It is similar to M. podolicus (p 119). The male genitalia are similar overall to other species of the M. fallax species group, but differ from all species particularly by the slender uncus. From the nearest species M. podolicus (Figure
BIN BOLD:ADB9039 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour M. podolicus is 6.3% (p-dist).
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).
Chilopselaphus balneariellus was described from an unstated number of specimens from France, Languedoc (
Chilopselaphus podolicus Toll, 1942: 170, pl. 13, figs 7–8, pl. 16, fig. 26.
Austria. 2 ♂, Burgenland, Weiden am See S, Zitzmannsdorfer Wiesen, 24.vi.1961, leg. F. Kasy (
Adult. Male (Figure
Female. Unknown.
Variation. The colour of the forewings varies from lighter to darker, depending on the amount of dark brown scales. Light specimens have more yellow between the veins. Worn specimens are generally lighter than fresh specimens. The hairs on the antennal scape are easily lost.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus podolicus is characterised by its long labial palps, slender wings and by the dark brown, longitudinal streak in the middle of the wing, which is absent in M. balneariellus (Figs
BIN BOLD:ADB8683 (n = 1). The distance to the nearest neighbour M. balneariellus is 6.3% (p-dist).
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.
Chilopselaphus podolicus was described from western Ukraine, Borszczow, Hlody (
Holotype ♂, “SW-KAZAKHSTAN 43°48'45"N 53°31'29"E Sengirkum sand, 70 m Terekurpa well 27.5.2011, K. Nupponen leg.” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus kazakhstanicus sp. n. is characterised by its light yellow, brownish, and blackish dusted forewings with pure white veins, giving it a striped look. It is very similar to M. albovenata (Figs
BIN BOLD:ADJ1190 (n = 1). The distance to the nearest neighbour M. albovenata is 5.1% (p-dist).
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.
Holotype ♂, “SPAIN.Islas Canarias Gran Canaria. Mogán, El Barranquillo Andres, 2 km. S. 24.vii.1995 K. Larsen” “GU 16/1454 ♂ P. Huemer” (
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus knudlarseni sp. n. is amongst the smallest species in the genus, with a reduced segment 3 of the labial palps, and only comparable with M. skulei sp. n. (Figs
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 M. knudlarseni sp. n. in the M. fallax species group is based only on the labial palps and the overall similarity of the male genitalia, whereas molecular data are unavailable for this species.
The M. majorella species group includes two species: M. majorella, and M. latiuncus sp. n.
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.
Diagnostic remarks. The M. majorella species group is defined by unique structures of the male genitalia, particularly the extraordinary large, sub-square uncus, the distinct gnathos hook, the short phallus with small sclerotised patch and the double-twisted interior sclerotisation of the ductus ejaculatorius. The two species in this species group are similar in the male genitalia. However, they differ considerably in phenotypic appearance. The systematic position of the M. majorella species group is uncertain and tentative due to the absence of females and supportive molecular data.
Megacraspedus imparellus (? var.) majorella Caradja, 1920: 117.
Lectotype ♂, “Alai” “LECTOTYPE Megacraspedus imparellus v. majorellus Car. ♂ des. Dr A. Popescu – Gori” “176434” “CIS-Korea Microlep. 4217 Megacraspedus majorella Car. – Alai, Himalaya K. T. Park ’96” (MNGA) [photographs examined]. Paralectotype ♂, same data, but without genitalia slide (MNGA) [photographs examined].
Adult. Male (Figure
Female. Unknown.
Variation. None from the few examined specimens.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus majorella is characterised by its light greyish brown forewings with three black dots and an interrupted black line along the termen. It is similar to M. imparellus (Figs
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.
Megacraspedus imparellus (? var.) majorella was described from two males from “Alai” (=Alai Mountains in Kyrgyzstan and Tajikistan), which were stated to be larger and having the two dots in the disc more separated compared with European M. imparellus (
Holotype, ♂, [Kazakhstan] “USSR 43°05'N, 77°15'E Kazakhstan, Zailiskiy Alatau, Alma-Atinskij Nat. P.” “1750 m, steppe slope/Carex creek, 14.vii.1990 ad luc., L Kaila & K Mikkola leg.” “Gen. prep. nr. 5324 ♂ O. Karsholt” (
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus latiuncus sp. n. is characterised by its light yellow, brownish and blackish dusted forewings with white veins. It is similar to M. longipalpella (Figs
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 Carex at an altitude of 1750 m.
The species name is a compound word derived from the Latin words latus (meaning wide) and uncus, referring to the broad uncus. The name is a noun in apposition.
The M. tenuignathos species group includes one species: M. tenuignathos sp. n.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Diagnostic remarks. The M. tenuignathos species group is defined by unique structures in the male genitalia. In particular the long and slender gnathos hook, almost extending to the anterior margin of the tegumen, and the strongly arched and evenly rounded posterior margin of the saccus as well as the distinct sub-apical tooth and supplementary small spines of the phallus are diagnostic.
The systematic position of the M. tenuignathos species group is uncertain and tentative due to the absence of females and supporting molecular data.
Holotype ♂, “MAROKKO Imilchil Hoher Atlas 2000 m Seeufer [lake shore] TF 11.08.1998 leg. Rolf Bläsius” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus tenuignathos sp. n. is among the largest species in the genus without a reduced segment 3 of the labial palpus. It is moreover characterised by its light yellow forewings with white veins edged with black without black dots. The male genitalia are unmistakable due to the long and slender gnathos hook, the strongly arched posterior margion of the saccus without medial emargination, and the characteristic spines of the phallus.
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 tenuis (meaning small) and gnathos, referring to the slender uncus. The name is a noun in apposition.
The M. glaberipalpus species group includes one species: M. glaberipalpus sp. n.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Diagnostic remarks. The M. glaberipalpus species group is defined by the unique structure of the labial palpus without a scale brush on segment 2. Furthermore the male genitalia exhibit some diagnostic features such as the presence of a short sacculus, the long saccus with a distinctly forked ridge and the broad and oblong sclerotisation of the ductus ejaculatiorius.
Holotype ♂, “Maroc, Haut Atlas, Toubkal Massif Tamatert near Imlil above Tamatert 27.vii.2008, leg. A. Steiner, 2400 m, 31°9'N, 7°53'W A. Steiner leg.” (
Adult. Male (Figs
Female (Figure
Variation. The amount blackish brown scales on the forewing varies, giving the wing either a yellowish or light brownish appearance. There is quite a large variation in the wingspan.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus glaberipalpus sp. n. is characterised by its light yellow to brownish forewings with two black spots. It is unique in Megacraspedus by the absence of a scale brush at segment 2 of the labial palpi. It otherwise resembles M. steineri sp. n. (Figure
BIN BOLD:ADF1285 (n = 1). The distance to the nearest neighbour, an unidentified Spilomelinae in BOLD, is 9.3%, the distance to the nearest congeneric species M. cerussatellus is 10.8% (p-dist).
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 glaber (= hairless or smooth) and palpus, referring to segment 2 of the labial palpi being without a scale brush. The name is a noun in apposition.
The M. imparellus species group includes one species: M. imparellus.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Diagnostic remarks. The M. imparellus species group is defined by diagnostic structures of the male genitalia such as the rounded shape of the uncus, the long and evenly curved gnathos hook, the presence of a short, digitate sacculus and in particular the very long and multiple contorted interior sclerotisation of the ductus ejaculatiorius. The female genitalia are characterised by the granulate microsculpture of segment VIII, and the ductus bursae and ductus seminalis, but females are unknown in several species groups and therefore this character may not be of diagnostic value.
Ypsolophus imparellus Fischer von Röslerstamm, 1843: 300, 303, pl. 100, figs 2a–d [on plate as ‘imparella’].
Megacraspedus
litovalvellus
Junnilainen, in
Paratypes of M. litovalvellus: Russia. 2 ♂, S-Ural, Orenburg oblast, 20 km S Pokrovka village, Schibendy valley, 6–7.vi.1998, leg. K. & T. Nupponen (
Adult. Male (Figs
Female (Figure
Variation.There is some variation in the amount of dark brown scales on the forewings. In some specimens they form a dark apical streak. Freshly emerged specimens are more greyish compared with specimens in collections. Specimens from Greece (Olympos) are slightly larger (wingspan 14–16 mm), slightly more broad-winged and with fewer dark scales on the forewing (thereby resembling M. leuca) but the reduction of dark scales may be due to worn specimens.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus imparellus is characterised by its light greyish brown forewings with three distinct black dots and an interrupted black line along termen. It normally has only a single pecten on the antennal scape, whereas the similar looking M. leuca (Figs
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 M. attritellus is 9.2% (p-dist).
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 M. litovalvellus was collected at artificial light at night and by sweeping before sunset, from late May to early August on chalk steppe (
Ypsolophus imparellus was described from two males and two females collected in June at Baden near Vienna, Austria (Fischer von Röslerstamm 1843). The excellent figures in the original description leave no doubt to the identity of this species. Megacraspedus litovalvellus was described from numerous specimens of both sexes collected in thesouthern Urals, Russia (
The M. consortiella species group includes eight species: M. multispinella, M. nupponeni sp. n., M. cerussatellus, M. attritellus, M. consortiella, M. pototskii sp. n., M. leuca, and M. orenburgensis.
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.
Diagnostic remarks. The M. consortiella species group is defined by diagnostic structures of the male genitalia, in particular the moderately large uncus, the long and slender, postbasally bent gnathos hook, and the large, oblong saccus with a long medial ridge. The female genitalia are similar overall to other species of Megacraspedus and mainly characterised by combined structures such the short apophysis posterior and anterior, the short, pointed sub-medial sclerites or membranous subostial area, and the broadly band-like connection of apophysis anterior with segment VIII. However, females are unknown in several species groups and therefore these characters may not be of diagnostic value.
Megacraspedus multispinella Junnilainen& Nupponen, 2010: 11, figs 19, 26.
Holotype ♂, “RUSSIA S-Ural Bashkiria Sakmara river Jantyshevo village 20.-21.6.1996 K. Nupponen, J.-P. Kaitila, J. Junnilainn & M. Ahola” “Prep. no. 03022406 ♂ Det. J. Junnilainen” “Megacraspedus aholai sp. n. ♂ Det. J. Junnilainen” “HOLOTYPE Megacraspedus aholai Junnilainen” “DNA sample KN00022 Lepid. Phyl.” (RCKN). Paratype. Russia. ♂, S-Ural, Cheliabinsk distr., near Moskovo vill., 18.vi.1998, leg. T. & K. Nupponen (RCKN). Non-type material. Russia. 19 ♂, 6 ♀, Tuva rep., 52°04'N, 94°22'E, Ust-Uljuk, steppe hills, 670 m, 3–5.vi.1995, leg. J. Jalava & J. Kullberg, genitalia slides GU 16/1418 ♂ Huemer, GU 16/1419 ♀ Huemer, genitalia prep. (in glycerin) (
Adult. Male (Figure
Female (Figure
Variation. There is a slight variation in the intensity of dark scales on the forewing.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus multispinella is characterised by the short segment 3 of the labial palpus, by the antennal scape with pecten having several hairs, and by the whitish grey forewings mottled with brownish- or blackish-tipped scales along apical margins. It resembles M. leuca (Figs
BIN BOLD:ACM0852 (n = 1) The distance to the nearest neighbour M. nupponeni sp. n. is 8.1% (p-dist).
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.
Megacraspedus multispinella was described from two males collected in the southern Ural Mountains, Russia (
Holotype ♂, “RUSSIA S-Buryatia 51°11–13'N 106°10–12'E 700 m, Hamar Daban mnts Murtoy river, Gusinoe Ozero vill. 6 km NW, forest steppe 27.5.2006 K. Nupponen leg.” “prep. No. 8/8.X.2006 K. Nupponen” “DNA sample KN00089 Lepid. Phyl.” (RCKN). Paratypes. Russia. 1 ♀, same data as holotype but genitalia slide 4/16.7.2017 K. Nupponen; 1 ♂, Buryatia rep., Chikoy valley, 10 km Novoselengnisk vill., 550–600 m, sand dunes/sandy steppe, 2.vi.2006, leg. K. Nupponen (RCKN).
Adult. Male (Figure
Female (Figure
Variation. Unknown.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus nupponeni sp. n. is characterised by its small size, its relatively short and broad forewings, the antennal scape with several hairs, and by its clear white forewings with scattered black scales and dots. It resembles some forms of M. attritellus (Figure
BIN BOLD:ACB0748 (n = 1). The distance to the nearest neighbour M. multispinella sp. n. is 8.1% (p-dist).
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.
Megacraspedus cerussatellus Rebel, 1930: (14).
Lectotype ♂, here designated, [Bulgaria] “Alibotusch [mountains] 1700 m 14.VII.29” “Megacraspedus cerussatellus Rbl Type ♂” “BC
Adult. Male (Figure
Female. Unknown.
Variation. Only one specimen was examined.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus cerussatellus is characterised by its short segment 3 of the labial palpus, the antennal scape with several hairs, and by its pure white forewings with scattered black scales and dots. It is most similar to M. multipunctellus sp. n. (p 167). The male genitalia are very similar to M. attritellus (Figure
BIN BOLD:ACA8764 (n = 1). The distance to the nearest neighbour M. attritellus is 7.6% (p-dist).
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.
Megacraspedus cerussatellus was described from several, mostly defective, males in poor condition, collected by A. K. Drenowski in the Alibotusch Mountains, S Bulgaria on 24.vii.1929 (
Megacraspedus attritellus Staudinger, 1871: 316.
Lectotype ♂, here designated, [Russia, Volgograd oblast] “Lecto-type” “Sarepta [Krasnoarmeysk] Chr.[istoph]” “Origin.” “Lectotype ♂ Megacraspedus attritellus Stdgr teste K. Sattler, 1986” “ex coll. Staudinger” “GU 16/1426 ♂ P.Huemer” (
Adult. Male (Figure
Female. Unknown.
Variation. There is variation in the amount of blackish brown scales on the forewing. Worn specimens become whitish.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus attritellus is characterised by segment 3 of the labial palpus being short, by the pecten of the antennal scape, and by its blackish brown forewings with a whitish longitudinal streak and three elongate black spots. Among species with a scale brush at segment 2 and short segment 3 of the labial palpus it resembles M. leuca most, but that species is larger, and has lighter forewings with indistinct black dots. It is furthermore similar to M. nupponeni sp. n. (p 135) and M. orenburgensis (p 144). The male genitalia are very similar to M. cerussatellus (Figure
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 M. leuca is 6.8% (p-dist).
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 (
Megacraspedus attritellus was described from four males collected by H Christoph in Sarepta (now Krasnoarmeysk), southern Russia (Staudinger 1871). A lectotype, already labelled as such by K Sattler, is here designated in order to fix the identity of the species and conserve stability of nomenclature.
Megacraspedus consortiella Caradja, 1920: 117.
Holotype ♂, [Kyrgyzstan] “Alai [mountains]” “HOLOTYPE Megacraspedus consortiella ♂ Car. DES. Dr. A. POPESCU-GORJ Romania” “176436” “CIS-Korea Microlep. 4218 Megacraspedus consortiella Car. – Alai, Himalaya K. T. Park ‘96” (
Non-type material. Kyrgyzstan. 1 ♂, Alai mts, Tengiz-Bai Gate, 2800–2900 m, 12.vii.2011, leg. A. Pototski (RCAP); 3 ♂, Alai mts, Ak-Bosogo, 2725 m, 31.vii.2010, leg. A. Pototski (RCAP,
Adult. Male (Figure
Female. Unknown.
Variation. In four of the examined specimens the black dot at end of the fold (which is present in the holotype) is almost obsolete and the hair at the base of the antennal scape is missing, but these specimens are rather worn.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus consortiella is characterised by the grey forewings with a black dot at the end of the cell, and by having most of the margin of the costa white. It is very similar to and hardly separable from M. pototskii sp. n. (p 141). The male genitalia are similar to M. cerussatellus (Figure
BIN BOLD:ADG5879 (n = 1). The distance to the nearest neighbour M. leuca is 6.6% (p-dist).
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.
Megacraspedus consortiella was described from one male in good condition from “Alai” (= Alai Mountains in Kyrgyzstan and Tajikistan) (
Holotype ♂, “KYRGYZSTAN Alai mts 3650 m 39°38'51.1"N 72°14'00.1"E Tengiz-Bai pass 23.7.2010 K. Nupponen & R. Haverinen leg.” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. There is a slight variation in the amount of light yellowish in the forewing, and there can be indistinct black dots fold, and in the middle of wing at 3/5. The few examined specimens are also variable in size.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus pototskii sp. n. is characterised by the light yellowish grey forewings with a black dot at the end of the cell, and by having most of the margin of the costa white. It is very similar to and hardly separable from M. consortiella (Figure
BIN BOLD:ADI8386 (n = 2). The intraspecific divergence of the barcode region is 0%. The distance to the nearest neighbour M. ibericus sp. n. is 9.1% (p-dist).
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 Megacraspedus from Asia available for our study.
Nothris leuca Filipjev, 1929: 9, pl. 2a, fig. 3.
Megacraspedus kaszabianus Povolný, 1982: 193, figs 1–2, syn. n.
Paratypes of M. kaszabianus. Mongolia. 1 ♂, Gobi Altaj aimak, Chasat chajrchan ul mts, cca. 20 km S Somon Zargalan, 2400 m, 15.vii.1966, leg. Z. Kaszab; 1 ♂, Südgobi aimak, Gurban Sajchan ul, 15 km S Dalanzadgad, 1750 m, 13.vi.1967, leg. Z. Kaszab; 1 ♂, 1 ♀, Südgobi aimak, Somon Bulgan Talyn bulag, 1350 m, 5.vii.1967, genitalia slides GU 16/1446 ♂ Huemer, GU 16/1447 ♀ Huemer, leg. Z. Kaszab (
Adult. Male (Figure
Female (Figure
Variation. The amount of brownish- or blackish-tipped scales of the forewing varies; they are normally scattered over the wing but can also form lines along the veins. The blackish brown dots on the forewing can be more or less distinct. One specimen (in good condition) is almost without dark scales.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus leuca is characterised by having the antennal scape with a pecten of several hairs, and by the having the cream whitish grey forewings mottled with brownish- or blackish-tipped scales. It resembles M. multispinella (Figs
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 M. skulei is 6.6% (p-dist).
Asian part of Russia, Mongolia.
Host plant and early stages are unknown.
Nothris leuca was described from one male collected at Munku-Sardyk (=Mönkh-Saridak) in the Sajan Mountains at the border between Russia and Mongolia (Filipjev 1929). The holotype in ZISP was studied and figured by
Megacraspedus kaszabianus was described from several hundred males collected in Mongolia by Z. Kaszab (
Megacraspedus orenburgensis Junnilainen & Nupponen, 2010: 13, figs 20–21, 27
Holotype ♂, “RUSSIA S-Ural Orenburg district Pokrovka village 20 km S Schibendy valley 21.6.1999 T. & K. Nupponen leg.” “Prep. no:03022405 Det J. Junnilainen” “Megacraspedus orenburgensis sp. n. ♂ Det J. Junnilainen” “HOLOTYPE Megacraspedus orenburgensis Junnilainen & K. Nupponen” (RCKN). Paratype. 1 ♂, same data as holotype, but 2.viii.2005, leg. K. Nupponen, genitalia slide no. 2/17.IV2006 K. (RCKN).
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus orenburgensis is characterised by the short segment 3 of the labial palpus and by the pecten of the antennal scape having many hairs. Among species with a scale brush on segment 2 and a short segment 3 of the labial palps it mostly resembles M. leuca (Figs
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 (
Megacraspedus orenburgensis was described from two males collected in the southern Urals, Russia (
The M. lagopellus species group includes five species: M. lagopellus, M. coleophorodes, M. feminensis sp. n., M. kirgizicus sp. n., and M. argyroneurellus.
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.
Diagnostic remarks. The M. lagopellus species group is mainly defined by the characteristic labial palpus and diagnostic structures of the male genitalia. The small and pointed uncus, the comparatively massive gnathos hook, the strongly arched posterior edge of the saccus, and in particular the broad valva with an apical spine are diagnostic. The highly adapted ovipositor in the female genitalia is only shared with M. kazakhstanicus sp. n., but females are unknown for the majority of Megacraspedus. This structure seems suitable for a specialised form of oviposition inside the host plant.
Megacraspedus lagopellus Herrich-Schäffer, 1860: 13, pl. [14], fig. 81.
Hungary. 1 ♂, Ofen [Budapest], 1868, leg. Anker, genitalia slide Mus. Vind. 16.656; 1 ♀, same data, but 1874, genitalia slide NM 16657 (all
Adult. Male (Figure
Female (Figure
Variation. The brownish part of the labial palpus can be more or less distinct, and the forewing colour can vary from whitish yellow to light yellow. Specimens tend to become greasy and the whitish parts of the head and body thereby turn brown.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus lagopellus is characterised by the long scale brush on segment 2 of the labial palps and by its light yellowish forewings with indistinct white on the veins. It resembles the larger M. argyroneurellus (Figs
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 M. feminensis sp. n. is 7.8% (p-dist).
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.
Megacraspedus lagopellus was described from an unstated number of specimens from Sarepta (now Krasnoarmeysk), southern Russia (Herrich-Schäffer 1860). We have not been able to trace any syntypes, but the identity of this species is fixed by Herrich-Schäffer’s figure (op. cit.: fig. 81). M. lagopellus is unusual among Megacraspedus because more females than males were available for study.
Chilopselaphus coleophorodes Li & Zheng, 1995: 1, 10, figs 1–4.
Holotype ♂, China, Liancheng, Yongdeng County, Gansu Province, 6-July-1985, Coll. Houhun Li, genitalia slide no L94136 (
Adult. Male (Figure
Female (Figure
Variation. According to the original description the colour of the forewings varies from grey-brown to grey-white.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus coleophorodes is characterised by its rather large size, the long scale brush on segment 2 of the labial palps, and by its yellowish forewings with white on the veins, but not on the costa. It resembles M. argyroneurellus (Figs
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.
Chilopselaphus coleophorodes was described from one male (holotype) from the Gansu province and 11 females from Ningxia Hui Autonomous Region, China (
Holotype ♀, “Kazakhstan 43°47'55"N 79°54'51"E Rahat Kuduk by Ili River shore tugai forest, 515 m 2.VI.2014 K. Nupponen & R. Haverinen leg.” “DNA Barcode
Adult. Male. Unknown.
Female (Figure
Variation. The colour of the upper and lower surface of the labial palpus varies from predominantly light yellow to white. Worn specimens become lighter.
Male genitalia. Unknown.
Female genitalia (Figure
Megacraspedus feminensis sp. n. is characterised by its rather large size, by the long, porrecting labial palps, and by its yellowish forewings with white veins. It resembles M. coleophorodes (Figure
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 M. lagopellus is 7.8% (p-dist).
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.
Megacraspedus feminensis sp. n. is one of the few species in the genus with predominant numbers of collected females, resembling other closely related species of the M. lagopellus species group.
Holotype ♂, “Kirgisien/ Prov. Batken, Turkestan Geb., S Zardaly, Stat. Korgon HP 19 1750 m 8.6.2010 GPS 39°57'52"N;70°58'36"E leg. Dr. C. Wieser Kärntner Landesmuseum” “KLM Lep 00111” “GU 16/1406 ♂ P. Huemer” (
Adult. Male (Figure
Female (Figure
Variation. There is a slight variation in the colour of the head, thorax, and tegulae: from almost white to light grey-brown. The single, soft hair at the base of the antennal scape is missing in several of the examined specimens (especially the males), but that may be because the specimens have been damaged.
Male genitalia (Figs
Female genitalia (Figure
Diagnosis. Megacraspedus kirgizicus sp. n. is characterised by its large wingspan, the very long labial palps, and by the light yellow forewings with a broad light grey streak from base to apex.
It resembles M. podolicus (Figure
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 M. argyroneurellus is 7.6% (p-dist).
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.
Megacraspedus argyroneurellus Staudinger, 1871: 316.
Lectotype ♂, here designated, [Russia, Volgograd oblast] “Lecto-type” “Sarepta [Krasnoamreysk] Chr.[istoph]” “Origin.” “Lectotype ♂ Megacraspedus argyroneurellus Stdgr teste K. Sattler, 1986” “ex coll. Staudinger” (
Adult. Male (Figs
Female (Figure
Variation. The size of the brownish area in the middle of the forewing varies and can extend into the apical area or alternatively be largely reduced. Specimens from the south-eastern part of the distribution range have no brown area in the middle of the forewing (see remarks below). The hair at the base of the antennal scape is easily broken.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus argyroneurellus is characterised by its large size, its long scale brush on segment 2 of the labial palps and by its yellowish forewings with silvery white veins and its white hindwings. It is similar to M. lagopellus (p 146), M. coleophorodes (p 148), and M. feminensis sp. n. (p 149). The male genitalia are similar to M. kirgizicus sp. n. (Figs
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 M. kirgizicus sp. n. is 7.6% (p-dist).
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.
Megacraspedus argyroneurellus was described from an unspecified number of specimens (probably more than one as the wingspan is given as “19–21 mm”) collected at Sarepta, now Krasnaormeysk, in south-western Russia by H. Christoph (Staudinger 1871). A lectotype, already labelled as such by K. Sattler, is here designated in order to fix the identity of the species and conserve stability of nomenclature.
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 M. violacellum species group includes two species: M. ibericus sp. n. and M. violacellum.
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).
Diagnostic remarks. The M. violacellum species group is defined by a combination of structures of the male genitalia, e.g., the large, nearly sub-square uncus, the abruptly bent gnathos hook, the broad valva, and particularly the suboval saccus and the rounded lateral hamps of the vinculum are characteristic.
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.
Holotype ♂, “SPAIN [prov.] Málaga 2 km NE Mijas, 560 m, 24.ix.2012 J. Tabell leg.” (
Adult. Male (Figure
Female. Unknown.
Variation. The species is variable in size. Worn specimens look light brownish.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus ibericus sp. n. is characterised by the long segment 2 of the labial palps, the blackish brown antennae, and by the brownish forewings with three black spots and a yellow fold dorsally edged black. It resembles M. spinophallus sp. n. (Figure
BIN BOLD:ACZ7298 (n = 1). The distance to the nearest neighbour M. violacellum is 7.5% (p-dist).
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.
Toxoceras violacellum Chrétien, 1915: 330.
Cauloecista chretienella Dumont, 1928: 34, figs 5A, B, syn. n.
Cauloecista halfella Dumont, 1928: 36, figs 5C, D, syn. n.
Mesophleps arnaldi Turati & Krüger, 1936: 76, pl. 11, fig. 27, syn. n.
Lectotype ♂, Toxoceras violacellum, here designated, “Toxoceras violacellum” “Gafsa 29.II.08” “TYPE” “Genitalia ♂ P. Viette Prèp. No. 2673” (
Lectotype ♂, Mesophleps arnaldi (without abdomen), designated by
Adult. Male (Figure
Female [based on original description and figure of lectotype] (Figure
Variation. The forewings vary from lighter to darker greyish brown, and there can be one or two additional small dots in the middle of the forewings. The specimen from Sbeidla is slightly more broad-winged and has the dorsal part of the forewing yellowish brown.
Male genitalia (Figure
Female genitalia. Undescribed. Unfortunately we did not obtain permission to examine the genitalia of the only available female.
Megacraspedus violacellum is characterised by its large size and its light grey-brown forewings with two brown streaks and two black dots. It resembles M. squalida (Figs
BIN BOLD:ADI7610 (n = 1). The distance to the nearest neighbour M. ibericus sp. n. is 7.45% (p-dist).
Libya, Tunisia.
The larva and pupa are described in details by
Toxoceras violacellum was described from an unstated number of specimens collected at Gafsa, Tunisia in November (
Cauloecista chretienella was described from an unstated number of specimens (“♂♂, ♀♀”) bred from larvae collected by Al-Mitlawi [“Metlaoui”], central Tunisia (
Cauloecista halfella was described from two females bred from larvae collected by Al-Mitlawi [“Metlaoui”], in central Tunisia (
Mesophleps arnaldi was described from two specimens collected by G. Krüger at Az Zuwaytīnah (‘Zuetina’), Libya (
A single male specimen from Sbeidla, Tunisia, coll. C. S. Larsen (
The M. squalida species group includes one species: M. squalida.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Diagnostic remarks. The M. squalida species group is defined by several unique and highly diagnostic structures in the male genitalia, in particular the shape of the valva with strong saccular bristles and the curved thorn-like process near base of valva. The female genitalia are characterised by the membranous ostium bursae, without delimiting sclerotisations, but females are unknown in several species groups.
Megacraspedus squalida Meyrick, 1926: 272.
Megacraspedus escalerellus Schmidt, 1941: 38, pl. 2, fig. 5, syn. n.
Holotype ♂, Megacraspedus escalerellus, [Spain] “Spain Montarco Madrid ix.1920 F. Escalera” “Genitalia slide Mus. Madrid 53788” (
Adult. Male (Figure
Female (Figure
Variation. There is only slight variation. The black dots in fold and at end of the cell are not always visible.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus squalida is characterised by its relatively large size and its blackish grey forewings with indistinctly lighter veins. The males, moreover, are recognizable by the thickened, black antennae. The species is somewhat similar to M. peyerimhoffi (p 181) and M. violacellum (p 157). The flap-like sclerite with thorn-like process in the male genitalia is unique in Megacraspedus. The female genitalia are unique in Megacraspedus with several diagnostic features such as the membranous area of the ostium bursae.
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 M. skulei sp. n. is 9.3% (p-dist).
Spain, Tunisia.
The larva feeds within a stem of Macrochloa tenacissima (Loefl. ex L.) (Poaceae) (
Megacraspedus squalida was described from one female collected by Ribbe in the Sierra Nevada, Spain (Meyrick 1926). The holotype and its genitalia were figured by
The ‘allotype’ of Cauloecista chretienella Dumont, 1928 has proved to belong to M. squalida (see remarks on M. violacellum). It was bred from Macrochloa tenacissima (Loefl. ex L.) (
The M. pentheres species group includes eight species: M. pentheres, M. steineri sp. n., M. gibeauxi sp. n., M. multipunctellus sp. n., M. teriolensis sp. n., M. korabicus sp. n., M. quadristictus, and M. eburnellus.
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.
Diagnostic remarks. The M. pentheres species group is defined by structures of the male genitalia, in particular the short and suboval, basally constricted uncus, the short and medially widened gnathos hook, the valva with usually separated sacculus, and the distally narrowing phallus with a sclerotised, denticulate ridge. The female genitalia are characterised by the comparatively large papilla analis, and the short apophysis posterior and anterior as well as segment VIII, but females are unknown in several other species groups.
Megacraspedus pentheres Walsingham, 1920: 10.
Paratype ♂, “Basses Alpes S FRANCE 22.viii.1913 Wlsm. 1913-390 95935” “Paratype” “Megacraspedus pentheres Wlsm. PARATYPE”, genitalia slide 33662 (
Adult. Male (Figure
Female. Unknown.
Variation. The examined specimens show only slight variation. One specimen has a fourth black dot in the forewing. The pecten is easily lost as demonstrated from a specimen with 3 hairs on one and 5 hairs on the other side.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus pentheres is characterised by light yellow-brown forewings with three black dots and black scales along termen. It closely resembles M. tristictus (Figure
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.
Megacraspedus pentheres was described from seven males collected by Walsingham at Annot, Alpes-de-Haute-Provence, France (Walsingham 1920). The label dates of the examined paratype disagree with the dates published in the original decription, as ’9–12.VIII.1913’.
Holotype ♂, “Maroc, Haut Atlas Toubkal Massif Oukaïmeden area piste Oukaïmeden-Tizi-n-Eddi 31°10'N, 7°50'W 3.viii.2008, 2815 m A. Steiner leg.” “GU 16/1421 ♂ Huemer” “DNA Barcode
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus steineri sp. n. is characterised by its light yellow forewing with three black spots. It has moreover a long scale brush at segment 2 of the labial palpi. The species is most similar to M. glaberipalpus sp. n. (p 128). The male genitalia are very similar to some other species of the M. pentheres species group with only small differences e.g., such as the medial ridge of the saccus or the absence of spines on the phallus.
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.
Holotype ♂, “Tozeur [south-western] TUNESIA 15.–29.x.1988 P. Sunesen col. K. Larsen” “GU 16/1442 ♂ P. Huemer” (
Adult. Male (Figure
Female. Unknown.
Variation. The intensity of the grey-brown scales is variable.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus gibeauxi sp. n. is characterised by its whitish forewing with grey-brown scales along the veins and by the absence of black dots on the forewing, and by the antennal scape having a pecten of several rather long hairs. The male genitalia are very similar to other species of the M. pentheres species group and differ only in subtle characters, in particular the distinct humps of the vinculum and the comparatively slender phallus.
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 “?Paltodora pectinatella 4923′” (in Chrétien’s handwriting) and “TYPE”. Chrétien apparently considered the specimens to belong to an undescribed species, but a description was not published.
Holotype ♂, “TURKEI Prov. Konya, 36°57'N 33°17'E Setravul Geçidi 1550 m 28 km S Karaman, 3.09.1983 LF leg. Werner Wolf” “GU 16/1455 ♂ Huemer” (ECKU). Paratype. Turkey. 1 ♂, same data as holotype, but without genitalia slide (
Adult. Male (Figure
Female. Unknown.
Variation. None.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus multipunctellus sp. n. is characterised by its short segment 3 of the labial palpus, the antennal scape with several hairs, and by its cream-white forewings with scattered black scales and dots. It is most similar to M. cerussatellus (Figure
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 multus (meaning many) and punctus (meaning dot), and the diminutive suffix –ellus. The name is a noun in apposition.
Holotype ♂, [Italy, Provinz Bozen-Südtirol] “ITALIA sept., Südtirol Montiggl, Kl. Priol 600 m, 30.8.1995 leg. Huemer” “BC
Adult. Male (Figs
Female. Unknown.
Variation. The forewing colour varies from yellowish (especially in the dorsal half) to light grey- brown with yellow only in the fold and along dorsum. Two examined specimens from Croatia have the forewing blackish brown with clear yellow longitudial streaks.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus teriolensis sp. n. is characterised by its light grey-brown forewings with four black dots, and a yellow streak in the fold. It is very similar to M. quadristictus (p 172). The male genitalia are very similar to several other species of the M. pentheres species group but differ in the particularly stout and well sclerotised uncus and other subtle characters. From the most similar M. quadristictus (Figs
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 M. quadristictus is 9.7% (p-dist).
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 M. teriolensis sp. n., M. korabicus sp. n., and M. quadristictus is almost entirely based on differences in the barcode, because morphological differences are minute.
Holotype ♂, “Macedonia NP [National Park] nMavrovo Korab [mountain], Korabska jezero, Kobilino pole, 2080–2180 m 20°34'55"E, 41°46'42"N 28.7.–1.8.2011 leg. Huemer & Tarmann” “Megacraspedus binotella det. P. Huemer, 2011” “P. Huemer GEL 1183 ♂” “BC
Adult. Male (Figure
Female (Figure
Variation. The yellowish dorsal part of the forewing can be more or less mottled with greyish scales. The hairs on the antennal scape are easily broken. Worn specimens become lighter.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus korabicus sp. n. is characterised by cream coloured forewing with four black dots. It is very similar to M. quadristictus (Figs
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 M. quadristictus is 10.1% (p-dist).
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.
Megacraspedus quadristictus Lhomme, 1946: 537.
Lectotype ♂, here designated, “quadristictus” “Cousson 11.8.03” genitalia slide PGCG 8018 (
Adult. Male (Figure
Female (Figure
Variation. The forewing colour varies from yellowish to greyish. A few specimens have an indistinct, dark, sub-costal patch near base. Worn specimens become lighter, and the black dots can become obsolete, and in such specimens the antennal pecten is often lost.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus quadristictus is characterised by its light grey-brown forewings with yellow veins and dorsum and four black dots, and the antennal scape with a pecten of fine hairs. It is very similar to M. teriolensis sp. n., but is on average larger and has more yellow on the forewing, especially along the veins. See also M. korabicus sp. n. (p 170). The male genitalia are very similar to several other species in the M. pentheres species group but differ by the particularly stout and well sclerotised uncus and other more subtle characters. From the most similar M. teriolensis sp. n. (Figs
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 M. teriolensis sp. n. is 9.7% (p-dist).
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.
Megacraspedus quadristrictus was described from an unstated number of specimens (“quelques exemplars”) from France, Digne-Le Cousson, in the collection of P. Chrétien (Lhomme 1946: 537). We have been able to examine photographs of syntypes from
Megacraspedus eburnellus Huemer & Karsholt, 2001: 238, figs 3, 8, 10, 13–14.
Holotype ♂, “Italia [prov. Verona] Mte Baldo Noveza 1300–1600 m M.6.66. leg. Burmann” “
Adult. Male (Figure
Female (Figure
Variation. The forewing colour varies from whitish cream to light brown. Worn specimens look whitish. The tip of segment 3 of the labial palps is sometimes black.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus eburnellus differs from most other European Megacraspedus species in lacking blackish brown or black scales, dots, or streaks on the forewings. It is externally similar to M. dolosellus (Figs
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 M. skulei sp. n. is 8.5% (p-dist).
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 M. longivalvellus species group includes two species: M. skulei sp. n. and M. longivalvellus.
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.
Diagnostic remarks. The M. longivalvellus species group is defined by the long and porrecti labial palpus with reduced segment 3, and in particular by several structures of the male genitalia such as the extraordinarily long and apically curved valva, the saccular bulge, the oblong saccus, and the long and slender phallus. The female genitalia are characterised by the comparatively large papilla analis, and the short apophysis posterior and anterior as well as segment VIII, and the membranous anteromedial edge of segment VIII. They are similar overall to e.g., M. peyerimhoffi and a few other species but females are unknown in several other species groups.
Holotype ♂, “SPAIN [prov.] Almeria Sierra de los Filabres Calar Alto, 2130 m N37.221523 W2.545495 5.7.2015 J. Tabell leg.” “DNA Barcode
Adult. Male (Figure
Female (Figure
Variation. There is some variation in size, even among specimens collected at the same date. The black dots on the forewing before apex can be absent. Strongly marked specimens have a black streak before the apex. Worn specimens become more greyish.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus skulei sp. n. is, together with M. knudlarseni sp. n. (p 122), the smallest species amongst Megacraspedus with a reduced segment 3 of the labial palps. It is similar to M. longivalvellus sp. n. (p 179). The male genitalia are almost unique within Megacraspedus and can be easily recognized by the extremely long and distally curved valva. They differ from the closely related M. longivalvellus sp. n. (Figs
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 M. longivalvellus sp. n. is 2.3% (p-dist).
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 Megacraspedus specimens used for our study.
The species status of the closely allied and allopatric M. skulei sp. n. and M. longivalvellus sp. n. is supported by morphology and DNA barcodes.
Holotype ♂, “Morocco [Middle Atlas] 1400–2000 m Azrou/Ifrane area 17–19.iv.1989 Zool. Mus. Copenh. Exp.” “GU 16/1422 ♂ P. Huemer” (
Adult. Male (Figure
Female. Unknown.
Variation. The amount of black scales and the distinctness of the black spots on the forewing are variable. The yellowish white tinge on the forewing is only present in fresh specimens.
Male genitalia (Figs
Female genitalia. Unknown.
Megacraspedus longivalvellus sp. n. is similar to M. skulei sp. n. (Figure
BIN BOLD:ADF1825 (n = 2). The intraspecific divergence of the barcode region is moderate with mean 0.9%. The distance to the nearest neighbour M. skulei sp. n. is 2.3% (p-dist).
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 longus and valva, indicating the characteristically elongated valva, and the diminutive suffix –ellus. The name is a noun in apposition.
DNA barcode divergence and diagnostic characters of adults including the male genitalia support a separate specific status of M. skulei sp. n. and M. longivalvellus sp. n.
The M. peyerimhoffi species group includes one species: M. peyerimhoffi.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Female genitalia. See species description.
Diagnostic remarks. The M. peyerimhoffi species group is defined by structures of the male genitalia, in particular the characteristic shape of the uncus and gnathos hook, the long valva without separated sacculus, and the sclerotised ridges of the saccus. The female genitalia are characterised by combined characters such as the short apophysis posterior and anterior and the long sub-medial sclerites of segment VIII, but females are unknown in several species groups and the diagnostic value of these structures is uncertain.
Megacraspedus peyerimhoffi Le Cerf, 1925: 12, figs 1–19.
Lectotype ♂, Megacraspedus peyerimhoffi, here designated, “TYPE” “Megacraspedus Peyerimhoffi ♂ Le Cerf” “MUSÉUM PARIS F. LE CERF” “chenille dans alfa Boaira-Sahari V-VI-24 P. de Peyerimhoff áclos Paris 12/13-VI-1925” genitalia slide PGCG 7964 (
Adult. Male (Figure
Female (Figure
Variation. There is only slight variation, apart from size. One female has a slender sub-costal line near base. Worn specimens become lighter.
Male genitalia (Figs
Female genitalia (Figure
Megacraspedus peyerimhoffi is characterised by its large wingspan and its elongate, greyish brown forewing without stripes or markings, apart from the slender white streak on the costa. It is often collected together with the somewhat similar to M. squalida (Figs
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 M. skulei sp. n. is 9.5% (p-dist).
Algeria, Spain.
The larva feeds within the lower part (above the earth surface) of a stem of Macrochloa tenacissima (L) Kunth (Poaceae), eating the pith and thereby hollowing the stem. It spins 4–7 opercula of silk within its mine. When fullfed it lines a chamber in the bottom of the mine with silk, gnaw an exit hole and pupates in the mine. The pupa is standing head upwards on excrements which are pressed together in the bottom of the mine. The exit hole is placed so that it is covered by a leaf sheat, which the hatching adult has to slide between to reach the outside (
Although Le Cerf (op. cit.: 21) stated that his observations on the behavior of the larvae of M. peyerimhoffi were incomplete, they are the most detailed given for any Megacraspedus species. The adults have been collected from early September to the beginning of November at altitudes from sea level to 1300 m.
Megacraspedus peyerimhoffi was described from one male and four females bred from larvae and pupae collected by M. de Peyerimhoff in the region of Bouira-Sahari (=Had sahari), northern Algeria (
The M. peslieri species group includes one species: M. peslieri sp. n.
External morphology. See species description.
Genitalia morphology. Male genitalia. See species description.
Diagnostic remarks. The M. peslieri species group is defined by a combination of characters such as the short and almost knife-shaped valva, the massive and smoothly sclerotised saccus without a medial ridge, and in particular the long thorn of the phallus extending from the middle to the apex which is a unique structure.
The systematic position of the M. peslieri species group is uncertain and tentative due to the absence of females and supportive molecular data.
Holotype ♂, “France Pyr.[enees]-Or.[ientales] Conat (Bel Loc) 13-IX-2017 880 m Peslier leg.” “P. Huemer GEL 1274 ♂” (
Adult. Male (Figure
Female. Unknown.
Variation. The specimen from Spain has a black streak in the fold.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus peslieri sp. n. is characterised by the cream-brown forewings with a black streak through the fold and a black dot at end of cell. It is similar to M. binotella (Figs
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 M. grisea species group includes three species: M. grisea, M. pacificus sp. n., and M. armatophallus sp. n.
The M. grisea species group is defined by following characters:
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.
Diagnostic remarks. The M. grisea species group is defined by several structures in the male genitalia, in particular the broad valva with a transverse scleorized ridge or flap, the large V-shaped saccus with a weakly developed longitudinal ridge and in particular the phallus with a prominent, weakly dentated sub-basal thorn or two rod-like, weakly dentated sclerotisations. The female genitalia are in particular characterised by the shape of the subostial sclerotisation of segment VIII and the longitudinal sclerotised zone venula of segment VIII. However, it has to be acknowledged that females are unknown in several other species groups.
Reichardtiella grisea Filipjev, 1931: 168, fig. 16, pl. 10, figs 5–7.
Syntype ♂, [China] “Reichardia grisea Flj. N. Filipjev det. [partially illegible]” “Para-lecto-type” “Paralectotype Reichardtiella grisea Filipjev teste. K. Sattler, 1978” “Alai valley, loc. Sary-tash Reihardt 27.VI.928 [transcribed from Cyrillic]” “Pres. by Zool. Inst. Leningrad B.M. 1978-417” “
Adult. Male (Figure
Female. Unknown.
Variation. Unknown.
Male genitalia (Figure
Female genitalia. Unknown.
Megacraspedus grisea is characterised by the almost plain dark grey forewings. Externally it mostly resembles M. fuscus sp. n. (p 67). The male genitalia are unmistakable in Megacraspedus, in particular due to the characteristic shape of the valva and the dentation of the phallus.
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 (
Reichardtiella grisea was described from several males in variable condition collected at Sary-Kol in the Chinese part of the Pamir Mountains at an altitude of 4300 m (
Holotype ♂, “29.VII.1963 Afghan.[istan] centr., O v [East of] Band-i-Amir, 3600 m Kasy & Vartian leg.” “Gen. Präp. Mus. Vind. 16.661 ♂” (
Adult. Male (Figure
Female (Figure
Variation. None from the few examined specimens.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus pacificus sp. n. is characterised by its light greyish brown forewings with three weak black dots and an interrupted black line along the termen. It is similar to M. imparellus (Figs
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 pacificus, in the sense of bringing peace to the type locality of Afghanistan.
Holotype ♂, “SO-AFGHANISTAN Safed Koh, S-Seite Shahidan, 2700 m 21.6.1966 H. G. Amsel leg.” “
Adult. Male (Figure
Female (Figure
Variation. The female has no pecten on the antennal scape, but as it is rather worn such may have been lost.
Male genitalia (Figure
Female genitalia (Figure
Megacraspedus armatophallus sp. n. is characterised by the greyish brown forewings with three weak black dots and an interrupted black line along the termen. It is similar M. pacificus sp. n. (Figs
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 armatus (meaning armed warrior) and phallus, indicating the weapon-like shape of the phallus.
The phylogeny of numerous Gelechiidae genera is still far from resolved and this also applies to Megacraspedus. The first comprehensive molecular study of the family by
The correct identification of Gelechiidae to species level is a difficult task, even in some already revised genera. Characters traditionally used to delineate taxa are mainly found in the genitalia and external morphology, but the degree of diagnostic features varies from group to group. In several genera even the genitalia morphology is only of limited value for species delimitation, as shown in a monographic review of Palearctic Gnorimoschemini (
Large intraspecific barcode divergence as observed in Megacraspedus is generally rare in Lepidoptera (
Similar to other Gelechiidae genera with brachypterous females (
Lepidoptera is one of the best known insect orders, and in Europe even the smaller so-called Microlepidoptera is considered relatively well known. Even though several new species of Lepidoptera from Europe are found and described every year it is very unusual to have 22 new European species in a single genus. When compared with tropical areas Europe is normally not considered to be a hotspot in biodiversity, but, contrary to general belief, developed and heavily-studied parts of the world are important reservoirs of unknown species, making Europe an unexpected frontier for biodiversity exploration (
We have examined a vast number of Megacraspedus specimens collected during our own fieldwork, and from private and institutional collections. The result is the first comprehensive revision of the genus. A considerable number of the species of Megacraspedus dealt with here, both new species and some described long ago, are only know from a few specimens or from one or two localities. It is therefore likely that further species remained unrecognized. It is our hope that the present revision will stimulate lepidopterists to give more priority to Megacraspedus and search for both unknown and already known species in new places, as well as for some of the many unknown females, and perhaps investigate the early stages and biology. This may also result in a better understanding of the reasons for the high genetic variation in some species.
Megacraspedus adults in dorsal view. 7 M. lanceolellus (Zeller, 1850) – male, Italy (
Megacraspedus adults in dorsal view. 13 M. lanceolellus (Zeller, 1850) – male, Spain (
Megacraspedus adults in dorsal view. 19 M. homochroa Le Cerf, 1932 – male, Morocco (
Megacraspedus adults in dorsal view. 25 M. similellus sp. n. – Holotype, male (RCJJ) 26 M. similellus sp. n. – Paratype, female (RCJJ) 27 M. golestanicus sp. n. – Paratype, male (
Megacraspedus adults in dorsal view. 31 M. dolosellus (Zeller, 1839) – male, Greece (
Megacraspedus adults in dorsal view. 37 M. dolosellus (Zeller, 1839) – male (form incertellus), Bulgaria (RCZT) 38 M. dolosellus (Zeller, 1839) – male, Greece (
Megacraspedus adults in dorsal view. 43 M. cuencellus Caradja, 1920 – male, Spain (RCZT) 44 M. bidentatus sp. n. – Holotype male, Spain (
Megacraspedus adults in dorsal view. 49 M. alfacarellus Wehrli, 1926 – Holotype, male, Spain (
Megacraspedus adults in dorsal view. 55 M. occidentellus sp. n. – Holotype, male, Portugal (
Megacraspedus adults in dorsal view. 61 M. devorator sp. n. – Paratype, male, Romania (RCJJ) 62 M. binotella (Duponchel, 1843) – male, Austria (RCEA) 63 M. binotella (Duponchel, 1843) – male, Austria (RCEA) 64 M. binotella (Duponchel, 1843) – female, Italy (
Megacraspedus adults in dorsal view. 67 M. barcodiellus sp. n. – Paratype, male, Macedonia (
Megacraspedus adults in dorsal view. 73 M. gallicus sp. n. – Holotype, male, Spain (
Megacraspedus adults in dorsal view. 79 M. numidellus (Chrétien, 1915) – male, Morocco (
Megacraspedus adults in dorsal view. 85 M. niphorrhoa (Meyrick, 1926) – male, Kazakhstan (RCKN) 86 M. niphorrhoa (Meyrick, 1926) – female, Russia (
Megacraspedus adults in dorsal view. 91 M. balneariellus (Chrétien, 1907) – female, Italy (RCEA) 92 M. podolicus (Toll, 1942) – male, Hungary (RCZT) 93 M. kazakhstanicus sp. n. – Holotype, male, Kazakhstan (RCKN) 94 M. knudlarseni sp. n. – Holotype, male, Spain (
Megacraspedus adults in dorsal view. 97 M. tenuignathos sp. n. – Holotype, male, Morocco (RCAW) 98 M. glaberipalpus sp. n. – Paratype, male, Morocco (
Megacraspedus adults in dorsal view. 103 M. multispinella Junnilainen & Nupponen, 2010 – male, Russia (
Megacraspedus adults in dorsal view. 109 M. consortiella Caradja, 1920 – male, Kyrgyzstan (RCKN) 110 M. pototskii sp. n. – Holotype, male, Kyrgyzstan (RCKN) 111 M. leuca (Filipjev, 1929) – male, Russia (
Megacraspedus adults in dorsal view. 115 M. lagopellus Herrich-Schäffer, 1860 – female, Hungary (
Megacraspedus adults in dorsal view. 121 M. argyroneurellus Staudinger, 1871 – male, Russia (
Megacraspedus adults in dorsal view. 127 M. violacellum (Chrétien, 1915) – Lectotype, female (halfella), Tunisia (
Megacraspedus adults in dorsal view. 133 M. multipunctellus sp. n. – Holotype, male, Turkey (ECKU) 134 M. teriolensis sp. n. – Paratype, male, Croatia (RCZT) 135 M. teriolensis sp. n. – Paratype, male, Slovenia (
Megacraspedus adults in dorsal view. 139 M. quadristictus Lhomme, 1946 – female, France (
Megacraspedus adults in dorsal view. 145 M. peyerimhoffi Le Cerf, 1925 – male, Spain (
Megacraspedus male genitalia. 228 M. imparellus (Fischer von Röslerstamm, 1843) – Austria, GEL 1195 P.H. (
Megacraspedus male genitalia. 231 M. multispinella Junnilainen & Nupponen, 2010 – Paratype, Russia, GU 16/1418 P.H. (
Megacraspedus male genitalia. 237 M. leuca (Filipjev, 1929) – Russia, GEL 1240 P.H. (
Megacraspedus female genitalia. 291 M. nupponeni sp. n. – Paratype, Russia, gen. slide 4/16.7.2017 K. Nupponen (RCKN) 292 M. leuca (Filipjev, 1929) – Paratype (kaszabianus), Mongolia, GU 16/1447 P.H. (
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 Lepidoptera, granted to Serge Peslier. Furthermore the authorities of the Parc National Mercantour, Parco Naturale Alpe Marittime, Parco Nazionale del Gran Sasso e Monti della Laga and Mavrovo National Park are gratefully acknowledged for support of field research.
Stefan Heim (
Distribution catalogue