©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at Atalanta (December 1999) 30(1/4): 87-117, colour plates VI, VII, Wurzburg, ISSN 0171-0079 A review of the East-Palaearctic taxa of the Melitaea didyma (Esper, 1779)-group 1. The M. ala Staudinger, 1881-M. chitralensis Moore, 1901-complex (Lepidoptera, Nymphalidae) by K. A. Kolesnichenko received 14.X.1999 Summary: The M. ala-M. chitralensis taxa complex Is distinguished on the base of the mor­ phology of the male genitalia. The specific status of M. enarea Fruhstorfer and M. kotshubeji Sheuuzhko is confirmed. Data on morphology, variation and distribution of the species of the complex in question are summarized. Three new subspecies are described: M. kotshubeji bun- deli, M. kotshubeji kugarti and M. enarea gromenkoi. The distinctive features of the complex of taxa comprising various forms of the Central Asian species M. ala Staudinger, 1881 - M. citralensis Moore, 1901 consist in the structure of the male genitalia (the ringwall1 is produced into a projection). Females of this complex have a more or less divided antevaginal plate (in M. didyma Esp. and similar species it is undivided). M. ala was described by Staudinger (1881) from Ala-Tau Mts. as a variety of M. didyma and was regarded either under that status (Alpheraky, 1881; Ruhl, 1895; Staudinger, 1901) or as a subspecies of M. didyma (Seitz, 1908; Bramson, 1910) up to the beginning of our century. The specific rank of M. ala was pointed out by Sushkin (1913) on the base of the male genital ar­ mature. The form bicolor Seitz (1908), described as an aberration of M. didyma, was attrib­ uted to M. ala also by this author. Sheuuzhko (1929) was the first who tried to make a review of the geographical variation of M. ala. Besides the typical form, he recognized ssp. bicolor (Inner Tian-Shan, Alexander Mts.), ssp. kotshubeji Sheuuzhko, 1929 (Peter the Great Mts.), included latonia Grumm-Grshimailo (a form of didymoides Eversmann which does not belong to the complex in question) and two aberrations: latemarginata Sheuuzhko, 1929 and immod- ulata Sheuuzhko, 1929. Later, the taxa kotshubeji and bicolor were also considered as subspe­ cies of M. ala (Bryk, 1940). During the following years a number of forms similar to this species was described by different authors: ninae Sheuuzhko, 1935 (with several forms), rosea Hig­ gins, 1938 (described as a separate species), allah Bryk, 1940, sheljuzhkoi Bryk, 1940, determinate Bryk, 1940, strandi Bryk, 1940, ella-claudia Bryk, 1940. The status of the taxon enarea Fruhstorfer, 1916 (Garm), which was described and consid­ ered later (Bryk, 1940) as a subspecies of M. didyma or as a subspecies of M. chitralensis (Hig­ gins, 1941), was unclear. At the same time Sheuuzhko (1929) described (also as a subspecies of M. didyma) two forms, similar to enarea: shugnana and ishkashima from the South-West Pamirs. He pointed out that the last one was very similar to M. chitralensis and females of shugnana were similar to females of M. ala. 1 The term ringwall (Ventralplatten—in Sushkin, 1913) was introduced by Zander (1903) for the designa­ tion of a chitinized plate (IX sternite) connecting valvae and saccus. Later, this term was accepted by the researchers of the genus Melitaea (Higgins, 1941). 87 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at M. chitralensis was described as a separate species and is known only from Chitral. We exam­ ined a small series (3 dd, 3 $9) of this species in the Zoological Institute of the Russian Acad­ emy of Sciences (St. Petersburg). These specimens are very similar externally to the photo of the holotype of M. chitralensis which is at our disposal (the type series is in the Natural History Museum, London), and the structure of the genital armature of the male corresponds to the male genitalia figured by Higgins (1941). In Bryk's review (1940) dedicated to the M. didyma-group only M. didyma and M. ala had specific rank and part of the taxa (chitralensis, ishkashima, shugnana, ninae, rosea) were not regarded at all. The revision of the genus Melitaea was published by Higgins (1941). He estab­ lished the synonymy and distinguished three species within the complex under present consid­ eration: 1. M. ala with ssp. bicolor, 2. M. chitralensis with ssp. enarea (= shugnana, = ish­ kashima) (however pointing out the differences in the structure of the male genitalia of the last one), modifications kotshubeji and permuta Higgins, 1941 (Hazret-Sultan Mts.), and 3. M. pseudoala Sheuuzhko, 1928 (= ninae, = rosea)2 on the base of external features and morphology of the male genitalia. The forms described by Bryk (1940) were dealt with by Higgins in 1955. He believed ella-clau- dia to be a synonym of pseudoala and classified the form sheljuzhkoi as M. ala bicolor. He con­ sidered the males of allah and strandi not to be correctly placed with M. ala and not to belong to the ala-chitralensis complex (he believed that allah represents a specimen of M. trans- caucasica Turati and strandi was a form near didymina Staudinger) but females of these two forms belonged to M. ala bicolor. The name determinata was the only one which had possible value and represented a modification which seemed to be common in the Eastern Tian-Shan, with characters somewhat intermediate between the typical form and bicolor. Later, Higgins (1981) united the taxa in question with other species of the didyma-group in the genus Didy- maeformia Verity with M. didyma as the type species. In spite of that, up to the present time the status of the taxa in question is unclear. This prob­ lem was reflected in the catalogues of Korshunov (1972) and Tuzov (1993), where some of the taxa were absent or attributed to other species. The analysis of a rather large material shows that we deal with five separate (but related) spe­ cies with rather constant characteristic features, which form a number of subspecies in differ­ ent parts of their distribution area. 2 The situation with M. ninae, described by Sheuuzhko from the western Tian-Shan as subspecies of M. ala is tangled. The specific rank of it was pointed out by Higgins (1941) under the name pseudoala (which he considered to be the oldest valid name). The form pseudoala was described by Sheuuzhko from Kamtshi (near Tashkent) as an aberration of a single female of M. didyma turkestanica Shel- juzhko with heavily suffused forewings with grey and red-brown hindwings. The name alboocellata is based upon the same specimen, and refers to white little spots inside orange submarginal macules on the under surface of hind-wing. This female stands as the type of both names, but the name pseudo­ ala comes first in the description. We examined the monotype of pseudoala + alboocellata (in the Zoo­ logical Museum of the Kiev University). Undoubtedly the form pseudoala is a melanic variety of M. di­ dyma. The name pseudoala was established in literature for this species (Higgins, 1955; Korshunov, 1972; Higgins, 1981; Tuzov, 1993) after Higgins' publication. The name ninae was correctly restored by Lukhtanov (Lukhtanov & Lukhtanov, 1994) but without indication of reasons for regarding the name pseudoala as a synonym. 88 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at The present study is based upon the collections of the following institutions: Zoological Mu­ seum of the Moscow State University, Zoological Institute of Russian Academy of Sciences (St. Petersburg), State Darwin Museum (Moscow), Zoological Museum of the Kiev University, Zoo­ logical Museum of Zoological Institute of Ukraine (Kiev), and collections of a number of ama­ teur collectors. The scheme of the wing pattern of M. ala is illustrated in fig. 1. In terminology of wing pattern, wing venation and genitalia we mainly follow Higgins (1941). Fig. 1: scheme of the wing pattern: M. ala ala Stgr. Mb - marginal band Sbs submarginal series Pds - postdiscal series Ds - discal series Bbs - black basal suffusion Osf - orange submarginal fascia v - vein. Abbreviations: FW - forewing HW -hindwing UPS - upperside UNS - underside UPF - upperside of forewing UPH - upperside of hindwing UNH - underside of hindwing Mts. - mountains coll. - collection ZMMU - Zoological Museum of Moscow State University ZISP - Zoological Institute of Russian Academy of Sciences (St. Petersburg) ZMKU - Zoological Museum of the Kiev University BMNH The Natural History Museum (London) 89 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at Key for the species of the M. o/o-complex based on external characters and morphology of the male genitalia: 1 Aedeagus enlarged in the central part. UNH black postdiscal lunules in the cellules between veins 3 and 5 outwardly concave. Ringwall produced distally into a large con­ ical or oval projection.. 2 Aedeagus slender, slightly curved. UNH black postdiscal lunules in the cellules be tween veins 3 and 5 inwardly concave. Ringwall produced distally into a rounded pro­ jection. 4 2 Ringwall produced distally into a big oval projection M. kotshubeji Sheljuzhko Ringwall produced distally into a conical projection.. 3 3 Distal projection of ringwall with a central apical notch. Aedeagus with a sharpened projection on the ventral side. UPS ground-colour yellow-orange. M. enarea Fruhstorfer Distal projection of ringwall without a central apical notch. Aedeagus without a sharpened projection on the ventral side. UPS ground-colour deep orange-red . M. chitralensis Moore 4 Ringwall produced distally into a small projection not extends to the tegumen. UPS ground-colour not pink . M. ala Staudinger Ringwall produced distally into a large projection extends to the tegumen. UPS ground-colour pinkish. M. ninae Sheuuzhko Key for the species of M. ala-complex based on external characters and morphology of the female genitalia: 1 1 Antevaginal plate deeply divided into lobes (figs. 11, 14).. 2 Antevaginal plate not deeply divided into lobes (figs. 10, 12, 13). 3 2 Antrum projected into the base of the antivaginal plate, forming a depression. Post- vaginal plate not wider than antevaginal plate, as a rule. UPH ground-colour deep orange-red. UPF greenish-grey suffusion well developed, as a rule M. kotshubeji Sheuuzhko Antrum not projected into the base of the antevaginal plate. Postvaginal plate wider than antivaginal plate. UPH ground-colour orange-yellow. UPF grey suffusion poorly developed, as a rule.. M. enarea Fruhstorfer 3 UNH black postdiscal lunules in the cellules between veins 3 and 5 inwardly concave M. ala Staudinger UNH black postdiscal lunules in the cellules between veins 3 and 5 outwardly con­ cave. 4 4 Postvaginal plate oval and wider than antivaginal plate. UPF grey suffusion well devel­ oped. UPH ground-colour deep red, submarginal and discal series well developed. M. chitralensis Moore Postvaginal plate trapezium-shaped. UPF grey suffusion not well developed. UPH ground-colour pinkish. UPH submarginal and discal series absent, as a rule. M. ninae Sheuuzhko 90 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at 1. Melitaea ala (Staudinger, 1881) Male: 38-44 mm. UPS ground-colour orange or orange-red; black marginal border wide (2 mm), fused with internervural marginal black spots. UPF submarginal series often complete and formed by black sharp lunules, often connected with the black marginal border; the post- discal series represented by one or two black spots near costa. UPF discal series regular, formed by black spots, sometimes fused near costa, on UPH it is often absent or represented by two spots: near costa and anal border. UPH submarginal series (if not reduced) is formed by black rounded lunules; wide black basal suffusion covers discal spot near anal angle. UNH postdiscal lunules forming the proximal edge of orange submarginal fascia in the cellules be­ tween veins 3 and 5 inwardly concave; there is a small orange macule in cellule between veins 2 and 1 near the base. Female: 38-43 mm. UPF ground-colour pale-yellow; black pattern and greenish-grey suffusion well developed. UPH more or less look like in male but black basal suffusion is wider and reaches submarginal lunules near anal angle. UNH orange submarginal fascia is represented by separate rounded macules indicated by black scales. Pale area between submarginal fascia and proximal black lunules is well developed. Proximal black lunules in the cellules between veins 3 and 5 inwardly concave. Male genitalia (fig. 2, A-C): Saccus more or less conical, with a depression in the apical part. Ringwall produced distally into a small rounded projection. Valva oval, slightly elongated, with a small spine on the dorsal surface in its distal part. Caudal process long with a row of small teeth on its ventral side. Large harpe without teeth on ventral surface. Aedeagus slender, slightly curved. Genitalia of males show no differences within the distribution area. Female genitalia (fig. 10): Postvaginal plate oval and not wider than antevaginal plate, as a rule; it is sclerotized in the central part near the outer side. Antevaginal plate strongly chitin- ized with an opening slightly dividing it into lobes. Antrum quadrangular, ductus narrow, with a sclerotized fork-shaped area. Distribution (map 1): East, West (Talassky Ala-Tau), North and Inner Tian-Shan, Dzhungarsky Ala-Tau Mts., Saur Mts. and South Altai (Lukhtanov & Lukhtanov, 1994). The record from Amdo (Higgins, 1941) is highly doubtful. We have no material from this locality. Geographical variation and subspecies: The variation of M. ala is great. A number of forms was described from different localities by different authors (Seitz, 1907; Sheuuzhko, 1928; Bryk, 1940). However the majority of these forms most likely reflects climatic and individual variation, and they are not likely to pretend to the subspecific status. 1a. Melitaea ala ala (Staudinger, 1881) (colour plate VI, figs. 1-4) M. dldyma var. ala Staudinger, 1881, Stettin, ent. Ztg. 42: 288 (Lepsa, Ala-Tau). M. ala Staudinger; Sushkin, 1913, Zeitschr. f. wiss. Insektenbiologie 9: 171, f. 5-6 (genital.). M. ala Staudinger; Wagner, 1913, Ent. Mitteil. 2: 91- 93 (Burhan, Usek). 91 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at M. ala ala Staudinger; Sheuuzhko, 1929, Mitth. Münch. Ent. Ges. 19: 368 (Dzhungarsky Ala- Tau, Tian-Shan oc.). M. ala ab. immodulata Sheuuzhko, 1929 (loc. dt.): 366 (Dzharkent). M. cla ala Staudinger; Bryk, 1940, Folia Zoologica et Hydrobiologica 10 (2): 346. M. ala allah Bryk, (loc. cit.): 346, Taf. 3, Flg. 23; Taf. 6, Fig. 57 ("Siberia"). M. ala Staudinger; Higgins, 1941, Trans, ent. Soc. Lond. 91 (7): 242-244, pl. 2 figs. 4, 10. M. ala Staudinger; Lukhtanov & Lukhtanov, 1994, Herbipoliana, Buchreihe zur Lepidoptero- logie, Bd. 3: 184-185, Taf. 36, Fig. 5, 6 (S. Altai). Type locality: Lepsa, Ala-Tau (Dzhungarsky Ala-Tau, Lepsy river). Male (colour plate VI, figs. 1, 2): 37-40 mm. UPS ground-colour deep orange-red, black pat­ tern often well developed: submarginal series complete on both wings, as a rule, outlined by black scales and connected with the black marginal border. UPF discal series represented by black spots, often fused near the costa. UPH discal series sometimes reduced. The UNH black postdiscal lunules forming the proximal edge of the orange submarginal fascia are connected with each other. UNH veins indicated by black scales. Female (colour plate VI, figs. 3, 4): 37-42 mm. UPF black pattern and greenish-grey suffusion well developed. UPH discal and submarginal series often complete. The UNH black postdiscal lunules forming the proximal edge of the orange submarginal fascia connected. UNH veins of­ ten indicated by black scales. Distribution: South Altai (Lukhtanov & Lukhtanov, 1994), Saur Mts. (Gemenay), Dzhungarsky Ala-Tau (Tekeli, Tentek river, Lepsy river, Kok-su river), Burchansarytau Mts. (Enbekshi, Burhan), Tyshkantau Mts. (Tyshkan, Usek river), Koiandy-Tau (Arasan), Horgos river, Altyn-Emel Mts. Taxonomic notes: The male specimens from the southern part of the Dzhungarsky Ala-Tau (Tyshkantau, Burchan-Sarytau ect.) have a partly reduced black pattern. Moreover, the speci­ mens with characters of ssp. bicolor are found among specimens with typical characters. How­ ever, the majority of the examined material tends to the nominate subspecies. The status of specimens recorded by Lukhtanov (Lukhtanov & Lukhtanov, 1994) from South Altai (Irtysh Valley, the mouth of river Buchtarma, southern parts of Narym and Kurtshum Mts. and Bukombai Mts.) is unclear. These specimens may be classified as ssp. bicolor, judging from the figures (Taf. 35, Fig. 5-6), but it was pointed out in the description, that the UNH veins are indicated by black scales. We did not examine material from this locality but there is a small series of M. ala (6 cfcf in the collection of S. Churkin) from Saur Mts at our disposal. These specimens are very similar externally to the typical form. The male of the form allah described by Bryk with the label "Siberia" is not likely to be classi­ fied as M. transcaucasica (as Higgins, 1955, suggested). We believe that this form belongs to M. ala and has characters of the nominate subspecies. The data of the label are probably wrong. Ab. immodulata, described by Sheuuzhko (1929) from three specimens, is a female variety with yellowish-orange ground-colour of both wings. 92 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at Fig. 2: male genitalia of M. ala ala Stgr. Dzhungarsky Ala-Tau, Kok-su river. A - General view from the ventral side (the shape of ringwall); B - Aedeagus; C - Valva, lateral view from the out­ side. Biology: Unknown. Flies in one generation in June-July in the mountains up to 1000-2000 m. In Altai found only in dry stony desert foothills up to 420-650 m (Lukhtanov & Lukhtanov, 1994). Material examined: 106 d'J' and 38 99 including the type of f. immodulata Sheljuzhko, 1929 (Dzharkentskij ujesd, Tyshkan - ZMKU). 1b. Melltaea ala blcolor {Seitz, 1907) (colour plate VI, figs. 5-7) A4. didyma ab. bicolor Seitz, 1907, Grossschm. Erde 1: 219, t. 66, f. 7-8 (Karagatai Mts.). A4, ala bicolor Seitz; Sushkin, 1913, Zeitschr. f. wiss. Insektenbiologie 9: 172. A4, ala bicolor Seitz; Sheuuzhko, 1929, Mitt. Münch. Ent. Ges. 19: 368 (Karagai-tau Mts, Na- ryn, Alexandri-mts.). A4, ala ab. latemarginata Sheuuzhko, 1929 (loc. cit.): 366-367, Taf. 28, Fig. 7-8 (Naryn). A4, ala bicolor Seitz; Bryk, 1940, Folia Zoologica et Hydrobiologica 10 (2): 347-348 (Karai- gatan Gbge. südl. Narynsk. Issykul mer.). A4, ala bicolor Seitz; Higgins, 1941, Trans, ent. Soc. Lond. 91 (7): 244, pi. 6, fig. 6, 12. Type locality: Karagatai Mts (? Naryn-Too Mts.). 93 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at Map 1 (distribution of M. ala Stgr.): • - M. ala ala Stgr.; © - M. ala bicolor Seitz; A M. ala with intermediate characters; O - M. ala Stgr. of unclear subspecific status. The name blcolor was introduced by Seitz (1907) for a female variety which had a pale UPF and red UPH. This name was used traditionally by different authors (Sushkin, 1913; Shel- juzhko, 1929; Bryk, 1940; Higgins, 1941) as subspecific for M. ala from southern parts of the area. We could not identify the type locality (Karagatai Mts.) of this form on any map. The au­ thors mentioned believed that the Karagaitau Mts. (old name of Naryn-Too Mts.) is the type lo­ cality of this form. Sheuuzhko (1928) found that females from Naryn. were similar to the form bicolor, figured in Seitz. The males of M. ala from Inner Tian-Shan have good external differ­ ences from the typical form. We consider it reasonable to use the name bicolor as subspecific. 94 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at Male (colour plate VI, figs. 5, 6): 37-42 mm. UPS ground-colour orange or orange-red. Black pattern partly reduced on both wings: UPF submarginal series often not complete, formed by black crescents not connected with the black marginal border, UPH submarginal series ab­ sent. UPF discal series formed by small black spots, UPH discal series usually reduced or repre­ sented by two spots (near the costa and near the anal angle). UPH black basal suffusion less developed than in the typical form. UNH black postdiscal lunules forming the proximal edge of the orange submarginal fascia are disconnected with each other. UNH veins not indicated by black scales. Female (colour plate VI, fig. 7): 40-42 mm. UPF black pattern and greenish-grey suffusion less developed than in the nominate subspecies. UPH discal and submarginal series often absent. UNH veins not indicated by black scales. Distribution: Kungey Ala-Tau Mts. (Chon-Urukty, Kok-Bulak, Chon-Aksu, Kegen, Cholpon-Ata, Kintyk, Grigorievka, Kelsai lake), Kirgizsky Mts (Chai-Sandyk, Ala-Archa, Chon-Kurchak, Usun-Gyr, Kara-Balta river, near Orto-Tokoi lake), Suusamyr valley, Suusamyr Mts. (Kokomeren river), Talassky Ala-Tau Mts. (Kara-Bura river), Terskey Ala-Tau Mts. (Pokrovka, Karakol, Chon- Kisylsu, Kere-Getash, Dzhety-Ogus, Ak-Suu), Baidullu Mts. (near Dolon pass), Moldo-Too Mts. (Korogo, Eki-Naryn, Kurtka river), Sonkel-Too Mts. (nearSon-Kel pass), Naryn-Too Mts. (Naryn). Taxonomic notes: In our opinion the specimens from Zailiisky Ala-Tau Mts. (Talgar, Turgen river, Ak-tuz, Almaatinka river, Medeo, Kok-Dzhailau, Issik river) and Ketmen Mts. (Bolshoi ketmen, Timerlik) have intermediate status between the typical form and ssp. bicolor. UPS black pat­ tern often similar to the typical form, but UNH veins not or less indicated by black scales. It is necessary to point out that the specimens with intermediate characters occur in the Kirgizsky Mts. (with the exception of the extremely eastern part of it). The specimens distributed in the western part of Chinese Tian-Shan (Kouldja—type locality of sheljuzhkoi Bryk— and Juldus) also have intermediate status between the typical form and ssp. bicolor (there are 8 dd from Juldus and 3 dd from Kouldja at our disposal from ZMMU and ZISP): UPH submarginal and discal series often represented, UNH veins slightly indicated by black scales. The forms described by Bryk (1941) from the extremely eastern part of Chinese Tian-Shan are of great interest. The form strandi (Nan-Shan(?), Urumtshi) may be classified as ssp. bicolor judging from the photo. We have no material of M. ala from this locality. The form determinata (Fu-Shu-Shi) is characterized by the large size of both sexes, UPS black pattern of the male is well developed, and UPH discal area of the female has no markings. We examined a little se­ ries (5 dcT and 12 99, in ZISP) of M. ala, taken by Groum-Grzhimailo in Bogdo-Ola Mts. Their subspecific status is unclear, because the material is not sufficient. The specimens examined look like the specimens from Kouldja and Juldus. We believe their characters to be intermedi­ ate between the typical form and ssp. bicolor. However, it is possible that the form deter­ minata has subspecific status. Identification should be confirmed by examination of supple­ mentary material from this locality. Biology: Unknown. Flies in one generation in June-July in the mountains up to 1000-2500 m in dry meadows and steppe-like slopes (Lukhtanov & Lukhtanov, 1994). Material examined: 310 dd and 137 99 including the type of f. latemarginata Sheuuzhko, 1929 (Naryn) (ZMKU). 95 ©Ges. zur Förderung d. Erforschung von Insektenwanderungen e.V. München, download unter www.zobodat.at 2. Melitaea kotshubeji (Sheuuzhko, 1929) Male: 38-47 mm. UPS ground-colour deep orange-red; black marginal border wide (2 mm), fused with internervural marginal black spots on both wings. UPF submarginal series often complete and usually formed by black sharp lunules, often connected with the black marginal border; discal series regular, formed by usually rather small black spots. UPH discal series of­ ten complete; submarginal series (if not reduced) formed by black sharp lunules connected with the black marginal border; wide black basal suffusion covers discal spot near anal angle. UNH submarginal orange fascia rather wide; postdiscal lunules forming the proximal edge of this fascia in cellules between veins 3 and 5 outwardly concave; small orange macules in cellule between veins 2 and 1 near the base absent. Female: 40-50 mm. UPF ground-colour variable: from pale-pink to pale-yellow; black pattern and greenish-grey suffusion well developed. UPH ground-colour similar to male; submarginal series (if not reduced) formed by black rounded or sharp-pointed lunules; discal series some­ times absent; wide black basal suffusion reaches submarginal lunules near anal angle. UNH orange submarginal fascia represented by separate rounded macules; the pale area between this fascia and proximal black lunules is often developed. UNH proximal black lunules in the cellules between veins 3 and 5 outwardly concave. Male genitalia (figs. 3-5, A-C): Saccus more or less conical with a depression in the apical part. Ringwall produced distally into a large oval projection with a little notch in the distal part. Valva wide and oval, caudal process long. Large harpe without teeth. Aedeagus straight, enlarged in the central part, a small spine may be present in its distal part near apex. Female genitalia (fig. 11): Postvaginal plate oval and sclerotized in the central part near the outer side. Antevaginal plate strongly chitinized and deeply divided into wide rounded lobes. The conical antrum projects into the base of the antevaginal plate. Ductus narrow, with a sclerotized fork-shaped area. Distribution: Peter the Great Mts., northern part of Hazreti-Sho Mts., western parts of Zaa- laisky Mts., southern slopes of Alaisky Mts., south-western slopes of Fergansky Mts., south­ eastern slopes of Chatkalsky Mts. Geographical variation and subspecies: only the nominate subspecies (Peter the Great Mts., Tuptshek) was known until now. Two new subspecies are described below. 2a. Melitaea kotshubeji kotshubeji (Sheuuzhko, 1929) (colour plate VI, figs. 8-11) M. didyma var. ala Groum-Grschimajlo, 1890, in Romanoff, Mem. Lep. 4: 430 (Touptshek). M. ala kotshubeji Sheuuzhko, 1929, Mitt. Münch. Ent. Ges. 19: 364, T. 27, F. 1-4 (Touptshek). M. ala kotshubeji Sheuuzhko; Bryk, 1940, Folia Zoologica et Hydrobiologica 10 (2): 348. M. chitralensis mod. kotshubeji Sheuuzhko; Higgins, 1941, Trans, ent. Soc. Lond. 106: 248, pi. 6, figs. 4, 10. M. ala bicolor Seitz; Shchetkin, 1981, Ent. Obozr. Tajikistana: 163-164 (Surkhob valley). 96