Remarkable confusion in some Western Palearctic Clepsis leads to a revised taxonomic concept (Lepidoptera, Tortricidae)

Abstract The taxonomy of some Palearctic species of the genus Clepsis Guenée, 1845 (Lepidoptera, Tortricidae), in particular C. neglectana sensu auctt. and C. consimilana sensu auctt., is revised based on combined characters of external and internal adult morphology, including everted vesicae in male genitalia, and DNA barcodes. Clepsis striolana (Ragonot, 1879), stat. rev., C. acclivana (Zerny, 1933), C. trivia (Meyrick, 1913), stat rev., and C. xylotoma (Meyrick, 1891), stat. rev. are resurrected from synonymy with C. neglectana (Herrich-Schäffer, 1851), C. semiana (Chrétien, 1915), stat. nov. is considered as a valid species; C. eatoniana (Ragonot, 1881), stat. rev., is resurrected from synonymy with C. consimilana (Hübner, 1817), and C. razowskii Gastón, Vives & Revilla, 2017, syn. nov. is synonymised with C. eatoniana.


Introduction
Clepsis Guenée, 1845 is a large genus in the tribe Archipini with 163 valid species described worldwide (Gilligan et al. 2014) and 32 species present in Europe (Aarvik 2013). It is likely that these numbers will change in the future because a recent phylogenetic study revealed that the genus is paraphyletic and its position within the tribe is not completely resolved (Dombroskie and Sperling 2013). As such, the genus is in need of revision and the status of some taxa assigned to the genus should be evaluated. As with many groups of Lepidoptera, characters of the male genitalia have been used almost exclusively to define the present-day species boundaries of the taxa comprising Clepsis. Many taxa were synonymised in the past decades because of their similarity in male genitalia, resulting in long synonymic lists for some species. Two of the most problematic species are C. consimilana (Hübner, 1817) and C. neglectana (Herrich-Schäffer, 1851), each with many synonymic taxa: 13 for C. consimilana and eight for C. neglectana (Gilligan et al. 2014). These species are widely distributed in the Western Palaearctic and parts of Asia (Razowski 2002), and appeared to be variable in wing pattern, size, and even genital morphology. DNA barcode sequences of these species from different parts of Europe revealed that there is a considerable genetic variability within each of the species, and the DNA sequences cluster into compact groups. These groups appear to correspond well with certain wing patterns, secondary sex characters (costal fold in males), and genital morphology. It appears that some previously described taxa in this group were erroneously synonymised because of an incorrect assessment of variability in the genitalia, and that these names should be reinstated. A few taxa (also previously synonymised) were described from a single male specimen and no further material has been collected. Their present status is unknown and these taxa are in need of thorough revision, which is currently not feasible because of difficulties in obtaining fresh material from their type localities. The purpose of the present paper is to revise some of the taxa previously in synonymy with C. neglectana and C. consimilana, and to discuss some questionable taxa.

Materials and methods
We examined ca. 150 specimens from various European collections. Data for the specimens and their repository are given in the results for each taxon treated. The genitalia were dissected and mounted following the methods of Robinson (1976) and Zlatkov (2011). The everted vesicae were drawn before mounting on a slide as described by Zlatkov and Huemer (2017). The female genitalia were submerged in euparal essence without deformation and drawn in a manner similar to the male genitalia. A three-dimensional perception of the studied structures was achieved by modifying a compound microscope Amplival (Carl Zeiss Jena) after Hammond (1996). Certain structures whose shape and size are not affected by preparation were measured. Angles were measured using a photograph of a protractor aligned digitally to photos, and the linear measurements were performed using ocular scales. Forewing length was preferred rather than wingspan, as the latter depends too much on the condition of the specimen. The terminology of the wing pattern and genitalia follows Kuznetzov (1978) and Razowski (2002). The classification of cornuti and microstructures of the signa are after Anzaldo et al. (2014) and Lincango et al. (2013), respectively.

Morphology of genitalia
The male genitalia of the studied species, especially the valva, are relatively complex. The female genitalia are simple, but there is some discrepancy in the terms used in literature. For the purpose of unification, we provide general schemes of male and female genitalia of the studied taxa. Male (Fig. 1). The uncus is a large wide plate, convex dorsally, densely setose on both dorsal and ventral surfaces. In some species its shape can vary depending on the amount of pressure applied during preparation of the slide, and thus should be interpreted with caution. The gnathos is large, with a relatively large medial part, plough-shaped in all taxa. Because it projects posteriorly, it is often bent to the left or right in microscope slides. The socius is small, membranous, setose, and teardrop-shaped. The valva is complex, with largely sclerotised basal half and membranous distal half. The costa is enlarged, swollen, with a large more or less z-shaped, tubular costal sclerite. The sclerite protrudes medially into a subspherical or elongated spinulate structure called labis (plural labides) (Razowski 1979) or processus basalis (Horak 1984). The labides are probably derivatives of the transtilla but since they are not connected with a median sclerite, the term transtilla is less appropriate in this case. They are covered with large sclerotised acanthae. The middle part of the costal sclerite medially transforms into a setose membrane. The ventral part of the costal sclerite ends into a hemispherical transparent protrusion on the medial surface of the valva surrounded by a wrinkled furrow. The sacculus is large, medially concave, sclerotised, with a large triangular basal half, an elongated apical half and a ventrally pointed process located at the transition between the two halves. The distal part of the valva is membranous and terminates into a rounded, more or less extended plate referred to as a brachiola. The medial surface of the membranous part of the valva is densely covered by setae and scales forming a tuft on a small protuberance in some species. Most of the scales can be easily removed during cleaning of the macerated genitalia, but they should not be confused with a tuft of firmly attached scales and setae in some species which is well preserved after such manipulations. There can be a row of several very large modified setae on the median surface of valvae. The setae are elliptical in cross-section, falcate, with curved tips and can be broken during preparation of genitalia, and perhaps during copulation (males with setae broken off were found), but their sockets still remain observable (Fig. 7F). Apart from the medial scales and setae, there is a large, membranous pad on the lateral ("external") surface of the valva bearing a tuft of long deciduous scales (possibly androconia), which are usually lost during dissection. The juxta is a wide nearly semi-circular plate with a median incision for articulation with the caulis. The phallus is slightly sinuate, pointed, predominantly membranous dorsally, with a long stout lateral process on the left side. The anterior portion of the phallus is bent at a large ventral angle. The vesica is comparatively simple, tubular, with a small basal widening and a thumb-like diverticulum apically. A few long aciculate deciduous cornuti are attached near the base of the diverticulum, adjacent to the gonopore.
Female (Fig. 8). The papillae anales and the apophyses are not specialised and they are similar in all the taxa studied. The sterigma is more-or-less broadly funnel-shaped, dorso-ventrally flattened, sclerotised, with a large excavation on the dorsal wall. It ends into a narrow, funnel-shaped colliculum comprising areas of sclerotised tanned plicate and thickened transparent cuticle. This part is referred to as antrum by some authors (Kuznetzov 1978), but the latter term is used mainly for the sclerotised ring or collar in Cochylini (Razowski 1970). Both structures are localised in the same region and are probably homologous (Horak 1984). The ductus seminalis is inserted at the transition between the colliculum and the ductus bursae. The ductus bursae is relatively long, with a smooth elongate sclerite referred to as cestum extending on most of its length and sometimes expanding onto the corpus bursae as well. The corpus bursae is ovoid with two types of signa: one plate-like consisting of sclerotised papillae, and a hookshaped one that is more typical of most Archipini with longitudinal rows of teeth and a large capitulum.

Taxonomy
The

Clepsis neglectana species group
The species in this group externally are similar to taxa in the C. consimilana group but are distinguished by the absence of large modified setae on the median surface of the valvae (Fig. 1).

Clepsis neglectana (Herrich-Schäffer, 1851)
neglectana Diagnosis. Externally, C. neglectana is similar to the other species in the C. neglectana species group (apart from C. striolana) and C. consimilana, but the markings are darker and the costal fold is rudimentary. The male genitalia are very close to C. striolana, with the most obvious difference in the shape of labis; additionally, the setal tuft of the valva is less compact and smaller, the sacculus is straighter, and the vesica most often bears a single cornutus. Clepsis neglectana differs from C. acclivana and C. trivia by the shape of the uncus and numerous characters on the valvae and phallus. Apart from the forewing pattern, the female differs from C. striolana by the presence of a transparent protrusion of the colliculum on the right side, and from C. trivia by the length and shape of the colliculum. In contrast to C. trivia, both C. neglectana and C. striolana lack a plate-like signum.
Description. Adult ( Fig. 2A-C). Sexual dimorphism not detected. Head. Vertex, frons, palps and antennae monochrome, covered with ochreous scales. Sensilla trichodea (often referred to as "cilia") on antennae denser and longer in males. Thorax dorsally, legs and tegula ochreous, thorax ventrally creamy. Forewing length in males 6.3-7.4 mm (mean 6.9, N = 10), in females 6.5-8.2 mm (mean 7.3, N = 4). Forewing elongated, with costa convex basally and slightly concave apically, costal fold rudimentary (Fig. 3A). Upperside background ochreous to ferruginous with darker transverse or reticulate pattern. Markings brown to grey brown: basal blotch usually ill-defined, expressed mainly at costa and dorsum; median fascia widened at the middle. Subapical blotch triangular, ill-defined, sometimes connected with the median fascia. Cilia concolourous or paler than background. Underside pale grey-brown, sometimes with ill-defined reticulate pattern and creamy longitudinal blotch in the distal half of costa. Hindwing grey on both sides with underside paler, cilia whitish with grey line. Abdomen grey. Male genitalia (Fig. 4A, B). Uncus ovoid, widening apically, rounded, gnathos relatively large, socius membranous. Valvae pointed dorsolaterally when mounted on slide. Costal sclerite of valva relatively narrow, with short elliptic labis covered with small acanthae and extended into triangular pointed medial process (Fig. 5A, B). Apical part of sacculus ca. 1.4× longer than basal part, both forming angle of 145-155°, saccular process pointed. Membranous part of valva with protuberance bearing tuft of firmly attached, relatively sparse scales and setae; its terminal part with concave dorsal and convex ventral margin, brachiola ill-defined, pointed dorsolaterally. Posterior part of phallus slightly bent dorsally, with lateral process as long as 0.23× distance between anterior opening and tip of phallus, straight, in single specimen apically bent dorsally. Anterior and posterior part of phallus form angle of 130-140°. Caulis large, L-shaped, parallel to coecum. Vesica bent at ca. 110° dorsally, with small basal widening and terminal diverticulum mediodorsally, slightly pointed to right (Fig. 6A). One long, slightly curved deciduous cornutus attached ventroterminally adjacent to gonopore (Fig. 7A); single specimen with two cornuti. Female genitalia ( Fig. 8A) with papillae anales not modified. Apophyses anteriores 1.3× longer than apophyses posteriores. Sterigma widened caudad, with shallow lateral sclerotised pockets cephalad and large excavation on the dorsal wall. Colliculum short, with length 0.14× length of ductus bursae, straight, funnel-shaped, with plicate longitudinal sclerotisation and lateral protrusion at right cranial end consisting of colourless thick cuticle. Ductus bursae long and narrow, emerging at left to cuticular protrusion, with cestum extending along cranial 0.9× of its length and expanding for short distance on corpus bursae. Ductus seminalis inserted dorsally at caudal end of ductus bursae. Corpus bursae ovoid, with large falcate signum with capitulum ( Fig. 9A).
Ecology. Moths were collected in July and the beginning of August. The larval host plant is stated as Fragaria (Razowski 2002) but due to repeated misidentifications this record as well as published habitats need verification.
Remarks. This is the oldest described taxon from the group. Many other taxa were subsequently synonymised, but at least three of them are species bona and three others are incertae sedis (see below).   Diagnosis. This is the only species of the C. neglectana group with a uniform wing pattern in both sexes. Unlike the males of C. neglectana, the costal fold is developed. The male genitalia are very similar to C. neglectana but the setal tuft of the valva is better developed, the uncus is slightly longer, the sacculus is more angled, the vesica bears two cornuti instead of one, and the shape of the labis is different. The female genitalia are also similar to C. neglectana, although some differences in the colliculum (the transparent cuticular protrusions) are present. The colliculum is much shorter and straighter in C. striolana than in C. trivia.
Preimaginal stages are unknown. Molecular data (Fig. 16). BIN: BOLD:AAM0282. The intraspecific average and maximum distances of the barcode region is unknown (N = 1). The minimum distance to the nearest BIN-sharing neighbour, C. neglectana, is 1.53%.
Distribution (Fig. 17). This species seems to be limited to the Alps: Switzerland, Austria and Italy.
Ecology. Moths were collected from late June to August on sparse scree along alpine rivers. The larval host plant is unknown. Remarks. The genitalia of this species resemble those of C. neglectana, only a few details in certain structures are different. The genetic distance (though small) and wing pattern both support the existence of two separate taxa.
Tortrix trivia Meyrick, 1913: 297 (Tunisia) Material examined. Holotype ♂, pinned, with 6 labels: "Tunis / 27.5 / Coll. O. Leonhard" "T / 29" "Tortrix / trivia Meyr. / type" "Holotypus" [ Diagnosis. The species is most similar to C. acclivana. Externally the males differ in their wing markings, which are better defined in C. acclivana. The wings in C. trivia are also more elongated. The gnathos of C. trivia has angled arms, the sacculus is straighter, the brachiola is displaced dorsally, the phallic process has a different orientation but the vesica is very similar to C. acclivana. The wing pattern of C. trivia females is similar to those of both the C. consimilana and C. neglectana groups, but is C. trivia is paler. Clepsis trivia has the longest colliculum among the known females of the C. neglectana group.
Description. Adult. Sexual dimorphism prominent. Male ( Fig. 2H-J). Head. Vertex pale fulvous, frons, palps and antennae with ochreous scales. Antennae with numerous sensilla trichodea as long as width of flagellomeres. Thorax dorsally fulvous, ventrally creamy, legs pale brown, tegula creamy with fulvous anterior part. Forewing relatively elongate, with length 6.1-7.9 mm (mean 7.3, N = 7). Costa convex basally and straight apically, with small costal fold extending from base to 0.4-0.5× length of costal margin (Fig. 3C). Background pale yellowish with ill-defined reticulate pattern. Basal blotch ill-defined, consisting of small transverse fulvous markings. Median fascia brown or fulvous, widened at middle. Subapical blotch triangular, ill-defined, connected with median fascia. Underside pale grey-brown with creamy area in distal half of costa. Cilia concolourous with background. Hindwing upperside pale grey, underside whitish, cilia white. Abdomen pale grey. Male genitalia (Fig. 4E, F). Uncus wide, more or less round apically, with parallel lateral margins, gnathos with relatively large median part and angled arms, socius membranous. Valvae pointed laterally or slightly dorsolaterally when mounted on slide. Costal sclerite wide, with elongated labis cov-ered with large acanthae (Fig. 5D). Apical part of sacculus 1.6× longer than basal part, both forming an angle of 150-160°, saccular process pointed, relatively small. Membranous part of valva with protuberance lacking tuft of setae but has deciduous scales, terminal part with small dorsal and large ventral curvature, brachiola prominent, pointed dorsally. Posterior part of phallus slightly sinuate, with lateral process 0.24× distance between anterior opening and tip of phallus, weakly curved dorsad or rarely parallel to tip. Anterior and posterior part of phallus form angle of 130°. Caulis small, diverging from coecum. Vesica bent at 110-130° dorsad, with basal widening and terminal diverticulum dorsally, slightly pointed to right (Fig. 6C). Three ventroapically located deciduous cornuti adjacent to gonopore (Fig. 7C). Female more unicolourous than male (Fig. 2K). Head as in male but sensilla trichodea less numerous and shorter. Thorax as in male but tegula fulvous. Forewing length 6.6-7.9 mm (mean 7.3, N = 5). Costa convex basally and slightly concave apically. Upperside background pale yellowish with pale fulvous reticulate pattern, without markings, underside pale grey-brown with creamy area in distal half of costa, cilia concolourous with upperside. Hindwings upperside pale grey, underside and cilia whitish. Abdomen grey. Female genitalia (Fig. 8C). Papillae anales not modified. Apophyses anteriores slightly longer (1.1×) than apophyses posteriores. Sterigma widened caudad, with small shallow lateral pockets cephalad and large excavation on dorsal wall, lamella antevaginalis narrow. Colliculum asymmetrical, with length 0.2× length of ductus bursae, funnel-shaped, bent to left, with plicate longitudinal sclerotisation, large elongated protrusion at right and a small one at left both consisting of colourless thick cuticle. Ductus bursae long and narrow, emerging dorsally of cuticular protrusions. Cestum expanding for short distance on corpus bursae and extending along cranial part of ductus bursae for 0.9× of its length. Ductus seminalis inserted dorsolaterally at caudal end of ductus bursae. Corpus bursae ovoid, with large falcate signum with capitulum and flat signum located near end of cestum consisting of sclerotised papillae (Fig. 9C).
Distribution (Fig. 17). Known from Tunisia (type locality) and Crete. Ecology. The moths fly in the middle of April to late May and in early November, which may indicate two generations per year.
Remarks. Meyrick (1913) described C. trivia from a single male. The name remained valid until Karisch and Blackstein (2014) dissected the holotype (by monotypy) and synonymised it with C. neglectana. Despite of proposed synonymy, they explicitly stated that C. acclivana and C. trivia are very similar to each other but differ from C. neglectana. We find very little support for synonymy between C. neglectana, C. trivia, and C. acclivana. The holotype of C. trivia has very similar wing pattern to the males collected in Crete, and the male genitalia (apart from the unstudied vesica of the holotype) appear identical, therefore the Cretan population can be assigned to C. trivia despite lacking barcode data for the holotype. There is a considerable DNA barcode gap between C. trivia (from Crete) and C. neglectana (from Europe) which also supports existence of two taxa. (Zerny, 1933), stat. rev. Zerny, 1933:108, pl. 1, fig. 11 (Lebanon) Material examined. Lectotype ♂ by designation of Razowski (1979) Diagnosis. Clepsis acclivana is most similar to C. trivia but the forewings are paler and wider with more distinct markings, the uncus is narrower, the median part of the gnathos is smaller and its arms are not angled, the sacculus is more curved, the labis is more massive and with shorter acanthae, and the apex of the phallic process is curved ventrolaterally instead of dorsally.

Cacoecia acclivana
Description. Adult. Sexual dimorphism unknown. Male (Fig. 2L, M). Head. Vertex pale fulvous, frons, palps and antennae with ochreous scales. Antennae with numerous sensilla trichodea as long as width of flagellomeres. Thorax dorsally pale fulvous, ventrally creamy, legs pale brown, tegula pale fulvous. Forewing with length of 7.8 mm (in both specimens), costa basally convex, apically straight, with costal fold extending from base to 0.4× length of costal margin (Fig. 3D). Upperside background pale yellowish with fulvous reticulate pattern. Markings ill-defined, consisting of brown and ferruginous scales: basal blotch faint, with remnants only at costa as dark line; median fascia narrow, more prominent at dorsum; subapical blotch reduced, dash-like. Cilia concolourous with background. Underside pale grey-brown, costal and terminal areas creamy with some reticulate pattern. Hindwings upperside monochrome pale grey, cilia concolourous, underside whitish. Abdomen pale grey. Male genitalia (Fig. 10A, B). Uncus round apically, with parallel lateral margins, gnathos relatively small, socius membranous. Valvae pointed laterally or ventrolaterally when mounted on slide. Costal sclerite of valva very wide, with short wide labis covered with small acanthae (Fig. 5E). Basal and apical parts of sacculus with equal length forming angle of ca. 140°, saccular process almost right-angled, relatively large. Membranous part of valva with protuberance devoid of tuft of setae but has deciduous scales; its terminal part with nearly symmetrical dorsal and ventral curvature, brachiola prominent, pointed laterally. Posterior part of phallus slightly sinuate, with lateral process as long as 0.29× distance between anterior opening and tip of phallus, apically bent ventrolaterally. Anterior and posterior part of phallus form angle of 130°. Caulis large, diverging widely from coecum. Vesica bent at ca. 110° dorsad, with basal widening and terminal diverticulum dorsally, slightly pointed to right. Three sockets of ventroapically located deciduous cornuti adjacent to gonopore are detectable (Fig. 6D).
Female unknown. Preimaginal stages unknown. Molecular data. Unknown. Distribution. Known from the type locality only: Lebanon (Fig. 17). Ecology. Not known.

Remarks.
Comparison of the wing pattern and genitalia of the lectotype and paralectotype of C. acclivana with the species considered above confirmed the assumption that it is a distinct species. The barcoding distance to the other species was not studied because of lack of fresh material, but the great similarity in all morphological characters of the two specimens convinced us that they can represent a distinct species, therefore we resurrect the initial status of the taxon acclivana synonymised by Razowski (1979) with C. neglectana.
Taxa incertae sedis. Without further morphological or genetic support the following species cannot be interpreted with certainty, but the synonymy with C. neglectana, C. acclivana or C. trivia does not seem justified for now. They demonstrate some morphological gap; at least the differences between them are not smaller than the differences with the above mentioned species of the group. Additional material and genetic study is necessary to solve their real status. (Kennel, 1901), stat. rev. Kennel, 1901: 227 (Algeria) Remarks. The species was synonymised with C. neglectana by Razowski (1979) based on examination of the genitalia of the female holotype (Fig. 2N). The slide subsequently was lost and we were unable to re-examine the genitalia. Regarding the type locality (Algeria) it seems unlikely that this taxon is conspecific with C. neglectana. (Chrétien, 1915 (Fig. 10C) are very similar to those of C. striolana. However, there is a clear difference in the wing pattern. Though the lectotype of C. unifasciana var. semiana is poorly preserved, some poorly defined markings can be detected (Fig. 2O). The physical distance between the populations of this taxon and C. striolana is considerable and it is very unlikely that var. semiana is conspecific. The synonymy between C. semiana and C. consimilana proposed by Obraztsov (1955) is not justified. Apparently, it is rooted in the initial incorrect assignment of var. striolana to Cacoecia unifasciana, but the latter turned out to be a junior synonym of Clepsis consimilana (see below). The taxon var. semiana was mechanically moved to the synonymic list of C. consimilana without studying the genitalia of the type specimen.

The Clepsis consimilana species group
This species group is characterised by the presence of large modified setae on the median surface of the valvae (Fig 1).
Description. Adult. Sexual dimorphism prominent. Male (Fig. 11A-F). Head. Vertex, frons and labial palps fulvous. Antennae with scapus and pedicellus ochreous, flagellum with fulvous brown-tipped scales and numerous sensilla trichodea as long as width of flagellomeres. Thorax. Dorsally fulvous, ventrally creamy, legs brownish. Tegula fulvous. Forewing relatively wide, with length 6.3-7.8 mm (mean 7.0, N = 13). Costa basally convex, apically straight, costal fold extending from base to ca. 0.4-0.6 of costa (Fig. 3E). Upperside background dark yellow with rusty reticulate pattern. Markings fulvous to brown: basal blotch atrophied, more prominent at dorsum forming dark dot; median fascia often with darker borders; subapical blotch triangular, often ill-defined. Cilia concolourous with background. Underside pale brown with creamy longitudinal blotch in the distal half of costal area. Hindwings upperside monochrome grey with paler cilia, underside with creamy costal half pale grey with scattered pale brown scales and monochrome pale grey dorsal half. Forewings with variable colouration, darker or paler, sometimes with reduced markings similarly to female (Fig. 11F). Abdomen. Grey. Male genitalia (Fig. 12A-E). Uncus with variable shape and median incision depending on preparation, more or less ovoid, widened distally, rounded. Gnathos plough-shaped. Socius small, membranous. Valvae pointed dorsolaterally when mounted on slide. Costal sclerite protruded medially into large triangular labis with elongated tip (Fig. 5F, G). Apical part of sacculus ca. 1.6× longer than basal part, both forming angle of 160-165°, saccular process small, curved, thornshaped. Distal part of valva membranous, widened apically and protruding into small brachiola, costal edge slightly convex, with longitudinal fold on the median surface bearing row of 5-8 large modified setae. Phallus smooth, coecum medially concave, basal part curved ventrad at ca. 140°, distal part smoothly curved dorsad, with small sharp tipped lateral process as long as 0.26× distance between anterior opening and tip of phallus, slightly curved laterally and not overpassing tip of phallus. Caulis small, adjoining coecum. Vesica cylindrical, curved at angle of 100-120° to phallus, with expansion basally and finger-like sinistrodorsal diverticulum (Fig. 13A, B). Three to four deciduous aciculate, robust, weakly sinuate, slightly flattened cornuti attached ventroapically (Fig. 7D). Gonopore located at left to cornuti sockets and diverticulum. Female darker than male, with uniform forewings (Fig. 11G, H). Head. Frons, vertex and labial palps fulvous to ochreous, antennae with fulvous brown-tipped scales and sparse sensilla trichodea shorter than width of flagellomeres. Thorax dorsally ochreous, ventrally creamy, legs brownish. Forewing length 6.3-7.6 mm (mean 6.8, N = 3). Upperside ground colour ochreous to rusty with darker reticulated pattern and more or less atrophied ill-defined markings. Basal blotch reduced to dark spot at dorsum, median fascia more or less prominent in costal and dorsal area or completely reduced. Cilia paler than background. Underside pale brown, costal and terminal areas paler or creamy with reticulate pattern. Hindwing upperside pale grey, underside with whitish or pale grey reticulate patterned costal half and monochrome pale grey dorsal half. Abdomen. Grey. Female genitalia with papillae anales not modified (Fig. 14A). Apo- physes anteriores 1.5× longer than apophyses posteriores. Sterigma widened caudad, with lateral sclerotised pockets cephalad, large excavation on the dorsal wall and wide v-shaped lamella antevaginalis. Colliculum large, with length 0.17× length of ductus bursae, slightly bent to left, funnel-shaped, with plicate longitudinal sclerotisation forming small process at left, and spherical protrusion at cranial end consisting of colourless thick cuticle. Ductus bursae long and narrow, emerging at left side of cuticular protrusion, with cestum extending along cranial 0.8× of its length. Ductus seminalis inserted dorsally at caudal end of ductus bursae. Corpus bursae ovoid, with large falcate signum with capitulum and flat signum located near end of cestum consisting of sclerotised papillae (Fig. 9D, F).
Preimaginal stages. Detailed descriptions of the ovum, larvae of all instars, and pupa are provided by Sheldon (1920) and Bradley et al. (1973). The chaetotaxy of the larva is described by Swatschek (1958).
Molecular data (Fig. 16). BIN: BOLD:AAC4212. The intraspecific average of the barcode region is 0.34%, the maximum distance 1.08% (p-dist) (N = 29). The minimum distance to the nearest neighbour, Clepsis eatoniana, is 2.25%. Distribution (Fig. 17). Europe, Asia Minor, Syria, European Russia, W Africa to Lebanon, Madagascar (introduced; the data come from the type specimen only), North America (introduced) (Bradley et al. 1973, Razowski 2002. Ecology. Moths are on the wing from May to October in shrubby habitats. The larvae feed on dead or withered leaves of Crataegus, Malus, Carpinus, Polygonum, Hedera, Lonicera, Ligustrum, Syringa, and overwinter in the third instar (Bradley et al. 1973).
Remarks. The collection of Jakob Hübner was acquired by Vincenz Abbate Edler von Mazzola in the early part of the 19 th Century, and the European material was deposited in the "Hof-Naturalien-Kabinett" in 1823 where most of the material was destroyed by fire during the Vienna Rebellion of 1848 (Calhoun 2003, Gilligan andWright 2013). Despite a personal search by PH we found no potential type material and conclude that the syntypes are lost or destroyed. As the boundaries of this taxon are revised, designation of a neotype is necessary to preserve the stability of nomenclature. The type locality is presumably Europe, as indicated by the title of the original description (Hübner 1817) and material may have originated from Germany as many other species described by Hübner. Here we designate as neotype a male specimen with an everted vesica and preserved cornuti, and wing pattern resembling the illustration in the work by Hübner (1817: pl. 38, fig. 239). Though not very detailed, this painting demonstrates some important differences with C. consimilana sensu auctt., namely the shape of basal fascia and costa, and questions the interpretation of the taxon. The painting either refers to another representative of the tribe Archipini or may be a result of an artistic decision. Without studying the type material any interpretation of the figure may be incorrect, therefore we prefer to preserve the present-day concept for C. consimilana. Obraztsov (1955) considered consimilana as a probable synonym of unifasciana. Bradley and Martin (1956) established the name consimilana as having priority over unifasciana, though they did not state explicitly this nomenclatorial act. We found support for this synonymy as the lectotype of Tortrix unifasciana appears conspecific with the neotype of C. consimilana. Razowski (1979) synonymised Tortrix eatoniana with C. consimilana (see below); this is not justified in our opinion. Some taxa from the synonymic list were not examined by us but, minding the minimal genetic diversity throughout Europe, we consider all of them correctly synonymised with C. consimilana. Cacoecia unifasciana var. semiana was erroneously regarded as a synonym of Clepsis consimilana (see above). (Ragonot, 1881), stat. rev. Ragonot, 1881: 231 (Portugal). Tortrix xylotoma Meyrick, 1891: 13 (Algeria) syn. nov. Clepsis razowskii Gastón, Vives & Revilla, 2017: 691 (Spain) syn. nov. non Clepsis consimilana (Hübner, 1817) Material examined. Lectotype ♂ by designation of Razowski (1979) Diagnosis. The wing pattern in males resembles those of C. trivia, C. acclivana, and C. consimilana. Clepsis eatoniana differs from C. consimilana, by the more yellowish instead of fulvous forewing ground colour. The most characteristic feature in the external morphology of C. eatoniana is the absence of a forewing costal fold, in con-trast to C. consimilana. The male genitalia are similar to those in C. consimilana but the valva is more slender, elongate distally, and the modified large setae are more numerous and more slender. The phallus is adorned with spines, with the keel and terminal process overpassing its tip; in C. consimilana the phallus lack lateral spines and a keel, and the phallic process is shorter. The caulis in C. eatoniana is larger, the vesica is bent at nearly a right angle to the phallus, its diverticulum has different location, and the cornuti are more slender and smoother. Females of C. eatoniana are not distinguishable externally from the females of C. consimilana but differ from C. striolana by the presence of two brown dots on the forewing. The female genitalia of C. eatoniana are similar to those in both the C. neglectana and C. consimilana species groups, but the protrusion on the right side of the colliculum is much larger and elongated, and the sterigma has larger lateral pockets.

Tortrix eatoniana
Description. Adult. Sexual dimorphism prominent. Male (Fig. 11I-K). Head. Vertex fulvous, frons and labial palps brown. Antennae with brown scales on scapus and fulvous scales on pedicellus and flagellum, and with numerous sensilla trichodea as long as width of flagellomeres. Thorax. Fulvous dorsally and creamy ventrally, legs brownish. Tegula fulvous with brown costal margin. Forewing with length 6.8-7.4 mm (mean 7.1, N = 3), elongated, costa convex at basal half, slightly sinuate apically. Costal fold lacking (Fig. 3F). Upperside background dark yellow with fulvous reticulate pattern more prominent in the paler subterminal area, cilia concolourous or paler. Basal blotch atrophied, with remnants of darker scales at costa and dorsum. Median fascia brown with lead refractive tint, almost interrupted at middle of median cell by yellowish scales, often ceasing at vein CuA. Subapical blotch brown, triangular, more or less well defined. Underside pale grey-brown with creamy longitudinal blotch in distal half of costal area. Specimens with paler and darker forewings and incomplete median fascia observed. Hindwing upperside monochrome pale grey with paler cilia, underside whitish with scattered pale grey-brown scales. Abdomen. Grey. Male genitalia (Fig. 12F,  15). Uncus variably shaped, more or less trapezoidal, slightly widened distally, rounded, with slightly convex distal edge which may look incised if excessive pressure is applied on coverslip. Gnathos plough-shaped. Socius small, membranous. Valvae pointed dorsolaterally when mounted on slide. Costal sclerite wide, protruded medially into large triangular labis with elongated spinulate tip (Fig. 5H). Basal and apical part of sacculus of nearly equal length, both forming angle of 150-160°, saccular process large, flat, triangular. Distal part of valva membranous, comparatively small, narrow, with parallel costal and saccular margins, apically triangular, without distinct brachiola, with longitudinal fold on the median surface bearing row of 8-12 large modified setae. Apical part may look as brachiola due to deformation during preparation. Phallus robust, coecum medially concave, basal part curved ventrad at ca. 140°, distal part smoothly curved dorsad, with wide keel on left side grading into large sharp tipped lateral process as long as 0.23× distance between anterior opening and tip of phallus, slightly curved laterally and dorsally, overpassing phallic tip. Several more or less prominent spines pointed caudad located at basal part of keel. Caulis large, widely separated from coecum. Vesica cylindrical, curved at 80-90° to phallus, with small expansion basally and fingerlike dorsal diverticulum (Fig. 13C). Three deciduous, slender, straight cornuti attached ventroapically (Fig. 7E). Gonopore located in straight line with cornuti sockets and diverticulum. Female darker than male (Fig. 11L). Head. Vertex rusty, frons and labial palps brown. Whole antennae with brown scales, sensilla trichodea sparse, shorter than width of flagellomeres. Thorax. Rusty dorsally and creamy ventrally, legs brownish. Tegula rusty with brown costal margin. Forewing length 6.7 mm (n = 1), with shape as in males, upperside uniformly rusty with ill-defined reticulate pattern more distinct in distal half and two brown dots at dorsum, underside mainly creamy with scattered pale brown scales, denser in middle of wing. Hindwing upperside grey brown, underside creamy with darker dorsal part. Abdomen grey.
Female genitalia (Fig. 14B). Papillae anales not modified. Apophyses anteriores 1.1× longer than apophyses posteriores. Sterigma relatively wide, widened cephalad, with large lateral sclerotised pockets, large excavation on dorsal wall and wide v-shaped median part of lamella antevaginalis. Colliculum large, with length 0.25× length of ductus bursae, asymmetrical, funnel-shaped, with well-developed sclerotisation and two evaginations: very large one at right and small one at left, both consisting of colourless thick cuticle. Ductus bursae long and narrow, emerging at left between outpocketings, with cestum extending along cranial 0.7× of its length. Ductus seminalis inserted dorsally at caudal end of ductus bursae. Corpus bursae ovoid, with large falcate signum with capitulum and flat signum consisting of sclerotised papillae located near end of cestum (Fig. 9E).
Ecology. The moths were collected in macchie habitat (Gastón et al. 2017) from April to the first half of September. The larval host plant is unknown. Remarks. The species was described by Ragonot (1881) after two males from Portugal. He emphasised its resemblance with C. consimilana. Razowski (1979) considered it conspecific with C. consimilana, probably relying on the superficial similarity of the modified setae of the valvae of these two taxa. The female remained unstudied until Gastón et al. (2017) described it again under the name C. razowskii. Tortrix xylotoma was sunk into C. neglectana by Razowski (1979). This synonym is not justified as the genitalia of the male lectotype (illustrated also by Clarke 1958: pl. 109) rather resemble C. eatoniana regarding the shape of labis, presence of modified setae on the valva, and morphology of the phallus. The preparation of the genitalia is poor, therefore the shape of uncus and valvae looks unusual. The wing pattern fits with C. eatoniana. After comparison of material from Spain, the lectotype of T. eatoniana, the lectotype of T. xylotoma, the original description of C. razowskii, and numerous photographs of specimens identified as C. razowskii by the authors of the latter taxon, we concluded that all these specimens are conspecific and the present valid name of the species should be C. eatoniana.

Discussion
The taxa treated in the present paper are not easily distinguished from each other because of prominent sexual dimorphism in some of them, variability in wing pattern and considerable similarity in the genitalia. The subtle differences in both male and female (when available) genitalia, secondary sex characters (male forewing costal fold) and barcode gap between the populations support presence of more than the two taxa currently recognised. The most defining characters are found in the genitalia. Two groups of taxa can be recognised: those with large modified setae on the median surface of the valvae in males, and those without such setae. Apparently, this was the main criterion for prior synonymisation resulting in only two species: C. consimilana (with setae) and C. neglectana (without setae). Further scrutinising of the genital characters revealed existence of several taxa, which can be treated as species. Here we propose two species groups named after the oldest described taxon in each of them. They should not be confused with the groups proposed by Razowski (1979) for subgeneric divisions of Clepsis, which appear hierarchically superior to our groups.
Since recently collected material for mtDNA sequencing was available for European populations only, the status of some taxa of the C. neglectana species group remains unresolved. Clepsis severana and C. semiana appear to be closely related considering the genital morphology, and their synonymy with some of the remaining species of the group cannot be excluded. They are known only from their holotypes (by monotypy), and moreover, the morphology of the vesica is inaccessible because of the prolonged embedding in microscope slide media (C. semiana) or the genitalia slide is missing (C. severana). On the other hand, each of them demonstrates some differences with all remaining members of the group, which seemingly are not lesser than those between well separated taxa as C. neglectana and C. trivia. Tortrix xylotoma is certainly a synonym of a member of the C. consimilana species group, namely C. eatoniana, but not of C. neglectana as it was treated up to now. The status of C. acclivana is more or less controversial because of lacking DNA barcode sequences, but the two male specimens with practically identical morphology, different to all the other taxa convinced us that they represent a distinct species.
The Cretan population of C. trivia is well separated genetically and morphologically from the other European species (C. neglectana and C. striolana), but demonstrates certain morphological similarity with the other Mediterranean taxon, C. acclivana, and is probably related to it. It should be emphasised that the Cretan specimens were assigned to C. trivia solely on male genitalia morphology without studying the vesica of the holotype and its DNA barcode and it may still represent a separate taxon.
The barcode distance between C. neglectana and C. striolana is comparatively small. The genital morphology of these two taxa also demonstrates considerable similarity, but they are well distinguished by wing pattern and costal fold. Apparently, C. striolana has a range limited in the region of the Alps. It can be assumed that these species had been isolated recently, maybe only after the last glacial period, and even continuing gene flow cannot be dismissed. Some taxa still in synonymy with C. neglectana may turn out to be species bona as well, e.g., Tortrix dorana from Eastern Kazakhstan.
The C. consimilana species group encompasses only two species and is separated from the C. neglectana group by considerable morphological and genetic gaps. The species have strikingly disproportional distribution ranges: C. eatoniana appears to be a Western-Mediterranean species distributed in the Iberian Peninsula and Algeria, and possibly part of France, but C. consimilana is distributed throughout the rest of Europe, part of Asia and has been introduced to the tropics (Madagascar) and North America. Some morphological variability in C. consimilana (the position of cornuti and other minor male genitalia characters) somewhat contradicts the genetic invariance within this species in Europe.
The overreliance or underestimation of certain genital characters can cause incorrect taxonomic interpretations, as it is demonstrated by the above mentioned taxonomic problems. Traditionally, the shape of uncus, gnathos and valvae is widely used in taxonomy of Lepidoptera, but our experience with other groups as well (Zlatkov and Huemer 2017) proved that these characters must be treated cautiously. Being complex three-dimensional structures, their visible shape depends on several factors: fixation before embedding in a permanent medium, thickness of the medium layer and pressure applied on the coverslip. The shape of uncus and distal part of valva are especially prone to deformation in the species treated here. Some relatively flat sclerites were found to be less affected and hence more useful for taxonomical purposes, e.g., the sacculus and in lesser extent the costal sclerite, including the labis, of the valva. The morphology of the vesica and phallus is believed to be very informative but must be studied before mounting on a slide. Moreover, these structures are considerably simplified in the taxa considered here and provide a relatively small number of characters. The female genitalia can also provide many characters but only in a comparatively limited region: the colliculum and part of the sterigma. In fact, these are the structures which interact directly with the male genitalia. It can be assumed that the distal part of the phallus is inserted into the colliculum and its lateral process is accommodated into the transparent cuticular extension at the right cephalic end, therefore they should be studied in comparative aspect to obtain additional taxonomical information. Some taxonomic problems in the C. neglectana and C. consimilana species groups had apparently ensued from ignoring female genitalia morphology or simply the lack of female specimens.
(Vienna, Austria). We thank also Joaquin Baixeras (València, Spain), Jason Dombroskie (Cornell, USA), and Erik J. van Nieukerken (Leiden, Netherlands) for helpful comments and suggestions on an earlier version of the paper.