Zootaxa 4571 (1): 099–110 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4571.1.6 http://zoobank.org/urn:lsid:zoobank.org:pub:EAE0ADD7-512A-4A99-8B58-6283B2FB0BDA

Neochloroglyphica, a new genus of from China (, Geometridae), with description of a new species

HONGXIANG HAN1, PEDER SKOU2 & RUI CHENG1,3 1Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China 2Aamosen 1, Ollerup, DK-5762 Vester Skerninge, Denmark 3Corresponding author. E-mail: [email protected]

Abstract

Neochloroglyphica gen. nov. and its type species N. perbella sp. nov. are described from Yunnan, China. Morphological characters and molecular phylogenetic analysis, based on one mitochondrial and three nuclear genes, support the hypoth- esis that Neochloroglyphica is a member of the tribe Neohipparchini, and that it is a sister genus to Chloroglyphica. Mor- phological characters, including those of the genitalia, are figured and compared with related genera, especially Chloroglyphica, Neohipparchus and Chlororithra. Diagnoses for the genus and the species are provided and illustrations of external features and genitalia are presented.

Key words: Neochloroglyphica, new genus, phylogeny,

Introduction

The subfamily Geometrinae, commonly known as emerald , is the fourth largest subfamily in Geometridae, with more than 2500 described species in approximately 270 genera worldwide (Scoble & Hausmann 2007). Considerable progress has been made in systematics and phylogeny within the Geometrinae. The major global taxonomic revision of Geometrinae was that of Prout (1912, 1934–38). Subsequently, many regional works on Geometrinae have been produced. For example, Inoue (1961) reviewed the Japanese Geometrinae and established ten tribes; Ferguson (1969, 1985) studied the Geometrinae of North America; Pitkin (1996) researched the Neotropical Geometrinae; Holloway (1996) reviewed the Bornean Geometrinae and divided the subfamily into two tribes: Dysphaniini and Geometrini; McQuillan & Edwards (1996) studied Australian Geometrinae; Viidalepp (1996) produced a checklist of Geometridae from the USSR; Hausmann (2001) investigated Geometrinae from Europe; Beljaev (2016) published a catalogue of the Geometrinae of the Russian Far East. Han & Xue (2011) studied Chinese representatives of the subfamily, and placed most genera into nine tribes. Among molecular phylogenetic studies, the most comprehensive taxon sampling at the tribal level was the research by Sihvonen et al. (2011), in which 27 geometrine species in 25 genera, representing 16 of 18 geometrine tribes sensu Forum Herbulot (2007) were included. Ban et al. (2018) revised the tribal classification of the Geometrinae based on 116 species belonging to 56 genera, which were mainly Palaearctic and Oriental, and proposed that the Geometrinae were composed of 13 tribes: Ornithospilini, Agathiini, Hemitheini, Dysphaniini, , Aracimini, Neohipparchini, Nemoriini, Synchlorini, Dichordophorini, Comibaenini, Timandromorphini and Geometrini. Through examination of recently collected specimens, the collections of IZCAS, ZFMK and PCPS, one undescribed geometrine species that could not be placed in any known genus were discovered. This study aims to describe the new genus based on its type species, to provide clear diagnostic characters in relation to other genera, and to evaluate the generic placement of the species.

Accepted by S. Erlacher: 11 Jan. 2019; published: 25 Mar. 2019 99 Material and methods

Specimens of the new species are deposited in the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IZCAS), Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK), and Private Collection of Peder Skou (PCPS). Morphological examinations. Dissection and preparation of genitalia slides were performed following the standard protocols (Robinson 1976). Genitalia were embedded in Canada balsam and mounted on slides. Terminology for wing venation followed the Comstock-Needham System (Comstock, 1918) as adopted for Geometridae by Scoble (1992) and Hausmann (2001), and that for the genitalia was based on Pierce (1914, reprint 1976), Klots (1970) and Nichols (1989). Moths were photographed with a digital camera (Canon Pc1057). Composite images were generated using Auto-Montage software version 5.03.0061 (Synoptics Ltd). The plates were compiled in Adobe Photoshop software 7.0. Ink (Adobe Systems Software Ireland Ltd). Molecular examinations. To explore the phylogenetic affinity of the new genus within the Geometrinae, molecular phylogenetic analyses were conducted based on 18 species belonging to 13 genera representing six geometrine tribes (Aracimini, Neohipparchini, Nemoriini, Comibaenini, Timandromorphini and Geometrini). Most data were from our recently published study (Ban et al. 2018), except that of Neochloroglyphica and new sequences of Chloroglyphica Warren. In total, fragments of one mitochondrial protein-coding gene (COI), and three nuclear protein-coding genes (CAD, GAPDH and RPS5) were sequenced. The methods of DNA extraction, amplification, sequencing and phylogenetic analyses follow Ban et al. (2018) and Cheng et al. (2016). Details of studied specimens, including GenBank accession numbers, are summarized in Table 1. A phylogenetic tree was reconstructed for the combined dataset using the maximum likelihood (ML) method in RAxML v7.2.6 (Stamatakis 2006). We adopted five exemplars representing the tribes Hemitheini and Pseudoterpnini as outgroups. For ML analyses, the dataset was partitioned by gene and the GTR+GAMMA+I model for each partition was used. All model parameters were estimated during the ML analysis. A rapid bootstrapping algorithm with a random seed value of 12345 (command -f a -x 12345) was applied with 1000 replicates. Based on the generic placement, pairwise distances within and between Neohipparchini species for the COI barcoding region (640 bp) were calculated, and a neighbour-joining (NJ) tree (Saitou and Nei 1987) was constructed based on the Kimura 2-parameter (K2P) method (Kimura 1980) using MEGA 6.0.

Results

Phylogenetic relationships. The analyses conducted in this study are based on the sequence data from one mitochondrial gene region (1371 bp of COI), and three nuclear gene regions (847 bp of CAD, 706 bp of GAPDH, 602 bp of RPS5). In the maximum likelihood analyses, Neochloroglyphica is placed in the tribe Neohipparchini, with strong bootstrap support, and appears as a sister-group to Chloroglyphica (Fig. 1). Pairwise distances of the barcoding region (COI) between the target species and the other genera in Neohipparchini show that genetic differentiation is not large in this group, ranging from 7.04% to 10.48% (Table 2). Pairwise distances between Neochloroglyphica and Chloroglyphica range from 7.04% to 7.96%, those between Neochloroglyphica and Neohipparchus range from 9.70% to 10.08%, and those between Neochloroglyphica and Chlororithra range from 9.89% to 10.47%. Tribal association of Neochloroglyphica gen. nov. Neochloroglyphica shares the following characters with other genera (Neohipparchus, Chloroglyphica, Chlororithra) of the Neohipparchini sensu Ban et al. (2018): antennae bipectinate in male and filiform in female; hind tibia of male dilated, with hair-pencil and two pairs of spurs, terminal extension present in Neochloroglyphica, Chlororithra, and some species of Neohipparchus; hind wing with a tail process or protrusion on the end of vein M3; the pair of setal patches on the male third sternite is present; male eighth segment modified in most species (Figs 15–17); coremata present; the aedeagus of Neochloroglyphica bears a pointed process, which is shared with Chloroglyphica (glaucochrista) and Neohipparchus (vallata, maculata, hypoleuca), though the processes are in a different position (Figs 12–14).

100 · Zootaxa 4571 (1) © 2019 Magnolia Press HAN ET AL. * * * * * * * * * * * * * * * * * * * * * * # GAPDH MG014860 MG014923 MG014961 MG014873 MG014908 MG014877 MG014943 MG014869 MG014964 GU829786 MG014897 MG014885 MG014894 MG014931 MG014957 MG014864 MH842270 MH842271 MG014865 missing MG014866 MG014867 MH842272 MH842273 MG014933 MG014932 missing MG014965 MG014966 * * * * * * * * * * * * * * * * * * * * * * + CAD MG015446 MG015507 MG015544 MG015457 MG015493 MG015462 MG015524 MG015454 MG015546 JF785211 MG015484 MG015472 MG015481 MG015514 MG015540 MG015449 MH842274 MH842275 missing MH842276 MG015451 MG015452 missing MH842277 MG015516 MG015515 MG015467 MG015547 MG015548 * * * * * * * * * * * * * * * * * * * * * * #

RPS5 MG015560 MG015621 MG015660 MG015573 MG015608 MG015578 MG015641 MG015569 MG015662 GU830656 MG015597 MG015587 MG015594 MG015629 MG015656 MG015563 MH842266 MH842267 missing missing MG015565 MG015566 MH842268 MH842269 MG015631 MG015630 MG015582 MG015663 MG015664 * * * * * * * * * * * * * * * * * * * * * * * # COI MG014733 MG014800 MG014842 MG014747 MG014785 MG014752 MG014821 MG014743 MG014845 GU828457 MG014774 MG014762 MG014771 MG014808 MG014838 MG014737 MH842261 MH842262 MG014738 MH842263 MG014739 MG014740 MH842264 MH842265 MG014810 MG014809 MG014757 MG014846 MG014847 Collecting locality China Yunnan, China Yunnan, China Hainan, China Guangxi, Sichuan, China China Yunnan, Tibet, China China Shaanxi, China Shaanxi, Finland China Yunnan, China Yunnan, China Yunnan, China Guizhou, China Fujian, Tibet, China Tibet, China Tibet, China China Yunnan, Sichuan, China Tibet, China China Shaanxi, China Yunnan, China Yunnan, China Shaanxi, China Hubei, China Shaanxi, China Hainan, China Gansu, Voucher code M 14087 M 10642 M 11070 M 19115 M 9394 M 22086 M 13773 M 5262 M 11258 MM01146 M 16550 M 10653 M 16556 M 19229 M 10029 M 17085 M 13771 M 17089 M 14493 M 17098 M 20637 M 4939 M 14495 M 14496 M 16577 M 16495 M 4786 M 8133 M 17041 from study the present are from others 2011, al. et Sihvonen + from Mutanen et al. 2010, al. et Mutanen from # Species distinctaria Chlorissa thetydaria Maxates Thalassodes immissaria para baba Herochroma viridata Dooabia taiwana Paramaxates nigromacularia chlorophyllaria papilionaria Geometra symaria Geometra fragilis Loxochila smaragdus Loxochila Mixochlora vittata viridiluteata Tanaorhinus glaucochrista g. Chloroglyphica glaucochrista g. Chloroglyphica glaucochrista g. Chloroglyphica grearia g. Chloroglyphica g.Chloroglyphica grearia Chlororithra fea Chlororithra missioniaria perbella Neochloroglyphica perbella Neochloroglyphica vallata Neohipparchus vallata Neohipparchus difficta Eucyclodes discolor Timandromorpha enervata Timandromorpha from Ban et al. 2018, al. Ban et from TABLE 1. List of specimens and GenBank accession numbers accession and GenBank ofList specimens 1. TABLE Tribe Hemitheini Hemitheini Hemitheini Pseudoterpnini Pseudoterpnini Aracimini Aracimini Comibaenini Comibaenini Geometrini Geometrini Geometrini Geometrini Geometrini Geometrini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Neohipparchini Nemoriini Timandromorphini Timandromorphini *

NEOCHLOROGLYPHICA, A NEW GENUS OF GEOMETRINAE Zootaxa 4571 (1) © 2019 Magnolia Press · 101 0.0000 0.0000 0.0016 0.0016 0.0016 0.0016 0.0532 0.0532 0.0515 0.0515 0.0532 0.0532 0.0514 0.0514 0.04960.0095 0.0778 0.0778 0.07590.0796 0.0777 0.0723 0.0723 0.07040.0741 0.0722 0.0047 0.0824 0.0824 0.08050.0824 0.0805 0.10080.0989 0.0805 0.0805 0.07870.0805 0.0787 0.09890.0970 0.0016 0.0881 0.0881 0.08620.1048 0.1028 0.10470.1009 0.0989 0.0989 0.0957 0.0957 0.09380.0991 0.0972 0.10080.0989 0.0878 0.0878 0.0758 17089 17089 17085 17085 17098 14493 13771 13771 glaucochrista glaucochrista glaucochrista glaucochrista grearia grearia

14495 14495 14496 14496 4939 20637 20637 Pairwise distance COIof the barcoding region based on Kimura-2-parameter model. Chloroglyphica glaucochrista Chloroglyphica glaucochrista Chloroglyphica Chloroglyphica glaucochrista Chloroglyphica Chloroglyphica glaucochrista Chloroglyphica Chloroglyphica glaucochrista Chloroglyphica Neochloroglyphica perbella Neochloroglyphica perbella 16495 16495 vallata Neohipparchus 16577 16577 vallata Neohipparchus Chlororithra missioniaria Chlororithra Chlororithra fea Chlororithra 4 5 7 8 1 2 3 6 9 10 2. TABLE 1 2 3 4 5 6 7 8 9 10 11

102 · Zootaxa 4571 (1) © 2019 Magnolia Press HAN ET AL. In the male genitalia, the costal base of the valva bears a stout hooked process in Neochloroglyphica (Fig. 7), a spinose cluster in Neohipparchus vallata (Fig. 9), a slender process in Chlororithra (Fig. 10), and a stout, nearly triangular process in Chloroglyphica glaucochrista Prout (Fig. 11), which might constitute synapomorphies. Although different from the type species of Chloroglyphica (Fig. 8), Chloroglyphica glaucochrista shares the presence of a small spine or process on the valval ventral margin with Neochloroglyphica.

FIGURE 1. Maximum likelihood tree based on the combined dataset for assessing the close affinity of Neochloroglyphica. Values on the left of nodes indicate bootstrap supports for major clades.

NEOCHLOROGLYPHICA, A NEW GENUS OF GEOMETRINAE Zootaxa 4571 (1) © 2019 Magnolia Press · 103 FIGURE 2. Neighbour-Joining (NJ) tree of selected Chinese Neohipparchini species based on Kimura-2-parameter model.

The well-sclerotized ovipositor lobes of the female genitalia of Neochloroglyphica are smooth, which are different from the oblique and papillate ones in most other Geometrinae (Holloway 1996). The ovipositor lobes are similar to those of members of the Neohipparchini (Chlororithra, Chloroglyphica, Neohipparchus) (Figs 19–23), and most genera of the Geometrini (Geometra Linnaeus, Loxochila Butler, Mixochlora Warren) (Figs 24–26). However, the ovipositor lobes are more modified into an acute, slender conical structure in some species of Neohipparchus and Chloroglyphica (Figs 20, 22). This combination of morphological and molecular data, provides sufficient evidence to place Neochloroglyphica in the tribe Neohipparchini.

Taxonomic accounts

Neochloroglyphica Han & Skou, gen. nov.

Type species. Neochloroglyphica perbella Han & Skou, sp. nov.

Gender feminine. Description. Head. Antennae shortly bipectinate, filiform at apex in male, filiform in female. Frons flattened, width similar to diameter of compound eyes. Labial palpus short, just extending beyond frons. Proboscis well developed. Chaetosemata small, with a few long setae. Vertex rough-scaled. Thorax. Foreleg with well-developed epiphysis; tibial spurs absent. Midleg with one pair of terminal tibial spurs. Hind tibia with two pairs of spurs in both sexes, dilated in male with hair-pencil and a short terminal extension. Wing shape. Apex of forewing blunt, that of hind wing rounded; outer margin of forewing slightly curved, that of hind wing forming a distinct protrusion at end of M3 (Figs 4, 5).

Venation (Fig. 3): Frenulum developed. Forewing: R1 free; R2-5 arising before upper angle of cell; M1 free; base of M2 close to M1, M3 and CuA1 separate; discocellulars deeply incurved between veins M2 and M3. Hind wing:

Sc+R1 close to upper vein of cell at one point near base; Rs and M1 free, M3 and CuA1 separate; discocellulars deeply incurved between veins M2 and M3. Male genitalia (Figs 7, 12). Uncus shortly digitiform, blunt terminally. Socii very long, expanded inwards at middle, narrow at base and tapering posteriorly. Gnathos with lateral arms longer than socii, medial process long,

104 · Zootaxa 4571 (1) © 2019 Magnolia Press HAN ET AL. slender and pointed. Valva with middle part expanded, tip pointed with a tiny tooth; costal base with a stout hooked process, tip spinose; subventral margin strongly sclerotized at middle, appearing as a sclerotized ridge, with three or four tiny teeth posteriorly. Transtilla as a pair of slightly sclerotized processes. Juxta as a rounded sclerite, not developed. Saccus small, protruding. Coremata present, very weak. Aedeagus slender, posterior half sclerotized laterally, with a pointed tooth at tip.

FIGURE 3. Venation of Neochloroglyphica perbella sp. nov.

Female genitalia (Fig. 18). Ovipositor lobes sclerotized and smooth, not papillate. Apophyses posteriores about five times the length of apophyses anteriores. Lamella postvaginalis an irregular sclerotized region. Antrum developed. Ductus bursae short, wrinkled, broadened at middle. Corpus bursae large, rounded; signum a tiny process. Etymology. The generic name references the close relation to the genus Chloroglyphica. Diagnosis. On the wing pattern, Neochloroglyphica is most similar to the genus Neohipparchus Inoue. The transverse lines are much slenderer than in most species of Neohipparchus (such as N. vallata Butler, N. vervactoraria Oberthür, N. verjucodumnaria Oberthür and N. maculata Warren), and lack the brown shading of the antemedial and postmedial lines. Compared to the type species of Neohipparchus, N. vallata Butler, the hind wing fringes of Neochloroglyphica only bear one brown tuft, whereas vallata has two. In the male genitalia,

NEOCHLOROGLYPHICA, A NEW GENUS OF GEOMETRINAE Zootaxa 4571 (1) © 2019 Magnolia Press · 105 Neochloroglyphica is characterized within the Neohipparchini by the presence of a small spine on the tip of the valva. Neochloroglyphica has a small uncus, developed socii, and comparatively simple valvae. Compared to the type species of Neohipparchus (Fig. 9), the uncus of Neochloroglyphica is quite small, but it is as large as the socii in N. vallata; the costa lacks a spinose cluster but bears a hooked process; the ventral margin of the valva bears a

FIGURES 4–10. FIGURES 4–5. Neochloroglyphica perbella gen. nov., sp. nov. 4, male, holotype, ZFMK; 5, female, paratype, IZCAS; 1.5x life size. FIGURE 6. Gongziding, Deqin, Yunnan, one type-locality of Neochloroglyphica perbella. FIGURES 7–10. Male genitalia of the Neohipparchini species. 7, Neochloroglyphica perbella, paratype; 8, Chloroglyphica variegata (Butler); 9, Neohipparchus vallata (Butler); 10, Chlororithra fea Butler. Scale bars = 1 mm.

106 · Zootaxa 4571 (1) © 2019 Magnolia Press HAN ET AL. FIGURES 11–17. Male genitalia, aedeagus and the eighth segment of the Neohipparchini species. 11, Chloroglyphica glaucochrista glaucochrista (Prout); 12, Neochloroglyphica perbella, paratype; 13, Chloroglyphica glaucochrista glaucochrista (Prout), 14, Neohipparchus vallata (Butler); 15, Neochloroglyphica perbella, paratype; 16, Chloroglyphica glaucochrista glaucochrista (Prout); 17, Chlororithra fea Butler. Scale bars =1mm. sclerotized ridge and spines, which are lacking in N. vallata; the lateral arms of the gnathos of Neochloroglyphica are much longer than those of N. vallata; the male eighth tergite has two small hooked lateral processes in Neochloroglyphica, while N. vallata has three developed processes; sternite 8 in the male is deeply concave and V- shaped in Neochloroglyphica, but unmodified in N. vallata. Neochloroglyphica also can be distinguished from Chloroglyphica variegata (Butler) (Fig. 8) and Chlororithra Butler (Fig. 10) by the small uncus, different costal base and ventral margin. Neochloroglyphica shares a modified eighth segment with Chloroglyphica glaucochrista Prout and Chlororithra, but can be differentiated by the following differences: the eighth tergite is shallowly curved in the very broad middle part and bears two lateral processes in Neochloroglyphica (Fig. 15), but two triangular processes are present in Chloroglyphica glaucochrista (Fig. 16) and are curved in a semicircular curve in Chlororithra fea (Fig. 17); the eighth sternite of Neochloroglyphica is less developed than in Chloroglyphica glaucochrista. In the female genitalia, Neochloroglyphica can be differentiated by the presence of the developed antrum.

Neochloroglyphica perbella Han & Skou, sp. nov.

Description. Head. Frons bluish green (light brown after discoloration). Labial palpus pale green, the third segment very short, not elongate in female. Vertex with anterior half white, posterior half bluish green. Thorax. Dorsal side of thorax concolorous with wings, deep bluish green, ventral side whitish. Wings. Forewing length: male 18.5–20 mm; female 18.5 mm. Wings deep bluish green, diffused with white

NEOCHLOROGLYPHICA, A NEW GENUS OF GEOMETRINAE Zootaxa 4571 (1) © 2019 Magnolia Press · 107 striations. Forewing with costa a narrow straw-yellow band, decorated with black spots. Antemedial line white, slender, sinuous, from basal quarter of costa to basal one third of inner margin; postmedial line white, almost straight, slightly bending inwards near costa; submarginal line indistinct, sinuous, whitish. Hind wing with postmedial line straight, slightly tapering towards anal margin; submarginal line white, wavy, more distinct than on forewing. Discal spot deep green on discocellulars. Fringes of both wings with basal half dark brown, terminal half whitish, dark brown on M3 end of hind wing. Underside of wings much paler than upperside, with forewing basal half and hind wing more whitish, streaks on upperside discernible. Abdomen. Dorsal side of abdomen deep bluish green, yellow after discoloration, with a row of white dots at middle. Third sternite of male abdomen with a pair of setal patches. Male eighth segment modified, a broad tergite bearing two short, pointed and curved lateral processes, a narrow sternite deeply concave at middle, V-shaped (Fig. 15).

FIGURES 18–26. Female genitalia of the Neohipparchini species. 18, Neochloroglyphica perbella, paratype, scale bar = 1mm; 19, Chloroglyphica variegata (Butler); 20, Chloroglyphica glaucochrista glaucochrista (Prout); 21, Neohipparchus vallata (Butler); 22, Neohipparchus hypoleuca (Hampson), 23, Chlororithra fea Butler, 24, Mixochlora vittata (Moore), 25. Loxochila kina (Swinhoe), 26. Geometra glaucaria Ménétriés.

108 · Zootaxa 4571 (1) © 2019 Magnolia Press HAN ET AL. Male genitalia (Figs 7, 12) and female genitalia (Fig. 18) are as in description of the genus. Etymology. The specific name is derived from the Latin word “perbellus”, which means very beautiful. Type material. Holotype, ♂ (ZFMK), CHINA: Yunnan: Li-kiang, Provinz Nord-Yuennan, 4000 m, 12.vii.1935, H. Höne. Paratypes: Yunnan: 10♂9♀ (ZFMK), Li-kiang (China), Provinz Nord-Yuennan, H. Höne, 3♂, 03.vi., 11.vi., 20.vi.1934, 3200–3500 m; 1♀, 12.vi.1934, 3200–3500 m; 2♂, 5.vii. (gen. slide 2378-DS), 21.vii.1934, 2400–2800 m; 2♀, 01.vii.1934 (gen. slide 2739-DS), 3200–3500 m; 1♀, 09.vii.1934, 3200–3500 m; 4♂, 14.v., 20.v., 28.v., 29.v.1935, 3200–3500 m; 1♀, 21.v.1935, 3200–3500 m; 1♂, 15.vi.1935, 4000 m; 1♀, 19.vi.1935, 4000 m; 1♀, 22.vi.1935, 2400–2800 m; 1♀, 12.vii.1935, 2400–2800 m; 1♀, A-tun-tse, (Nord Yünnan), mittlere Höhe (ca. 4000 m), 02.viii.1936, H. Höne; 1♂ (IZCAS), Lijiang, Yushuizhai, 2680 m, 21.vi.2009, coll. Yang Chao; 1♂ (IZCAS), Lijiang, Yulongshan, 2800 m, 17.vii.1984, coll. Chen Yixin; 1♂ (IZCAS ex. ZFMK), Li- kiang, Provinz Nord-Yuennan, 2400–2800 m, 20.VI.1935, H. Höne; 3♀ (two in IZCAS, one in PCPS), Deqin, Yunling, Gongziding, 2914 m, 18.vii.2014, coll. Li Xinxin and Pan Xiaodan; 1♂ (IZCAS), Yangbi, 25.v.1983, coll. Long Yongcheng; 1♂ (PCPS), Benzilan (right coast of Jinshajiang river), 2100–2500 m, 8–14.VI.2006; 4♂ (PCPS), Env. Bailakon Pass, 3400–3600 m, 25.V.–7.VI.2006 (two dissected); 1♂ (PCPS), Yuhy (Wuhu) vill., 30 km NW Lijiang, 2700 m, 7.VII.2009, leg. V. Kononenko. Note. In the collection of IZCAS, there are four female specimens collected from the same locality, and unilateral legs of two specimens were taken off and preserved in anhydrous ethanol, numbered as IOZ LEP M 14495, 14496. Sequences were successfully obtained from the two specimens. Only specimen 14495 (Fig. 5) was spread while specimen 14496 remained papered. At present we are unfortunately unable to locate the latter, and therefore do not include it in the type series, though we are certain that it is in somewhere in the Lepidoptera collection of IZCAS. Diagnosis. As in the diagnosis of the genus. Distribution. China (Yunnan).

Acknowledgements

We express our sincere thanks to Li Xinxin and Pan Xiaodan for collecting the fresh specimens of the new species, to Ban Xiaoshuang for carrying out the molecular work, and to Yang Chao for dissecting the new species. We thank Dieter Stüning (ZFMK) for kindly loaning specimens and providing the data of the new species. We are grateful to Anthony Galsworthy for correcting the English and providing valuable comments. We sincerely appreciate Dr David Plotkin (Gainesville-Florida, USA), Dr Shipher Wu (Biodiversity Research Center, Academia Sinica, Taiwan, China) and an anonymous referee for constructive comments on the manuscript. This project was supported by the National Science Foundation of China (No. 31872967, 31672331, 31702041), and by a grant (No. Y229YX5105) from the Key Laboratory of the Zoological Systematics and Evolution of the Chinese Academy of Sciences.

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