Trans. lepid. Soc. Japan 49 (3): 177-193, June 1998.

A review of Erebia ligea-complex (Lepidoptera, Satyridae) from Eastern Asia

V. V. Dubatolov 1), Y. P. Korshunov 1), P. Yu. Gorbunov 2), O. E. Kosterin 3), A. L. Lvovsky 4)

1) Siberian Zoological Museum, Institute of Animal Systematics and Ecology, Siberian Division of Russian Academy of Sciences, Frunze street, 11, Novosibirsk, 630091, Russia.

2) Institute of Plant and Animal Ecology, Uralian Division of Russian Academy of Sciences, March 8 street, 202, Ekaterinburg, 620144, Russia.

3) Institute of Cytology and Genetics, Siberian Division of Russian Academy of Sciences, Lavrentiev avenue, 10, Novosibirsk, 630090, Russia.

4) Zoological Institute, Russian Academy of Sciences, Universitetskaya Naberezhnaya, 1, St.-Petersburg, 199034, Russia.

ABSTRACT: It is shown that East Asia is inhabited by two similar species of Erebia (ligea Linnaeus, 1758) group, well differing in the male genitalia structure as well as a number of the wing pattern details. They are E. ligea Linnaeus, 1758 and the second sympatric species, which has a priority name Erebia ajanensis Menetries, 1857, stat. nov., nec. Eiffinger, 1907, nec. Warren, 1936, nec. Kurentzov, 1950, 1970; and on a preliminary review seems to be represented by three subspecies: the nominotypical, known by a type series from Ayan (the coast of the Sea of Okhotsk in the Khabarovskii Krai Province, Russia), ssp. kosterini P. Gorbunov, Korshunov et Dubatolov, 1995 (=Erebia kosterini) in the southern Magadan Province, Russia, and ssp. arsenjevi Kurentzov, 1950 (=Erebia ligea arsenjevi) in more southern regions of the Russian Far East, Korea, N and NE? China. The lectotypes of E. ajanensis, E. ligea arsenjevi (that is E. ajanensis arsenjevi) and of Erebia ligea koreana Matsumura, 1928 have been designated, as well as that of Erebia eumonia Menetries, 1859. The latter taxon was described by a heterogeneous series including both species, E. ligea and E. ajanensis. After lectotype designation the name eumonia became the valid one for an eastern subspecies of E. ligea, ranging from the Altai Mts. (except for the north part) to the Magadan Province of Russia, Japan and Korea. All the subspecies of the true E. ligea described from this vast territory, except for E. ligea sachalinensis Matsumura, 1919, E. l. rishirizana Matsumura, 1928 and E. ligea takanonis Matsumura, 1909, are considered as synonyms to E. ligea eumonia.

Key words: Review, synonymy, Satyridae, Erebia, Erebia ligea, lectotype, distribution, East Asia, Siberia, Far East.

In 1994, while working on the book by Yu. P. Korshunov and P. Yu. Gorbunov (1995) on the butterflies of the Asian part of Russia, P. Gorbunov found out that specimens collected by O. E. Kosterin in the Magadan Region, earlier identified as "Erebia ligea", differ substantially from the mentioned species in the male genitalia structure. After discussing systematic position of these specimens the authors decided that they comprise a new species, and later described as Erebia kosterini P. Gorbunov, Korshunov et Dubatolov (Korshunov and Gorbunov, 1995). They differ from a common E. ligea by stick-like valvae, without a ledge and a sharp decrease in their thickness.

Later similar butterflies were collected by V. Baglikov in the southern Sikhote-Alin' Mountains (Primorye territory, or the Primorskii Krai Province of Russia). Taking into account these new materials, it became necessary to study thoroughly the systematics of all East Asian taxa on the E. ligea group.

We have studied collections of Siberian Zoological Museum (SZMN) of Institute of Animal Systematics and Ecology, Siberian Division of the Russian Academy of Sciences (Novosibirsk), the Institute of Plant and Animal Ecology, Uralian Division of the Russian Academy of Sciences (Ekaterinburg), and the Zoological Institute (ZIN) of the Russian Academy of Sciences (St.-Petersburg), including the type materials. Thanks to the courtesy of Mr. Hiroshi Yoshimoto we were provided with a photograph of the type specimen of Erebia ligea koreana Matsumura, preserved at the University of Hokkaido (Sapporo).

The first taxon of this group described from East Asia was Erebia ligea ajanensis, described by E. Menetries (1957) on the material collected by I. Wosnesensky from Ayan (recently - the northern Khabarovskii Krai Province, Russia). Later, after receiving of further materials, he redescribed it but as an independent species under the name E. eumonia Menetries, 1859. This description was based on collections of L. Schrenck from the Hadshi Bay "de la bai d'Hadshi, sur les c˘ tes de la Mandshourie, par le 49░ de latitude" (now being Sovetskaya Gavan', Russia) and mentioned Erebia ligea ajanensis only as a synonym. In another work (Menetries, 1859b) he again gave Erebia eumonia with a synonym E. ligea ajanensis and enumerated all the available materials from Ayan, the Hadshi Bay, and one more specimen from Nikolaevsk. The same work contains a picture of E. eumonia showing undoubtedly an eastern type of E. ligea.

We have studied the type specimens of both taxa described by E. Menetries, Erebia ligea ajanensis and E. eumonia, preserved in the collection of ZIN. The type series of Erebia ligea ajanensis is represented by two females. These females are characterised by all the characters of the species described by us as Erebia kosterini (see below), so: Erebia ajanensis Menetries, 1857, stat. nov. = Erebia kosterini P. Gorbunov, Korshunov et Dubatolov, 1995, syn. n.

The type series of E. eunomia, according to the original description, consisted of "17 individuals". It turned to be heterogeneous as being represented now only by two males of E. ligea and one female of E. ajanensis available in ZIN collection. The sex of the specimen depicted in Menetries (1859b) was not specified there and can hardly be undoubtedly inferred from the picture as such (the habitus of the wing pattern resembles a male, while a well developed inner white band on the hind wing underside rather suggests a female). Neither of the three specimens existing at present corresponds to the picture in individual details of the pattern. Since the Menetries' picture shows undoubtedly an E. ligea specimen, we have chosen a specimen of E. ligea (a male) as a lectotype, so giving the name "eumonia" priority to the eastern subspecies of this species, earlier known as E. ligea ajanensis auct. nec Menetries, 1857 (Kurentzov, 1950, 1970; Tuzov, 1993; Korshunov and Gorbunov, 1995). The paralectotype of E. eumonia referring to E. ajanensis, belongs to a subspecies which received the name in 1950 after the description of Erebia ligea arsenjevi Kurentzov, 1950.

The diagnostic characters of the similar species Erebia ajanensis and Erebia ligea are summarised in the following table (see also Fig. 3):

Character

Erebia ligea

Erebia ajanensis

1. internal margin of the fulvous band on the fore wing

Straight or almost straight

with an angular incision

2. male androconial scales

Prevail on an entire surface of a dark sex-brand, well visible against the light

found only at the vein A on the fore wing, with a sex-brand vague, weakly visible against the light

3. the number of ocelli on the fore wing

2-5

4-6

4. an imaginary straight line drawn through the centres of ocelli between the veins M1 - M2 and M2 - M3 of the fore wing

Crossing the ocellus between the veins Cu1 -Cu2 and the wing margin at the anal angle

crossing the ocellus between the veins M3 - Cu1 and the outer wing margin

5. the ocellus between the veins Cu1 - Cu2 on the fore wing

not larger than that between the veins M2 - M3

Larger than that between the veins M2 - M3

6. the ocellus between the veins R5 - M1

Absent

Present in more than half of specimens

7. an angle formed by the row of ocelli on the hind wing, with the apex at the ocellus between the veins M3 - Cu1

very blunt, more than 140░

much less blunt, about 110-130░

8. the ocellus between the veins M3 - Cu1 on the hind wing

always much less than the two adjacent ones

Equal or slightly less than the two adjacent ones

9. the ocellus between the veins M2 - M3 on the hind wing

absent in more than half of specimens

always present

10. the valva in the male genitalia

thick but narrowing drastically, with a ledge, in the last fourth of the length

thin, tapering smoothly to the apex

11. the dorsal margin of the valva

dentate on more than a half of its length

dentate on less than a half of its length

In general, by the male genitalia structure E. ajanensis (Figs 18-19, 25-32) somewhat resembles a North American species Erebia vidleri Elwes, 1898 (Fig. 24) belonging to a special species group (Warren, 1936; Kogure and Iwamoto, 1992). Besides good differences between these species by the wing pattern, those by the male genitalia structure can also be found as follows:

Character

E. ajanensis

E. vidleri

The shape of the valva

the distal half is about evenly thin, proximally the valva smoothly becomes thicker

the distal one-third is thin, proximally the valva smoothly becomes thicker

ratio of the thickness of the valva at the base, at the middle, and at the apex

2:1:1

4:2:1

the teeth on the distal half of the valva

large, of uneven sizes

small, of an even size

a narrowing of the valva proximally of the most proximal tooth

small but present

absent

the valva and tegumen+uncus

form an acute angle

are about parallel

Below a preliminary review is given of a both two species subspecies level taxa which inhabit Asia east of 120░ E.

Erebia ajanensis Menetries, 1857, stat. nov.

Range (Fig. 16): Russia: the southern Magadan Region and the northern Khabarovskii Krai Province: mostly along the coast of the Sea of Okhotsk, and Bureinskie Gory [Bureinskie Mountains]; the Sikhote-Alin' Mountains within the southern Khabarovskii Krai and Primorskii Krai Provinces. North Korea. North and, probably, North-Eastern China.

Erebia ajanensis ajanensis Menetries, 1857, stat. nov. (Fig. 1)

Erebia ligea ajanensis Menetries, 1857, Enumeratio Corporum Anim. Mus. imp. Acad. Sci. Petropolitanae (Lepid.) (2): 104.

Type locality: "Ajan" [now - Ayan, Kabarovskii Krai, Russia].

Type materials. Lectotype - (Fig. 1), with labels: "Ajan." (handwriting on a red paper); "coll. Acad./Petrop." (printed on a white paper); "Wosnesensky leg." (handwriting on a white paper by V. V.Dubatolov); "Lectotypus Erebia / ligea var. ajanensis / Men. / design. Dubatolov 1997" (on a red paper); "Erebia / ajanensis / ajanensis Men. / V.V.Dubatolov det. VI 1997". Paralectotype: 1 , "657" (printed on a white paper); "k. Ershova." (printed on a white paper; in Russian); "Ajanensis / Menet. Ajan." (handwriting on white paper; label with a double black margin"; "Wosnesensky leg." (handwriting on a white paper by V. V. Dubatolov); "Paralectotypus Erebia / ligea var. ajanensis / Men. / design. Dubatolov 1997" (on a red paper); "Erebia / ajanensis / ajanensis Men. / V. V. Dubatolov det. VI 1997". All in ZIN.

Range. Known only on the type series from the environs of Ayan (a coast of the Sea of Okhotsk), the northern Khabarovskii Krai Province, Russia.

A differential diagnosis is based on the two known females only. From females of other subspecies they differ by a small size, the fore wing length being 20-21 mm, and three relatively large ocelli on the fore wing upperside, that between the veins M2 and M3 containing a white dot, there is also a black dot between the veins M1 and M2. There are four ocelli with white pupils on the hind wing underside, these ocelli are small, their diameter not exceeding a half of the distance between the veins. The fulvous band on the fore wing upperside is comparatively wide, its narrowing between the veins M3 and CuA is weak, the ratio of the width of the band at this point to the fore wing length being approximately 0.16 in the two specimens.

Erebia ajanensis kosterini P. Gorbunov, Korshunov et Dubatolov, 1995, stat. nov. (Figs 2-3)

Erebia kosterini Korshunov, P. Gorbunov, 1995, Butterflies Asian Part Russia: 123, 188: fig. 253, 190: fig. 253.

Erebia (ligea): Kosterin, 1994, Actias 1: 75, 78, nec Linnaeus, 1758.

Type locality: the Khidzha River, Koni Peninsula, Magadan Province, Russia.

Type materials. Holotype - (Fig. 2), Magadan Province, Koni Peninsula, the low reaches of the Khindzha River, 20.VII.1989 (O. Kosterin leg.). Paratypes: 2 , 1 , Koni Peninsula, the low reaches of the Khindzha River, 20.VII.1989 (O. Kosterin leg.); 1 (allotype) (Fig. 3), the same locality, 20.VII.1989 (O. Kosterin leg.). All in SZMN.

Additional materials. Magadan Province: 9 , 4 , the settlement Palatka, 10-19.VII.1993, 18-19.VII.1995, 22.VII.1996 (V. Baglikov leg.).

Range. the southern Magadan Province, Russia.

A small subspecies, the fore wing length being 10-21 mm in males, 23 mm in females. Both sides of the hind wing have four blind (in males) or white-centred (in females) ocelli of a considerably elongate (oval) shape. The fulvous band on the fore wing upperside is comparatively wide, the narrowing between the veins M3 and CuA is weak, mainly in males; the ratio of the width of the band at this point to the fore wing length being 0.18-0.20 in males and 0.17-0.20 in females.

Note. Now we only provisory accept E. ajanensis kosterini as a separate subspecies, since in the above mentioned series of 9 males and 4 females, collected by V. Baglikov at the settlement Palatka, the Magadan Region, 5 specimens have perfectly round ocelli on the wing underside while other 8 specimens have elongate ocelli. The size of these butterflies is the same as in the typical kosterini and ajanensis, the fore wing length being 19.5-23.2 mm in males and 20.0-23.3 mm in females. Additional materials from the type locality of E. ajanensis kosterini is necessary for a determination of its subspecies independence. If there would be specimens with round ocelli then this taxon should be synonymized to the nominotypical subspecies E. ajanensis ajanensis. Now we suppose the population from Palatka as representing a transitory zone between the subspecies, one of us (P. Gorbunov) thinking that there is too little evidence of subspecies differentiation in E. ajanensis.

Habitat. On the Koni Peninsula the butterflies were found in the valleys of the Khindzha and Burgauli Rivers, covered with a forest-tundra-like landscape formed by alternating tall bushes of Dushekia fruticosa and Pinus pumila, heath-like openings with various Ericaceae and Empetrum dominating in the herbage, and polydominate herb meadows (Kosterin, 1994).

Erebia ajanensis arsenjevi Kurentzov, 1950, stat. nov. (Fig. 4-6)

Erebia ligea arsenjevi Kurentzov, 1950, Ent. Obozr. 31(1-2): 243-244; Azarova, 1986, Syst. Ecol. Cheshuekr. Daln. Vost. SSSR: 123; Kogure, Iwamoto, 1992, Yadoriga (150): 17; Tuzov, 1993, Syn. List Butterflies ex-USSR: 31; Korshunov, Gorbunov, 1995, Butterflies Asian Part Russia: 122.

Erebia ligae arsenievi [sic]: Kurentzov, 1970, Butterflies Far East USSR: 62.

Type locality: "Ist. Kolumbe" (in Russian) [head-water of the Kolumbe River, Nature Reserve "Sikhote-Alinskii", Krasnoarmeiskii raion (district), Primorskii Krai Province, Russia].

Type materials. A good picture of a type specimen (syntype) of Erebia ligea arsenjevi and its labels ("20 VII 34 / Ist. Kolumbe"; "Syntypus Erebia ligea / arsenjevi Kurenzov, / 1950") is given in the work of M. Kogure and Y. Iwamoto (1992: p. 17, f. 8). According to this picture, the specimen possesses all the characters of the species Erebia ajanensis. Besides, in the first description it was stated that: "The valvae in the latter race [a Sakhalin race] are thicker, with a sharp ledge in the proximal part; in a new subspecies they are thin and has no such a ledge". We choose the depicted specimen as a lectotype. It is preserved in the Institute of Biology and Pedology, Far Eastern Division of the Russian Academy of Sciences (Vladivostok).

Additional materials. China: 1 , between Peking (=Beijing) and Lake Dolon-nor. Korea: Ryanggangdo: 1 , Paekimjun sangnone (Fig. 6: 1), 8.VII.1985 (from Im Chong An); 2 , Paekimjun (Fig. 6: 2-3), 11.VII.1985 (from Im Chong An). Russia: Primorye (Primorskii Krai Province): 7 , Sidemi (or Sidimi in another transliteration; now - Narva, Khasan district), 1897 (Jankowsky leg.); 7 , St. Ilya Mt., Spasskii Uezd [district], 31.VII-1.VIII.1926 (Djakonov et Filipjev leg.); 6 , Chuguevskii Raion [district], the headwater of the Sokolovka River, [approximately 20 km SE of Chuguevka], 15-31.VII.1974 (V. Kuznetsov leg.); 1 , 2 , Mt. Oblachnaya, S slope, 26.VII.1982 (Beljaev leg.); 1 , near the settlement Krasnorechenskii, 13.VII.1992 (V. Baglikov leg.); 1 , Anuchinskii Raion [district], [30 km N of Chernyshevka], village LZP-3, VI.1980 (V. Bakurov leg.); 1 , the middle flow of the Iman River (now - the Bolshaya Ussurka River), 20.VII.1934 (Kurentzov leg.). Khabarovskii Krai Province: 1 , Nikolaevsk-na-Amure, 12.VIII.1854 (?Schrenck leg.); 1 , Nature Reserve "Komsomol'skii", the cordon Zolotoi, 11.VII.1988 (V. Olschwang leg.); 2 , the Sikhote-Alin' Mts., the gold mine Zolotoi, 28.VI.1982 (E. Novomodnyi leg.); 1 , the railway station Vysokogornaya, 10.VII.1974 (A. Barkalov leg.); 1 (the paralectotype of Erebia eumonia Men.), Bai Hadschi [Sovetskaya Gavan'] (Schrenck leg.); 2 males, Bureinskie Gory [the Bureinskie Mountains], [the headwater of the Verkhnyaya Bureya River], the zimovie [winter house] Medvezhye, 26.VII.1977 (E. Novomodnyi leg.).

Range. Russia: the southern Khabarovskii Krai Province north to the Bureinskie Mountains and the Amur River mouth; the Primorskii Krai Province (Primorye territory). North Korea. North, and probably, North-Eastern China.

The subspecies is characterised by an on average greater size, the fore wing length being 22-26 mm in males and 22-24 mm in females. The fulvous band on the wing underside is considerably narrower than in the above mentioned subspecies. The ocelli on both sides of the hind wing are roundish and large, their diameter being noticeably larger than the half of the distance between the veins. They are blind in males and centered with very small white dots in females. As different from two former subspecies the fulvous band on the fore wing upperside, mainly in males, has a strong narrowing between the veins M3 and CuA, so that the ratio of the width of the band at this point to the fore wing length varies from 0.09 to 0.14 in males and from 0.16 to 0.18 in females.

Note: Describing this taxon Kurentzov (1950) compared it with "Erebia ligea ajanensis". It turned out, however, that he applied the latter name to the butterflies belonging to E. ligea indeed (to its eastern subspecies), while, as we have shown, the type series of the taxon ajanensis in fact belongs to the other species. So the differential diagnosis by A. Kurentzov is applicable in general to the species E. ajanensis (=E. ligea arsenjevi sensu Kurentzov, 1950) as differing from E. ligea (=E. ligea ajanensis sensu Kurentzov, 1950), but not to distinguishing subspecies E. ajanensis s. str. and E. a. arsenjevi. Of course, we can only provisory state that the enumerated above materials belong to a subspecies differing from the nominotypical one, which is known by two females only. However, easily noticeable differences in general size and in relative size of the ocelli on the hind wing, as well as a great distance between the sites, give us reasons for such a conclusion.

Erebia ligea (Linnaeus, 1758).

Range (Fig. 2). Western and Eastern Europe, North-Eastern Kazakhstan, Russia (European part, Siberia, and the Far East, except for the north; Sakhalin), Northern Mongolia, North-Eastern and North China (Kogure, Iwamoto, 1992: 18, pl. 1, fig. 4; 1993: 30), Korea, Japan (Rishiri, Hokkaido, Honshu).

Erebia ligea eumonia Menetries, 1859, stat. nov. (Fig. 7, 14)

Erebia eumonia Menetries, 1859, Bull. phys.-mat. Acad. Sci. St. Petersb. 17: 216; In Schrenck, L. v., Reisen und Forschungen im Amur-Lande 2 (1): 34-35, pl. 3, fig. 4.

Erebia ajanensis: Graeser, 1888. Berl. ent. Z. 32: 33-105.

Erebia ligea ajanensis: Eiffinger, 1907, in Seitz, A. [Ed.] Gross -Schmett. Erde 1: 108, pl. 37, row g; Warren, 1936, Monograph Genus Erebia: 51-52, pl. 58, figs 549, 550, 553-555, genit. pl. 24, fig. 245; Kurentzov, 1950, Ent. Obozr. 31(1-2): 243; 1970, Butterflies Far East USSR: 62, fig. 60-1 (fig. 32); Sedykh, 1979, Ent. Obozr. 58(2): 294; Kogure and Iwamoto, 1992, Yadoriga (150): 17, 18, pl. 1, fig. 6; Korshunov and Gorbunov, 1995 Butterflies Asian Part Russia: 122.

Erebia ligea takanonis: Doi, 1919, Chosen Iho 58-59: ???, nec Matsumura, 1909.

Erebia ligea koreana Matsumura, 1928, Insecta matsum. 2(4): 194-195, syn. n.; Kurentzov, 1970, Butterflies Far East USSR: 62-63; Lee, 1982, Butterflies Korea: 80, pl. 52, figs. 185A-C; Tuzov, 1993, Syn. List Butterflies ex-USSR: 31.

Erebia ligea koreana, ab. hakutozana Matsumura, 1928, Insecta matsum. 2: 195 [infrasubspecific; unavailable].

Erebia ligea kamtschadalis Goltz, 1932, Dt. ent. Z. Iris 47: 114, syn. n.; Korshunov and Gorbunov, 1995, Butterflies Asian Part Russia: 122.

Erebia ligea ssp.: Kogure and Iwamoto, 1992, Yadoriga (150): 18, pl. 1, fig. 3.

Erebia ligea: Kogure, 1985, Yadoriga (122): 5, pl. 1, fig. B, C.

Type locality: "Bai Hadschi" [now - Sovetskaya Gavan', Khabarovskii Krai Province, Russia].

Type materials. Lectotype of Erebia eumonia - (Fig. 7) with labels: "Bai Hadschi" (handwriting on a red paper); "Schrenk." (printed on a white paper); "coll. Acad. / Petrop." (printed on a white paper); "Lectotypus Erebia / eumonia Men. / design. Dubatolov 1997" (on a red paper); "Erebia / ligea / eumonia Men. / V. V. Dubatolov det. VI 1997". Paralectotypes: 1 (Fig. 14) with the same labels, except for: "Paralectotypus Erebia / eumonia Men. / design. Dubatolov 1997"; 1 [=ajanensis], with same labels as male, except for: "Erebia / ajanensis arsenjevi / Kurentz. / V. V. Dubatolov det. VI 1997". All in ZIN. Lectotype of Erebia ligea koreana (Fig. 8) with the labels figured on fig. 9. It is preserved in the Hokkaido University, Japan.

Additional materials. Amurskaya Province: 6 , a western border of the Nature Reserve "Zeiskii", the Erakingra River, 12- 20.VII.1977 (L. Morozova leg.). Khabarovskii Krai Province: 2 , Okhotsk, 23.VII.1987 (V. Nebaikin leg.); 1 , the same locality, 8.VIII.1987 (V. Nebaikin leg.); 10 3 , Shantarskie Isl., Bolshoi Shantar Is., 18.VII-8.VIII.1925 (Dulkeit leg.); 1 , the same locality, 22.VII.1911 (Soldatov leg.); 1 1 , Nikolaevsk-na-Amure (Graeser leg.?, Erschoff collection); 1 , Sukhanovka (Low Amur), 6.VIII.1910 (Soldatov leg.); 7 3 , the same locality (Graeser leg., coll. Dieckmann); 1 , the Nature Reserve "Komsomolskii", 3.VII.1988 (V. Olschwang leg.). Primorskii Krai Province: 4 , near the settlement Krasnorechenskii, 12-13.VII.1992 (V. Baglikov leg.). Yakutia: 3 , near Olekminsk, 10.VI-17.VII.1910 (Kharitonov leg.); 3 , the same locality, Nakhtuiskoe, 5.X(!?).1916 (Gubelman leg.); 1 , the Nature Reserve "Olekminskii", the Tuolba River, 1.VI.1988 (Golyakov leg.); 1 , 273 km from Ust-Maiskii (=Ust-Maya), 26.VII.1928 (Kyakshto leg.). Magadan Province: 1 , Omulevskie Mts., the Avr rivulet, a left tributary of the Omulevka River, 10.VII.1974 (V. N. Dubatolov leg.); 1 , 15 km SW of Verkhnyaya Buyunda, 8.VII.1981 (A. Jakimavic us); 3 , the settlement Palatka, 14.VII.1993, 16.VII.1995 (V. Baglikov leg.); 1 , the settlement Ola, 19.VII.1993 (V. Palekha leg.); 1 , the settlement Karamken, 8.VIII.1993 (V. Palekha leg.); 1 , a paratype of E. kosterini (!), Koni Peninsula, the low reaches of the river Khindzha, 20.VII.1989 (O. Kosterin leg.). Kamchatka: 1 , a ravine on the western slope of Mt. Nachikskaya, 17.VII.1959 (Kurentzov leg.); 1 , the Nature Reserve "Kronotskii", the caldera of the Uson volcano, 11.VIII.1985 (A. Lvovsky leg.); 1 , the same Nature Reserve, Dolina Geizerov [the valley of geysers], 7-13.VIII.1985 (A. Lvovsky leg.); 4 , a coastal hilly Empetrum tundra at the settlement Ozernovskii, 11-12.VIII.1991 (O. Kosterin leg.); 3 , Bolsheretsk, Apacha, 16.VII.1983 (V. Olschwang leg.); 1 male, Ganaly, 16.VII.1983 (V. Olschwang leg.); 1 , the same locality, 2.VIII.1961 (N. Violovich leg.).

Range. Russia: the southern Magadan Province, including the Kolyma River upper flow, Kamchatka, S Yakutia, the Amurskaya Province, the Khabarovskii Krai Province (a coast of the Sea of Okhotsk, the mountains northwards of the Amur River lower flow), Primorskii Krai Province (the Sikhote-Alin' southwards to the settlement Krasnorechenskii), Chitinskaya Province (except for SE), Buryatia, Tuva, Southern Krasnoyarskii Krai Province (the eastern part of the East Sayan), Khakasia, Altai Mts., except for the northern part. E Kazakhstan (the Altai Mts. within the Vostochno-Kazakhstanskaya (East-Kazakhstan) Province). Mongolia: Mongolian Altai, East Sayan, North Hangai, Hentei, recorded for the following aimaks: Bajan-Olgij, Uvs, Dzavchan, Chovsgol, Archangaj, Bulgan, Tov, Chentij (Korshunov, Soljanikov, 1976; Korshunov, 1977). Korea: Gaemagoweon Plateau area in Pyeongannamdo, Hamgyeongnamdo, Hamgyeongbudgo Provinces (Lee, 1982) and, probably, others in north part of the country. According to M. Kogure and Y. Iwamoto (1993), North and North-Eastern China (Hebei, Henan, Shaanxi, Shanxi, Nei Mongol, Jiling, Liaoning, Heilongjiang).

As different from dark butterflies of a East European / West Siberian subspecies E. ligea kamensis Krulikowsky, 1909 with reddish-brown bands on the wing upperside and much reduced white band on the hind wing underside, which inhabit the southern West Siberia, including the Kuznetskoe Upland (including the elevation of Salairskii Kryazh, the elevation of Gornaya Shoria and the Kuznetskoe Alatau Mts.) east to the environs of Krasnoyarsk, the E. l. eumonia is characterised by brownish-yellow bands on the wing upperside and a well developed band on the hind wing underside, composed of white spots (Korshunov, Gorbunov, 1995). In general, the butterflies of the latter subspecies are very variable in shape and expression of the fulvous band and ocelli on the wing upperside, as well as of the wing underside pattern. So, the specimens from Kamchatka, described as E. l. kamtschadalis Goltz, 1932 are within the range of variation of E. l. eumonia.

Erebia ligea sachalinensis Matsumura, 1911

Erebia ligea, var. sachalinensis Matsumura, 1911, Thousand Insects Japan. (Addit.) 3: 525, pl. 37, figs. 4-4a.

Erebia ligea sachalinesis: Hori and Tamanuki, 1937, Rep. Saghalien Cent. exp. Stn 19: 126-128, pl. 3, f. 2; Kurentzov, 1970, Butterflies Far East USSR: 62-63, fig. 60-2; Kogure and Iwamoto, 1992, Yadoriga (150): 17, 18, pl. 1, fig. 5.

Erebia ligea, ab. murasei Matsumura, 1928, Insecta matsum. 2: 195 [infrasubspecific; unavailable].

Erebia ligea takanonis: Tuzov, 1993, Syn. List Butterflies ex-USSR: 31; Korshunov and Gorbunov, 1995, Butterflies Asian Part Russia: 122.

Type locality: "Saghalien (Motodomari)" [now - most probably, settlement Vostochnyi, Makarovskii district in Sakhalin, Russia].

Type material: not studied.

Materials. Russia: the Sakhalin: 8 1 , no more precise locality (Suprunenko leg.); 3 , Yuzhno-Sakhalinsk vicinity, 20-30.VII (A. Basarukin leg.).

Range. Russia: Sakhalin.

Butterflies of this subspecies rather resemble those of the above described one by the wing upperside, but the fulvous band being much darker, near to the extent observed in E. ligea kamensis. The character is well seen from the nice colour photograph in the paper of Kogure and Iwamoto (1992: pl. 1, fig. 5). The most remarkable difference consists in blue, not white, colour of the pupils in the ocelli, if they are present. This character is shared by subspecies from Sakhalin (ssp. sachalinensis) and Hokkaido (ssp. rishirizana).

Erebia ligea rishirizana Matsumura, 1928 (Figs 10-11)

Erebia ligea rishirizana Matsumura, 1928, Insecta matsum. 2(4): 195; Kawazoe and Wakabayashi, 1977, Color. Illust. Butterflies Japan: 273, pl. 60, fig. 2a-2b; Kurentzov, 1970, Butterflies Far East USSR: 63.

Erebia ligea rishirizana ab. daisetsuzana Matsumura, 1928, Insecta matsum. 2(4): 195 [infrasubspecific; unavailable].

Erebia ligea daisetsuzana Kurentzov, 1970, Butterflies Far East USSR: 63.

Erebia ligea takanonis, ab. junsaiensis Matsumura, 1928, Insecta matsum. 2: 194[infrasubspecific; unavailable].

Type locality: Oshidomari, Rishiri Is., Japan.

Type material: not studied.

Materials. Japan, Hokkaido: 1 1 , Kato-gun, Kamishihoro-cho, Mt. Nishikumanoshiri, 28.VII.1994 (K. Maeda leg.).

Range. Japan: the Rishiri Island, the mountains of the central part of Hokkaido (Fukuda et al., 1984).

This subspecies is similar to the previous one by the wing upperside, but the fulvous band is much lighter, as in E. ligea eumonia. The difference again consists in blue, not white, colour of the pupils in the ocelli.

Erebia ligea takanonis Matsumura, 1909 (Figs 12-13)

Erebia ligea takanonis Matsumura, 1909, Ent. Z. 23: 91; Warren, 1936, Monograph Genus Erebia: 52; Kawazoe and Wakabayashi, 1977, Color. Illust. Butterflies Japan: 273-274, pl. 60, figs. 2d, 2e?, 2f?.

Erebia ligea kisokomana Murayama, 1964, Z. wien. entomol. Gess. 49: 37, pl. 4, figs. 10-13. Kawazoe and Wakabayashi, 1977, Color. Illust. Butterflies Japan: 273-274, pl. 60, fig. 2c.

Type locality: Honshu (Yatsugatake 7000 Fuss).

Type material: not studied.

Materials. Japan, Honshu, 1 1 , Yamanashi Pref., Kitakoma-gun, Oaza-mura, Mt. Yatsugatake, Daimnsawa, 6. VIII. 1979 (M. Hosotani leg.)

Range. Japan: the mountains of the central part of Honshu (Fukuda et al., 1984).

It is characterized by a very wide and light fulvous band on the wing upperside. E. l. kisokomana Murayama was described also from Honshu (Kisokomagadake Range in Nagano). It is very similar to E. l. takanonis. But, according to the original diagnosis, in this subspecies the fulvous band of the fore wing is even more widened, as compared with takanonis, which leaves a narrower brownish-black outer margin. Based on the available to us illustrations of E. l. kisokomana (Murayama, 1964: T. 5, f. 10-13; Kawazoe and Wakabayashi, 1977: T. 60, f. 2c) and E. l. takanonis (Kawazoe and Wakabayashi, 1977: T. 60, f. 2d) as well as the materials studied we consider that the differences between them are within the rank of infrasubspecies variation.

Acknowledgements

The authors express their sincere gratitude to Mr. H. Yoshimoto (Tokyo, Japan) for useful discussion and for providing us with a photo of the type species of E. l. koreana and a number of very important literature sources. They are also much grateful to Mr. Kazunobu Maeda (Sapporo, Japan) for specimens of E. l. rishirizana from Hokkaido and E. l. takanonis from Hoshu, to Dr. E. A. Beljaev (Vladivostok, Russia) for a search for a specimen which could be designated as the lectotype E. ajanensis arsenjevi in Kurentzov's collection at the Institute of Biology and Pedology, Vladivostok and to Dr. Midori Kogure for his reprints on Erebia problem. Besides, we are indebted to all the colleagues who collected the materials comprising the base of the present work. The work by Dr. A. L. Lvovsky was supported by the Russian Grant GNTP, "Bioraznoobrazie", No 76 of 1997.

REFERENCES

Azarova, N. A., 1986. Tipovoi material A. I. Kurentsova po bulavousym cheshuekrylym (Lepidoptera, Rhopalocera) v kollektsii Biologo-pochvennogo instituta DVNTs AN SSSR [The type materials of A.I. Kurentzov on Lepidoptera, Rhopalocera in the collection of the Institute of Biology and Pedology, Far-East Scientific Centre of Acad. Sci. of USSR]. Sistematika I ekologiya cheshuekrylykh Dal'nego Vostoka SSSR [Systematics and Ecology of Lepidoptera of the Far East of USSR]: 121-128. DVNTs AS USSR, Vladivostok (in Russian).

Doi, H., 1919. A list of butterflies from Korea. Chosen Iho 58: 115-128; 59: 90-92.

Eiffinger, G., 1907. 12. Gattung: Erebia Dalm. In Seitz, A. [Ed.], Die Gro▀ -Schmetterlinge fer Erde. 1: 94-114, pls 35-37. Stuttgart.

Fukuda, H., Hama, E., Kuzuya, T., Takahashi, A., Takahashi, M., Tanaka, B., Tanaka, H., Wakabayashi, M. and Y. Watanabe, 1984. The life histories of butterflies in Japan. Vol. 4, XXII+373 pp., 64 pls. - Hoikusha, Osaka. (in Japanese).

Goltz, [H. D.], 1933. Alte und neue Erebienformen. Dt. Ent. Z. Iris. 47: 114-116.

Graeser, L. 1888. Beitrage zur Kenntniss der Lepidopteren-Fauna des Amurlandes. Berl. ent. Z. 32: 33-105.

Hori, M. and K. Tamanuki, 1937. Insect-fauna of Saghalien. Pt. I. Butterflies. Lepidoptera-Rhopalocera. Rep. Saghalien Cent. exp. Stn 19, 12, 224 pp., 8 pls. (In Japanese).

Kawazoe , A. and M. Wakabayashi, 1987. Coloured illustrations of the butterflies of Japan. 423 pp., 72 pls. Hoikusha, Osaka (in Japanese).

Kogure, M., 1985. Some problems about the Genus Erebia. Yadoriga (122): 2-18 (in Japanese).

Kogure, M. and Y. Iwamoto, 1992. Illustrated catalogue of the genus Erebia in Color (I). Yadoriga (150): 2-33 (in Japanese).

Kogure, M. & Y. Iwamoto, 1993. Illustrated catalogue of the genus Erebia in Color (II). Yadoriga (154): 2-38 (in Japanese).

Korshunov, Yu. P. 1977. Bulavousye cheshuekrylye (Lepidoptera, Rhopalocera) Mongol'skoi Narodnoi Respubliki, II [Diurnal butterflies (Lepidoptera, Rhopalocera) of the Mongolian People's Republic, II]. Nasekomye Mongolii. 5: 649-681. Nauka, Leningrad. (in Russian).

Korshunov, Yu. and P. Gorbunov, 1995. Dnevnye babochki asiatskoi chasti Rossii. Spravochnik [Butterflies of Asian Part of Russia. A Guide]. 202 pp. Ekaterinburg, (in Russian).

Korshunov, J. P. and V. P. Soljanikov, 1976. Diurnal butterflies (Lepidoptera, Rhopalocera) of the Mongolian People's Republic // Nasekomye Mongolii 4: 403-458. Nauka, Leningrad. (in Russian).

Kosterin, O. E., 1994. Butterflies (Lepidoptera, Diurna) of the Koni Peninsula (Magadan Region). Actias 1: 77-81.

Kurenzov [Kurentzov] A. I. 1950. New species of (Lepidoptera) from Sikhote-Alin. Ent. Obozr. 31(1-2): 243-246 (in Russian).

Kurentzov, A. I., 1970. Bulavousye cheshuekrylye Dal'nego Vostoka SSSR. Opredelitel'. [The butterflies of the Far East USSR]. 163 pp., 14 pls. Nauka, Leningrad. (in Russian).

Matsumura, S. 1909. Die Danaiden und Satyriden Japans. Ent. Z. 23: 91-92.

Matsumura, S. 1919. Thousand Insects of Japan (Addit.). 3.

Matsumura, S., 1928. New butterflies especially from the Kuriles. Insecta matsum. 2(4): 191-201.

Menetries, E., 1857. Enumeratio Corporum Animalibum Musei imperialis Academiae Scientiarum Petropolitanae (Lepid.) Vol. 2. 6+66+112+144 pp., pls 7-14. Petropoli.

Menetries, E., 1859a. Lepidopteres de la Siberie orientale et en particulier des rives de l'Amour. Bull. phys.-mat. Acad. Sci. St.-Petersb. 17: 212-221.

Menetries, E., 1859b. Lepidopteres de la Siberie orientale et en particulier des rives de l'Amour. In Schrenck, L. v., Reisen und Forschungen im Amur-Lande. 2(1): 1-75, pls 1-5.

Murayama, S., 1964. Neue Tagfalterformen aus Japan und Korea. Z. wien. ent. Ges. 49: 35-37, pl. 4.

Sedykh, K. F., 1979. Cheshuekrylye (Lepidoptera, Macrolepidoptera) fauny Kamchatki i prilegayushchikh oblastei [Macrolepidoptera of Kamchatka and neighbouring regions]. Ent. Obozr. 58(2): 288-296 (in Russian).

Tuzov, V. K., 1993. The synonymic list of the butterflies from the ex-USSR. 73 pp. Rosagroservice, Moscow.

Warren, B. C. S., 1936. Monograph of the genus Erebia. 407 p., 104 pls. British Museum (Natural History), London.

Legends to the figures

Figs 1-15. Erebia ajanensis and Erebia ligea.

          

1 - E. ajanensis ajanensis Menetries, female, lectotype of Erebia ajanensis Menetries.

          

2 - E. ajanensis kosterini P. Gorbunov, Korshunov et Dubatolov. Holotype of Erebia kosterini P. Gorbunov, Korshunov et Dubatolov.

          

3 - Ditto. Paratype (allotype) of Erebia kosterini P. Gorbunov, Korshunov et Dubatolov.

          

4 - E. ajanensis arsenjevi Kurentzov, , Primorye, Oblachnaya Mt.

          

5 - Ditto. , Primorye, Oblachnaya Mt.

          

6 - Ditto. , North Korea, Paekimjun.

          

7 - E. ligea eumonia Menetries. Lectotype of Erebia eumonia Menetries.

8 - Ditto. Lectotype of E. ligea koreana Matsumura.

9 - Labels of lectotype of E. ligea koreana Matsumura.

          

10 - E. ligea rishirizana Matsumura. , Hokkaido, Mt. Nishikumanoshiri.

          

11 - Ditto. , Hokkaido, Mt. Nishikumanoshiri.

          

12 - E. ligea takanonis Matsumura. , Honshu, Mt Yatsugatake.

          

13 - Ditto. , Hokkaido, Mt. Nishikumanoshiri.

          

14 - E. ligea eumonia Menetries. Paralectotype of Erebia eumonia Menetries.

          

15 - E. ajanensis arsenjevi Kurentzov. Paralectotype of Erebia eumonia Menetries.

Fig. 16 - A distribution map of E. ajanensis and E. ligea.

Fig. 17 - A scheme of the wing pattern of E. ligea (a) and E. ajanensis (b). Numerals correspond to the specific characters in the table.

Figs 18-24. Male genitalia structure of Erebia ajanensis and Erebia ligea.

18 - E. ajanensis kosterini P. Gorbunov, Korshunov et Dubatolov. Holotype of Erebia kosterini P. Gorbunov, Korshunov et Dubatolov.

19 - E. ajanensis arsenjevi Kurentzov, Primorye, Dal'negorsk.

20 - E. ligea eumonia Menetries, Buryatia, Northern Transbaikalia, Kichera.

21 - ditto, Khabarovskii Krai, Nature Reserve "Komsomol'skii".

22 - E. ligea ssp. (after: Kawazoe , Wakabayashi, 1976: 268).

23 - E. ligea eumonia Menetries, (after Kurentzov, 1970: 61, fig. 59-6. E. ligea ajanensis sensu Kurentzov).

24 - E. vidleri Elwes (after Warren, 1936: pl. 30, fig. 292).

Fig. 25-36. Valvae of E. ajanensis and E. ligea:

25-30 - E. ajanensis arsenjevi Kurentzov, Primorye, the Sokolovka River;

31-32 - Ditto, North Korea, Paekimjun;

33 - E. ligea rishirizana Matsumura, Hokkaido, Mt. Nishikumanoshiri;

35 - E. ligea eumonia Menetries. Lectotype of Erebia eumonia Menetries.

36 - Ditto. Paralectotype of Erebia eumonia Menetries.

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