A Systematic Revision of Charissa, Subgenus Pterygnophos Wehrli, 1951, with Description of a New Species (Lepidoptera: Geometridae)

Zootaxa 4341 (3): 400–418 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4341.3.4 http://zoobank.org/urn:lsid:zoobank.org:pub:FF11A5C7-2DAD-444C-8520-6098932AE208 A systematic revision of Charissa, subgenus Pterygnophos Wehrli, 1951, with description of a new species (Lepidoptera: Geometridae)

SVEN ERLACHER1,3,4, LAURA MARRERO PALMA1,2 & JOSEPHA ERLACHER3 1 Museum für Naturkunde, Moritzstraße 20, D-09111 Chemnitz, Germany. E-mail: [email protected] 2 Calle Giralda 13, E-41930 Bormujos (Sevilla), Spain 3 Further Straße 18, D-09113 Chemnitz, Germany 4Corresponding author

Abstract

The subgenus Pterygnophos Wehrli, 1951 within the genus Charissa Curtis, 1826 nomen protectum (= Hyposcotis Hüb- ner, [1825] nomen oblitum) is taxonomically revised based on morphology and DNA barcoding. The subgenus comprises four species in total which are presented in detail. Diagnostic characters are depicted and keys to the species based on the morphology of male and female genitalia are provided. Males and females of each species and their genitalia are illustra- ted. The distribution of all species is described and figured on a map, and a neighbor joining tree based on DNA barcoding of 17 specimens is presented. Charissa (Pterygnophos) beljaevi spec. nov. from Mongolia is described as new. A neotype for Gnophos creperaria Erschoff, 1877, and lectotypes for Gnophos deliciaria shantungensis Wehrli, 1953, Gnophos dorkadiaria Wehrli, 1922, Gnophos ochrofasciata Staudinger, 1895, and Gnophos finitimaria Fuchs, 1899 are designated. The following synonyms are recognized: Gnophos finitimaria Fuchs, 1899 syn. nov. is a synonym of Gnophos ochrofasciata Staudinger, 1895 and Gnophos deliciaria shantungensis Wehrli, 1953 syn. nov. is a synonym of Gnophos agnitaria Staudinger, 1897.

Key words: Gnophos, Hyposcotis, Charissa (Pterygnophos) beljaevi spec. nov., DNA barcoding, morphology, synonyms, taxonomy, Asia

Resumen

El subgénero Pterygnophos Wehrli, 1951 dentro del género Charissa Curtis, 1826 nomen protectum (= Hyposcotis Hüb- ner, [1825] nomen oblitum) es revisado taxonómicamemte basándose en morfología y DNA barcoding. El subgénero comprende cuatro especies en total, las cuales están presentadas en detalle. Los caracteres diagnósticos han sido descritos y se adjuntan claves para su determinación basadas en la morfología de los aparatos genitales masculinos y femeninos. Ejemplares de machos y hembras de cada especie han sido ilustrados. La distribución de todas las especies está descrita e ilustrada en mapas adjuntos. Charissa (Pterygnophos) beljaevi spec. nov. de Mongolia se describe como nueva. Han sido designados un neotipo para Gnophos creperaria Erschoff, 1877, y lectotipos para Gnophos deliciaria shantungensis Weh- rli, 1953, Gnophos dorkadiaria Wehrli, 1922, Gnophos ochrofasciata Staudinger, 1895, y Gnophos finitimaria Fuchs, 1899. Los siguientes sinónimos son reconocidos: Gnophos finitimaria Fuchs, 1899 syn. nov. es un sinónimo de Gnophos ochrofasciata Staudinger, 1895 y Gnophos deliciaria shantungensis Wehrli, 1953 syn. nov. es un sinónimo de Gnophos agnitaria Staudinger, 1897.

Introduction

Charissa Curtis, 1826 is a species-rich genus within the Ennominae, the largest subfamily of Geometrid moths. Most of the species of Charissa occur in the Palearctic region. They look externally very similar due to their adaptation to rocks, and this is the reason why they are generally considered to be hard to determine. For this purpose it is often necessary to analyze their genitalia, but even on that basis finding of differences between species

400 Accepted by A. Saldaitis: 2 Aug. 2017; published: 1 Nov. 2017 is sometimes difficult. Until now, about 30 species have been newly transferred from Gnophos Treitschke, 1825 to Charissa (Parsons et al. 1999). The taxon Pterygnophos Wehrli, 1951 was originally proposed as a subgenus of Gnophos by Wehrli (1951). The type species is Gnophos ochrofasciata Staudinger, 1895. It was again proposed when the former French article was translated into German and published in Wehrli (1953). Subsequently, Pterygnophos was raised to generic rank by Vojnits (1975) and so the taxon was retained by Viidalepp (1988, 1996). In his study on the Gnophos-group, Sauter (1990) treated Pterygnophos along with other subgenera of Gnophos as a subgenus of Charissa. However, Parsons et al. (1999) retained the species of Pterygnophos within Gnophos and listed the subgenus as a synonym of the latter, as they did with all previously described subgenera, without giving any reasons. Finally, Beljaev (2016) treated Pterygnophos as a subgenus of Charissa and transferred the three known Pterygnophos species to Charissa without emphasizing this action as a taxonomic act. Members of Pterygnophos are distributed across Central Asia and the Far East. Apart from some short comments and somewhat ambiguous illustrations by Viidalepp (1975, 1988), there has been no revision of this group since Wehrli (1953). Recently collected material of Charissa spp. from Mongolia and the Russian part of the Altai Mountains has revealed difficulties in identification which necessitate a revision of the subgenus Pterygnophos. Some previous researchers have not succeeded in establishing a clear taxonomic order in the subgenus because of a number of taxonomic problems, for example: the type series of Gnophos ochrofasciata contained specimens of another species (C. agnitaria); the genitalia of some types were stored in glycerol or even in wax; the genitalia of the holotype of “Gnophos erschoffi” Wehrli, 1922 (an unnecessary replacement name for Gnophos creperaria Erschoff, 1877) were poorly prepared by a previous colleague (Pl. 2, Fig. 3e); and last but not least females have been almost impossible to associate with conspecific males and even with subgenera. Our studies have concluded that the subgenus Pterygnophos comprises four species. Morphological characters are described in detail, and keys for determination of all species are given. One species is described as new. Additionally, DNA barcoding was used to support the results obtained by morphological examination.

Methods

Morphology. Moths were identified on the basis of both external and internal morphological characters. Genitalia preparation was conducted as described in Trusch & Erlacher (2001). Photographs of the genitalia were taken with the stereomicroscope Nikon SMZ1000 and a connected Nikon D90 camera. In order to obtain optimally focused images several photographs with different focal planes were combined by means of the freely available software CombineZP. Photos were enhanced and arranged to plates with Adobe® Photoshop® software. The following external characters were measured: wingspan (widest distance between forewing margins in set specimens); forewing length (length of the costa from the base to the apex); forewing width (longest line of the forewing parallel to the body in set specimens). Terminology of male and female genitalia is explained in Fig. 1. The following genitalia characters were measured (see Fig. 2): in the male genitalia—total length (tl; distance from the posterior end of the uncus to the anterior end of the saccus); length of ventral valva margin (vm.l; straight measured distance from the base of the strongly sclerotized valva margin to the distal end of the valva process); aedeagus length (ae.l); in the female genitalia—total length (tl; distance from the posterior end of the ovipositor to the anterior end of the corpus bursae); antrum length (an.l; distance from the ostium to the anterior end of the strongly sclerotized antrum); antrum width (an.w; widest lateral extension of the antrum); length of the spined longitudinal folding (slf.l; distance from the antrum-bursa junction to the anterior end of the strongly sclerotized folding). DNA analysis. All specimens were processed at the Canadian Centre for DNA Barcoding (CCDB, Guelph) to obtain DNA barcodes using the standard high-throughput protocol as described in Ivanova et al. (2006) and deWaard et al. (2008) (regularly updated protocols used at the CCDB can also be found at http://www.ccdb.ca/ resources.php). PCR amplification with a single pair of primers consistently recovered a 658 bp region near the 5’ end of the mitochondrial cytochrome c oxidase I (COI) gene that includes the standard 648 bp barcode region for the animal kingdom (Hebert et al. 2003). DNA extracts are stored at the CCDB. Complete specimen data including images, voucher deposition, barcode identification number, sequence process ID, barcode index number (BIN),

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 401 GPS coordinates, sequence and trace files can easily be accessed in BOLD (Barcode of Life Data System, Ratnasingham & Hebert 2007; Ratnasingham 2017) in the public data set DS-PTERYGNO (https://doi.org/ 10.5883/DS-PTERYGNO). 17 specimens were successfully sequenced and analyzed with BOLD’s tools (http:// www.boldsystems.org). Gnophos furvata [Denis & Schiffermüller], 1775, was used as outgroup. A neighbor joining tree (Saitou & Nei 1987) with all barcoded species of this article has been made with BOLD, analyzing the nucleotide sequences with a pairwise distances model and including fragment lengths of >400 bp. Genetic distances between species were obtained from a barcode gap analysis of full-length barcode fragments using the analytical tools of BOLD and are reported as minimum pairwise distances, Kimura 2 parameter (Kimura 1980). Abbreviations. The examined specimens are deposited in the following institutional and private collections. Abbreviations of institutional insect collections follow Parsons et al. (1999) and Evenhuis (2017), with priority to Parsons et al. (1999).

FIGURE 1. Structures of the genitalia in Charissa, subgenus Pterygnophos, based on the example of C. (P.) beljaevi spec. nov. a)—male, b)—female. ab: appendix bursae; ae: aedeagus; an: antrum; ap: apex; c: cornutus; cb: corpus bursae; co: costa; gn: gnathos; ju: juxta; op: ovipositor; pvp: postvaginal plate; sa: saccus; slf: spined longitudinal folding; sd: sclerotized valva dilation; un: uncus; vm: ventral valva margin; vp: valva process.

Institutional collections:

BMNH Natural History Museum, London, United Kingdom IZCAS Chinese Academy of Sciences, Institute of Zoology, Beijing, China MNC Museum für Naturkunde Chemnitz (coll. S. Erlacher), Germany MNHU Museum für Naturkunde, Berlin, Germany ZFMK Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany ZRAS Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia ZSM Zoologische Staatssammlung, Munich, Germany

GJK J. Gelbrecht, Königs Wusterhausen, Germany LHS H. Löbel, Sondershausen, Germany MRJ R. Müller, Jena, Germany RUK U. Ratzel, Karlsruhe, Germany SBD B. Schacht, Dahlewitz, Germany SGA G. Sircoulomb, Anquetierville, France SMM M. Sommerer, Munich, Germany SPV P. Skou, Vester Skerninge, Denmark

Further abbreviations: barcode id. = barcode sample identification number; BIN = barcode index number of BOLD systems; E = east of; env. = environment; gen. prep. = genitalia slide number; lux = light-trap; m = meters above sea level; Mts. = Mountains; N = north of; Prov. = Province of; S = south of; W = west of.

FIGURE 2. Measured genitalia structures based on the example of C. (P.) agnitaria a)—male, b)—female. ae.l: aedeagus length; an.l: antrum length; an.w: antrum width; slf.l: length of longitudinal spined folding; tl: total length; vm.l: length of ventral valva margin. Scale bar: 1 mm.

Results

Checklist of Charissa, subgenus Pterygnophos

Charissa Curtis, 1826 nomen protectum = Hyposcotis Hübner, [1825] nomen oblitum Subgenus Pterygnophos Wehrli, 1951 1. Charissa (Pterygnophos) agnitaria (Staudinger, 1897) = Gnophos deliciaria shantungensis Wehrli, 1953 syn. nov.

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 403 = Gnophos dorkadiaria Wehrli, 1922 = Gnophos sphalera Wehrli, 1938 2. Charissa (Pterygnophos) beljaevi spec. nov. 3. Charissa (Pterygnophos) creperaria Erschoff, 1877 = Gnophos erschoffi Wehrli, 1922 (unnecessary replacement) 4. Charissa (Pterygnophos) ochrofasciata (Staudinger, 1895) = Gnophos finitimaria Fuchs, 1899 syn. nov.

Key to Charissa (Pterygnophos) species: male genitalia structures (Abbreviations in brackets refer to Fig. 1a)

1 Sclerotized dilation present close to base of costa (sd) ...... 2 - Without sclerotized dilation close to base of costa ...... 3 2 Sclerotized dilation (sd) with serrate keel, juxta (ju) almost symmetrical ...... C. (P.) beljaevi spec. nov. - Sclerotized dilation swollen with minute spines, juxta usually distinctly asymmetrical ...... C. (P.) creperaria 3 Valva processes (vp) about half of costa length, length of ventral valva margin (vm) 2.2–2.9 mm, without costal thorns...... C. (P.) agnitaria - Valva processes about 1/5 of costa length, length of ventral valva margin 2.0–2.3 mm, with costal thorns ...... C. (P.) ochrofasciata

Key to Charissa (Pterygnophos) species: female genitalia structures (Abbreviations in brackets refer to Fig. 1b)

The structure of the female genitalia is not basically different from some species of Charissa, subgenus Dysgnophos Wehrli, 1951. Occasionally, it can be difficult to judge whether a specimen under examination is a female of Pterygnophos or of Dysgnophos. For this reason, using the provided key assumes that the female in question actually belongs to Pterygnophos.

1 Postvaginal plate (pvp) posteriorly almost not extending the antrum ...... C. (P.) beljaevi spec. nov. - Postvaginal plate strongly extending the antrum ...... 2 2 Antrum comparatively short (1.57–1.83 mm) and broadly funnel-shaped (antrum width: 0.87–1.00 mm). . . C. (P.) creperaria - Antrum longer (1.77–2.91 mm) and slender (antrum width: < 0.82 mm) ...... 3 3 Antrum length > 2.2 mm and length of spined longitudinal folding (slf) > 1.3 mm ...... C. (P.) agnitaria - Antrum length 1.8–2.3 mm and length of spined longitudinal folding (slf) 1.1–1.3 mm ...... C. (P.) agnitaria or C. (P.) ochrofasciata Shape and measured values of genitalia structures are too similar and widely overlapping, making a diagnosis ambiguous. Fur- ther information, for example origin of the specimen, should be taken into account and, if possible, comparative material should be used.

Charissa Curtis, 1826 nomen protectum (= Hyposcotis Hübner, [1825] nomen oblitum)

Charissa Curtis, 1826, British entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland. John Curtis, London: object number 105.—Type species: Geometra obscurata [Denis & Schiffermüller], 1775, originally designated as “Geometra obscuraria Hüb.” (unjustified emendation). Hyposcotis Hübner, [1825], Verzeichniß bekannter Schmettlinge[sic!], Jakob Hübner, Augsburg: 314.—Type species: Geometra mucidaria Hübner, [1799], by subsequent designation by Prout (1904: 122).

Diagnosis. The genus Charissa is characterized by the following autapomorphic character states:

– Juxta divided into two arms, without terminal spines – Costa more sclerotized than the rest of the valva, pilose but with not more than one or two spines – Ductus bursae (= part between antrum and base of ductus seminalis) not present or extremely short – Bursa copulatrix with sclerotized longitudinal folds

404 · Zootaxa 4341 (3) © 2017 Magnolia Press ERLACHER ET AL. Moths belonging to Charissa are externally very similar to those belonging to Gnophos, Gnophopsodos Wehrli, 1945, Rhiphignophos Wehrli, 1951 and several other genera, which are just as perfectly adapted to rocks. Most of the species of Charissa are medium sized moths with medial lines and discal spots often distinct. The antennae of males range from filiform to not more than shortly bipectinate, both states occurring even in the same subgenus (e.g. Cnestrognophos Wehrli, 1951). Hind tibiae are clavately thickened. The wing venation is very variable in the genus, even within a single species, and not at all constant as indicated in Sauter (1990). Unlike the situation with external characters, the species of Charissa can be recognized by their male and female genitalia (see list of autapomorphies). Distribution. The genus Charissa currently comprises about 70 species occurring across the Palearctic ecozone and the Arabian Peninsula. One member of the subgenus Dysgnophos Wehrli, 1951 also occurs in the Nearctic. Remarks. Objectively, there are some problems in using the name “Charissa”. In his bibliographical account of the entomological works of Jacob Hübner, Hemming (1937) concluded that the pages 305–431 of Hübner’s “Verzeichniss bekannter Schmettlinge[sic!]” including the generic name Hyposcotis most likely appeared in 1825. On this basis, he initiated an “Opinion” by the International Commission on Zoological Nomenclature to establish his findings in order to stabilize nomenclature (ICZN 1943). When Sauter (1990) proposed the genus name Charissa Curtis, 1826 for several subgenera and species formerly belonging to Gnophos Treitschke, 1825, he obviously overlooked that Hyposcotis Hübner, [1825] should have been used, although Fletcher (1979) already considered both Hyposcotis and Charissa as valid. However, since their type species, Geometra mucidaria Hübner, [1799] and Geometra obscurata [Denis & Schiffermüller], 1775 respectively, are currently treated as congeneric (e.g. Parsons et al. 1999), Charissa and Hyposcotis must be considered to be synonyms. For Hyposcotis, to our knowledge, there are only four sources in which it has been used according to Article 23.9.1.1. ICZN after 1899 (Prout 1904, Oberthür 1913, Prout 1912–1916, Prout 1914). Apart from the general list in Fletcher (1979) (according to Article 23.9.6. ICZN), more than 100 years have passed without Hyposcotis being used as a genus name. On the other hand, many authors have increasingly used Charissa in their publications since Sauter (1990) and now the name is widely adopted, as exemplified in Erlacher & Erlacher (2017). Nevertheless, the name Hyposcotis instead of Charissa was recently used by two Turkish colleagues (e.g. Kemal & Koçak 2015; Kemal & Koçak 2016). Even if this seems unnecessary, it is in accordance with a strict interpretation of the ICZN (2017). To avoid taxonomic instability and confusion amongst the scientific community it was decided to refer this matter to the International Commission of Zoological Nomenclature for a ruling under the plenary power (Erlacher & Erlacher 2017). According to § 23.9.3. ICZN the prevailing usage of Charissa Curtis, 1826 is to be maintained while the case is under consideration (nomen protectum).

Subgenus Pterygnophos Wehrli, 1951

Pterygnophos Wehrli, 1951: Lambillionea, 51: 25.—Type species: Gnophos ochrofasciata Staudinger, 1895.

Description. Medium sized moth species of high variability in coloration within species, upper side of wings with range of colors from dark-brown to light-gray or cream, underside generally pale and dull, from brownish to grayish-white; wing pattern with medial lines and discal spot sometimes distinct, sometimes mingling with background; medial lines serrate or indicated by dots; hindwing termen slightly curved; fringes with alternating dark and light patches; antennae filiform in both sexes. Male genitalia (Pl. 2). Aedeagus straight and stout, more or less abruptly narrowing at proximal end, with a cornutus shaped as plate with sclerotized swelling (with exception of C. (P.) agnitaria); valva with a single ventral truncate, sclerotized process; distal part of costa (cucullus) with setae, costal thorns absent (with exception of C. (P.) ochrofasciata); juxta long, asymmetric (with exception of C. (P.) beljaevi spec. nov.), widened at its distal part and divided up to the base; gnathos and uncus small, and strongly sclerotized. Female genitalia (Pl. 3). Antrum funnel-like and well sclerotized; corpus bursae with strongly sclerotized, spined longitudinal folding, and laterally with appendix bursae (dome-like, ruffled structure housing a set of strong and large dentes). Diagnosis. The subgenus Pterygnophos is characterized by the following autapomorphic character state:

– Valva bipartite with a single truncate, sclerotized process

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 405 Moths of the subgenus Pterygnophos are externally not certainly distinguishable from other Charissa species, especially from those of the subgenera Dysgnophos and Trilobignophos Wehrli, 1951. However, the species of Charissa (Pterygnophos) can clearly be recognized by their male genitalia, which are characterized by bipartite valva with the above-mentioned single ventral process. The species of Charissa (Dysgnophos) and Charissa (Trilobignophos) have undivided valva without sclerotized processes. Furthermore, the latter are characterized by several basally bulbous cornuti inside the aedeagus, as already stated in the original description of the subgenus by Wehrli (1951).

FIGURE 3. Neighbor joining tree for all species of Charissa, subgenus Pterygnophos, on the basis of DNA barcoding (pairwise distances model for COI-5P marker) with Gnophos furvata ([Denis & Schiffermüller], 1775) as outgroup. Terminal branches provided with species name, barcode id., barcode length, origin, and BIN number from BOLD.

406 · Zootaxa 4341 (3) © 2017 Magnolia Press ERLACHER ET AL. Distribution. Species of the subgenus Pterygnophos occur across Asia from Kyrgyzstan in the West to Korea in the East. The north-south-extension ranges from Krasnoyarsk in Siberia to the central regions of China. They live at altitudes up to 2700 m. Distributions of the species are mapped in Fig. 5. Biology. There are no descriptions of larval stages and the natural food plants are unknown. The flight periods last from June to August, possibly in one generation, and in one species (C. (P.) agnitaria) from the middle of May to the beginning of October. Remarks. Due to morphological correspondence of the male and female genitalia with the autapomorphic character states of the genus Charissa, the taxon Pterygnophos is treated as a subgenus of the latter. DNA barcoding analyses with BOLD (http://boldsystems.org) strengthen this assumption. A neighbor joining tree for all known species of the subgenus Pterygnophos based on DNA barcoding is depicted in Fig. 3. Genetic distances to the nearest neighbor of the Pterygnophos species resulting from an analysis of DNA barcodes are shown in Tab. 1. Since it appears to be difficult, even for experts, to determine species of the subgenus Pterygnophos only verified material was used. Thus, gaps in distribution were rather accepted than incorrect identifications. The actual distribution of the species belonging to the subgenus Pterygnophos may therefore be wider than currently known.

TABLE 1. Genetic distances (minimum pairwise distances, Kimura 2 parameter) to the nearest neighbor of four species of Charissa, subgenus Pterygnophos resulting from a barcode gap analysis of full-length barcode fragments using the analytical tools of BOLD (COI 5’, 658 bp). C. (P.) ochrofasciata C. (P.) creperaria C. (P.) beljaevi spec. nov. C. (P.) agnitaria 2.35% 2.83% 4.96% C. (P.) beljaevi spec. nov. 3.79% 2.66% C. (P.) creperaria 2.02%

1. Charissa (Pterygnophos) agnitaria (Staudinger, 1897) (adults Pl. 1, Figs 1a–1d; male gen. Pl. 2, Fig. 1e; female gen. Pl. 3, Figs 1f, 1g)

Gnophos agnitaria Staudinger, 1897, Deutsche Entomologische Zeitschrift Iris, 10: 62, pl. 3, fig. 40.—Locus typicus: Russia, Askold Island. Deposition of holotype: MNHU. Examined. Gnophos deliciaria shantungensis Wehrli, 1953 syn. nov., Die Spanner des palaearktischen Faunengebietes. In: Seitz, A. (Ed.): Die Gross-Schmetterlinge der Erde, Supplement zu Band 4, Alfred Kernen, Stuttgart: 593.—Locus typicus: China, Prov. Shandong [Shantung], Tai–Shan, 1550 m. Deposition of lectotype: ZFMK. Examined. Gnophos dorkadiaria Wehrli, 1922, Deutsche Entomologische Zeitschrift Iris, 36: 20, pl. 1, fig. 16, pl. 2, fig. 26, 42.—Locus typicus: Kukunoor [China, Qinghai Lake]. Deposition of lectotype: ZFMK. Examined. “Gnophos perdita” Staudinger, 1897, Deutsche Entomologische Zeitschrift Iris, 10: 61.—Lapsus calami of Gnophos paerlita Butler, 1886: 389; misidentified. Gnophos sphalera Wehrli, 1938, Mitteilungen der Münchner Entomologischen Gesellschaft, 28: 89.—Locus typicus: Russia, Vladivostok. Deposition of holotype: ZFMK. Examined.

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 407 Lectotype ♂ of Gnophos deliciaria shantungensis Wehrli, 1953 (hereby designated): [China:] ‘Tai–shan. (1550 m). | Prov. Shantung, China. | 3.5.1934. H. Höne.’, ‘v. shantungensis | W[e]h[r]li. | Gnoph[os]. | deliciaria | Oberth[uer]. | Dr. Wehrli ♂ Type’, ‘GlobInG | specimen ID: | 83 | Exemplar + Etiketten | dokumentiert | specimen + label | data documented’, ‘SE–706 ♂ | gen. prep. | Erlacher, 2016’, ‘LECTOTYPE ♂ | Gnophos deliciaria shantungensis | Wehrli, 1953 | Designated by | Erlacher, Marrero & Erlacher (2017)’, ZFMK.—Paralectotypes 2♀ of Gnophos deliciaria shantungensis Wehrli, 1953 (hereby designated): China: 1♀, Prov. Shandong [Shantung], Tai–Shan, 1550 m, 12.v.1934, leg. H. Höne, gen. prep. SE–707; 1♀, same locality, 1934, leg. H. Höne, gen. prep. Bonn–5 / ZFMK–69 (marked as “holotyp” by mistake), ZFMK. Holotype ♀[sic!] of Gnophos sphalera Wehrli, 1938 (Pl. 1, Fig. 1b): [Russia:] ‘Wladiwoastock’, ‘5.6. [18]77’, ‘sphalera | Wehrli | Gnophos | Type.’, ‘Gnophos perlita, Butl[er] | (agrees with sp. from Kala– | pani but not so well with | others from Dharmsala)’, ‘GlobInG | specimen ID: | 0005 | Exemplar + Etiketten | dokumentiert | specimen + label | data documented’, ‘[gen. prep. Wehrli–]7168 ♂[sic!]’, ‘HOLOTYPE ♀ | Gnophos sphalera Wehrli, 1938’, ZFMK. Further material. China: 1♂, Beijing Shi, Donglingshan, Xiaolongmen, barcode id. BC–AxYi–0109, BIN BOLD:AAC5997, IZCAS. 1♂, Tibet, Qinghai Lake [Kuku Noor], coll. A. Bang-Haas; 1♀, same locality, 1898, leg. Rückbeil, gen. prep. SE–711, MNHU. 1♂, same locality, 1894, leg. Rückbeil, gen. prep. JH–265; 1♂ same data, gen. prep. SE–574; 1♂, same data, gen. prep. SE–575; 1♂, same data, gen. prep. SE–576; 1♂, Prov. Shandong, Tai–Shan, 1550 m, 04.x.1934, leg. H. Höne, gen. prep. Rezbanyai-Reser No. 7614; 1♂, Prov. Shaanxi [“Sued–Shensi”], Taibaishan [“Tapaishan”], Qinling [“Tsingling”], ca. 1700 m, 9.ix.1936, leg. H. Höne, gen. prep. SE–708; 1♀, same locality, 18.ix.1935, leg. H. Höne, gen. prep. SE–709; 1♂, same locality, 21.ix.1935, leg. H. Höne; 1♀, same locality, 03.vi.1936, leg. H. Höne; 1♂, Prov. Shanxi, Mian Shan, 1500 m, 07.vi.1937, leg. H. Höne, gen. prep. SE–583; 1♀, same locality, 04.vi.1937, leg. H. Höne, gen. prep. SE–584, ZFMK. 1♂, Tibet, Qinghai Lake [Kuku Noor], leg. R. Tancré , gen. prep. SE–712; 1♀, Prov. Hebei, Baiqi, Chongli, 1350 m, 41.05°N / 115.317°E, 23.ix.2006, leg. C. Wang, barcode id. BC–ZSM–Lep–13095; 1♀, same locality, 19.v.2007, leg. C. Wang, barcode id. BC–ZSM–Lep–13107; 1♀, same data, barcode id. BC–ZSM–Lep–13101; 1♀, same data, barcode id. BC–ZSM–Lep–13104; 1♀, same locality, 10.v.2010, leg. C. Wang, barcode id. BC–ZSM–Lep–12977; 2♀, same locality, 28.v.2006, leg. C. Wang, barcode ids BC–ZSM–Lep–13050 and BC–ZSM–Lep–13046; 1♀, Prov. Hebei, Gaojiaying, 40.8833°N / 114.95°E, 15.v.2007, barcode id. BC–ZSM–Lep–13509; 1♂, Dayushu, Yanqing, Beijing Shi, 40.3833°N / 115.95°E, 520 m, 12.ix.2007, leg. C. Wang, barcode id. BC–ZSM–Lep–15635; 1♀, same locality, 23.ix.2006, leg. C. Wang, barcode id. BC–ZSM–Lep–13415; 1♀, same locality, 13.ix.2007, leg. C. Wang, barcode id. BC–ZSM–Lep–15647, ZSM. Korea: 1♂, South–Korea, Seoul, Kalinowalski, Mus. de Varsovie, gen. prep. JH–151, barcode id. SE–MNC–Lep–01291, BIN BOLD:AAC5997; 1♀, North–Korea, Seishin–Olto [= Ch’eongjin], ca. 41°47’N / 129°47’E, leg. A. Kricheldorff, gen. prep. SE–572, ZFMK. Mongolia: 1♂ (syntype of Gnophos ochrofasciata Staudinger, 1895, misidentified), 1894, leg. Lederer, gen. prep. SE–566, MNHU. 1♀ (Pl. 3, Fig. 1g), Prov. Ömnögovi, 20 km E Sevrei Sum, Zoolongii nuruu, Shain Khökh, Khuushuue, 23.–24.viii.1997, leg. P. Gyulai & A. Garai, gen. prep. SE–580, barcode id. SE–MNC–Lep–01152, BIN BOLD:AAC5997, SMM. 1♀, Prov. Govi Altai, 45 km SE Biger, 1850 m, 05.vi.2004, leg. A. Saldaitis, gen. prep. SE–582, barcode id. SE–MNC–Lep–01121 and SE–MNC–Lep–01268, BIN BOLD:AAC5997, SPV. 1♂, Terelj, ca. 80 km NW Ulan Bator, 1500–1600 m, e.o. 29.ix.1983, leg. H. Salpeter, gen. prep. / barcode id. DNATAX– 2317; 1♀ (Pl. 1, Fig. 1d), same locality, e.o. 05.ix.1983, leg. H. Salpeter, gen. prep. / barcode id. DNATAX–2318; 1♀, same locality, e.o. 15.x.1983, barcode id. SE–MNC–Lep–00142; 1♀, same locality, e.o. 09.ix.1983, barcode id. SE–MNC–Lep–00143, GJK. Russia: 1♂, South Siberia, Kyka village, Cita locality, 25.v.–10.vi.1999, leg. J. Miatleuski, gen. prep. JH–271, SGA. 1♂ (Pl. 2, Fig. 1e), Primorsky Krai, Ussuriysk district, 42 km SW Ussuriysk, Krounovka River, 30.v.–01.vi.2003, leg. E.A. Beljaev, gen. prep. JH–150, barcode id. SE–MNC–Lep–01061, BIN BOLD:AAC5997; 1♀, same data, gen. prep. SE–579, barcode id. SE–MNC–Lep–01062, BIN BOLD:AAC5997, SPV. 1♀, Prov. Tuva, Sayan Mts., Turan, 2000 m, vii., gen. prep. SE–573; 1♂, without data, gen. prep. Wehrli–601, ZFMK. 1♂, Askold, gen. prep. Wehrli–7136, MNHU. Description. Adults (Pl. 1, Figs 1a–1d). Measurements. Wingspan: 27–33 mm (♂), 30–33 mm (♀); forewing length: 14–17 mm (♂), 16–18 mm (♀); forewing width: 9–11 mm (♂), 10–11 mm (♀). Wings. Coloration varies between different shades of cream and grayish-brown, variably patterned intense dark grayish-brown.

408 · Zootaxa 4341 (3) © 2017 Magnolia Press ERLACHER ET AL. Male genitalia (Pl. 2, Fig. 1e). Total length: 2.29–2.78 mm; length of ventral valva margin: 2.21–2.92 mm; uncus and gnathos small, blunt-ended and strongly sclerotized; juxta distinctly asymmetric; no costal thorns developed; aedeagus length: 2.34–3.13 mm; aedeagus without cornutus. Female genitalia (Pl. 3, Figs 1f, 1g). Total length: 4.6–6.6 mm; antrum length: 1.8–2.91 mm; antrum width: 0.63–0.81 mm; length of spined longitudinal folding: 1.12–1.64 mm. Diagnosis. Males of C. (P.) agnitaria are determinable by the length of the sclerotized ventral valva margin (2.21–2.92 mm), which is shorter in the three other species of the subgenus (1.61–2.35 mm). This differentiation can be done by sight with no need for actual measuring. The only species that could resemble C. (P.) agnitaria in this specific character is C. (P.) creperaria, when there is no size comparison. But the latter species has a swollen sclerotized dilation close to its costa covered with minute spines, which is absent in C. (P.) agnitaria. Typical females of C. (P.) agnitaria are remarkable for their slender-looking and long antrum (1.8–2.91 mm) and also the long spined longitudinal folding (1.12–1.64 mm). In some specimens the measurements do not lead to sure distinction between the species, because the measurement-ranges of different species slightly overlap. Especially females of C. (P.) ochrofasciata are sometimes very similar to those of C. (P.) agnitaria. Unfortunately, in this case there is no sure distinction by use of female genitalia structures. C. (P.) beljaevi spec. nov. differs from C. (P.) agnitaria in the postvaginal plate, which strongly extends the antrum in C. (P.) agnitaria, while it not extends the antrum in C. (P.) beljaevi spec. nov. Furthermore, the antrum of the latter is clearly shorter with 1.37 mm at maximum, whereas the antrum length of C. (P.) agnitaria amounts to 1.8 mm at minimum, as mentioned above. Females of C. (P.) creperaria have the same postvaginal plate as C. (P.) agnitaria females, but cannot be confused with them due to their remarkably broad, funnel-shaped antrum. Distribution (Fig. 5). This species is distributed from Central Asia to the Far East, and from South Siberia to China. In Kyrgyzstan it occurs around the Issyk–Kul Lake, in Mongolia north of Ulan Bator, in the Govi Altai Province SE Biger and in the Ömnögovi Province 20 km E Sevrei District. In China the species is distributed around the Qinghai Lake in Qinghai Province, as well as in the Provinces of Shaanxi, Shanxi, Hebei including the Municipality of Beijing, and Shandong. In Russia it is to be found near Lake Baikal, on Askold’s Island around Vladivostok and in Primorsky Krai around Ussuriysk. Charissa (P.) agnitaria is also known from North and South Korea. The moths occur at an altitude range from 520 m to 2000 m. Biology. The flight period of C. (P.) agnitaria lasts from the beginning of May to the beginning of October, possibly in some regions in two generations. Breeding of this species was successful with Taraxacum officinale and Artemisia vulgaris as food plants for the caterpillars (J. Gelbrecht and B. Müller in lit.). Remarks. Due to the highly variable habitus of the adults it had not been possible to match males of C. (P.) agnitaria to the female holotype. Checking the locations as well as DNA barcode analyses have helped to confirm the conspecificity of male and female specimens of the species. Gnophos deliciaria shantungensis Wehrli, 1953 syn. nov. is recognized to be conspecific with Charissa (Pterygnophos) agnitaria (Staudinger, 1897). Based on available molecular data the nearest species to C. (P.) agnitaria is C. (P.) ochrofasciata with a COI distance of 2.35% (see also Tab. 1 and Fig. 5). The morphology of male and female genitalia also indicates a close relationship between these species.

2. Charissa (Pterygnophos) beljaevi spec. nov. (adults Pl. 1, Figs 2a–2d; male gen. Pl. 2, Fig. 2e; female gen. Pl. 3, Fig. 2f)

Locus typicus: Mongolia, Selenge Province, Amarbayasgalant Monastery. Deposition of holotype: MNC.

Material examined. Type material. Holotype ♂ (Pl. 1, Fig. 2a; Pl. 2, Fig. 2e): ‘Mongolia: Selenge | Amarbajasgalant Mo- | nastir (Felssteppe) | 49°56’N / 103°5715”E | 1060 m; 07.vii.2004 | leg. Karel Černý’, ‘barcode id. SE–MNC–Lep–01147’, [BIN BOLD:ABW0084], ‘JH–154 ♂ | gen. prep. | Harmeling, 2013’, ‘HOLOTYPE ♂ | Charissa | (Pterygnophos) beljaevi | Erlacher, Marrero & Erlacher 2017’, MNC. Paratypes 2♂, 3♀: Mongolia: 1♂, Prov. Selenge, Amarbayasgalant Monastery, 49°56’N / 103°57’15”E, 1060 m, 07.vii.2004, leg. Karel Černý, gen. prep. SE–577, barcode id. SE–MNC–Lep–01148, BIN BOLD:ABW0084; 1♀ (Pl. 1, Fig. 2c; Pl. 3, Fig. 2f), same data, gen. prep. SE–578, barcode id. SE–MNC–Lep–01290, BIN

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 409 BOLD:ABW0084, SMM. 1♂ (Pl. 1, Fig. 2b), Ulan Bator [Urga], gen. prep. JH–157; 1♀, same data, gen. prep. SE– 585; 1♀ (Pl. 1, Fig. 2d), same data, gen. prep. SE–601, MNHU. Description. Adults (Pl. 1, Figs 2a–2d). Measurements. Wingspan: 31–35 mm (♂), 37 mm (♀); forewing length: 15–17 mm (♂), 17–19 mm (♀); forewing width: 10–11 mm (♂), 9–12 mm (♀). Wings. Ground coloration of different shades of light-brown with diffuse gray-brown pattern; discal spot dark gray-brown, sometimes only faintly contrasting with the background; medial lines weakly defined, indicated by dots or even not visible; underside likewise variable, pale-cream and light-gray, antemedial line absent, postmedial line visible or missing. Male genitalia (Pl. 2, Fig. 2e). Total length: 2.31–2.30 mm; length of ventral valva margin: 1.61–1.84 mm; uncus and gnathos both blunt and strongly sclerotized, gnathos slender, hook-like; juxta almost symmetrical; sclerotized dilation close to base of costa with serrate keel, not swollen; aedeagus length: 2.04–2.24 mm; aedeagus with cornutus. Female genitalia (Pl. 3, Fig. 2f). Total length: 4.4–5.71 mm; antrum length: 1.35–1.37 mm; antrum width: 0.77–0.87 mm; length of spined longitudinal folding: 0.94–1.12 mm; antrum relatively short and stout, postvaginal plate almost not extending the antrum. Diagnosis. The male genitalia of C. (P.) beljaevi spec. nov. may resemble those of C. (P.) creperaria and C. (P.) ochrofasciata. Both Charissa (P.) beljaevi spec. nov. and C. (P.) creperaria have a sclerotized dilation close to the base of the costa but in C. (P.) beljaevi spec. nov. this structure is not swollen and has a serrate keel, while it is bloated and covered with minute spines in C. (P.) creperaria (Fig. 4). Charissa (P.) ochrofasciata resembles C. (P.) beljaevi spec. nov. in the shape of the short valva process but besides the fact that the above mentioned valva sclerotization is absent in C. (P.) ochrofasciata, the species can be recognized by its costal thorns, situated proximal to the setae covered apex. Regarding the female genitalia, identification of C. (P.) beljaevi spec. nov. is possible on the basis of the antrum, which is shorter than in the other species (antrum length: 1.35–1.37 mm at maximum) and the postvaginal plate is not strongly extending the antrum as it is in the female genitalia of the other species of the subgenus Pterygnophos. Distribution (Fig. 5). The holotype and two paratypes of C. (P.) beljaevi spec. nov. originate from the Monastery of Amarbayasgalant located in Selenge Province (1060 m) and three further paratypes have been captured around Ulan Bator, both in Mongolia. Biology. All known specimens have been collected in the beginning of July. Remarks. Males and females of the new species were assigned to each other by DNA barcoding analyzed with BOLD (http://www.boldsystems.org). Based on available molecular data the nearest species to C. (P.) beljaevi spec. nov. is C. (P.) creperaria with a COI distance of 2.66% (see also Tab. 1 and Fig. 5). The morphology of male and female genitalia also indicates a close relationship between these species. Etymology. The new species is dedicated to our colleague and friend Eugeniy A. Beljaev, Vladivostok, Russia, a leading authority in Geometrid moths of the Far East, in recognition of his consistent great willingness to help.

FIGURE 4. Detail of valva with arrows pointing at diagnostic characters. A—C. (P.) beljaevi spec. nov.; B—C. (P.) creperaria.

3. Charissa (Pterygnophos) creperaria Erschoff, 1877 (adults Pl. 1, Figs 3a–3d; male gen. Pl. 2, Figs 3e, 3f; female gen. Pl. 3, Fig. 3g)

Gnophos creperaria Erschoff, 1877, Horae Societatis Entomologicae rossicae, 12: 337, St. Petersburg.— Locus typicus: Russia, Radde. Deposition of neotype: ZFMK. Examined.

Material examined. Type material. Neotype ♂ of Gnophos creperaria Erschoff, 1877 (Pl. 1, Fig. 3a; Pl. 2, Fig. 3e) (according to Article 75 and 76 ICZN hereby designated): [Russia:] ‘Creperaria Ersch[off] | ♂ 31.7.[18]86 | Raddeffka [Radde]’, ‘[Wehrli–]543 | ♂ | creperaria | ?’, ‘erschoffi | W[e]h[r]li. | Gnophos | abgebildet Seitz IV. | Suppl. fig. | Dr. Wehrli’, ‘GlobInG | specimen ID: | 85 | Exemplar + Etiketten | dokumentiert | specimen + label | data documented’, ‘NEOTYPE ♂ | Gnophos creperaria Erschoff, 1877 | Designated by | Wehrli (1922)’, ZFMK. Further material. Mongolia: 1♀, Prov. Khentii, Khentii Mts., 04.vii., coll. O. Bang-Haas; 1♂, Ulan Bator, 30.vii., gen. prep. SE–597; 1♀, same data, gen. prep. SE–611, MNHU. 1♂, Prov. Selenge, Ero–Gol River, 49°48’N / 106°35’E, 650 m, 28.–30.vii.2002, leg. A. Saldaitis, gen. prep. JH–272, SGA. 1♀, Prov. Ömnögovi, “Gurvansaikhan Mts.”, Eliin pass, 22.–23.vii.1997, leg. P. Gyulai & A. Garai, gen. prep. SE–581, barcode id. SE– MNC–Lep–01149, BIN BOLD:ACE3591, SMM. 1♀, Prov. Ömnögovi, ca. 75 km W Dalanzadgad, 43.613045°N / 103.77472°E, 2300 m, 09.vii.2001, leg. K. Nadrowski (Nadrowski & Albrecht 2007), gen. prep. SE–661, barcode id. SE–MNC–Lep–01150, BIN BOLD:ACE3591; 1♀ (Pl. 1, Fig. 3d), Ulan Bator, e.o. 08.xi.1987, leg. Peter Salk, gen. prep. DNATAX–02316, barcode id. SE–MNC–Lep–00144, BIN BOLD:AAI7416; 1♂, same data, e.o. 08.x.1987, leg. P. Salk, gen. prep. DNATAX–2315, barcode id. SE–MNC–Lep–00145, BIN BOLD:AAI7416, ZSM. Russia: 1♂, South–Siberia, Prov. Tuva, Eastern Sayan Mts., Tunkinskie Gol'tsy Mts., Mt. Kulugaisha, near

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 411 Mondy, 1700 m, 07.viii.2000, lux, leg. L. Lehmann, gen. prep. JH–285; 1♂, same data, barcode id. SE–MNC– Lep–00260, BIN BOLD:AAI7416, MNC. 1♂, Siberia, Yablonoi Mts., leg. Dörries, gen. prep. SE–598; 1♀, same data, gen. prep. SE–599; 1♀, same data; 1♂, Region Jewish Autonomous Oblast, Radde, 31.vii.1886, leg. R. Tancré, gen. prep. SE–600; 1♀, Sayan Mts., Baranovskiy, leg. Dörries; 1♀ (syntype of Gnophos turfosaria Wehrli, 1922, misidentified), before iv.1912, gen. prep. JH–159, MNHU. 1♂, Buryatia, W Lake Baikal, Primorskii Chrebet, 60 km SE Irkutsk, near Malo Goloustnoye, 52°18’038’’N / 105°18’177’’E, 700–790 m, 02.–12.vii.2015, lux, leg. A. Steidel; 2♀ (1♀ Pl. 1, Fig. 3c), Buryatia, Sayan Mts., 400 km W Lake Baikal, 24 km N Sagan Shuluta, 1850–1950 m, 14.–18.vii.2015, lux, leg. A. Steidel & O. Czadek, MRJ. 1♂, Schawyr, E Tannuola, 2500 m, vi., gen. prep. JH–143; 1♀ (Pl. 3, Fig. 3g), Sayan Mts., Tunkinskie Gol'tsy Mts. [“Tunkinsk Weisgbg.”], SW Irkutsk, 2000 m, vii., SE–571; 1♂ (Pl. 1, Fig. 3b), Prov. Irkutsk, Sayan Mts., Tunkinskie Gol'tsy Mts. [“Tunkisk Weisberge”], 2000 m, vii., gen. prep. JH–153; 1♂ (Pl. 2, Fig. 3f), Prov. Tuva, Sayan Mts., Tunkinskie Gol'tsy Mts., Turan, 2000 m, vii., gen. prep. JH–152, ZFMK. Description. Adults (Pl. 1, Figs 3a–3d). Measurements. Wingspan: 28–33 mm (♂), 31–33 mm (♀); forewing length: 15–18 mm (♂), 15–17 mm (♀); forewing width: 10–11 mm (♂), 10–12 mm (♀). Wings. Upper side with a range of colors from dirty-white to dark brown; wing pattern with medial lines and discal spot sometimes clearly defined and rich in contrast, sometimes almost missing or blurring with an unevenly darkened background; underside dull, cream or light-gray, faintly patterned. Male genitalia (Pl. 2, Figs 3e, 3f). Total length: 2.23–2.56 mm; length of ventral valva margin: 1.92–2.16 mm; uncus and gnathos small, strongly sclerotized; sclerotized, swollen dilation close to base of costa covered with minute spines; juxta with degree of asymmetry variable; aedeagus length: 2.10–2.36 mm; aedeagus with cornutus. Female genitalia (Pl. 3, Fig. 3g). Total length: 4.57–5.41 mm; antrum length: 1.57–1.83 mm; antrum width: 0.87–1.00 mm; length of spined longitudinal folding: 0.86–1.12 mm. Diagnosis. Males of C. (P.) creperaria can be confused with male specimens of C. (P.) agnitaria because of the shape of the ventral valva with its long process, and with males of C. (P.) beljaevi spec. nov. because of the presence of a sclerotized dilation close to the base of the costa. For distinction see diagnosis of the respective species. The female genitalia of C. (P.) creperaria are characteristic because of the short and posteriorly remarkably broadened antrum (antrum width of C. (P.) creperaria: 0.87–1.00 mm; other species of the subgenus Pterygnophos: 0.87 mm at maximum), thus forming a wide funnel. Distribution (Fig. 5). Charissa (P.) creperaria is known from southern Siberia where it has been found in the Sayan Mountains, near Schawyr east of Tannuola and around Radde (Radeffka) located in the Jewish Autonomous Oblast. In Mongolia the species occurs around Ulan Bator, in the Khentii Mountains of Khentii Province, in Selenge Province at Ero–Gol River and in South Mongolia (Ömnögovi Province) both from the Gurvansaikhan Mountains and from 75 km E Dalanzadgad. The vertical distribution range extends from 650 m (Ero–Gol River) to 2500 m (Schawyr). Biology. The flight period lasts from June to the beginning of August. Breeding of C. (P.) creperaria was successful with withered leaves of Taraxacum officinale as food plant for the caterpillars (B. Müller in lit.). Remarks. Alpheraky (1892) had already stated that the female holotype of Gnophos creperaria was not present in the Erschoff collection, St. Petersburg. It could also not be found in ZRAS today (Mironov in lit. 2016). Therefore it is to be assumed that the holotype was destroyed. A neotype for Gnophos creperaria Erschoff, 1877 is designated with the purpose of clarifying its taxonomic status according to Article 75 and 76 ICZN. Wehrli’s description of “Gnophos erschoffi” explicitly referred to Gnophos creperaria which he already recognized as unidentifiable (Wehrli 1922). This should be interpreted as an invalid attempt to designate a neotype: “Ich ziehe deshalb den Namen für die fragwürdige Art ein und setze an deren Stelle eine klar umschriebene Species […]” (Wehrli 1922: 19). In accordance with Recommendation 75D of Article 75.7 ICZN it was decided to give preference to this earlier invalid “neotype” although it does not come as nearly as practicable from the original type locality. The assumption by Vojnits (1975) that “Gnophos erschoffi” is a synonym of “Dysgnophos benepunctarius” without examination of type specimens is preposterous and must be rejected. Based on available molecular data the nearest species to C. (P.) creperaria is C. (P.) ochrofasciata with a COI distance of 2.35% (see also Tab. 1 and Fig. 5), whereas the morphology of male and female genitalia indicates a close relationship between C. (P.) creperaria and C. (P.) beljaevi spec. nov.

412 · Zootaxa 4341 (3) © 2017 Magnolia Press ERLACHER ET AL. PLATE 1. Adults of Charissa, subgenus Pterygnophos. 1—C. (P.) agnitaria: 1a—♂, 1b—♂, 1c—holotype ♀, 1d—♀; 2—C. (P.) beljaevi spec. nov.: 2a—holotype ♂, 2b—paratype ♂, 2c—paratype ♀, 2d—paratype ♀; 3—C. (P.) creperaria: 3a— neotype ♂, 3b—♂, 3c—♀, 3d—♀; 4—C. (P.) ochrofasciata: 4a—lectotype ♂, 4b—♂, 4c—paralectotype ♀, 4d—♀. Undersides of specimens are denoted by ‘*’. Scale bar: 10 mm.

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 413 PLATE 2. Male genitalia of Charissa, subgenus Pterygnophos, with arrows pointing at diagnostic characters; valvae and aedeagus. 1e—C. (P.) agnitaria; 2e—C. (P.) beljaevi spec. nov. (holotype); 3e—C. (P.) creperaria (lectotype), 3f—C. (P.) creperaria; 4e—C. (P.) ochrofasciata (lectotype). Scale bar: 1 mm.

414 · Zootaxa 4341 (3) © 2017 Magnolia Press ERLACHER ET AL. PLATE 3. Female genitalia of Charissa, subgenus Pterygnophos, with arrows pointing at diagnostic characters. 1f—C. (P.) agnitaria (holotype), 1g—C. (P.) agnitaria; 2f—C. (P.) beljaevi spec. nov. (paratype); 3g—C. (P.) creperaria; 4f—C. (P.) ochrofasciata. Scale bar: 1 mm.

4. Charissa (Pterygnophos) ochrofasciata (Staudinger, 1895) (adults Pl. 1, Figs 4a–4d; male gen. Pl. 2, Fig. 4e; female gen. Pl. 3, Fig. 4f)

Gnophos ochrofasciata Staudinger, 1895, Deutsche Entomologische Zeitschrift Iris, 8: 362, pl. 6, fig. 15.—Locus typicus: Mongolia, Uliassutai. Deposition of lectotype: MNHU. Examined. Gnophos finitimaria Fuchs, 1899 syn. nov., Jahrbücher des Nassauischen Vereins für Naturkunde, 52: 149.—Locus typicus: Russia, Krasnojarsk. Deposition of lectotype MNHU. Examined.

Material examined. Type material. Lectotype ♂ of Gnophos ochrofasciata Staudinger, 1895 (Pl. 1, Fig. 4a; Pl. 2, Fig. 4e) (hereby designated): [Mongolia:] ‘Sibiria | (Uliassatai)’, ‘Cotype | ochrofasciata St[audin]g[e]r. ♂’, ‘SE–595 ♂ | gen. prep. | Erlacher, 2013’, ‘LECTOTYPE ♂ | Gnophos ochrofasciata Staudinger, 1895 | Designated by | Erlacher, Marrero & Erlacher (2017)’, MNHU.—Paralectotypes 3♂, 3♀ of Gnophos ochrofasciata Staudinger, 1895: Mongolia: 1♂, Prov. Zavkhan, Ulias(sa)tai, gen. prep. Wehrli–7111; 1♂, same locality, gen. prep. JH–156; 1♂, same locality, gen. prep. SE–596; 1♀ (Pl. 1, Fig. 4c), same locality, gen. prep. SE–569; 1♀, same locality, gen. prep. SE–586; 1♀, same locality, MNHU. Lectotype ♂ of Gnophos finitimaria Fuchs, 1899 (hereby designated): [Russia:] ‘Krasnojarsk | Ost–Sibirien | Juli | 1880 | Archlow’, ‘Cotype | finitimaria Fuchs ♂’, ‘Finitimaria Fs.’, ‘SE–568 ♂ | gen. prep. | Erlacher, 2013’, ‘LECTOTYPE ♂ | Gnophos finitimaria Fuchs, 1899 | Designated by | Erlacher, Marrero & Erlacher (2017)’, MNHU.—Paralectotype 1♂ of Gnophos finimitaria Fuchs, 1899 (hereby designated): Russia: East–Siberia, Krasnojarsk, vii.1880, gen. prep. SE–567, MNHU.

REVISION OF CHARISSA, SUBGENUS PTERYGNOPHOS Zootaxa 4341 (3) © 2017 Magnolia Press · 415 Further material. Mongolia: 1♂, Govi Altai, S Mongolian Altai Mts., Alag Kharkhan Mts., 45°31’56.3’’N / 94°04’02.3’’E, 2500 m, 05.–07.vii.2004, leg. A. Saldaitis, gen. prep. JH–155, barcode ids. SE–MNC–Lep–01120 and SE–MNC–Lep–01267, BIN BOLD:ABZ7842, SPV. Russia: 1♂ (Pl. 1, Fig. 4b), Buryatia, Sayan Mts., 400 km W Lake Baikal, 24 km N Sagan Shuluta, 1850–1950 m, 14.–18.vii.2015, lux, leg. A. Steidel & O. Czadek, gen. prep. SE–737; 1♀, same data, gen. prep. SE–739, MRJ. 16♂, 2♀, Altai Republic, 3 km NW Kuray, steppe slopes (“Steppenhänge“), 1650 m, 29.vii.2011, leg. B. Schacht; 22♂, 1♀, Altai Republic, Aktash, 1300–1600 m, 30.vii.– 04.viii.2011, leg. B. Schacht; 1♂, same locality, 27.–29.vii.2011; 5♂, same locality, 17.–22.vii.2009, leg. B. Schacht; 1♀, same data, gen. prep. SE–404, barcode id. SE–MNC–Lep–00978, BIN BOLD:ACF0610; 2♂, same locality, 1600–2000 m, 17.vii.2010, leg. B. Schacht; 6 ♂, same locality, 06.–15.vii.2010, leg. B. Schacht; 1♂, Altai Republic, Aktash, Tourist Basis, 25 km NW Inja Kartun River, 670 m, 13.vi.2012, leg. B. Schacht, SBD. 1♂, Altai Republic, 3–5 km W Aktash, 12.–17.vii.2010, leg. H. Löbel; 1♀, same data, gen. prep. SE–405, barcode id. SE– MNC–Lep–00979, BIN BOLD:ACF0610; 1♀, Altai Republic, Altai Mts., Kuraiskij Khrebet, Aktash, 1365 m, 50°19’N / 87°36.5’E, ♀: 28.vii.2011, e.o. 08.vi.2012, gen. prep. JH–257; 1♀ (Pl. 1, Fig. 4d; Pl. 3, Fig. 4f), same data, e.o. 09.vii.2012, leg. J. Gelbrecht, gen. prep. JH–258; 1♀, Altai Republic, Altai Mts., ca. 10 km SE Aktash, 50°15’25’’N / 87°42’48’’E, 1480 m, 29.vii.2011, leg. J. Gelbrecht & E. Schwabe, gen. prep. JH–260, LHS. 1♂, Altai Republic, SE Altai, Kosh–Agachsky District, Kuraisky Ridge, 5 km E Chagan–Uzun, 50°24’27’’N / 87°35’50’’E, 2130 m, 08.vii.2012, leg. A. Volynkin, gen. prep. SE–570; 2♂, Altai Republic, Ulagan N Aktash, Aigulaksky Ridge, 50°19’N / 87°35’E, 1400 m, 3.–5.viii.2011, leg. A. Volynkin; 4♂, same locality, 11.– 13.viii.2001, leg. A. Volynkin; 4♂, same locality, 5.vii.2012, leg. A. Volynkin, RUK. Description. Adults (Pl. 1, Figs 4a–4d). Measurements. Wingspan: 30–34 mm (♂), 32–34 mm (♀); forewing length: 16–17 mm (♂), 18–19 mm (♀); forewing width: 10–11 mm (♂), 11–12 mm (♀). Wings. Mongolian specimens ocher-colored, slightly scattered with grayish-brown scales, forming patches distal of the medial band; Russian specimens of yellowish-white ground color with silver-gray pattern; medial band framed by silver-gray bands; underside similar to the respective upper side color and pattern, but paler. Male genitalia (Pl. 2, Fig. 4e). Total length: 2.23–2.50 mm; length of ventral valva margin: 2.0–2.35 mm; uncus and gnathos small, strongly sclerotized; juxta asymmetric; costal thorn developed proximal to setae-covered apex; aedeagus length: 2.18–2.55 mm; aedeagus with sclerotized cornutus. Female genitalia (Pl. 3, Fig. 4f). Total length: 5.33–6.13 mm; antrum length: 1.77–2.12 mm; antrum width: 0.73–0.78 mm; length of spined longitudinal folding: 1.18–1.26 mm. Diagnosis. Male genitalia of C. (P.) ochrofasciata are similar to those of C. (P.) beljaevi spec. nov. For distinction see diagnosis of the latter. The female genitalia resemble those of C. (P.) agnitaria. It seems that in most specimens of C. (P.) ochrofasciata the antrum is slightly shorter than in C. (P.) agnitaria but this is unlikely to provide a sure diagnosis. See also diagnosis of C. (P.) agnitaria. Distribution (Fig. 5). In Mongolia C. (P.) ochrofasciata occurs around Uliastai, capital of Zavkhan Province, and in the Govi Altai Province in the Alag Kharkhan Mountains (southern part of Mongolian Altai Mountains). The species is further known from Russia, where it is distributed around Aktash, NW Kuray and in the Kosh– Agachsky District, all located in the Altai Republic, as well in the Republic of Buryatia near Mondy, and in Krasnojarsk. It occurs up to altitudes of 2500 m. Biology. The flight period lasts from the beginning of June to the end of July. As in the other species of the subgenus Pterygnophos no more is known about the biology of C. (P.) ochrofasciata. Unlike in C. (P.) agnitaria and C. (P.) creperaria, the caterpillars of C. (P.) ochrofasciata rejected Taraxacum officinale but accepted Artemisia absinthium. The caterpillars overwinter (B. Müller in lit.). Remarks. Gnophos finitimaria Fuchs, 1899 syn. nov. is recognized to be conspecific with Charissa (Pterygnophos) ochrofasciata (Staudinger, 1895). Based on available molecular data the nearest species to C. (P.) ochrofasciata is C. (P.) creperaria with a COI distance of 2.02% (see also Tab. 1 and Fig. 5), whereas the morphology of male and female genitalia indicates a close relationship between C. (P.) ochrofasciata and C. (P.) agnitaria.

We are grateful to John Chainey, Natural History Museum, London, United Kingdom; Wolfram Mey, Museum für Naturkunde, Berlin, Germany; Dieter Stüning, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany; Vladimir Mironov, Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia; Axel Hausmann, Zoologische Staatssammlung, Munich, Germany; Jörg Gelbrecht, Königs Wusterhausen, Germany; Hans Löbel, Sondershausen, Germany; Rando Müller, Jena, Germany; Ulrich Ratzel, Karlsruhe, Germany; Bernd Schacht, Dahlewitz, Germany; Guy Sircoulomb, Anquetierville, France; Peder Skou, Vester Skerninge, Denmark, and Manfred Sommerer, Munich, Germany, for their generosity in loan of material. We are also grateful to Paul D.N. Hebert, Biodiversity Institute of Ontario, University of Guelph, Canada, and his competent teams at the Canadian Centre for DNA Barcoding (CCDB, University of Guelph), for sequencing and for access to the BOLD informatics platform; DNA analyses were supported by Genome Canada, the Ontario Ministry of Research and Innovation and Natural Science and Engineering Research Council of Canada (NSERC) in the framework of the International Barcode of Life (iBOL) program. Once more we would like to thank Axel Hausmann for giving advice and support in many situations. We would also like to thank Jörg Gelbrecht, Vladimir Mironov, Manfred Sommerer, and especially Evgeny Beljaev, Institute of Biology and Soil Science, Laboratory of Entomology, Vladivostok, Russia, as well as an anonymous reviewer for their useful comments on the manuscript. Special thanks to Sir Anthony C. Galsworthy, London, United Kingdom, who assisted in checking the English. This research received support from the SYNTHESYS Project (www.synthesys.info) financed by the European Community Research Infrastructure Action under the FP6 “Structuring the European Research Area” program.

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