Extensive generic descriptions and diagnoses of Holarctic species of the Agrotis fatidica (Hübner, 1824) species-group have been published in several reviews (von Mentzer and Moberg 1987; Fibiger 1990, 1997; Nupponen et al. 2001; Fibiger et al. 2010), and are not repeated here.

Our study is based on about 1,500 specimens of the Agrotis fatidica species-group. Most of the material was traditionally set and dried, although a few specimens are pinned but remained unset. Genitalia preparations followed standard techniques for Noctuoidea, including everting the vesica.

Full-length lepidopteran DNA barcode sequences are a 658 base-pair (bp) long segment of the 5’ terminus of the mitochondrial COI gene (cytochrome c oxidase 1). DNA extracts were prepared from a dried leg of each of 22 specimens of the Agrotis fatidica species-group from Europe (12 A. fatidica, 2 A. proverai Zilli, Fibiger, Ronkay & Yela, 2010, 5 A. mayrorum sp. n., 3 A. mazeli sp. n., see Table 1.) and 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 deWaard et al. (2008). We also used 3 sequences of Agrotis luehri von Mentzer & Moberg, 1987 from BOLD [sample ID´s NHMO-DAR-9557, NHMO-DAR-9557, CNCNoctuoidea1317] for our analysis. All sequences are 658 bp long, except for 4 sequences of A. fatidica ranging from 600 bp to 657 bp and one sequence of A. mazeli with 581 bp. Sequences were submitted to GenBank; further details including complete voucher data and images can be accessed in the public dataset “Lepidoptera of the Alps - Agrotis fatidica species-group” dx.doi.org/10.5883/DS-AGROFATI in the Barcode of Life Data Systems (BOLD; Ratnasingham and Hebert 2007). Degrees of intra- and interspecific variation in the DNA barcode fragment were calculated under Kimura 2 parameter (K2P) model of nucleotide substitution using analytical tools in BOLD systems v3.0. (http://www.boldsystems.org). A neighbour-joining tree of DNA barcode data of selected taxa was constructed using Mega 6 (Tamura et al. 2013).

Representative samples of already described European taxa of the Agrotis fatidica group and their Asiatic relatives were studied and photographed. Material was examined from several European museums and private collections (see abbreviations below). All specimens dissected during the studies were photographed in colour, as were their genitalia slides. The habitus of the specimens was photographed with a Nikon D90 camera; the images of the genitalia slides were taken with a Nikon Eclipse 80i photomicroscope with a Nikon DS-Fi2 digital camera. All images are preserved in the photographic catalogue of the Heterocera Research Team, Budapest.

BMNH The Natural History Museum (formerly British Museum, Natural History), London, United Kingdom

CNC Canadian National Collection of Insects, Arachnids and Nematodes

HNHM Hungarian Natural History Museum, Budapest, Hungary

LMK Landesmuseum Kärten, Klagenfurt, Austria

MNHU Museum für Naturkunde, Humboldt-Universität, Berlin, Germany

NHMO Natural History Museum, University of Oslo, Norway

NHMW Naturhistorisches Museum Wien, Austria

RCBS Research Collection Bjarne Skule, Veksø, Denmark

RCTM Research Collection Toni Mayr, Feldkirch, Austria

RNS Royal Natural History Museum (Naturhistoriska Riksmuseet), Stockholm, Sweden

TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria

ZMHU Museum für Naturkunde – Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin, Germany

ZMUC Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark

ZSM Zoologische Staatssammlung, Munich, Germany

Agrotis fatidica fatidica (Hübner, 1824)

incurva Herrich-Schäffer, 1851

monedula Dannehl, 1925

Agrotis fatidica luehri von Mentzer & Moberg, 1987 stat. n.

Agrotis proverai Zilli, Fibiger, Ronkay & Yela, 2010

Agrotis mayrorum sp. n.

Agrotis mazeli sp. n.

Agrotis trifurcula (Staudinger, 1892)

Agrotis innocens (Boursin, 1967)

Agrotis characteristica Alphéraky, 1892

bombycia Eversmann, 1851

robusta Eversmann, 1856, nec Blanchard, 1852

sajana Corti, 1932 (= sajana Bang-Haas, manuscript name)

robustana Poole, 1989

Agrotis iremeli Nupponen, Ahola & Kullberg, 2001

Agrotis ruta (Eversmann, 1851)

xylographa Boursin, 1947

?Agrotis patula (Walker, 1857) (status unclear)

septentrionalis Möschler, 1862

kurodakeana (Matsumura, 1927)

subinformis (Bryk, 1941)

Figs 1–8; Gen. figs 1–2, 16

Noctua fatidica was described from an unspecified number of male specimens and figured in colour (Hübner 1824, pl. 152, figs. 704–705). The type-locality is vague ([Europe]) although the traditional concept of A. fatidica is based on specimens from the Alps, which is the major source of specimens in collections from the European Continent (see, for instance, Fibiger 1990, 1997; Fibiger et al. 2010).

The type material is considered lost (Horn and Kahle 1935–1937) and we here designate a neotype from the eastern Alps in order to maintain stability of the nomenclature and ground the traditional interpretation of “fatidica”. Such a designation follows the qualifying conditions of section 75.3 of the ICZN Code, 4^th edition. We select an Austrian specimen, which is barcoded and which has a sequence that fits well with the DNA barcodes of the other specimens we have investigated. The sequence is in fact identical to those of four specimens.

The neotype is a male and bears the following labels: “AUSTRIA occ. | Nordtirol | Rettenbachtal | 2700 m, 5.9.1991 | leg. Burmann et al.”; “BC TLMF Lep 00184” (green label). The neotype and its labels are illustrated in Figs 1–2.

Apart from the Alps, published records of A. fatidica-like taxa have come from the Pyrenees, the Balkan Peninsula (Albania, Macedonia and Bulgaria) and the Caucasus region. Furthermore, two old specimens from the Frivaldszky collection (HNHM), labelled as “Corsica”, are practically identical with those of the specimens from the French and Ligurian Alps. The report of “A. fatidica” from Corsica is probably based on these, almost certainly mislabelled, specimens, and the occurrence of a fatidica-group species in the island of Corsica has never been confirmed (cf. Brusseaux and Nel 2004).

Von Mentzer and Moberg (1987) excluded the presence of A. fatidica from the Ural region; however, the A. fatidica specimen illustrated in Noctuidae Europaeae 1: pl. 7, fig. 2, (Fibiger 1991) is from the Urals. Moreover, an allied species, A. iremeli, is present in that region.

Agrotis fatidica var. monedula is just an individual, dark form from the Ortler Region (South Tyrol) and has no taxonomic relevance. Contrary to the statement of Poole (1989), the type-locality is mentioned in the description.

Agrotis incurva, a junior synonym of A. fatidica, was described from three highly variably coloured and shaped females and an unstated number of males collected on the high peaks of the Tyrolean Alps by Stenz (Herrich-Schäffer 1852). In the original description, brachyptery in the female sex was considered as probably the result of malformation.

It remains unclear as to whether some or all the eastern European and Asiatic populations currently referred to A. fatidica, either named (e.g. bombycia Eversmann, 1851; possibly also sajana Corti, 1932) or unnamed (e.g. the Balkanic, Caucasian and Tibetan populations), are actually conspecific with the Alpine ones or not. Material from the Altai Mts which corresponds with A. fatidica in external morphology falls within the range of intraspecific DNA barcode variation and is therefore considered as conspecific. However, the barcodes of certain Agrotis specimens, identified by their external features as A. trifurcula (Staudinger, 1892) and A. ruta (Eversmann, 1851), show no divergence from the barcodes of the European populations of A. fatidica (see also under A. trifurcula and A. patula).

Clearly, subspecific division of A. fatidica requires further study, especially with respect to the south-eastern and eastern European populations which are poorly represented in collections.

Figs 9–16

Originally, A. luehri was considered to be the Arctic sister-species of A. fatidica in which the male has shorter and broader-based lamellae of the antenna, and the female is longer-winged and with thinner antennomeres. It was described from Norway and compared with a long series of “A. fatidica” specimens from both Norway and the Alps (von Mentzer and Moberg 1987). Subsequently, however, all Norwegian populations of “A. fatidica” were said to represent A. luehri (Fibiger 1997, in Noctuidae Europaeae 3: 81). This situation is questionable as it implies that the latter has been diagnosed also in comparison to itself. The distinction of A. luehri from the alpine A. fatidica was accepted by Fibiger et al. (2010) on the basis of the characters noted above, but the logical incongruence relating to the Norwegian populations can only be explained by admitting in A. luehri a broader range of variation in morphological characters than that fixed in the original description.

The DNA barcode of 3 specimens from Norway show, however, no difference compared with those of 12 specimens from the eastern Alps (see Table 1). As it seems unlikely that two species occur together in Norway, we consider all the DNA barcodes to be sequences of A. luehri. There is some variation in DNA barcodes of A. fatidica (within some 0.4%), but the three known DNA barcodes of A. luehri nest within this range, matching exactly those of certain specimens of A. fatidica from Austria and Switzerland.

Thus, irrespective of the disputed diagnostic characters in morphology, the COI sequences support the conspecificity of A. luehri with A. fatidica. The detailed investigation of the Scandinavian specimens show, however, differences in the colouration and the wing shape between the nordic and the alpine populations (the ssp. luehri is darker and more unicolorous than the ssp. fatidica, especially the hindwings are more dark grey suffused, the melanistic specimens appear more frequently; the forewings of the ssp. luehri are narrower and more elongated). These differences and the long and complete isolation of A. fatidica and A. luehri support their distinction at subspecies level, A. fatidica luehri stat. rev. (see also Aarvik et al. 2017).

Figs 17–20; Gen. fig. 3

The taxon is endemic to Central Italy (Abruzzo region; on the massif of Gran Sasso d’Italia), and is the only population of the A. fatidica species-group outside of the Alps ever found on the Italian Peninsula. Only males have been collected, and it seems likely that the females of this isolated species are brachypterous, as is the case in several other members of the species-group. The male genitalia and various other structural characters correspond with typical A. fatidica and its close relatives. Furthermore, DNA barcodes show a high degree of similarity to typical A. fatidica from the Eastern Alps, with only ca. 0.5% divergence (n=2), and they belong to the same BIN. The genetic distance from A. mayrorum and A. mazeli is much greater, minimally 1.55% and 1.71%, respectively.

The habitus of the moths from the Abruzzi region (especially the narrower forewings and the stronger and more prominent noctuid pattern), some allometric differences, and some incongruence in the variation of colours between areas of forewing compared with A. fatidica and A. mayrorum from the Alps, indicate that the Central Italian population has differentiated sufficiently to qualify as a distinct species.

Figs 21–32; Gen. figs 4–7, 17

Certain features of the SW Alpine (and the two “Corsican”) specimens, distinguish them from the typical eastern Alpine A. fatidica, as well as from the A. characteristica species-complex and A. trifurcula. The main diagnostic features are the proportionally longer and slenderer branches of the male antenna, the pale hindwing of the individuals, the enhanced pale lining of the cubital vein of forewing, the generally obsolete postmedial line, and the somewhat shiny flat, not fluffy scaling. These features are partly shared with examples from Bulgaria and specimens of A. mazeli from the NE Pyrenees, while the fatidica-like moths occurring in the Ural region have generally dark hindwings, and their forewing pattern also resemble more closely the typical eastern Alpine A. fatidica.

In the male genitalia, the valvae of A. mayrorum are comparatively the longest and narrowest within the European species of the fatidica complex, having a less dilated distal section, only slightly convex distal dorsal margin and reduced membranous ventral edge along the sacculus. The clasper (harpe) is short and broad, being distinctively shorter than in A. mazeli and on average shorter and apically more rounded than in A. fatidica; the juxta is also somewhat larger and broader, more pentagonal than in A. fatidica and A. proverai; and the dentated plate of the phallus is stronger and more densely spiny in A. mayrorum than in the related species.

The female genitalia of A. mayrorum has, in comparison with A. fatidica, proportionally a longer ovipositor, longer apophyses anteriores and a longer appendix bursae; while the ovipositor, the ductus bursae, and the appendix bursae are proportionally longer than in A. mazeli.

The species status is strongly supported by genetic distances from other taxa of the A. fatidica species-group (see below).

The isolated populations occurring in the high mountains of the Balkans in Bulgaria, Macedonia and Albania appear from their external morphological features to be more similar to the south-western Alpine A. mayrorum than to the geographically closer eastern Alpine A. fatidica.

The few specimens known from the Caucasus (Daghestan) show transitional features, with rather long branches of the male antennae and greyish suffused but still paler hindwing than in the typical A. fatidica populations.

The detailed taxonomic analysis of these populations and their biogeographical constraints requires further research and we cannot identify them with confidence as A. mayrorum.

Figs 33–40; Gen. figs 8–9, 18

The Pyrenean populations represent a common lineage with the SW Alpine A. mayrorum, sharing the features of the male antennae and the paler (generally white or whitish) hindwing which distinguish the two new species from A. fatidica. The specific differences between A. mayrorum and A. mazeli can be found in the wing shape, the different form of the postmedial line, the shape and size of the dark dots following the terminal line and the colouration of the forewing underside. These are the same characters in which A. mayrorum and A. fatidica differ (besides the group features mentioned above).

The male genitalia of A. mazeli differ from those of A. fatidica and A. mayrorum by the short and basally broad valvae with the very typical, wide membranous ventral marginal section bordering the saccular part. The cucullus is broad with a finely pointed apex, the clasper (harpe) is rather long and slender, and the dentate subbasal plate of the vesica is weaker than in the closely related European taxa.

The female genitalia of A. mazeli have longer apophyses anteriores and a somewhat shorter appendix bursae than those of A. fatidica; the ovipositor and the ductus bursae are proportionally shorter, and the appendix bursae is shorter in A. mazeli than in A. mayrorum.

The species status is strongly supported by genetic distances from other taxa of the A. fatidica species-group (see below).

Figs 41–46; Gen. figs 10–11

The species was described from Mongolia; it is widespread and locally common in Central Asia. Certain authors synonymised it with A. fatidica (see e.g. Poole 1989), despite the clearly recognisable differences in the external and genital morphology (see the Figs 1–8, 41–46; Gen. figs 1–2, 10–11). The species has not been treated in detail in the European literature. Some of the last catalogues (Fibiger 1990, 1997; Fibiger et al. 2010; Witt and Ronkay 2012) consider it as a distinct species while Kononenko (2005, Noctuidae Sibiricae Vol. 1) and a recent monograph about the Noctuidae fauna of the western Altai Mts (Volynkin 2012) mention it as A. fatidica.

Our investigations on the DNA barcodes of certain Agrotis species occurring in the Altai Mts showed, surprisingly, no genetic distance between A. trifurcula and A. fatidica, indicating possible introgression (Huemer et al. 2017).

Figs 47–50; Gen. fig. 12

This taxon was described by Boursin from three male specimens collected in the central region of Mongolia by Dr. Z. Kaszab. The specimens of this short series differ conspicuously from the sympatrically occurring and widespread A. trifurcula by their pinkish-shaded pale ochreous-grey ground colour of both wings and the reduced pattern.

The genitalia differences between the holotype of A. innocens (see Boursin 1967) and the studied specimens of A. trifurcula were subtle in the original study since the everted vesica of the two taxa were not examined. The lack of the female and the absence of A. innocens in the large samples of A. trifurcula from various places of Mongolia and the surrounding areas implied the possible conspecificity of the two taxa, especially as there are no remarkable differences in the male antennal characters. The study of the vesica of the two species revealed notable differences in the configuration of the vesica. The vesica of A. innocens is clearly longer than that of A. trifurcula, medially slightly dilated and distally evenly tapering, and is without a thickening at the nearby ductus ejaculatorius, which is typical of A. trifurcula (see the Gen. figs 10–12). Thus, A. innocens is considered here as a species distinct from A. trifurcula.

Gen. fig. 15

After over a century of confusion regarding the identity of A. characteristica and A. robusta, and of these with other taxa of the A. fatidica species-group (e.g. trifurcula Staudinger, 1892, sajana Corti, 1932) (Corti 1932 in Corti and Draudt 1931–1933; Kozhanchikov 1937; von Mentzer and Moberg 1987), these two names were recently synonymised, thus making robustana Poole an unnecessary replacement name for robusta Eversmann, preoccupied by Blanchard (see Kononenko et al. 1998; Kononenko 2005). Nevertheless, there is evidence that two concepts are still mixed under A. characteristica: that adopted by Fibiger (1990: pl. 1) in Noctuidae Europaeae Vol. 1., referring to a form with comparatively short lamellae in the male antenna and non-brachypterous females which occurs in the Urals, and an Asiatic insect, referred to by Kononenko et al. (1998), which has longer lamellae, and fully-winged females. In passing, it may be noted that Corti (1931–1933: pl. 5, row l) illustrated a brachypterous female of “A. robusta” while stating in the text that the female is fully developed and larger than the male. Unfortunately, the type specimen of characteristica is a female, which prevents an undisputed morphological assessment of its relationship to robusta. On the other hand, it seems likely that these two names may actually refer to the same Asiatic species, as their type localities suggest ([Inner Mongolia]: Ordos and Southern Altai, respectively). If so, the populations of the “Ural form” still remained unnamed.

As a close relative of Agrotis ruta (Eversmann, 1851), this species is also a member of the A. fatidica species-group which, on grounds of larval morphology, reveals a relationship with the A. segetum ([Denis & Schiffermüller], 1775) species-group, notably A. clavis (Hufnagel, 1766) (Nupponen et al. 2001). The large size and the dark reddish-brown ground colour of A. iremeli make it unlikely to be confused with its congeners. It is most like A. ruta, the ground colour of which is however always grey and less monotonous, more mottled with darker and paler areas, and has darker, more sharply defined stigmata, and a more wedged subterminal line. Some forms of A. cinerea ([Denis & Schiffermüller], 1775) might be mistaken for A. iremeli, particularly some females, but as a rule they are smaller, with more weakly crenulated antemedial line of forewing, absolutely tiny orbicular stigma, if any, smaller reniform and no claviform.

A. iremeli has been collected solely in restricted biotopes with xeric, rocky montane tundra, either in clearings within taiga forest or on the peaks of the mountains of the South Urals, at an altitude of 1300–1400 m. It appears to be endemic to the Southern Urals, in essentially similar habitats of the Northern Urals where only A. ruta was found (Fibiger et al 2010, citing K. Nupponen).

Figs 51–56; Gen. figs 13–14

The only Holarctic member of the species-group. Its Nearctic populations have long been distinguished under the name A. patula. The type-locality of the species is poorly defined (“eastern Siberia”). The species appears to be distributed from the Ural Mountains through Central and eastern Asia to the Pacific area and in vast parts of North America. It is highly variable in external features, with several, often remarkably dissimilar, forms and local populations. Recent molecular studies (Zahiri et al. 2014) show, however, a high consistency within the Canadian populations whereas material from the Altai Mts is distant and clusters nearest to A. proverai. Further in depth studies are necessary for the Palaearctic populations.

The species was described from the Rocky Mountains (USA), but several large European collections house specimens under this name from Labrador and from the eastern side of the mountains of North America (named by Möschler as A. septentrionalis in 1862) and from the northern part of Pacific Asia (Japan: “Euxoa” kurodakeana Matsumura, 1927; Russia, Kurili Islands: Feltia subinformis Bryk, 1941). All these taxa have been recently considered to represent a single species, A. ruta (Kononenko 2005; Lafontaine and Schmidt 2010), mainly due to their very similar DNA barcodes. The fact that the DNA barcodes of certain closely related taxa of the A. fatidica group (e.g. A. trifurcula and A. ruta) are identical or very similar suggests that the ruta–patula complex needs to be reviewed.

The species was described from a single male specimen from Transbaikalia when our knowledge of the fatidica-group species was limited. At that time, separation of A. xylographa from A. ruta seemed reasonable. Subsequent study based on much more material of ruta shows the great external variation of the Asiatic populations of A. ruta (s. str.), as well as its wide range from the Ural region throughout Siberia and Mongolia to the Pacific area. The holotype of A. xylographa falls within the morphological and geographical range of A. ruta. Accepting the statement by Kononenko (1990; 2005), we treat A. xylographa as a junior synonym of A. ruta. Agrotis xylographa does, however, differ from the Asian Pacific specimens of A.“patula”.

25 specimens belonging to 4 species were successfully sequenced (Table 1) and group in 4 different clades (Fig. 61). Maximum intraspecific distances are ranging between 0% and 0.51% whereas minimum distances to the nearest neighbour are between 0.46% and 1.12% (Table 2). Agrotis mazeli sp. n. and A. mayrorum sp. n. have unique BINs (Barcode Index Number) in BOLD (Ratnasingham and Hebert 2013), whereas A. fatidica and A. proverai share their BIN.

The vast majority of European Noctuinae species are well-defined by morphological characters such as phenotypical appearance and genitalia structures. However, remarkable exceptions are found in some genera, e.g. Diarsia Hübner, 1821, Xestia Hübner, 1818, Euxoa Hübner, 1821, and Agrotis Ochsenheimer, 1816, all of which include cryptic and/or disputed taxa. The Agrotis fatidica-group belongs to such a disputed species-complex reflected by unstable taxonomic treatment in various monographic papers (e.g. Fibiger 1997; Fibiger et al. 2010; Gyulai et al. 2017; Hreblay et al. 1998; Hreblay and Ronkay 1998; Kononenko 2005; Nupponen et al. 2001; Volynkin 2012), and through the description of two new species from Europe during the last few decades, namely A. proverai from Italy (Fibiger et al. 2010) and A. luehri from Norway (von Mentzer and Moberg 1987). In this paper, we have examined the congruence of these classical and morphologically-driven species concepts using molecular data, which has resulted in a reformulated picture of the taxonomy.

We treat A. luehri as a Scandinavian subspecies of A. fatidica, a decision that means the species fatidica is now perceived to have an arctic-alpine distribution. By contrast, the southwestern alpine and Pyrenean populations previously treated as A. fatidica are separated from our revised concept of the species both by morphology and by molecular data of the DNA barcode region. We have described these residual alpine populations of ‘A. fatidica’ into two new species — A. mayrorum and A. mazeli. DNA barcoding resulted in excellent support for delimitation of the two alpine species with a distinct barcode gap of about a minimum of 1.7% and a very low intraspecific divergence. A. proverai from the Central Italian Apennines, however, which can be separated from its alpine relatives by morphology, is only weakly separated from typical A. fatidica by a barcode gap of about 0.5% and both species share their BIN. We consider even this low but constant distance as interspecific, a hypothesis which is supported by the generally low divergence in the entire species-group. Examples are the morphologically readily separated A. ruta and A. robusta from North America, with only small distances of roughly 1% to taxa of the A. fatidica species-group from Europe.

Of relevance here is that the A. fatidica species-group is considered by authors to be most closely related to the A. segetum species-group, notably to A. clavis, according to their larval morphological features (Nupponen et al. 2001). However, our molecular data (DNA barcodes) do not confirm this statement, and place the A. fatidica-group in the Palaearctic fauna nearest to the A. vestigialis- and the A. cinerea-groups.

All western Palaearctic species treated here show an allopatric pattern of distribution. Agrotis mayrorum (SW Alps), A. mazeli (Pyrenees) and A. proverai (Apennines) occur in well separated mountain systems, while A. fatidica in Europe occurs in the Alps and Scandinavia, with closely related or conspecific populations from the Balkans and from Ural to Central Asia which need further investigation. It seems likely that the species was widely distributed in the tundra belt during glacial periods and was able to spread quickly from hypothetical refugia perhaps north of the Alps or from Eastern Europe and Asia to formerly glaciated regions in the North and South after the last cold period. Similar movements are known in other species of Lepidoptera such as Hepialus humuli (Linnaeus, 1758) (Hepialidae) (Simonsen and Huemer 2014). The present disjunct distribution pattern is interpreted as a consequence of warmer postglacial periods which induced an increasing isolation of populations.

The distribution of A. fatidica in the Alps is discontinuous and patchy. It is widely distributed in the central eastern Alps from western Styria (south-eastern Austria) to Graubünden (eastern Switzerland), but almost completely absent from the northern and southern mountain ridges with e.g. only a single population known from the Italian Dolomites. In the western Alps just a few Swiss (Wallis) localities are reported (Wyman et al. 2015) and none southwards (Parenzan and Porcelli 2005–2006), indicating a considerable distribution gap from A. mayrorum (see Fig. 60). It seems likely that separation of these two species from a common ancestor may date back several glacial periods (see also Huemer and Hausmann 2009).

Surprisingly the nearest species, in terms of the DNA barcode, to typical A. fatidica is not A. mayrorum from the SW Alps but A. proverai from the Apennines. One might have expected that gene flow between populations from the SW Alps and the Apennines would be much more likely than from the eastern Alps to central Italy due to almost continuous mountain systems connecting the Apennines with the SW Alps. However, the barcode distance of A. proverai and A. mayrorum with 1.55% is considerably greater than that between A. fatidica and A. proverai (ca. 0.5%), thus indicating a much longer isolation of these taxa. A similar pattern of distribution is documented for only few alpine species, e.g. Aspilapteryx spectabilis Huemer, 1994 (Gracillariidae), which is only known from the eastern Alps and the Central Italian Apennines but absent from the SW Alps (Huemer 2011). However, in the absence of molecular evidence these widely separated populations of this latter species are considered conspecific.