Taxonomic review of the genus Mithuna Moore (Lepidoptera: Erebidae: Arctiinae: Lithosiini), with descriptions of two new genera and seventeen new species

Israel Journal of entomology, Vol. 54, pp. 113–165 (14 December 2025) DoI: 10.5281/zenodo.17927846; Issn (online) 2224-6304 received: 22 october 2025 / revised: 10 november 2025 / accepted: 8 December 2025 urn:lsid:zoobank.org:pub:1BC21B27-5ee9-4BDB-9e74-a6aa1DDD10e5 Taxonomic review of the genus Mithuna Moore (Lepidoptera: Erebidae: Arctiinae: Lithosiini), with descriptions of two new genera and seventeen new species Anton V. Volynkin 1 & Karel Černý 2 1190 Ridgemoor Road, Leominster, HR6 8UN, United Kingdom. E-mail: monstruncusar[email protected] 2Tiergartenstrasse 27, Innsbruck, A-6020, Austria. E-mail: [email protected] ABSTRACT the oriental footman-moth genus Mithuna moore, 1878 is reviewed. two new genera and 17 new species are described: Eilemithuna gen. n. (type species: Ilema transducta de Joannis, 1930); Mithunoconosia gen. n. (type species: Mi thuna clivusa Bucsek, 2012); Mithuna armata sp. n. (ne India), Mithuna atkin soni sp. n. (ne India), Mithuna bolovena sp. n. (laos), Mithuna bucseki sp. n. (thailand, laos and Vietnam), Mithuna flabellata sp. n. (thailand), Mithuna fletcheri sp. n. (ne India), Mithuna lamdonga sp. n. (Vietnam), Mithuna meghalaya sp. n. (ne India), Mithuna ochrocephala sp. n. (thailand), Mithuna phahompoka sp. n. (thailand), Mithuna pianma sp. n. (China, yunnan), Mithuna robusta sp. n. (thailand), Mithuna securis sp. n. (Vietnam), Mithuna suthepia sp. n. (thailand), Mithuna swanni sp. n. (myanmar), Mithuna thaica sp. n. (thailand), and Mithuna wilemani sp. n. (taiwan). seven new combinations are introduced: Pseudopelosia fuscivena (Hampson, 1896), comb. n., Mithuna bilineata (Bucsek, 2020), comb. n., Eilemithuna dimidilinea (Černý, 2009), comb. n., Eilemithuna transducta (de Joannis, 1930), comb. n., Mithunoconosia clivusa (Bucsek, 2012), comb. n., Mithunoconosia pulverea (Bucsek, 2012), comb. n. and Mithunoconosia strigifera (Hampson, 1900), comb. n. lectotype is designated for Mithuna arizana Wileman, 1911. adults and male and female genitalia of all species considered are illustrated. KeyWorDs: Biodiversity, Eilemithuna, footman moth, lectotype, lithosiina, Mithunoconosia, oriental region, new combination, Pseudopelosia, south asia, taxonomy. INTRODUCTION the oriental footman-moth genus Mithuna moore, 1878 belongs to the subtribe lithosiina and was erected to solely include Mithuna quadriplaga moore, 1878 from Indian Himalaya. subsequently, another three species were described in the genus: M. fuscivena Hampson, 1896 from sri lanka, M. strigifera Hampson, 1900 from northeast India and M. arizana Wileman, 1911 from taiwan Island. about a century later, seven more species were included in the genus: M. quadriplagoides Holloway, 2001 from Borneo Island; M. dimidilinea Černý, 2009 from Thailand; M. clivusa Bucsek, 2012, M. flavia Bucsek, 2012, M. jendeki Bucsek, 2012 and M. pulverea Bucsek, 2012 from Western malaysia and M. tranthiedui Dubatolov 114 Israel Journal of entomology, Vol. 54, 2025 & Bucsek, 2016 from Vietnam (Holloway 2001; Černý & Pinratana 2009; Bucsek 2012; Dubatolov & Bucsek 2016), of which M. jendeki was transferred to the genus Euconosia Watson, 1980, two years after the description was published (Bucsek 2014). additionally, five Mithuna-looking species were described in the genus Teulisna Walker, 1862 (Černý & Pinratana 2009; Dubatolov & Bucsek 2016; Kirti & singh 2016; Bucsek 2020) and were recently assigned to Mithuna by Volynkin et al. (2025). further studies of the unidentified Mithuna materials deposited in various collections led the authors of the present paper to the discovery of additional 17 unknown species, which are described in the present paper as species new to science. the known and newly discovered species of Mithuna can be subdivided into nine groups. after the detailed examination of their genitalia structures and external morphology, it became clear that two of those groups display fundamental mor phological differences from others in Mithuna and represent distinct lineages, for which two new genera are erected in the present paper. In addition, ‘Mithuna’ fuscivena Hampson, 1896 was found to be non-congeneric with the type species of Mithuna and the newly described genera and is therefore transferred to the genus Pseudopelosia Krüger, 2015 in the present paper. MATERIALS AND METHODS acronyms of the depositories used: BmnH – see nHmuK; CaV – research collection of anton Volynkin (leominster, uK); CKC – research collection of Karel Černý (Innsbruck, Austria); mfn – museum für naturkunde, Berlin, germany; MNHN – Muséum national d’Histoire naturelle, Paris, France; mWm/Zsm – museum Witt münchen / Zoologische staatssammlung münchen, munich, germany; nHmuK – natural History museum, london, uK; oumnH – oxford university museum of natural History, oxford, uK; snm – slovak national museum, Bratislava, slovakia; sZmn – siberian Zoological museum of the Institute of systematics and ecology of animals, novosibirsk, russia. abbreviations used: aV – genitalia slide prepared by a. Volynkin, Ht – holotype, LT – lectotype, PT – paratype. the genitalia were dissected applying standard methods of preparation (lafontaine & mikkola 1987; Kononenko 2010), then stained with eosin y or Chlorazol Black and embedded in euparal on microscope slides, or preserved in glycerol in microvials pinned under the specimens. the photos of adults were taken using a nikon D3100/af-s or nikon D7500 camera equipped with a nikkor 18–55 mm or sigma 105 mm f2.8 eX Dg macro os lens while the photos of genitalia were taken using the aforementioned cameras attached to a microscope with an lm- VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 115 scope adapter. All pictures were processed using the Adobe Photoshop CC 2018 software. the male and female genitalia terminology follows Volynkin (2024). for the holotype label citations, information provided in quotation marks is transcribed verbatim. Different labels are separated by a slash (‘/’) while the different lines of the same label are separated by a vertical bar (‘|’). any additional data are provided in square brackets. the content of the labels of additional specimens examined including paratypes is edited or translated in accordance with english gram mar and standardised. In the species and genus accounts, the abbreviations ‘sp. n.’ and ‘gen. n.’ are omitted to make the text easier for reading. Images of specimens deposited in the nHmuK are used with permission and are copyright of the trustees of the natural History museum, london and made available under the CC 4.0 CC-By license (https://creativecommons.org/licenses/ by/4.0). TAXONOMY order lepidoptera linnaeus, 1758 family erebidae leach, 1815 subfamily arctiinae leach, 1815 tribe lithosiini Billberg, 1820 subtribe lithosiina Billberg, 1820 genus Mithuna moore, 1878 Mithuna moore, 1878: 21. (type species: Mithuna quadriplaga moore, 1878, by monotypy.) Diagnosis: species of the genus are quite uniform externally and characterised by the brown colouration and fuscous and diffuse forewing pattern consisting of a medial line, a small discal spot and a subterminal line interrupted into irregular patches. the male genitalia, although diverse in the valva shape, are characterised by the combination of the following features: (1) the juxta is plate-like with apicolateral plates wrapping phallus laterally and (in certain groups) dorsally; the juxta structure is considered herein as characteristic of the genus; (2) the costa bears a small processus momenti; (3) the valval section of the transtilla is fold-shaped and membranous (in certain species weakly sclerotised); the editum and tendon are absent; and (4) the dorsal section of the valvula is well-sclerotised and in certain species extended into an apical process. In the female genitalia, the genus is characterised by the short ductus bursae, which is sclerotised in most species, the long utricular or saccate corpus bursae and the short conical or semiglobular appendix bursae situated postero-laterally or postero-ventrally. Redescription: sexual dimorphism limited. antenna setose ciliate in both sexes with shorter cilia in females. Body and forewing ground colour from ochreous brown to deep brown or reddish brown. Costal margin of forewing convex postmedially; anal margin convex antemedially; outer margin slightly convex; apex triangular or rounded; tornus rounded. forewing markings fuscous, diffuse; pattern 116 Israel Journal of entomology, Vol. 54, 2025 consisting of medial transverse line, discal spot (reduced in many species) and subterminal line. latter interrupted into irregular patches. Cilia fuscous. Hindwing pale ochreous, in many species suffused with greyish brown. Male genitalia. uncus proximally swollen and distally tapered; in certain species with tiny claw-shaped tip. scaphium from setose to plateor string-like. subscaphium setose. arms of tegumen moderately broad and sclerotised. Vinculum as long as scaphium or longer, with narrow but well-sclerotised arms, ventrally rec tangular or u-shaped, with short saccate intravincular corema sparsely covered with short hairs. Valva diverse in shape, from narrow and strongly elongate with almost parallel margins to trapezoidal or almost elliptical. Basis valvae narrow, in certain groups with jugum distalis originating from proximal dorsal corner of valva and directed ventro-distad. Costa narrow, moderately sclerotised, stretching along the dorsal margin of valva or short and occupying only its basal section; bearing short process in certain species; short processus momenti present. Valval section of transtilla elongate fold-like, stretching along ventral margin of costa, membranous or weakly sclerotised; tendon absent. Valvula lobular or distally tapered, with secondary sclerotisation, densely setose or bearing sclerotised apical pro cess in certain species. lamella centralis reduced but in certain species present as weakly sclerotised transverse area. sacculus diverse in length with distal process of various shapes, or without it in certain groups. Juxta broad, moderately sclerotised with membranous medio-ventral section; dorsal section with apicolateral processes wrapping phallus laterally and (in certain groups) dorsally and bearing cluster of denticles, or clawor bird wing-shaped process in certain species. Phallus cylindrical, distally dilated in certain species; in most species with string-like longitudinal lateral sclerotised fold. Vesica diverse in shape, with semi globular, conical or utricular diverticula; in certain groups bearing cornuti, ser rulate plates or clusters of denticles or areas of graniculi. Vesica ejaculatorius without elasma. Female genitalia. Ovipositor short, broad conical. Papilla analis trapezoidal with rounded corners, setose. Pseudopapilla present, short, triangular and setose. Glan dula present. apophyses rod-like, well-sclerotised. 8th sternite with postvaginal plate; in certain species additional sclerotisations present lateral and posterior to the postvaginal plate. Ductus bursae diverse in structure: tubular or vase-shaped, sclerotised or membranous, rugose or smooth; in certain species with anterioventral sclerotised pocket. Corpus bursae elongate or saccate, membranous or with sclerotised and rugose posterior section; in certain species with scobination. appendix bursae short, semiglobular or conical, membranous, gelatinous or weakly sclerotised, situated postero-laterally or postero-ventrally. 7th sternite moderately sclerotised, in certain species bearing subostial rugose sclerotisation or small lateral gelatinous pockets. Distribution: the genus is widespread in the oriental realm from Himalaya to the Pacific coast, reaching Malay Peninsula and Borneo Island in the south. VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 117 Notes: Besides the taxa placed in Mithuna and the new genera Eilemithuna gen. n. and Mithunoconosia gen. n. in the present paper, Mithuna fuscivena Hampson, 1896 was originally described in Mithuna and has been kept in the genus since (Holloway 2001; Bucsek 2012). However, Holloway (2001) discussed the substantial morphological differences of fuscivena from the typical Mithuna and noted that “they are probably not congeneric”. the examination of the genitalia structures of fuscivena (illustrated by Holloway 2001: fig. 39) led the authors of the present paper to the conclusion that it is morphologically close to the genus Pseudopelosia Krüger, 2015 (illustrated by Krüger 2015: figs 91, 92, 248, 345), to which the species is transferred herein: Pseudopelosia fuscivena (Hampson, 1896), comb. n. Species content of Mithuna: the genus Mithuna in its current concept comprises more than two dozens of externally similar species with diverse male and female genitalia structures, which are herein subdivided into seven species-groups: the M. tongdzuythanhi species-group M. flava (Dubatolov & Bucsek, 2016) M. securis sp. n. M. thaica sp. n. M. tongdzuythanhi (Dubatolov & Bucsek, 2016) the M. arizana species-group M. arizana Wileman, 1911 M. bilineata (Bucsek, 2020), comb. n. M. bolovena sp. n. M. ochrocephala sp. n. M. tenebrosa (n. singh & Kirti, 2016) M. wilemani sp. n. the M. quadriplaga species-group M. armata sp. n. M. atkinsoni sp. n. M. fletcheri sp. n. M. meghalaya sp. n. M. phahompoka sp. n. M. pianma sp. n. M. quadriplaga moore, 1878 M. robusta sp. n. M. swanni sp. n. the M. tranthiedui species-group M. flabellata sp. n. M. lamdonga sp. n. M. tranthiedui Dubatolov & Bucsek, 2016 the M. quadriplagoides species-group M. quadriplagoides Holloway, 2001 118 Israel Journal of entomology, Vol. 54, 2025 the M. mithunoides species-group M. bucseki sp. n. M. mithunoides (Černý, 2009) the M. varia species-group M. suthepia sp. n. M. varia (Bucsek, 2020) Incertae sedis ‘Mithuna’ flavia Bucsek, 2012 Mithuna tongdzuythanhi species-group Diagnosis: the male genitalia of the species-group are characterised by the heavily sclerotised and upcurved distal saccular process, the distally dilated phallus with a protruding lateral crest, and the vesica armed with a thorn-shaped subapical cornutus. In the female genitalia, the appendix bursae is well-sclerotised. Mithuna tongdzuythanhi (Dubatolov & Bucsek, 2016) figs 1–4, 69, 105 Teulisna tongdzuythanhi Dubatolov & Bucsek, 2016: 229, figs 2, 3, 21. (type locality: “Central Vietnam, Vinh Phuc Prov., Tam Dao, 21°27’N 105°38.37’E, 900 m”.) Diagnosis: the forewing length is 10.0–11.0 mm in males and 11.0–12.0 mm in females. Mithuna tongdzuythanhi is externally reminiscent of M. thaica and M. bilineata, but distinguished by the somewhat larger size and paler hindwing. reliable identification requires the examination of the genitalia structures, which are clearly different in all three species. the genitalia structures of M. tongdzuythanhi are similar to M. flava, from which the former differs externally in the somewhat larger size, the more elongate forewing, the darker, brown forewing ground colour (it is brownish ochreous in M. flava) and the medial line more strongly curved in the cell. unlike in M. flava, the male genital capsule of M. tongdzuythanhi has a thinner uncus, a longer juxta, longer and narrower valvae with shorter and less upcurved distal saccular processes. the phallus of M. tongdzuythanhi is longer and narrower than in M. flava. the vesica of M. tongdzuythanhi can be readily distinguished from M. flava by the narrower and shorter main chamber, the longer, narrower and more heavily granulose largest diverticulum and the shorter and thinner subapical cornutus. the female genitalia of M. tongdzuythanhi differ from M. flava in the longer and narrower ductus bursae, the shorter and ovoid corpus bursae (it is tubular posteriorly and globular anteriorly in M. flava) and the longer, utricular appendix bursae, which is conical in M. flava. Holotype (examined from photographs): Vietnam: ♂ “26–28.iii.2012 | Central Vietnam, | Vinh Phuc Prov., | Tam Dao, 21° 27’ N | 105° 38.37’E, 900 m, | V.Zolotuhin leg.”, gen. prep. by V. Dubatolov (preserved in sugar on a piece of paper pinned under the specimen) (sZmn). Additional material examined: Vietnam: 4♂ 8♀, Tam Dao, 60 km NW Hanoi, 21°34'N 105°20'E, 950 m, 17.x.1994, V. Sinyaev, gen. slide No. ZSM Arct. 20191105♂ (MWM/ZSM); 1♂ 1♀, same VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 119 Figs 1–18. adults of Mithuna spp. Depositories of the specimens: 1, 2, 10 and 17 in CKC; 3–5 in mfn; 6–9, 11, 12, 14–16 and 18 in mWm/Zsm (9, 16 and 18: ex CKC); 13 in nHmuK. 120 Israel Journal of entomology, Vol. 54, 2025 locality as previous but 1200 m, 1–15.xi.1992, Sinyaev & Simonov (MWM/ZSM); 1♂ 1♀, Tam Dao, 950 m, 11.x.1984, K. Spitzer, gen. slide Nos: AV7838♂, AV7839♀ (CKC); 1♂ 1♀, same data as previous but 16.x.1984, gen. slide Nos: AV7840♂, AV7841♀ (CKC); 1♂ 1♀, Tam Dao, 50 km N Hanoi, 900 m, 23.iii.1995, W. Mey, gen. slide Nos: NKMBAV010♂, NKMBAV011♀ (MfN); 1♂, Mt Fansipan (North), Chapa, primary forest, 22°17"N 103°44"E, 1600 m, 25–30.iii.1995, V. sinyaev & a. schintlmeister (mWm/Zsm). Thailand: Chiang Mai Prov.: 1♂, Mt Doi Phahompok, 10 km W of mae ai, 1500 m, 26.ii.1998, m. Hreblay & Cs. szabóky, gen. slide no. Zsm arct. 20191104♂ (MWM/ZSM); 1♂, Chae Son NP, 18°51'33"N 91°22'03"E, 1496 m, 9.vi.2005, K. Černý, gen. slide No. AV7827♂ (CKC); 1♂, Doi Pha Hom Pok, 20°02'54"N 99°09'49"E, 1400 m, 28–29.xi.2005, K. Černý, gen. slide No. AV7828♂ (CKC); 1♀, Fang, Doi Pha Hom Pok, 20°06'16"N 99°07'46"E, 2110 m, 23–24.v.2011, K. Černý, gen. slide No. AV7833♀ (CKC); 1♂ 1♀, Doi Inthanon NP, 18°30'59"N 98°28'13"E, 1416 m, 6–7.vi.2005, K. Černý, gen. slide Nos. AV7829♂, AV7830♀ (CKC); 1♀, Fang Distr., Doi Ang Khang, 29°54'11"N 99°02'32"E, 1425 m, 23–27.vii.2005, T. Ihle, gen. slide Nos. AV7834♀ (CKC); 1♂ 1♀,4 km SE of Pang Faen, 1100 m, 14.xi.1999, M. Hreblay (MWM/ZSM); 1♂ 1♀, 4 km SE of Pang Faen, 1100 m, 14.xi.1999, M. Hreblay (MWM/ZSM); 1♂ same locality as previous, 18.i.2004, P. Hentcshel & A. Szabó (MWM/ZSM). Nan Prov.: 1♂, 30 km E Pua, 1700 m, 10.xi.1999, M. Hreblay (MWM/ZSM). Mae Hong Son Prov.: 1♂, 21 km NW of Pai, 1360 m, 7.ii.1998, M. Hreblay & Cs. Szabóky (MWM/ZSM). India: 1♂, Khasis. Nat. Coll. / Doncaster Private coll., Purch. 1927, gen. slide No. AV8780♂ (OUMNH). Distribution: the species is currently known from northern Vietnam (lao Cai and Tuyen Quang Provinces) (Dubatolov & Bucsek 2016), Northern Thailand (Mae Hong Son, Chiang Mai and Nan Provinces) and Northeast India (Meghalaya) (pre sent study). Mithuna flava (Dubatolov & Bucsek, 2016) figs 5, 6, 70, 106 Teulisna flava Dubatolov & Bucsek, 2016: 230, figs 4, 22 (type locality: “north Vietnam, thanh Hoa Prov., Thuong Xuan Distr., Xuan Lien NR [National Reserve], 19°52’N 105°14.28’E, 130 m”). Diagnosis. the forewing length is 9.0–9.5 mm in males and 11.0 mm in females. Mithuna flava is morphologically similar to M. tongdzuythanhi and the detailed comparison is provided above in the diagnosis of the latter species. Holotype (examined from photographs): Vietnam: ♂ “29–30.xi.2012 | North Vietnam, Thanh Hoa | Prov., Thuong Xuan Distr., | Xuan Lien NR, anthropog. | 19°52’N 105°14,28’E, 130 m | V.Zolotuhin leg.”, gen. prep. by V. Dubatolov (preserved in sugar on a piece of paper pinned under the specimen) (sZmn). Additional material examined: Vietnam: 1♂, Tam Dao, 50 km N Hanoi, 23.iii.1995, 900 m, W. Mey (MfN); 2♂ 1♀, Mt Fansipan (North), Chapa, primary forest, 22°17'N 103°44'E, 1600 m, 25–30.iii.1995, V. Sinyaev & A. Schintlmeister, gen. slide Nos ZSM Arct. 2021494♂, 2021495♀, 2021660♂ (AV) (MWM/ZSM); 2♂ 1♀, BachMa Nat. Park, 16°10'N 107°54'E, 1200 m, 26.vii– 6.viii.1996, V. Sinyaev & E. Afonin, gen. slide Nos ZSM Arct. 2021499♂, 2021500♂, 2021501♀ (aV) (mWm/Zsm). Distribution. the species is currently known from Vietnam (thanh Hoa, lao Cai and Tuyen Quang Provinces) (Dubatolov & Bucsek 2016). Mithuna thaica sp. n. figs 7, 8, 71, 72, 107 LSID: urn:lsid:zoobank.org:act:C96958eB-25D9-4391-B494-442eaC06eCBf. VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 121 Figs 19–33. adults of Mithuna spp. Depositories of the specimens: 19 and 20 in CaV; 21 in oumnH; 22, 26, 29–31 and 33 in nHmuK; 23, 24 and 28 in mWm/Zsm; 25 and 28 in CKC; 32 in mfn. 128 Israel Journal of entomology, Vol. 54, 2025 provided above in the diagnosis of the latter species. Compared with the sympatric M. arizana, M. wilemani is markedly smaller and has a shorter forewing with a less convex anal margin and more diffuse medial line. the male genital capsule of M. wilemani can be easily distinguished from M. arizana in the distally narrower and straighter uncus, the shorter vinculum, the proximally narrower valva with a medially convex dorsal margin forming a collis, the densely setose apical section of the valva and the less upcurved distal saccular process. the phallus of M. wilemani is considerably narrower and shorter than in M. arizana. In the vesica, the main chamber of M. wilemani is shorter and narrower, the largest (distal) diverticulum is markedly shorter and narrower and the distal section is shorter and narrower than the corresponding structures of M. arizana. Female. unknown. Holotype: Taiwan: ♂, “Taiwan, Taoyuan Co. | Tungyenshan Recr[eation] Ar[ea] | ca. 24°50’N / 121°24’E, | 900 m, 21.–24.ix.2002 | leg. W. Schacht et al.” / “Slide | AV7846♂ | A. Volynkin” (MWM/ Zsm, ex CKC) (figs 18, 79). Paratypes: Taiwan: 1♂, Kuantaochi, Nantou [County], Formosa, 9.viii.1972, S. Yamane / Inoue Coll. B.M.1 992–71, NHMUK010292626 (NHMUK); 1♂, Wushe, Nantou [County], Formosa, 27.vii.1973, y. shibata / Inoue Coll. B.m.1 992–71, nHmuK010292627 (nHmuK). Distribution: the new species is endemic to taiwan Island. Note: It is likely that this species was first described as Mithuna arizana ab. parva Wileman, 1911, which is an unavailable name (ICZn 1999: art. 10.2, 45.5 and 45.6.2). However, as the primary types of ab. parva remain undissected, it is impossible to check this assumption. Mithuna bolovena sp. n. figs 19, 20, 80, 112 LSID:urn:lsid:zoobank.org:act:8C0849C9-DfCD-495e-83aC-9D96f017CB37. Etymology: The species name is derived from the Boloven Plateau, at the western end of which the new species has been found. the name is a noun in apposition. Diagnosis: the forewing length is 7.5 mm in the male holotype and 9.0 mm in the female paratype. Mithuna bolovena is externally reminiscent of M. bilineata but differs in the smaller size, the paler, more ochreous forewing ground colour and the somewhat more elongate forewing apex. the male genital capsule of M. bolovena is similar to M. wilemani but distinguished by the proximally broader uncus, the broader and more sparsely setose valvula bearing a tiny sclerotised thorn apically (it is apically rounded in M. wilemani) and the broader, more upcurved and apically pointed distal saccular process, which is apically rounded in M. wilemani. the phalli of the two species are very similar. In the vesica, M. bo lovena differs from M. wilemani in the narrower main chamber, the unilobate and curved distal diverticulum (it is straight and bilobate in the congener) and the some what narrower apical diverticulum. as the female of M. wilemani is unknown, the female genitalia of M. bolovena were compared with M. tenebrosa, from which VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 129 the new species can be distinguished by the more heavily sclerotised apophyses, the shorter and more heavily sclerotised 8th abdominal segment, the markedly shorter intersegmental membrane between the 7th and 8th abdominal segments and the narrower corpus bursae with a smaller appendix bursae. Holotype: Laos: ♂ “10–20.viii.2014, | South Laos, | Champasak prov., | 27 km ENE of Pakse, | near Tad Fane waterfall, | 15°11’01.1”N 106°07’40.5”E, | 950 m, local collector leg. | Coll. A.V. Volynkin” / “Slide | AV8792♂ | A. Volynkin” (MWM/ZSM, ex CAV) (Figs 19, 80). Paratypes: Laos: 2♀, same data as in holotype, gen. slide No. AV8793♀ (CAV). Distribution: the new species is currently known only from its type locality in southern Laos (Champasak Province). Mithuna tenebrosa (n. singh & Kirti, 2016) figs 21–25, 81–84, 113 Teulisna tenebrosus n. singh & Kirti in Kirti & singh, 2016: 163. (type locality: “assam, Jatinga” [NE India].) Diagnosis: the forewing length is 9.0–11.0 mm in males and 11.0–11.5 mm in females. Mithuna tenebrosa is externally reminiscent of M. tongdzuythanhi but differs in the forewing having a less elongate apex and an indistinct medial line. the male genitalia of M. tenebrosa are most similar to the taiwanese M. wilemani but differ clearly in the strongly asymmetrical valvae, of which the left one is extremely elongate (ca. twice as long as the right one) and nearly straight while the right valva is medially somewhat upcurved. additionally, the uncus of M. tenebrosa is proximally broader than in M. wilemani, the apical section of the valva is narrower and weaklier setose and the distal saccular process is thinner and stick-shaped whereas it is thumb-shaped in M. wilemani. the phallus of M. tenebrosa is slightly broader than in M. wilemani. the vesica configurations of the two species are similar but unlike in M. wilemani, the largest (distal) diverticulum of M. tenebrosa has larger lobes and the apical section has an additional lateral diverticulum. as the female of M. wilemani is unknown, the female genitalia of M. tenebrosa were compared with the most similar M. arizana instead, and M. tenebrosa differs from the latter in the longer anterior apophysis, the shorter ductus bursae with a heavily sclerotised cup-shaped posterior section and the narrower corpus bursae. the intersegmental membrane between the 7th and 8th abdominal segments of M. tenebrosa is the longest in the genus. Material examined: Bhutan: 1♂, Bhutan, Dudgeon, 95203, 2500’, 17.viii.[18]95 (NHMUK). India: 1♂, W[est] B[engal], Darjeeling, 5 km oberh. [above] Rambi, 800 m, 30.vii.1990, W. Thomas, gen. slide No. ZSM Arct. 2021491♂ (AV) (MWM/ZSM); 1♂, [Meghalaya] Khasis, x.1894, Nat. Coll., gen. slide No. AV8779♂ (OUMNH); 2♂ 1♀, [Meghalaya] Khasis, Nat. Coll. / Doncaster private coll., Purch. 1927, gen. slide No. AV8782♀ (OUMNH). Nepal: 1♂ 1♀, Milke Danda, Nesum, 1500 m, 2.viii.2000, Csővari & Hreblay, gen. slide Nos: ZSM Arct. 2021485♂, 2021486♀ (AV) (MWM/ ZSM); 2♂ 2♀, Annapurna Himal, 1 km N of Besisahar, 28°14’N 84°23’E, 850 m, 5.vi.1996, Hreblay & Szaboky, gen. slide Nos: ZSM Arct. 2021487♂, 2021488♀, 2021489♂, 2021490♀ (AV) (MWM/ Zsm). Thailand: Chiang Mai Prov.: 1♂, Fang Distr., Doi Pha Hom Pok, 20°02'54"N 99°09'49"E, 1400 m, 28–29.xi.2005, K. Černý, gen. slide No. AV7844♂ (CKC); 1♀, Doi Inthanon NP, 18°30'59"N 130 Israel Journal of entomology, Vol. 54, 2025 98°28'13"E, 1416 m, 6–7.vi.2005, K. Černý, gen. slide No. AV9284♀ (CKC); 1♂ Mt Doi Inthanon NP, 2300 m, 28.i.2000, M. Hreblay & A. Szabó (MWM/ZSM); 1♂ 4 km SE of Pang Faen, 1100 m, 14.xi.1999, M. Hreblay (MWM/ZSM); 1♂ 1♀, Doi Suthep, 18°47.974'N 98°54.901'E, 1087 m, at mV light, 1.viii.2006, g. martin, BmnH(e) 2006-128, nHmuK010916156, nHmuK010916157 (nHmuK). Distribution: the species is currently known from northeast India (north of West Bengal, meghalaya) (Kirti & singh 2016), nepal (Kishida 1995, as M. quad riplaga), Bhutan and Northern Thailand (Chiang Mai Province) (present study). Notes: (1) Both generic names, Teulisna (the original combination of the species) and Mithuna are feminine. the species name is a latin adjective and its original spelling ‘tenebrosus’ has a masculine ending, which is against article 31.2 of ICZn (1999). although provisions of this article seem to be widely ignored by the lepidopterological community (e.g., sommerer 2002; Welter-schultes 2012; nieukerken et al. 2019), the ending of the species name is herein corrected to the feminine gender with the authorship of the taxon being preserved, to comply with articles 33.2 and 34.2 of ICZn (1999). (2) the species was described with the male genitalia having a left valva broken off (although the illustrated specimen had it), which made the main diagnostic character of the male genitalia being not mentioned the original description (Kirti & singh 2016). (3) the species varies in its size and the shade of the forewing ground colour but the genitalia structures are rather invariable throughout its distributional range, only the length of the left valva somewhat varies in the thai populations (figs 83, 84). Mithuna quadriplaga species-group Diagnosis: the male genitalia of the species-group are characterised by the welldeveloped distal saccular process and the flattened juxta with a weakly sclerotised medio-ventral section and well-sclerotised apico-lateral lobes bearing sclerotised plates in most species. the female genitalia have a heavily sclerotised and rugose ductus bursae with sclerotised margins of the ostium, a strongly elongate, narrow and membranous corpus bursae and a membranous appendix bursae. Mithuna quadriplaga moore, 1878 figs 26–28, 85, 114 Mithuna quadriplaga Moore, 1878: 21, pl. s: fig. 9. (Type locality: “Darjiling” [NE India, West Bengal, Darjeeling].) Diagnosis: the forewing length is 10.5–11.0 mm in males and 12.5–13.0 mm in females. Mithuna quadriplaga is morphologically similar to the allopatric M. swanni and the detailed comparison is provided below in the diagnosis of the latter species. Lectotype (designated by Hampson (1900) as “type”): India: (Fig. 26): ♀, “Darjiling | 21” / “Moore Coll. | 94–106.” / red ring “type” label / Qr-code label with unique ID “nHmuK010401697” (nHmuK). VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 131 Figs 69–72. male genitalia of Mithuna spp. Depositories of the dissected specimens: 69 and 72 in CKC; 70 and 71 in mWm/Zsm. 132 Israel Journal of entomology, Vol. 54, 2025 Additional material examined: India: 1♂, W[est] B[engal], Darjeeling, Manjitar, 700 m, 20.vii.1990, W. Thomas, gen. slide No. ZSM Arct. 20191102♂ (AV) (MWM/ZSM); 1♂ 1♀, Sikkim, Pe ma yangtse, 2000 m, 23–28.vii.1990, W. Thomas, gen. slide Nos: AV7847♂, AV7848♀ (CKC); 1♀, [Meghalaya] Khasis, Nat. Coll. / Doncaster private coll., Purch. 1927, gen. slide No. AV8781♀ (oumnH). Nepal: 1♂ 1♀, Annapurna Himal, Ulleri, 1900 m, 28°23'N 83°43'E, 3.x.1994, Csorba & Ronkay, gen. slide Nos: ZSM Arct. 2021481♂ 2021482♀ (AV) (MWM/ZSM); 1♂, Langtang, 9 km S Dhunche, 28°04'N 85°14'E, 2110 m, 23.ix.1994, Csorba & Ronkay, gen. slide No. ZSM Arct. 2021483♂ (AV) (MWM/ZSM); 1♂ 1♀, Langtang, 1.5 km NE Dhunche, 28°06’N 85°18’E, 1950 m, 24.ix.1994, Csorba & Ronkay, gen. slide No. ZSM Arct. 2021484♀ (AV) (MWM/ZSM). Distribution: the species is known from nepal and northeast India (sikkim, north of West Bengal and meghalaya) (moore 1878). Notes: (1) moore (1878) described the species from an unspecified number of specimens preserved in his and o. staudinger’s collections (currently housed in nHmuK and mfn respectively). subsequently, Hampson (1900) cited only one female from moore’s collection as “type”, which, according to the article 74.5 of ICZn (1999), should be considered as lectotype designation. (2) unfortunately, only a single female from Khasi Hills was available for this study. this specimen has female genitalia very similar to the Himalayan populations and is therefore considered herein as belonging to M. quadriplaga. Mithuna swanni sp. n. figs 29–31, 86, 115 LSID: urn:lsid:zoobank.org:act:CB4f786e-3093-44e3-91e8-D54661D350CC. Etymology: the species is named after Captain arthur e. swann, who collected not only the type series of the new species but numerous other Heterocera in northern myanmar in 1921–1923. Diagnosis: the forewing length is 10.5–11.0 mm in males and 12.0 mm in the female. Mithuna swanni is externally very similar to M. quadriplaga and M. pianma and identification requires the examination of the genitalia structures. the male genitalia of M. swanni are similar to M. quadriplaga, but distinguished by the broader uncus, the more heavily setose scaphium, the smaller claw-shaped processes of the apico-lateral plates of the juxta and the longer and unilobate diverticulum of the vesica, which is bilobate in M. quadriplaga. In the female genitalia, M. swanni differs from M. quadriplaga in the broader ostium bursae, the shorter ductus bursae and the longer corpus and appendix bursae. Holotype: Myanmar: ♂, [North Myanmar, Kachin State, Myitkyina District] “Upper Burma: | Htawgaw. | 6000 ft. | Capt. a.e. swann” / “Brit. mus. | 1923–488.” / Qr-code label “nHmuK010292605” (nHmuK) (figs 29, 86). Paratypes: Myanmar: 2♂ 1♀, same data as in holotype, NHMUK010292606, NHMUK010292607, nHmuK010292610 (nHmuK). Distribution: the new species is currently known only from its type locality in northern myanmar (eastern Kachin state). VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 133 Figs 73–76. male genitalia of Mithuna spp. Depositories of the dissected specimens: 73 and 75 in mWm/Zsm (73: ex CKC); 74 in snm (after Bucsek 2020); 76 in CKC. 134 Israel Journal of entomology, Vol. 54, 2025 Mithuna atkinsoni sp. n. figs 32, 33, 87, 116 LSID: urn:lsid:zoobank.org:act:0B0710a9-0a8a-455B-8890-ffeB731C96DD. Etymology: the new species is named after William stephen atkinson (1820– 1876), the British lepidopterist, whose collection of Indian lepidoptera served as the basis for the description of numerous new taxa including the current one. Diagnosis: the forewing length is 14.5–15.0 mm in males and 16.5 mm in the female. Mithuna atkinsoni is the largest species in the genus. It is most externally similar to M. meghalaya, from which M. atkinsoni differs in the more elongate forewing and the medial line angled in the cell whereas it is nearly straight in M. meghalaya. the male genital capsule of M. atkinsoni is most reminiscent of M. pianma with its setose apex of the valva but can be easily distinguished by the narrower uncus, broader u-shaped sacculus (it is rectangular in M. pianma), the more strongly upcurved valva with a more densely setose apex and the broader juxta with larger sclerotised apico-lateral plates. unlike in M. pianma, the vesica of M. atkinsoni has a shorter but broader basal section with a dorsal cluster consisting of larger denticle-like cornuti, a markedly broader ventral diverticulum covered with graniculi and bearing a cluster of denticle-like cornuti laterally (it is situated apically in M. pianma), the presence of a utricular distal diverticulum and the apical section having a short bilobate diverticulum. the female genitalia of M. atkinsoni differ from M. pianma in the trapezoidal postvaginal plate (it is elliptical and connected to additional areas of sclerotisation laterally and posteriorly), the more rugose ductus bursae lacking the anterior ventral pocket and the narrower corpus bursae. Holotype: India: ♂, [Northeast India] “Darjeeling | 1864” / “coll. Atkinson” / “Coll. | Staudinger” / “Slide | NKMBAV012♂ | A. Volynkin” (MfN) (Figs 32, 87). Paratypes: India: 1♂ 1♀, Sikkim, Möller, 1887 / Collectio H.J. Elwes / Rothschild Bequest B.M. 1939-1, nHmuK010292452, nHmuK010292608 (nHmuK). Distribution: the new species is known from northeast India (north of West Bengal and sikkim). Mithuna pianma sp. n. figs 34, 35, 88, 117 LSID: urn:lsid:zoobank.org:act:e3eaf528-42a6-4B80-82Ba-C50C5264B520. Etymology: The species name is derived from the town of Pianma, the type lo ca lity of the new species. the name is a noun in apposition. Diagnosis: the forewing length is 11.0 mm in the male holotype and 12.5 mm in females. Mithuna pianma is externally very similar to M. quadriplaga and M. swanni and identification requires the examination of the genitalia structures. In the male genital capsule, M. pianma differs from the aforementioned congeners in the VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 135 Figs 77–80. male genitalia of Mithuna spp. Depositories of the dissected specimens: 77 and 79 in mWm/Zsm, ex CKC; 78 in CKC; 80 in CaV. 136 Israel Journal of entomology, Vol. 54, 2025 more heavily sclerotised scaphium, the narrower vinculum, the medially narrower valva with a broader, apically rounded and densely setose apex (it is apically pointed and weakly setose in M. quadriplaga and M. swanni), the more upcurved distal saccular process and the smaller apico-lateral plates of the juxta lacking processes. the phallus of M. pianma is somewhat shorter than in M. quadriplaga and M. swanni. In the vesica of M. pianma, the diverticulum bears an apical cluster of denticle-like cornuti, which is absent in both congeners. additionally, the diver ticulum of M. pianma is unilobate whereas it is bilobate in M. quadriplaga. the female genitalia of M. pianma differ from M. quadriplaga and M. swanni in the less rugose and markedly narrower ductus bursae bearing an anterior ventral pocket and the posteriorly broader corpus bursae. the detailed comparison with M. atkinsoni is provided above in the diagnosis of the latter species. Holotype: China: ♂, [China, Yunnan Prov., Nujiang Lisu Autonomous Prefecture, Lushui City, Pianma Town] “Upper Burma: | Hpimaw Fort, | Nr. Myitkyina. | 8000 ft. 13.viii.1923. | Capt. A.E. swann.” / “Brit. mus. | 1923–488.” / Qr-code label “nHmuK010292603” (nHmuK) (figs 34, 88). Paratypes: China: 2♀, same data as in holotype, NHMUK010292604, NHMUK010292609 (nHmuK). Distribution: the new species is currently known only from its type locality in Yunnan Province of China, near the border with Kachin State of Myanmar. Mithuna phahompoka sp. n. figs 36, 37, 89, 118 LSID: urn:lsid:zoobank.org:act:e2284eDB-2f86-4eeD-8eaa-040f726921ea. Etymology: The species name is derived from the Mount (Doi) Phahompok, where the new species is found. the name is a noun in apposition. Diagnosis: the forewing length is 11.5 mm in the male holotype and 12.5 mm in females. Mithuna phahompoka is externally similar to M. quadriplaga, M. swanni, M. pianma and M. fletcheri, but can be distinguished by the nearly straight medial line of the forewing, which is angled in the cell in the similar congeners. the male and female genitalia of M. phahompoka are most similar to M. fletcheri and the detailed comparison is provided below in the diagnosis of the latter species. the vesica configuration of M. phahompoka is also very similar to M. pianma having, however, a largely different male genital capsule, from which that of M. phahompoka differs in the broader uncus, the longer and broader juxta, the shorter vinculum, the markedly shorter and broader valva with a proximally broader dorsal section having an upcurved, more tapered and smooth distal part (it is rather trapezoidal and densely setose in M. pianma) and the broader sacculus with a considerably longer, hook-shaped distal process, which is blade-shaped and smoothly upcurved in M. pianma. the phallus of M. phahompoka is longer and somewhat broader than in M. pianma. In the vesica of M. phahompoka, the proximal cluster of cornuti is shorter than in M. pianma, the largest diverticulum is somewhat broader and the apical section has a small semiglobular diverticulum, VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 137 Figs 81–84. male genitalia of Mithuna tenebrosa. Depositories of the dissected specimens: 81 and 82 in mWm/Zsm; 83 in nHmuK; 84 in CKC. 144 Israel Journal of entomology, Vol. 54, 2025 Paratypes: Thailand: 1♂, same data as in holotype, gen. slide No. AV9280♂ (CKC); 3♂, Chiang Mai Prov., Chae Son NP, 18°51'33"N 91°22'03"E, 1496 m, 9.vi.2005, K. Černý, gen. slide Nos: AV7821♂ AV9282♂, AV9283♂ (CKC). Distribution: the new species is known from northern thailand (Chiang mai Province). Mithuna tranthiedui species-group Diagnosis: the male genitalia of the group are characterised by the pyramidal juxta (similar to the M. quadriplagoides species-group, whereas in other congeners it is flattened), the distally dilated valva with very short apical and distal saccular processes and the broad vesica bearing robust cornuti. the female genitalia of the M. tranthiedui species-group are similar to the M. tongdzuythanhi species-group with its sclerotised posterior end of the corpus bursae. However, in the M. tranthiedui species-group the corpus bursae is more saccate than in the M. tongdzuythanhi species-group and its posterior area of sclerotisation is longer and broader. Mithuna lamdonga sp. n. figs 45, 46, 95, 121 LSID: urn:lsid:zoobank.org:act:a267D24D-e3e6-4f6D-966D-05C8963ae2e8. Etymology: The species name is derived from Lam Dong Province of Vietnam, where the new species is found. the name is a noun in apposition. Diagnosis: the forewing length is 9.5 mm in both sexes. Mithuna lamdonga is externally similar to M. robusta and M. flabellata but can be easily distinguished by the smaller size, the narrower forewing and the paler, greyish ochreous ground colour of the forewing, which is ochreous brown in M. robusta and brown in M. flabellata. the male genital capsule of M. lamdonga is reminiscent of M. tranthiedui but is distinct in the longer uncus and the markedly longer and broader valva with broader and triangular apexes of the dorsal and saccular sections (they both are apically rounded in M. tranthiedui). the vesica configuration of M. tranthiedui remains unstudied but it was possible to compare the cornuti of the two species: M. lamdonga has two robust thorn-shaped cornuti and one serrulate plate whereas in M. tranthiedui, there are two serrulate plates and two large claw-shaped cornuti originating from the same heavily sclerotised base. as the female of M. tranthiedui remains unknown, the female genitalia of M. lamdonga were compared with M. flabellata (see below under the diagnosis of the latter species). Holotype: Vietnam: ♂, “S Vietnam | Đa Lat, Datan La 1380 m, | 11°54’10,6”N 108°26’59”E, | 29.ix.2013 leg. K. Černý” / “Slide | AV7837♂ | A. Volynkin” (MWM/ZSM, ex CKC) (Figs 45, 95). Paratype: Vietnam: ♀, same data as in holotype, gen. slide No. AV9286♀ (CKC). Distribution: the new species is currently known only from its type locality in southern Vietnam. VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 145 Figs 97–100. male genitalia of Mithuna spp. Depositories of the dissected specimens: 97 in sZmn (photo by V. Dubatolov); 98 in mfn; 99 in CKC; 100 in mWm/Zsm. 146 Israel Journal of entomology, Vol. 54, 2025 Mithuna flabellata sp. n. figs 47, 48, 96, 122 LSID: urn:lsid:zoobank.org:act:0920ae08-fBBf-4a38-8457-2C71a9C2417C. Etymology: the species name is a latin adjective meaning ‘fan-shaped’ and refers to the fan-shaped valva of the new species. Diagnosis: the forewing length is 10.0–10.5 mm in males and 10.0 mm in the female. Mithuna flabellata is externally reminiscent of M. robusta but can be dis tinguished by the somewhat smaller size and darker, brown forewing ground colour, which is ochreous brown in the congener. reliable identification often requires the examination of the genitalia structures, which are substantially diffe rent in the two species in their uncus, valva and vesica configurations. the male genital capsule of M. flabellata is most reminiscent of M. lamdonga but differs clearly in the markedly broader and longer valva with a sacculus having a medial ventral protrusion and bearing a broader distal protrusion-like process. the phallus of M. flabellata is proximally broader than in M. lamdonga. unlike in M. lamdonga, the vesica of M. flabellata has a lateral subbasal (proximal) diver ticulum displaced more distally and bearing a cornutus (vs. a serrulate plate in the congener), a broader and bilobate distal (largest) diverticulum bearing a shorter and more curved cornutus and a shorter apical section lacking the cornutus. In the female genitalia, M. flabellata is distinct from M. lamdonga in the longer post vaginal plate, the broader and more heavily sclerotised ductus bursae lacking the anterior ventro-lateral pocket, which is present in M. lamdonga, the anteriorly broader corpus bursae with a sclerotised postero-lateral area having a lateral protru sion and the more heavily sclerotised and conical appendix bursae, which is se mi globular in the similar congener. additionally, the anterior section of the corpus bursae of M. flabellata bears weak graniculi whereas that of M. lamdonga is membranous. Holotype: Thailand: ♂, “N. Thailand | Chiang Maiprov, | Chae Son NP, 1496 m | 18°51’22”N 91°22’3’’E | 9.vi.2005 leg. K. Cerný [recte: Černý]” / “Slide | AV7820♂ | A. Volynkin” (MWM/ZSM, ex CKC) (figs 47, 96). Paratypes: Thailand: 1♀, same data as in holotype, gen. slide No. AV7822♀ (CKC); 1♂, Chiang Mai Prov., Doi Inthanon NP, 18°30'59"N 98°28'13"E, 1416 m, 6–7.vi.2005, K. Černý, gen. slide No. AV7823♂ (CKC). Distribution: the new species is known from northern thailand (Chiang mai Province). Mithuna tranthiedui Dubatolov & Bucsek, 2016 figs 49, 97 Mithuna tranthiedui Dubatolov & Bucsek, 2016: 230, figs 6, 24. (type locality: “Central Vietnam, Gia Lai Prov., K’Bang Distr., Dak Roong Comm., vill[age]. Kon Loc, Kon Ka Kinh NP [National Park], 1050 m, 14°42.602’ N 108°39.062’ E”.) VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 147 Figs 101–104. male genitalia of Mithuna and Eilemithuna gen. n. spp. Depositories of the dissected specimens: 101 in snm (after Bucsek (2020)); 102 in nHmuK; 103 in mWm/Zsm; 104 in CKC. 148 Israel Journal of entomology, Vol. 54, 2025 Diagnosis: the forewing length is 10.5 mm in the male holotype. the species was described from a single worn specimen reminiscent of M. flabellata, from which M. tranthiedui differs in the darker hindwing. the male genitalia of M. tranthiedui are similar to M. lamdonga and the detailed comparison is provided above in the diagnosis of the latter species. Female. unknown. Holotype (examined only on photographs): Vietnam: ♂, “14–19.III.2012 | Central Vietnam, Gia Lai Prov | K’Bang Distr., Dak Roong | Comm., vill. Kon Loc, Kon Ka | Kinh NP, 14°42.602’ N 108° | 39.062’e, 1050 m | V.Zolotuhin leg.”, gen. prep. by V. Dubatolov (preserved in sugar on a piece of paper pinned under the specimen) (sZmn) (figs 49, 97). Distribution: the species is currently known only from its type locality in Central Vietnam (Gia Lai Province) (Dubatolov & Bucsek 2016). Mithuna quadriplagoides species-group Diagnosis: see below under the only species included in the group, M. quadriplagoides. Mithuna quadriplagoides Holloway, 2001 figs 50, 51, 98, 123 Mithuna quadriplagoides Holloway, 2001: 304, pl. 2: fig. 7 (moth), fig. 74 (♂ genitalia). (Type locality: [Malaysia, Borneo Island] “Sarawak: Gunong Mulu Nat. Park … Mulu, 1790 m”.) Diagnosis: the forewing length is 11.0–11.5 mm in males and 11.5 mm in females. Mithuna quadriplagoides is externally reminiscent of M. meghalaya but is somewhat smaller and has a more rounded forewing apex and a darker hindwing suffused with greyish brown. the male genital capsule of M. quadriplagoides is distinct from other congeners in the solid, nearly elliptical valva lacking processes but with a shallow subapical depression ventrally and the apico-lateral plates of the juxta modified into long bird wing-shaped processes bearing clusters of robust spines distally. the structure of the valva is similar to the M. mithunoides speciesgroup with its medial transverse area of sclerotisation ventrally connected with sacculus, which is possibly a weakly sclerotised lamella centralis. the phallus of the species is s-like curved medially and has a heavily sclerotised lateral stringlike fold, which is similar to the M. quadriplaga species-group. the vesica is broad, almost globular with a broad area of graniculi, which is unique within the genus. In the female genitalia, M. quadriplagoides differs from its congeners in the sclerotised, dorso-ventrally flattened and smooth ductus bursae (it is rugose in the M. quadriplaga species-group) and the strongly elongate and narrow corpus bursae, which is reminiscent of the M. quadriplaga species-group but has a weakly sclerotised posterior and a weakly granulose anterior sections, whereas the corpus bursae is membranous in the M. quadriplaga species-group. Holotype: Malaysia: ♂, [Borneo Island] “Sarawak: Gunong | Mulu Nat. Park | R.G.S. Exped. 1977–8 | J.D. Holloway et al. | B.m. 1978–206” / “site 1. January | Camp 4, mulu | 1790 m. 452463 | lower VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 149 Figs 105–108. female genitalia of Mithuna spp. Depositories of the dissected specimens: 105 in mfn; 106 and 107 in mWm/Zsm; 108 in CKC. 150 Israel Journal of entomology, Vol. 54, 2025 montane (moss) | forest. mV – canopy.” / “Holotype | Mithuna | quadriplagoides | Holloway” / round label with red ring “Holo | type” / blue label “Arctiidae | genitalia slide | No. 4708♂” / QRcode label “nHmuK 010916196” (nHmuK) (fig. 50). Paratype: Malaysia: ♀, [Borneo Island] Sarawak: Gunong Mulu Nat. Park, R.G.S. Exped. 1977–8, J.D. Holloway et al., B.m. 1978–206 / site 2, January, Camp 4, mulu, 1790 m, 452463, lower montane (moss) forest. Act[inic light]–understorey / Paratype Mithuna quadriplagoides Holloway / Paratype [round label with yellow ring], NHMUK010292611 (NHMUK). Additional material examined: Malaysia: 1♂, Borneo, Mt Kinabalu, Mesilau, 2000 m, 14–17. xi.2006, LF [light trap], W. Mey & K. Ebert leg. / West Mesilau River, Suspension Bridge (MfN); 1♂, Borneo, Sabah, Kinabalu NP, HQ, 4–6.xi.2003, LF [light trap], W. Mey, gen. slide No. NKMB AV009♂ (MfN). Distribution: the species is endemic to Borneo Island (Holloway 2001). Mithuna mithunoides species-group Diagnosis: the male genitalia of the species-group are characterised by the combination of the following features: (1) the juxta is broad, flattened and has small apico-lateral plates bearing spurs; (2) the costa is short, s-shaped and lacks a jugum distalis; (3) the distal section of the valva is formed by the secondary sclerotised valvula; (4) the sacculus is short and lacks processes, distally fused with a weak transverse sclerotisation presumably representing a lamella centralis; and (5) the vesica lacks the distal (largest) diverticulum and has an enlarged apical section having a large bilobate diverticulum bearing a single robust cornutus. In the female genitalia, the following features are characteristic of the speciesgroup: (1) the ductus bursae is broad, dorso-ventrally flattened, well-sclerotised with postero-lateral heavily sclerotised plates; (2) the posterior section of the corpus bursae has one or two broad areas of sclerotisation and a longitudinal membranous fold ventrally; and (3) the anterior section of the corpus bursae is densely scobinate. Mithuna mithunoides (Černý, 2009) figs 52, 53, 99, 124 Teulisna mithunoides Černý in Černý & Pinratana, 2009: 135, pl. 27, fig. 271. (Type locality: “SW Thailand, Chumphon, Pa Toh, Ban Lang Tang, 162 m, 9°46’5”N 98°46’59’’E”.) Diagnosis: the forewing length is 7.5–8.0 mm in males and 8.5 mm in females. Mithuna mithunoides is morphologically similar to M. bucseki and the detailed comparison is provided below in the diagnosis of the latter species. Holotype: Thailand: ♀, “SW Thailand | Chumphonprov., Pa Toh distr., | Ban Lang Tang, 162 m | 9°46’5”N 98°46’59’’ E | 19., 21.xi.2006 leg. K. Cerný [recte: Černý]” / red label “Holotype | Teulisna | mithunoides | Karel Černý, 2009” / “Slide | AV8073♀ | A. Volynkin” (MWM/ZSM, ex CKC) (Figs 53, 124). Paratypes: 1♂ 1♀, SW Thailand, Ranong Prov., Ranong, 10°01'32"N 98°40'13"E, 380 m, 3–4. xii.2005, K. Černý, gen. slide Nos AV8070♂, AV8071♀ (CKC). Additional material examined: Cambodia: 1♂, Mondolkiri Prov., Seima Biodiversity Conservation Area, between Seima and O’Rang, 12°15'44"N 107°03'49"E, 360 m, 27–29.i.2006, G. Csorba & G. Ronkay, gen. slide No. ZSM Arct. 2021505♂ (MWM/ZSM). Thailand: 1♀, Saraburi [Nakhon Nayok Prov.], Khao Yai NP, Mt Khao Kaew [Khao Khiao], 1244 m, 14°21'56"N 101°24'[00"]E, 31.v.2005, K. Černý, gen. slide No. AV7831♀ (CKC). VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 151 Figs 109–113. female genitalia of Mithuna spp. Depositories of the dissected specimens: 109 and 111 in CKC; 110 and 113 in mWm/Zsm; 112 in CaV. 152 Israel Journal of entomology, Vol. 54, 2025 Distribution: the species is known from southern and Central thailand (Chumphon, Ranong and Nakhon Nayok Provinces) (Černý & Pinratana 2009), Western Malaysia (Perak) (Bucsek 2016) and Cambodia (Bae et al. 2016). the record from Central laos (Bucsek 2020) belongs to M. bucseki (see below). Mithuna bucseki sp. n. figs 54, 55, 100, 125 LSID: urn:lsid:zoobank.org:act:1423Cf26-ee57-4a83-a198-BD666C5314D5. Etymology: the new species is named after Karol Bucsek, renowned expert in the asian arctiinae, who greatly contributed to the knowledge of the lithosiini of southeast asia. Diagnosis: the forewing length is 8.5–9.0 mm in males and 9.0 mm in females. Mithuna bucseki is externally indistinguishable from M. mithunoides and iden tification requires the examination of the genitalia structures. the male genital capsule of the new species differs clearly from M. mithunoides in the shorter uncus, the dorsally narrower juxta bearing smaller spurs, the broader valva having a longer distal dorsal section with a falcate tip and the ventral process directed ventrad (it is directed ventro-distad in M. mithunoides) and the ventrally protruding distal sec tion of the sacculus. the phallus of M. bucseki is distally narrower than in M. mithunoides. In the vesica, M. bucseki is distinguished from M. mithunoides by the smaller but more heavily granulose subbasal diverticulum, the narrower main chamber (medial section) having two short diverticula (absent in M. mithunoides) and the somewhat narrower apical diverticulum bearing a smaller cornutus. the female genitalia of M. bucseki differ from those of M. mithunoides in the anteriorly narrower ductus bursae with markedly larger postero-lateral sclerotised plates and the posterior section of the corpus bursae having two broad sclerotised areas separated ventrally by the longitudinal membranous fold, whereas M. mithunoides has one sclerotised area on the left side with a broader longitudinal membranous fold edging it on the right side. additionally, the anterior section of the corpus bursae of M. bucseki is less posteriorly constricted than in M. mithunoides and bears markedly finer scobination. Holotype: Thailand: ♂, “Thailand | Changwat [Province] Chiang Mai | 4 km SE of Pang Faen | 1100 m, 14.xi.1999 | leg. Márton Hreblay” / “Slide | ZSM Arct. | 2021502♂ | A. Volynkin” (MWM/ Zsm) (figs 54, 100). Paratypes: Laos: 72♂ 79♀, 10–20.viii.2014, Champasak Prov., 27 km ENE of Pakse, near Tad Fane waterfall, 15°11'01.1"N 106°07'40.5"E, 950 m, local collector, gen. slide Nos: AV7343♂, AV8787♀ (CaV). Thailand: 1♀, same data as in holotype, gen. slide No. ZSM Arct. 2021503♀ (AV) (MWM/ ZSM); 1♂, Chiang Mai Prov., Doi Suthep, 18°47.974'N 98°54.901'E, 1087 m, 1.viii.2006, G. Martin [leg.], at MV light, BMNH(E) 2006128, NHMUK010916155 (NHMUK). Vietnam: 1♂, Bao Loc (sec. forest), Rung Cat Tien, 11°32'N 107°48'E, 1500 m, 10–20.xii.1992, Sinyaev & Simonov, gen. slide No. ZSM Arct. 2021504♂ (AV) (MWM/ZSM). Distribution: the new species is currently known from northern thailand (Chiang Mai Province), Central and Southern Laos (Khammouane and Champasak Pro VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 153 Figs 114–119. female genitalia of Mithuna spp. Depositories of the dissected specimens: 114 in mWm/Zsm; 115–117 and 119 in nHmuK; 118 in CKC. 160 Israel Journal of entomology, Vol. 54, 2025 in the flat and rugose postvaginal plate (it is medially swollen in E. dimidilinea), the markedly narrower and shorter gelatinous antrum-like posterior section of the ductus bursae, the considerably broader and ventrally swollen posterior sclerotised end of the corpus bursae, the narrower corpus bursae bearing finer scobination and the smaller appendix bursae. In addition, the 7th abdominal sternite of E. transducta is anteriorly sclerotised and has lateral gelatinous pockets, which are absent in E. dimidilinea. Holotype (by monotypy): examined from the photograph deposited in mnHn (https://www.mnhn. fr/en/databases). Material examined: China: 1♂, [Yunnan Prov., Nujiang Lisu Autonomous Prefecture, Lushui City, Pianma Town] Upper Burma, Hpimaw Fort, nr Myitkyina, 8000 ft., 13.viii.1923, Capt. A.E. Swann [leg.], “Brit. Mus. 1923–488”, NHMUK010292602 (NHMUK). Myanmar: 1♂, Upper Burma, Htawgaw, 6000 ft., Capt. A.E. Swann [leg.], NHMUK010292601 (NHMUK). Vietnam: 1♂, Mt Fan sipan, near Chapa, 22°20'N 103°40'E, 16[00]–1800 m, secondary forest, v.1995, local collectors, ex coll. Schintlmeister, gen. slide No. ZSM Arct. 20191106♂ (AV) (MWM/ZSM); 1♂ 1♀, Mt Fansi pan (North), Chapa, primary forest, 22°17'N 103°44'E, 1600 m, 20–30.iv.1995, V. Sinyaev & local collector, gen. slide No. ZSM Arct. 2021496♀ (AV) (MWM/ZSM). Distribution: The species is known from Northern Vietnam (Lao Cai Province) (de Joannis 1930), northern myanmar (eastern Kachin state) and southwest China (northwestern Yunnan Province) (present study). Eilemithuna dimidilinea (Černý, 2009), comb. n. figs 63–65, 104, 130 Mithuna dimidilinea Černý in Černý & Pinratana, 2009: 151, pl. 31: fig. 306. (Type locality: “N Thailand, Chiang Mai, Fang, Doi Pha Hom Pok, 2050 m, 20°07’30”N 99°08’49’’E”.) Diagnosis: the forewing length is 9.5–10 mm in males and 11.0 mm in females. the detailed comparison with E. transducta is provided above in the diagnosis of the latter species. Holotype (examined): Thailand: ♂, “N. Thailand | Chiang Maiprov., Fang distr. | Doi Pha Hom Pok, 2050m | 20°07’30”N 99°08’49’’ E | 5–6.v.2006 leg. K. Cerný [recte: Černý]” / red label “Holotype | Mithuna | dimidilinea | Karel Černý 2009” / QRcode label “NHMUK 010889679” (NHMUK) (Fig. 63). Paratypes: 19♂ 10♀, same data as in holotype, gen. slide No. AV7835♂ (CKC); 1♂ 2♀, same locality and collector as previous but 21.iv.2006 (CKC). Additional material examined: Thailand: 1♀, same data as in holotype but 23–24.v.2011, gen. slide No. AV7836♀ (CKC). Distribution: the species is currently known only from its type locality in northern Thailand (Chiang Mai Province) (Černý & Pinratana 2009). genus Mithunoconosia gen. n. LSID: urn:lsid:zoobank.org:act:ea185aDe-270C-444C-aD48-D428064f0f19. Type species: Mithuna clivusa Bucsek, 2012, by present designation. Etymology: the generic name is a combination of the genus-group names Mithuna and Euconosia. the gender is feminine. VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 161 Diagnosis: unlike in Eilemithuna gen. n., the male of Mithunoconosia has setose ciliate antenna (similar to Mithuna whereas it is serrulate setose ciliate in Eilemithuna), a narrower and distally trapezoidal forewing with a cluster of andro conia in the folded cell (it has more rounded corners and a flat cell lacking an droconia in the similar genera) and an irregular forewing pattern consisting of areas of brown and blackish suffusion and blackish spots in the cell, at the costa and the terminal line interrupted into irregular dashes. In the male hindwing, the cell and the area along the costa are pale ochreous yellow, whereas in Mithuna and Eilemithuna the hindwing is more unicolorous. species of the new genus are most externally reminiscent of Euconosia but can be distinguished by the smaller size, the shorter antenna and the distally narrower forewing; the male and female genitalia of the two genera are fundamentally different in many aspects (for Euconosia, see Holloway (2001)). the male genital capsule of Mithunoconosia differs from both Mithuna and Eilemithuna in the following features: (1) the vinculum has a corema densely covered with long flattened scales whereas the corema of Mithuna is sparsely covered with weak hairs and, in Eilemithuna gen. n. the corema is absent; (2) the juxta bears a robust process directed dorsad, whereas in Mithuna the juxta is flat and has apico-lateral plates and in Eilemithuna gen. n. the juxta consists of two narrow stripes and is apically fused with the proximal section of the sacculus by the annelifer; (3) Ventrally, the juxta is extended into a heavily sclerotised caulis, which terminates with two weakly sclerotised and rugose ligulose processes supporting the phallus ventrally (a feature unique for the genus); (4) the lamella centralis is broad, heavily sclerotised and ventrally strongly articulated with the base of the distal saccular process; and (5) the valvula is large, lobular and weakly sclerotised (similar to Eilemithuna ) whereas in Mithuna the valvula has a heavy secondary sclerotisation. In the vesica, Mithunoconosia has areas of robust spinulose scobination, which are absent in Mithuna and Eilemithuna the female geni talia of Mithunoconosia are characterised by the combination of the following features: (1) the ductus bursae is broad, well-sclerotised and has a flattened dorsal wall while its ventral wall is swollen, pyramidally protruding ventrad and has a deep V-shaped notch in the ventral margin of the ostium bursae; (2) the posterior section of the corpus bursae is weakly sclerotised rugose and scobinate; (3) the anterior section of the corpus bursae bears a longitudinal ribbon-shaped signum, which is absent in Mithuna and Eilemithuna; and (4) the appendix bursae is reduced and the ductus seminalis originates from the anterior part of the rugose posterior section of the corpus bursae dorsally. additionally, the 7th abdominal sternite of Mithunoconosia bears large lateral gelatinous pockets (absent or small in Mithuna and Eilemithuna) and has a u-shaped ventral fold anteriorly of the ostium bursae; the pockets likely receive the distal saccular processes during copulation while the latter fold possibly serves as a structure receiving the process of the juxta. Description: Adults (figs 67, 68). sexual dimorphism limited: female with broader forewing than in male. antenna setose ciliate in both sexes. Body ochreous brown; 162 Israel Journal of entomology, Vol. 54, 2025 male abdomen fuscous medially and pale ochreous distally. forewing elongate, narrow and distally trapezoidal. forewing ground colour ochreous or ochreous brown. In male, cell with longitudinal fold invaginated ventrad and bearing elongate cluster of androconial scales. forewing pattern brown or fuscous, consisting of intense suffusion along veins and in postmedial area and irregular spots in the cell and at costal margin. terminal line thin and interrupted. Hindwing grey brown with ochreous terminal line; in male area along costal margin and in cell ochreous, with elongate narrow subbasal cluster of hair-shaped scales stretching along vein Cu. Male genitalia (fig. 131). uncus slender, apically pointed. tuba analis membra nous. tegumen short with moderately broad and weakly sclerotised arms. Vinculum equal to or somewhat longer than tegumen, more heavily sclerotised and u-shaped, with medial saccate corema bearing elongate and flattened scales. Valva broad, lobular. Basis valvae narrow but heavily sclerotised, with short jugum distalis dorsally. Costa broad and moderately sclerotised, occupying proximal half or two-thirds of the dorsal margin of valva. Valvula broad, membranous and weakly setose. Both, diaphragmal and valval sections of transtilla membranous. lamella centralis broad, well-sclerotised, smooth, ventrally fused with base of distal saccular process. sacculus heavily sclerotised, with large upcurved distal process having elongate setose area along its dorsal side and bearing robust spikeshaped tip. Juxta trapezoidal with membranous medial ventral section, bearing Figs 131, 132. Mithunoconosia clivusa (type species of Mithunoconosia gen. n.), male (131) and female (132) genitalia. the dissected specimens are deposited in mWm/Zsm. VOLYNKIN & ČERNÝ: TAXONOMIC REVIEW OF MITHUNA 163 medial heavily sclerotised process directed dorsad. Caulis heavily sclerotised, occupying whole ventral wall of anellus, apically bearing two weakly sclerotised and rugose tongueshaped processes supporting the phallus ventrally. Phallus cy lindrical, almost straight, with short conical and apically rounded coecum. Vesica with several conical or saccate diverticula bearing clusters of graniculi, spinulose scobination and short spine-like cornuti. Female genitalia (Fig. 132). Ovipositor narrowconical. Papilla analis trapezo idal with rounded corners, weakly setose. apophyses elongate and thin, anterior one shorter than posterior one. Ductus bursae broad, well-sclerotised, with flattened dorsal and swollen and ventrally protruding ventral walls; ventral margin of ostium bursae with deep V-shaped notch. Corpus bursae subdivided into two sections by medial constriction. Posterior section weakly sclerotised, rugose and scobinate. anterior section of corpus bursae membranous, ovoid and bearing longitudinal ribbon-shaped signum. appendix bursae reduced; ductus seminalis originating from anterior part of posterior section of corpus bursae dorsally. 7th abdominal ster nite with large lateral gelatinous pockets and u-shaped ventral fold anteriorly of ostium bursae. Distribution: the genus is distributed from northeast India to Indochina reaching Malay Peninsula in the south. Species included: at the moment, the new genus comprises three valid species: M. clivusa (Bucsek, 2012), comb. n., M. pulverea (Bucsek, 2012), comb. n. and M. strigifera (Hampson, 1900), comb. n. CONCLUSIONS as a result of the present review, the genus Mithuna has been found to be surprisingly diverse. the number of known species in the genus has increased sig nificantly—from 10 known species, of which four are transferred to other genera in the present paper—to 28, including 17 newly described species. of the new species, 12 are described from the mountains of northern Indochina, which is a well-known biodiversity hotspot. In terms of the number of species and major lineages of Mithuna, this area is most diverse suggesting that it may be the centre of origin of the genus. the results of the present study highlight the limitations of our knowledge of the true biodiversity of the tropical lichen moths, and it is likely that further new species of the genus Mithuna await discovery. ACKNOWLEDGEMENTS the senior author expresses his sincere thanks to the following colleagues for their kind assistance during visits to the collections under their care: axel Hausmann, mei-yu Chen and ulf Buchsbaum (Zsm); James Hogan (oumnH); alberto Zilli and geoff martin (nHmuK); and Wolfram mey (mfn). Vladimir Dubatolov (novosibirsk, russia) is acknowledged for the pictures of the holotype of Mithuna tranthiedui. the authors are also indebted to scott miller (national museum of natural History, Washington, DC, usa) and an anonymous reviewer for their valuable comments to the manuscript. 164 Israel Journal of entomology, Vol. 54, 2025 REFERENCES Bae, Y.S., Bayarsaikhan, U. & Kimsun, C. 2016. Biodiversity of Cambodia. Tiger Moths (Insecta, Lepidoptera). national Institute of Biological resources, Incheon. 303 pp. Bucsek, K. 2012. Erebidae, Arctiinae (Lithosiini, Arctiini) of Malaya Peninsula-Malaysia. Institute of Zoology sas, Bratislava. 170 pp. Bucsek, K. 2014. 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