Taxonomic and morphological survey of the Lygephila lusoria (Linnaeus, 1758) species-group with description of a new species (Lepidoptera, Erebidae, Toxocampinae)

Abstract The taxa of the Lygephila lusoria (Linnaeus, 1758) species-group are revised. The genital features of all known taxa are described and illustrated with special reference to the structure of vesica. The male genitalia of L. pallida pallida (Bang-Haas, 1907) are described and illustrated for the first time. L. pallida subpicata (Wiltshire, 1971) is treated here as a species, L. pallida subpicata (Wiltshire, 1971), stat. n., distinct from L. pallida. A new species, L. minima, sp. n.,from South Russia is described. Illustrations of the holotype and its genitalia are provided; a diagnostic comparison with L. pallida is given. L. alaica Remm, 1983 is included in the L. lusoria species-group for the first time.


Materials and methods
Male and female genitalia were dissected and mounted in euparal on glass sides. Photos of genitalia where made by Svitlana Pekarska using microscope Nikon SMZ745T and camera Moticam 2500. Photos of imago where taken by the author using camera Nikon D3000/Sigma 105, f/2.8.

Description of the Lygephila lusoria species-group
Head and body greyish brown with frons and collar chocolate brown. Forewing in general broad, elongated with pointed apex, greyish brown with indistinct transverse lines, orbicular stigma dot-like, reniform stigma more or less triangular, black, sometimes with sharp extension at inner corner and satellite streak-like spots on outer margin; hindwing with wide outer band and visible discal spot. The first characterisation of the genitalia structure of the L. lusoria species-group was given by Babics and Ronkay in 2009 and this characterisation is to be revised and amended. Some of the previously mentioned characters, such as the strong, sabre-shaped uncus being broadened in third quarter, and having an acute tip, the elongated valva with more or less parallel margins, the wide, funnel-shaped ostium bursae, and the membranous, elliptical corpus bursae are shared features with the Lygephila lubrica species-group. Therefore, these characters cannot be considered as distinctive features for the L. lusoria group. The autapomorphies of this group can be found in the shape of the ampulla, the aedeagus, the vesica structure and some of the specific parameters of the ostium bursae. The ampulla is tapered with a long, skewed base, which is comparable in length in practically all members of the group. The aedeagus is short and wide; the vesica has six or seven diverticula, the subbasal diverticulum is well developed with a ridge-like cornutus complex. The female genitalia are characterised by the markedly asymmetrical ostium bursae, in comparison with the other species groups of the genus.

Lygephila lusoria lusoria
Male genitalia (Figs 25,39,40). Uncus stem narrow and relatively long, dilated distally with fine tip, scaphium membranous with sclerotized plate on subscaphium; valva elongated, narrowed at base, apex rather acute; ampulla spine-like, almost straight, not reaching apex of valva, its base asymmetrical. Aedeagus straight, tubular, slightly dilated at carina with heavily sclerotized field on it. Vesica globular, everted forward and recurved laterally; medial part membranous; subbasal diverticulum oblate with heavily sclerotized crest contacting carina; 1 st medial diverticulum small; 2 nd and 3 rd  medial diverticula elongated, tube-like, rising from extension of main vesica chamber located opposite to each other; 4 th medial diverticulum on opposite side topped with large, rounded, plate-like cornutus with two teeth; 1 st terminal diverticulum tapered with large basal swelling; 2 nd terminal diverticulum tapered, bordering 2 nd medial diverticula, bearing three small pockets; terminal tube membranous with weak scobination at end near gonopore (starting point of ductus ejaculatorius), opening point of terminal tube located at base of medial part of vesica near carina. Female genitalia (Figs 76,77). Ovipositor relatively short, broad; papillae anales hairy with long setae on apical edges. Apophyses anteriores slender, apophyses posteriores thin with acute tips, longer than apophyses anteriores. Antrum tapering, ostium bursae broad with acute lateral edges, posterior margin incised producing shallow triangular cleft with almost straight margins; ductus bursae large, wide with coarse well-sclerotized wrinkles laterally. Appendix bursae small with ductus seminalis located near ductus bursae. Corpus bursae membranous, large, ellipsoidal.
Distribution. West Palearctic. In Europe it ranges from Spain to Bulgaria, from Ukraine to south Russia and western Kazakhstan (Uralsk). All earlier records for Asia Minor refer to L. amasina, whereas the records from north Caucasus and Transcaucasia belong to L. minima sp. n. ( Barbara, Granada, 1800m, 30.vi.19941 ♀, Spanien, Granada, Sierra Nevada, Pico Valeta, 2500m, 4.7.1987 Note. The name of this taxon is unavailable from Rambur, 1866, and there is some debate as to the correct authorship and date of this subspecies. This issue will be dealt with in a separate publication.

Lygephila lusoria glycyrrhizae
Diagnosis. This taxon was downgraded to a subspecies of L. lusoria by Bertaccini et al. (2008). It is interesting that, despite the remarkable external differences between L. l. lusoria and L. l. glycyrrhizae, no valuable differences can be recognised in the male and female genitalia of the two taxa. The most significant distinctive feature of L. l. glycyrrhizae is, in comparison with L. l. lusoria, the small size of the genitalia of both sexes. The genitalia of the Spanish moths are approximately 1.3 times smaller than those of L. l. lusoria from Central and Eastern Europe, Crimea and Urals. In addition, there are a few hardly recognisable differences in the shape of uncus, valva and aedeagus: L. l. glycyrrhizae has somewhat shorter uncus stem and valvae with costal dilatation medially and less curved aedeagus, whereas the plan of the female genitalia of the two taxa is practically the same. (Figs 26,41,42). Uncus stem narrow and relatively short, dilated distally with fine tip; scaphium membranous with sclerotized plate on subscaphium; valva elongated, narrowed at base, margins not parallel due to large costal dilatation medially, valval apex rather acute; ampulla almost straight, spine-like with symmetrical base. Aedeagus a straight tube with heavily sclerotized field on carina. Vesica globular, everted forward and recurved laterally; medial part membranous; subbasal diverticulum oblate with heavily sclerotized crest contacting carina; 1 st medial diverticulum small; 2 nd and 3 rd medial diverticula elongated, tube-like, rising from extension of main vesica chamber, located opposite to each other; 4 th medial diverticulum on opposite side topped with large, rounded, plate-like cornutus with two teeth; 1 st terminal diverticulum tapered, with large basal swelling; 2 nd terminal diverticulum bears three small pockets; terminal tube membranous with weak scobination at distal end near gonopore (starting point of ductus ejaculatorius); opening point of terminal tube located at base of medial part of vesica near to carina. Female genitalia . Ovipositor relatively short, broad, papillae anales hairy with long setae on apical edges. Apophyses anteriores slender, apophyses posteriores longer than apophyses anteriores, thin with acute tips. Antrum tapering, ostium bursae broad with acute lateral edges, posterior margin incised showing shallow triangular cleft with almost straight margins; ductus bursae large, wide with coarse well-sclerotized wrinkles laterally. Appendix bursae small with ductus seminalis located near ductus bursae. Corpus bursae membranous, large, elongated, ellipsoidal.

Male genitalia
Distribution. Spain. Diagnosis. L. amasina distinguishing from similar L. lusoria lusoria by more contrast wing pattern and somewhat longer, sometimes with acute apex of inner corner of the reniform stigmata. In genital structures it differs from L. lusoria by broader uncus, longer, thinner ampulla reaching apex of valva, not sharp lateral edges of antrum and ovoid corpus bursa.
Vesica small, globular, everted laterally; medial part membranous; heavily sclerotized crest with ridge-like cornutus complex based on elongated oblate diverticulum-like subbasal hump; 1 st medial diverticulum medium-sized, 2 nd medial diverticulum much larger, located on the opposite side of vesica with sclerotized area on the top; 1 st terminal diverticulum two-chambered, one of them elongated tapering, another globular; 2 nd terminal diverticulum tapering with acute top; 3 rd terminal diverticulum situated in the same line with 2 nd medial diverticulum; opening point of terminal tube located at base of medial part of vesica near to carina; terminal tube membranous with narrow sclerotized crest at base and weak scobination at distal end near gonopore (starting point of ductus ejaculatorius). Female genitalia (Fig. 81). Ovipositor short, papillae anales hairy with long setae on apical edges. Apophyses anteriores slender, apophyses posteriores thin, 1.6 times longer than apophyses anteriores. Antrum infundibuliform, asymmetrical, with heavily sclerotized elongated plate dorsally; ostium bursae broad, posterior margin gently concave; ductus bursae practically absent. Appendix bursae indistinct. Corpus bursae membranous, ovoid.
Distribution. Turkey, Lebanon and Israel. Diagnosis. L. colorata differ from somewhat externally similar L. amasina by more elongated forewing with pointed apex. In male genitalia it differ from congeners by very wide, massive valva, strong, thick ampulla. Female genitalia characterised by deeply concave posterior margin of antrum.
Distribution. North-western Pakistan. Diagnosis. Distinguishable from similar species only by genitalia characters. It differ from L. subpicata by shorter spine-like ampulla not reaching the valval edges and from L. minima sp. n. by narrower valva, longer ampulla and absents of sclerotization on top of the 2 nd medial diverticulum.
Male genitalia (Figs 31,(52)(53)(54)(55)(56)(57)(58)(59). Clasping apparatus somewhat asymmetrical (right valva narrower). Uncus stem narrow, short, dilated distally, with fine tip; scaphium membranous with weakly sclerotized plate on subscaphium; valva elongated, narrowed at base with rather acute apex; ampulla spine-like, slightly curved towards costa, finely pointed, does not reaching apex of valva. Aedeagus short, slightly curved medially, with heavily sclerotized convex field on carina and spinulose area on lamina. Vesica globular, everted forward and recurved laterally; medial part membranous; heavily  bered, one of them elongated tapering, another globular; 2 nd terminal diverticulum tapered; 3 rd terminal diverticulum elongated with very wide base and curved tapered part; opening point of terminal tube located at base of medial part of vesica near carina, terminal tube membranous with narrow sclerotized crest at base and weak scobination near gonopore (starting point of ductus ejaculatorius). Female genitalia (Fig. 83). There were no females with type labels or from the same collecting place as the cotype in the MNHU collection. Taking into consideration that the exemplar from Palandöken, Turkey is the most similar in male genitalia structure to the cotype specimen (the two slides are almost fully agree with each other) one can conclude that the female specimen from the same site would represent the female sex of L. pallida. Ovipositor short, papillae anales small, hairy with long setae on apical edges. Apophyses anteriores slender with fine tip, apophyses posteriores thin, somewhat longer than apophyses anteriores. Antrum U-shaped, asymmetrical, ostium bursae broad, posterior margin deeply concave with large prolongation of posterior end on one side; ductus bursae small, practically absent. Appendix bursae small. Corpus bursae membranous, ovoid.
Distribution. Central and eastern Turkey.

Lygephila fereidun Wiltshire, 1961
http://species-id.net/wiki/Lygephila_fereidun Fig. 18 Taxonomy. This taxon, described from the Elburz Mountains, Northern Iran, is known only from the holotype (coll. BMNH). In the original description the color was characterized as pale straw and the wing pattern close to the Spanish species glycyrrhizae. The diagnostic comparison was made with L. craccae ([Denis & Schiffermüller], 1775) and L. lusoria only, whereas a comparison with another similar species, L. pallida, was neglected. The original description contains the following text about the clasping apparatus structure (Fig. 28): "The harpe [ampulla], longer than that of craccae, is nevertheless shorter than that of lusoria." Comparative analysis of the ampullar length (shorter than that of lusoria) given by Wiltshire, makes it possible to conclude that the L. fereidun is different from the L. amasina and L. subpicata, because they have longer ampullae that reach the costal margin of the valva. So, by this feature L. fereidun could be compared only with L. pallida, the ampulla of which is rather shorter than that of L. lusoria and other members of its species group. Vesica structure in the original description is characterized as follows: "The vesica contains similar elements to those of lusoria but the proximal scobinated field is shorter and the five or six teeth on the distal chitinous lump are larger and more like cornuti than in lusoria." However, the only sclerotized cornutus formation illustrated in the original drawing looks similar to that of L. subpicata, but L. subpicata has two heavilysclerotized crown-like cornuti on the top of subbasal and 2 nd medial diverticula. The above-mentioned contradictions in the original description thereby make it impossible to clarify the taxonomical situation of this taxon without a study of the genitalia of the holotype, the preparation of which is opaque and requires specific recovery treatment. Based on the currently known characters L. fereidun is most likely an aberrant specimen of L. pallida.
Distribution. Northern Iran. Etymology. The name "minima" refers to the small size of the moth in contrast to the largest representative of the genus, Lygephila maxima (Bremer, 1861).
Diagnosis. The new species resembles L. pallida by its small size and pale brown ground color of the forewing. L. minima differs from the related species by its better developed noctuid pattern, more rounded wings and pale grey-brown ground color of the forewings. Apical dilatation of uncus wide, valva wide with rounded apex, ampullar tip not sharp, 1 st medial diverticulum reniform; 2 nd medial diverticulum hemispherical, membranous, without sclerotized areas, whereas L. pallida has narrower dilatation at the top of the uncus, longer, narrower valva with acute apex, fine tipped ampulla, 1 st medial diverticulum very wide at base, swelling-like; 2 nd medial diverticulum large, tubular, with sclerotized area on the top.
Description. Male (Figs 21-24). Wingspan 33 mm, length of forewing 17 mm. Head and collar coffee brown. Palpi short, relatively narrow, beige; antenna filiform. Thorax and abdomen beige. Forewing beige with silver shining, irrorated with a few blackish-brown scales; forewing short, wide; costa straight; outer margin rounded; wing pattern indistinct: basal, subbasal and antemedial lines hardly recognisable; medial line represented by large costal patch and some darker spots medially; postmedial line indistinct; subterminal line curved, composed by blackish-brown scales; terminal line marked by large triangular patches; cilia long, uniformly light brown; orbicular stigma dot-like, as coffee-brown colored as V-shaped reniform; claviform stigma indistinct. Hindwing beige brown, discal spot narrow. Female unknown.
Male genitalia (Figs 35-38, 60-67). Uncus stem short, broadly dilated distally with fine tip; valva short, wide, rounded apically with rather parallel margins in distal two-thirds, slightly narrower at base; ampulla spine-like with long base and pointed tip which does not reaching margin of valva. Aedeagus short, curved medially, with heavily sclerotized field on carina and spinulose area on lamina. Vesica globular, everted forward and recurved laterally; medial part membranous; basal cornutus ridge interrupted without sclerotized base, subbasal diverticulum medium sized; 1 st medial diverticulum large, reniform; 2 nd medial diverticulum hemispherical; 3 rd medial diverticulum tapered, 1 st distal diverticulum large, subconical, 2 nd distal diverticulum with wide base and crooked tip; opening point of terminal tube located at base of medial part of vesica, terminal tube membranous with sclerotized ribbon at base and weak scobination at end near gonopore.
Distribution. The species is known from south Russia, Stavropol region and Armenia.  Abadeh, nördl Didegan, 2000m, 9.6.19692 ♂♂ Iran, Prov. Fars, Zagros Mts, Ardakan, 2500-3000 m, 18.VI.2010 Benedek & T. Hácz, coll. P. Gyulai, slide Nos: OP2002m, OP2003m. Diagnosis. Lygephila subpicata differs from its sister species, L. pallida in the length and shape of the ampulla, and in vesica and aedeagus structure. L. subpicata has a much longer, curved ampulla, which reaches apex of valva and costal margin; subbasal diverticulum large with crown-like cornutus on top, tapering part of 1 st terminal diverticulum small, short and narrow, 2 nd medial diverticulum with crown-like cornutus on top, carinal extension practically absent. In comparison, L. pallida has shorter, less curved, finely pointed ampulla that does not reach apex of valva, a small, oblate subbasal diverticulum with a long, heavily-sclerotized, ridge-like cornutus complex that is a continuation of the carina.
Distribution. North and western Iran.
Diagnosis. Lygephila moellendorffi is known only from two males representing the type series. The photo of the paralectotype was illustrated in Kononenko et al. (1998); the genitalia of the paralectotype was first illustrated by Kononenko and Han (2007). Surprisingly, this species is confusingly similar to L. subpicata, displaying no differential features comparing the habitus and the genitalia structures of the two species. The simi-larly elongated forewings with pointed tips have the same pattern, especially the triangular reniform stigma with sharp extension on the inner corner and satellite streak-like spots are practically identical in the two taxa. The common features of the male genitalia are the similar shape of uncus and valvae with the similarly sized and shaped ampulla being also located subapically and reaching the apical valval margins. Both species have short and relatively wide aedeagus and vesica with characteristic subbasal and 2 nd medial diverticula topped by crown-like cornuti; the terminal diverticula are also similar. This striking resemblance suggests that they represent the same species, but the great distance between their ranges does not support this conclusion.
Distribution. North Korea. Diagnosis. Lygephila alaica should be attributed to the L. lusoria species-group on the basis of both the external and genital diagnostic characters. The elongated forewing with pointed apex is similar to those of all species of this species-group, particularly to eastern representatives, L. colorata and L. subpicata. The spine-like ampulla with long skewed base, the short and wide aedeagus, the characteristic vesica structure, especially the presence of the well-developed subbasal diverticulum with large cornutus in the male genitalia and the heavily sclerotized, funnel-shaped antrum with strongly asymmetrical ostium bursae in the female genitalia indicate the close relationship with the L. lusoria species-group.
Distribution. Central Asia -Tajikistan and Uzbekistan.