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NOTE: the text corresponds to Korshunov & Gorbunov, 1995) only.


FAMILIA NYMPHALIDAE Swainson, 1827



  The butterflies of intermediate or, less frequently, small or 
large size, with f.w.l. of 13-48 mm. In colouration red and 
broun colours usually predominate. The fore legs are reduced to 
the so-called "brushes" and useless for walking. The veins at 
the wing bases are never swallen. Imagines of some species are 
capable of far migrations. 
  The larvae usually bear spines or processes; sometimes they are 
gregarious, especially in earlier instars. The pupae are angular 
or with some prominences, they are always suspended 
head-downwards by the tail (with the aid of hooks on the 
cremaster).
  The family is distributed throughout the world and embraces 
about 2500 species, mostly tropical, somewat more than 100 
species inhabit the Asian Russia. 


SUBFAMILIA APATURINAE Tutt, 1896


GENUS SEPHISA Moore, 1882.
Type species: Limenitis dichroa Kollar, [1844].

  The genus includes three species from E. and S. Asia. 


112. Sephisa dichroa (Kollar, 1844).

TYPE LOCALITY: the West Himalaya.

RANGE: S Primorye, Korea, Central, East, South-East and South 
China, the Hymalaya. A local species. 

HABITAT: primary fir/broad-leaved mixed or oak forests on hill 
slopes; only some imagines penetrate into the valley broad-
leaved forests, where the males can congregate on river and 
brook banks and at pools on roads. The females keep to tree 
canopy and rarely descend to the ground. 

FLIGHT PERIOD: milddle July/late August. 

PREIMAGINAL PHASES: studied in Primorye (A. Danchenko). 
Foodplants: Quercus mongolica. Larva: gren with two brounish-red 
processes, set with sharp spinules, on the head 

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-42 mm. Wing upperside ochre-orange
in males and whitish in females, with an ornament of black spots of 
different sizes; fore wing outer margin substantially concave. 

GEOGRAPHICAL VARIATION: Primorye is inhabited by ssp. princeps 
Fixsen, 1887. 


GENUS DILIPA Moore, 1857.
Type species: Apatura? morgiana Westwood in Doubleday, [1850]

  The genus includes two species from E Asia.

- Dilipa fenestra (Leech, 1891).

TYPE LOCALITY: China: Omei-Shan.

RANGE: E and NE China, Korea. On 23rd April 1991 at the town 
Ussurisk (S Primorye) a fresh individual of this species, 
probably a migrant, was observed among a group of Roddia l-album 
(Masui, Inomata, 1993). 

PREIMAGINAL PHASES: Foodplant: Celtis, which is absent from Primorye.

PECULIAR TRAITS OF IMAGO: F.w.l.: 30-38 mm. Wing upperside is 
ochre-orange, with a dark border and separate spots, in males 
and brown with yellow and red spots in females. On hind wing 
underside there is a narrow straight stripe going from midle 
fore margin to anal angle. 


GENUS AMURIANA Korshunov et Dubatolov, 1984.
Type species: Adolias schrenckii Menetries, 1859.

  A monotypical genus confined to the Amur basin.


113. Amuriana schrenckii (Menetries, 1859).

TYPE LOCALITY: Priamurye: the mountains of Bureya.

RANGE: Priamurye (from the Bureya to the Goryun Rivers), 
Primorye, the neighbouring regions of NE China. 

HABITAT: polydominant broad-leaved and mixed montane forests up 
to the dark-needle taiga belt; the butterflies keep to river and 
brook banks, roads, edges, and open stand of trees. 

FLIGHT PERIOD: early July/late August or early September. 

PREIMAGINAL PHASES: Foodplants: in Priamurye - Ulmus propinqua 
and U. laciniata (Greaser, 1888), in Priamurye - Carpinus 
cordata (Kurentzv, 1970). Larva: light-green, with three pairs 
of dorsal processes and two caudal spines. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 42-50 mm. Wing upperside is 
dark-brown, almost black, with white spots not fused into bands; 
hind wing underside has two vast light-blue areas on a brick-red 
background. 

ETYMOLOGY: Leopold Ivanovich Schrenk (1826-1894) - a famous Russian 
naturalist, botanist, and zoologist, an explorer of the Far East. 


GENUS APATURA Fabricius, 1807.
Type species: Papilio iris Linnaeus, 1758.

  F.w.l.: 30-47 mm. Wing upperside of brown tones with white or 
fulvous spots and bands, in males with a strong iridescent 
violet flush; fore wing outer margin slightly concave; hind wing 
has an oclleus at anal angle. 
  Eggs: greenish, hemispheric, ribbed lengthwise; laid singly or 
by small batches on leaves, branches, or trunks of the 
foodplants, which are Salicaceae trees. Young larva is
dark-coloured, it lives in a shelter made out of silk-spun leaves 
and then hibernates. Last instar larva is green with a peculiar 
pattern of yellowish slanting streaks on sides; head 
bears a pair of horny processes; end of the body with two 
spikes. Pupa: greenish, smooth, its head and tail ends being 
similarly pointed, there are two small horns on head; usually 
it is  suspended on a leaf of the foodplant. Imagines, especially 
males, are attracted by rotting organic residues, such as 
carrion, excrements, fallen fruits, where they often congregate 
into small swarms. In sunny weather they are also gathered on the 
damp ground. Females keep to the tree canopy ans so are seldom 
seen. 
  A Palaearctic genus including about 10 species; our species have 
similar ranges in the temperate Eurasia with large gaps in the 
central part of the continent.


114. Apatura iris (Linnaeus, 1758).

TYPE LOCALITY: Germany, Egland.

RANGE: Europe, Middle and South Ural, Tyumen' Region 
(Yarkovskii, Tobol'skii and Tyumenskii Districts) and, after a 
tremendous gap, E Zabaikalye, Priamurye, Primorye, Central and 
North-East China, Korea, Japan. 

HABITAT: willow stands in various type woods. 

FLIGHT PERIOD: late June/middle September.

PREIMAGINAL PHASES: Foodplants: Salix (S. caprea and others), 
Populus. Eggs are laid on leaf upperside, up to 4 m above the 
ground. Larva: green, covered with yellowish dots, with two red 
spikes on rear segment and light-green horns on head. The four 
fore segments with lengthwise yellow streak on either side, 
further there are five slanting streaks of the same colour, the 
first of which being the longest. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 33-38 mm in males, 36-47 mm in 
females. Spots and bands on fore wing upperside are usually 
white, in Far-Eastern specimens often yellowish; outer edge of 
white band on hind wing has a sharp tooth. 

GEOGRAPHICAL VARIATION: In the western part of the range the 
nominotypical subspecies is distributed, while ssp. amurensis 
Stichel in Seitz, 1908 inhabits the eastern part. 


115. Apatura ilia (Denis et Schiffermuller, 1775).

TYPE LOCALITY: Austria: Vienna.

RANGE: Europe, south Ural, and then E Zabaikalye, Priamurye, 
Primorye, Sakhalin, S Kuriles, Central and NE China, Korea, 
Japan. 

HABITAT: river and brook banks, wood edges, open poplar/willow, 
broad-leaved and mixed forests, riparian bird cherry/willow 
thickets. 

FLIGHT PERIOD: middle June/middle September.

PREIMAGINAL PHASES: Foodplants: Populus tremulae, P. nigra, 
Salix. Larva: resembles A. iris but lateral streaks margined 
with red, head horns longer, black or brownish at front side. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-41 mm. On wing upperside 
spots and bands are white or yellowish (in an European form 
clyte Denis et Schiffermueller, 1775  and form praeclara 
Moltrecht, 1927 in the Far East). On hind wing the white band 
outer margin straight or smoothly curved. 

GEOGRAPHICAL VARIATION: The western part of the range is 
inhabited by the nominotypical subspecies, in the southern Far 
East there ranges ssp. ussuriensis Kurentzov, 1937, difering by 
on average more stretched out and pointed fore wings and usually 
well expressed fulvous spots along outer margin on hind wing 
upperside. 


116. Apatura metis Freyer, 1829.

TYPE LOCALITY: Hungary.

RANGE: SE Europe, the south of W Siberia (the Irtysh valley, 
from the village Kachiry in the Pavlodar Region of Kazakhstan to 
the city Tobol'sk), and, after a gap, SE Zabaikalye, Priamurye 
(from the Zeya to  the Goryun Rivers), Primorye, NE China, 
Korea, Japan. 

HABITAT: willow and poplar riparian associations. 

FLIGHT PERIOD: middle July/late August. 

PREIMAGINAL PHASES: Studied in Japan (Fukuda et al., 1982). 
Foodplants: Salix and Populus. Larva hibernates at the third 
instar in bark crevices on foodplant trunks. Pupa: light-green 
with yellowish streaks along wing case margins and along a 
convex ventral side. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 26-39 mm. All spots and bands 
on wing upperside are ochre-yellow or fulvous; on hind wing the 
band outer edge as a rule with blunt fractures or straight. 

GEOGRAPHICAL VARIATION: ssp. irtyshika Korshunov, 1982 ranges in 
West Siberia, it differ from East-European butterflies by wider 
fulvous submarginal bands; ssp. substituta Butler, 1873 inhabits 
E Transbaikalia and the southern Far East, it is characterized 
by a darker ground colour, a distinct pattern on hind wing 
underside, and a well expressed black ocellus at the anal angle. 
In Priamurye and Primorye A. metis tends to form local forms to 
some extent approaching A. ilia. According to V.V. Dubatolov's 
opinion, it was such forms which have been described as the 
following subspecies of other species: abramovi Kurentzov, 1970, 
described as subspecies of A. iris from the Lower Priamurye (the 
Goryun River) and the coastal regions of the Sikhote-Alin' 
mountain chain (the Samarga River), and krylovi Kurentzov, 1937, 
described as subspecies of A. ilia from the northern coastal 
regions of th e Sikhote-Alin'. A form with much widened fulvous 
bands is not frequent in S Primorye. From the Kunashir the taxon 
doii Matsumra, 1928 has been descried. 


GENUS ATHYMODES Moore, [1896].
Type species: Atyma nycteis Menetries, 1859.

  A monotypical East-Asiatic genus.


117. Athymodes nycteis (Menetries, 1859).

TYPE LOCALITY: the Amur.

RANGE: Priamurye (from the Shilka and Argun' Rivers junction 
downstream to the Goryun River), Primorye, NE China, N Korea. 

HABITAT: valley broad-leaved forests and montane mixed forests 
with participation of the elm. 

FLIGHT PERIOD: middle July/middle August. 

PREIMAGINAL PHASES: Foodplants: Ulmus propinqua (Kurentzov, 
1970). Larva: greyish-brown with three pairs of triangular white 
spots on the bak; head with long processes of the ground colour 
. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 32-39 mm. Wing upperside dark-
brown with white bands; wing underside brownish, there is no 
ocellus at hind wing anal angle; fore wing outer margin slightly 
concave. 



SUBFAMILIA LIMENITINAE Butler, 1869.


GENUS SEOKIA Sibatani, 1943.
Type species: Limenitis pratti Leech, 1890.

  A monotypical East-Asiatic genus.


118. Seokia pratti (Leech, 1890).

TYPE LOCALITY: Central China.

RANGE: The Sikhote-Alin' mountain chain northwards to the Terney 
District (the Shumnyy spring), N, E and Central China, N Korea. 
A local species. 

HABITAT: wide and windless valleys in the belt of montane Korean 
stone pine/broad-leaved mixed forests at 400-1000 m above sea 
level on smooth elevations. Imagines have a strong flight; 
females mostly keep to tree crowns. 

FLIGHT PERIOD: prolonged, middle July/September. 

PREIMAGINAL PHASES: studied in Primorye by M.M. and M.A. 
Omel'ko. Foodplant: Pinus koraiensis. Eggs: roundish, ochre-
coloured, fine dimpled; laid singly on the needle tips in lower 
and middle parts of crowns of the Korean stone pine trees of 
different ages. The larva hatches on 10-12 day, it is ochre-
coloured with a black glossy head and longitudinal rows of short 
bristles. It eats the pine needles in the evening or at night; 
after moults it eats its left skins. The larva feeds until 
middle November, hinernates at the third instar, continues 
feeding on the first half of April and pupates in early June on 
branches in the close vicinity of the place of feeding. Mature 
larva: up to 40 mm long, with dark-brown sides and a dark-sandy 
back; the 2nd, 3rd, 5th, 7th, 9th, and 11th segments dorsally 
bear pairs of spines 3.5-4 mm long; the other segments - pairs 
of prominences about 1 mm; the head is olive-grey with a pair of 
short horns. It camuflates well on a pine branch. Pupa: about 27 
mm long, dark-grey with a darker head and the two fore segments 
of the thorax; the 4th-6th abdominal segnments bear dorsally 
pairs of button-like tubercles. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 30-37 mm. Wing upperside dark-
brown with a transversal row of red spots and,proximaly of it, a 
row of white ones; hind wing underside has a narrow red 
postdiscoidal band. 

SYTEMATIC NOTE: Primorye is inhabited by ssp. eximia Moltrecht, 
1909 (= Ussuriensis efremovi Nekrutenko, 1960). Its 
conspecificity to S. pratti was stated by Prof. T. Fujioka. 



GENUS LIMENITIS Fabricius, 1807.
Type species: Papilio populi Linnaeus, 1758.

  F.w.l.: 22-45 mm. Wing upperside dark-brown, almost black, 
with white bands. Fore wing with discoidal vein. Females larger 
than males, with white marking wider. Flight period usually in 
middle summer, in our climate in a single brood. Similarly to 
Apatura species, these butterflies are attracted by rotting 
organic matter and, in hot weather, by wet ground, where they 
form small swarms. In the majority of species males are seen 
more frequently than females, which behave more secretly. 
  The larvae of our species develop on Lonicera, Spiraea, less 
frequently on Salicaceae, hibernate at the third or fourth 
instar. Young larvae are dark, at the last instar as a rule 
green with paired processes or the back. Pupae bear a blunt 
prominence on the 2nd abdominal tergte and a pair of short horns 
on head. 
  A Holarctic genus with about 50 species, mostly in E Asia.


119. Limenitis populi (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia, in Siberia as north as the middle 
taiga belt. 

HABITAT: river banks, wood edges and open stands of different 
kinds of forests. The butterflies migrate actively and so may 
occur in various open landscapes. 

FLIGHT PERIOD: late June/early August. 

PREIMAGINAL PHASES: Foodplants: Populus - Populus tremula, P. 
nigra, P. alba reported for S Ural, P. tremula - for Siberia, P. 
amurensis, P. koreana, P. maximoviczi - for the Far East, rarely 
Salix. Eggs: greenish, covered with tiny light knobs and hairs, 
laid singly, usually at the tips of the foodplant leaves about 
2-4 m above the ground. Youg larva: brown with tiny warts o each 
segment. It lives in a semi-rolled leaf, with edges drawn 
together by slik threads so that the resulted tube is opened at 
the leaf base. In autumn the larva attaches the leaf to the 
hibernating bud and overwinter in it. Mature larva: light- or 
dark-green with large brown areas and two rows of fleshy 
tubercles set with short hairs on 2nd, 3rd, 5th, 7th and 11th 
segments, besides, a pair of much greater spiny processes on the 
mesothorax protrudes forwards of the brown head. Pupa: 
yellowish-brown with dark-grey brands of different sizes and a 
glossy orange drop-like projection on the back; it usually hangs 
on a central vein of a semi-rolled leaf or on thin twigs. Its 
phase lastst for 10-14 days 

PECULIAR TRAITS OF IMAGO: F.w.l.: 33-45 mm. Hind wing upperside 
has a submarginal row of orange-red lunular spots; the wing 
underside ground colour is brick-red; white bands sometimes much 
reduced (f. tremulae Esper, 1798). 

GEOGRAPHICAL VARIATION: Butterflies from Ural and Siberia, 
attributed to ssp. enapius Fruhstorfer, 1908, described from 
Altai (?= eumenius Fruhstorfer, 1908) difer from European ones 
by a somewhat widened white pattern and expansion of dark and 
greyish-blue elements over fulvous ones on hind wing underside. 
Ssp. ussuriensis Staudinger, 1887 is known from Primorye and 
Priamurye, which differs from the former only by some further 
widening of the white pattern. 


120. Limenitis moltrechti Kardakov, 1928.

TYPE LOCALITY: Primorye: the Narva Bay.

RANGE: Middle Priamurye (not upstream of the Minor Hinggan 
Mts.), Primorye, adjacent regions of NE China and N Korea. A 
local species. 

HABITAT: edges and open stands in mixed and decisuous forests 
with a rich bush understorey. 

FLIGHT PERIOD: middle July/middle August. 

PREIMAGINAL PHASES: Foodplants: Lonicera praeflorens, L. 
chrysantha (Kurentzov, 1970). The larvae were observed from late 
May to late June. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 28-35 mm. On fore wing 
upperside cell contains one elongate white spot; hind wing 
upperside with a submarginal band composed of light lunules. 

ETYMOLOGY: A.K. Moltrecht- an amateur entomologist who travelled 
in Primorye in 1903-1930. 


121. Limenitis sydyi Kindermann in Lederer, 1853.

TYPE LOCALITY: W Altai: the settlement Ust'-Bukhtarminsk.

RANGE: W Altai: the upper Irtysh basin upstream of the city 
Ust'-Kamenogorsk, on Russian territory reported for the environs 
of the town Zmeinogorsk and the settlement Kolyvan, then, after 
a great range gap, E Zabaikalye, Priamurye, the southern ranges 
of the mountains of Bureya, Primorye, Central and NE China, 
Korea. 

HABITAT: in Altai, where this species is rare: book valleys and 
mountain slopes at 200-800 m above sea level with honeysuckle 
shrubbery, from where the butterflies often penetrate to open 
places. At hot daytime they keep to wet round, rest on stones 
and bushes, their feeding was observed on Spiraea crenata, 
Heracleum dissectum, etc. In the southern Far Eas: montane oak 
woods, valley broad-leaved, mixed or, rarely, small-leaved 
forests. The imagines often visit the flowers of Sorbaria 
sorbifolia and Spiraea. 

FLIGHT PERIOD: middle June/middle July in W Altai; early 
July/middle August in the Far East. 

PREIMAGINAL PHASES: studied in W Altai by P. Gorbunov and Y. 
Shevnin. Foodplants: Lonicera tatarica in W Altai; in the Far 
East Spiraea media was reported (Kurentzov, 1970).  Mature 
larva: light breen with a wide dark-green band along the back 
and a pattern of blak dots and strokes; there are 10 lengthwise 
rows of branchy spines: three rows of light-coloured spines on 
either side and four rows on the back: two external ones consist 
of long (6-7 mm) dark spines, two internal - of short spinules. 
Head light-brown with black eyes and a black triangle between 
them; set with light spinules, head top bears longer black 
spines. Thoracic legs black. Pupa: 22 mm long, nacreous with 
four rows of black spots on abdominal segments and along outer 
margins of wing cases; placed on thin foodplant branches. Its 
phase lasts for 10-12 days. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 26-31 mm. On fore wing 
upperside cell  either contains a diffuse elongate white spot or 
lacks it; in Far Eastern specimens white band on hind wing about 
5 mm wide. 

GEOGRAPHICAL VARIATION: ssp. sydyi is known from W Altai, iIn 
the Far East there ranges ssp. latefasciata Menetries, 1859 with 
a much widened white pattern. 


122. Limenitis camilla (Linnaeus, 1763).

TYPE LOCALITY: Germany.

RANGE: Europe, S Ural (the basins of the Belaya and Sakmara 
Rivers) and, after a tremendous gap, Middle and Lower Priamurye, 
Primorye, the Sakhalin, the Kunashir, NE China, Korea, Japan. 

HABITAT: in South Ural: locally on wood edges and open slopes 
with the honeysuckle bushes; in the Far East: mixed and valley 
broad-leaved forests. 

FLIGHT PERIOD: middle June/middle July in South Ural, middle 
July/late August in the Far East. 

PREIMAGINAL PHASES: studied in Europe (Buckler, 1886; other 
authors). Foodplants: various species of Lonicera: L. tatarica, 
L. xylostelum reported for S Ural (Migranov, 1991); L. maackii, 
L. gibbiflora for the Far East (Kurentzov, 1970). Eggs: pale-
green with 5-6-angled facets, beset with tiny hairs; laid singly 
on foodplant leaf pperside. Larva at the first three instars is 
brown and carries a peculiar shelter made out of its frass. Four 
instar larva: green, speckled with numerous white marks, with a 
light spiracular streak containing yellow spots on either side; 
on the back it bears two rows of spiny processes, black on 
thorax and brown on abdomen, those on segments 2, 3, 5, 10, and 
11 being mc larger; head brown, with minute spines. Pupa: green 
or olive-brown with brown spots and metallic brands, it bears a 
large blueish projection on the back of the first abdominal 
segment and a smaller one of the same colour on the head; thorax 
is humped on the back in front of it. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 23-30 mm. On fore wing 
upperside white band split at the middle owing to absence of a 
spot in one space; cell contains no white spot or has a small 
whitish spot at discal vein. 

GEOGRAPHICAL VARIATION: the nominotypical subpecies ranges in 
the western part of the range; ssp. japonica Menetries, 1857 (= 
angustata Staudinger, 1887), differing by a narrower white 
pattern, inabits the southern Far East, the Sakhalin and 
Kuriles. 


123. Limenitis amphissa Menetries, 1859.

TYPE LOCALITY: the mountains of Bureya.

RANGE: Priamurye (from the Zeya to the Goryun River), Primorye, 
Central and NE China, Korea. 

HABITAT: valley broad-leaved and mixed forests. 

FLIGHT PERIOD: late June/middle August.

PREIMAGINAL PHASES: Foodplants: Lonicera maakii is reported 
(Graeser, 1888). The larvae were observed in May and June. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 25-32 mm. On fore wing 
upperside contains two white spots, distal of which stretched 
lengthwise; on hind wing underside there are no black spots at 
wing base. Similar species: Limenitis helmanni, L. doerriesi. 


124. Limenitis doerriesi Staudinger, 1892.

TYPE LOCALITY: Primorye.

RANGE: Primorye, N China, N Korea. A local species. 

HABITAT: open stands in montane mixed and deciduous forests, 
sometimes at rock outcrops. 

FLIGHT PERIOD: July-August. 

PREIMAGINAL PHASES: Foodplants: Lonicera praeflorens is known 
(Kurezov, 1970). 

PECULIAR TRAITS OF IMAGO: F.w.l.: 23-32 mm. On fore wing 
upperside cell conains two white spots, area between them 
noticeably suffused by red scales; white bands wider than in L. 
helmanni, about 5 mm on hind wing in males; hind wing underside 
with black spots at wing base, as diffrent from L. amphissa. 

ETYMOLOGY: Fritz Doerries (1851-1949), a naturalist from Hamburg 
who, together with two his brother, collected butterflies in 
Primorye, Priamurye, and Zabaikalye in 1877-1886. 


125. Limenitis helmanni Kindermann in Lederer, 1853.

TYPE LOCALITY: W Altai: the settlement Ust'-Bukhtarmisk.

RANGE: W and N Altai Mts., the so-called Kuznetskoe upland, 
including almost lowland area at the upper Ob' River right bank 
at Barnaul, Tal'menka, Novosibirsk, the mountains of East 
Kazakhstan (westwards to the North Tien-Shan), E Transbaikalia, 
the southern ranges of the mountains of Bureya, Primorye, 
Central and NE China, Korea, reported for Japan. 

HABITAT: in the western part of its range the species 
locally occur at riparian thickets of honeysuckle, bushy hills 
and mountain slopes, on meadows in relic  forest associations 
with the participation of linden (Tilia sibirica); in the Far 
East these butterflies inhabit various biotopes, mostly mixed and 
deciduous forests, reaching the tree-line in the mountains. 

FLIGHT PERIOD: late June/late July or early August in the 
western range,  early July/early September in the Far East. 

PREIMAGINAL PHASES: Foodplants: Lonicera tatarica in Altai, L. 
maacki in the Far East (Graeser, 1888). According to 
observations of P. Gorbunov in W Altai, mature larva resembles 
that of Limenitis camilla, it is bright-green with greenish-
white stripe on either side of abdomen and whitish-green 
prolegs; segments 2, 3, 5, 10, and 11 bear light-green or 
reddish spiny processes 2-3 mm long; head whitish with four 
vertical reddish-brown stripes, set with minute spinules two of 
which, above eyes, ared ark and longer. For ouoation the larva 
usually attaches itself on leaf underside to the main vein. 
Pupa: 19 mm long, green, with stripes on either sides of the 
abdomen, drop-like tubercle on the back and horns on the head 
reddish. Its phase lasts for 10 days. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 24-32 mm. Cell on  fore wing 
upperside contains two white spots and no red suffusion; white 
bands rather narrow, about 3-4 mm on hind wing in males; hind 
wing underside with black spots at wing base. Similar species: 
Limenitis doerriesi, L. amphissa. 

GEOGRAPHICAL VARIATION: ssp. helmanni ranges in South Siberia; 
ssp. duplicata Staudinger, 1892 - in the southern Far East. The 
taxon pryeri Moore, 1877 refers to the Chinese fauna. 


126. Limenitis homeyeri Tancre, 1881.

TYPE LOCALITY: the Amur.

RANGE: Middle and Lower Priamurye (from the Bureya to the Goryun 
Rivers), Primorye, Central and NE China, Korea. Rather a rare 
species. 

HABITAT: intermontane valleys and open tree stands in montane, 
the Korean stone pine/broad leaved or dark-needle, forests. 

FLIGHT PERIOD: late June/early August. 

PREIMAGINAL PHASES: Foodplants: Lonicera. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 22-29 mm. Cell on fore wing 
upperside contains two white spots; white bands are narrow, 
about 1.5-2 mm wide on hind wing in males, band on fore wing 
split into separate spots. 

ETYMOLOGY: E.F. Homeyer - a zoologist, explorer of the Far East, 
E and S Sibiria. 


GENUS NEPTIS Fabricius, 1807.
Type species: Papilio aceris Esper, [1783].

  F.w.l.: 22-45 mm in our species. Wings substantially elongate, 
their upperside dark-brown, almost black, with white or yellow 
bands and separate spots; fore wing has no discoidal vein, its 
outer margin usually slightly convex. Similarly to Limenitis 
species, in hot weather these butterflies congregate on damp 
places on banks or roads in the woods and are attracted by 
rotting organic matter and flowing out tree sap, and, in a hot 
weather, by wet ground, where they form small swarms. The 
butterflies have a peculiar soaring mode of flying. 
  Eggs laid singly, usually on leaf tips of the foodplants. The 
larva lives solitarily in a shelter made out of a leaf as 
follows: it eat the incisions from either side of a leaf to the 
central vein and drow the edges of the resulting lobes together 
with silken threads. The larvae hibernate at the  third or 
fourth instars; they are variously coloured and bear pairs of 
processes on 2nd, 3rd, 5th and 11th abdominal segments. Pupa: 
short and stout, with convex wing cases and usually a pair of 
short horns on the head. 
  The genus includes more than 100 species from Eurasia, Africa 
and Australia, the majority of which inhabit East and South 
Asia. 


127. Neptis sappho (Pallas, 1771)
(= aceris Esper, 1783; hylas auct. nec Linnaeus, 1758).

TYPE LOCALITY: Lower Povolzhye [the lower Volga basin].

RANGE: The temperate Eurasia, in Siberia as north as the 
southern taiga belt; Japan. 

HABITAT: meadow patches, open tree stands, wood edges, and river 
valleys in the various type forests, mostly open. These 
butterflies tend to rest on broad leaves of herbs and often 
visit the inflorescences of Apiaceae for feeding. 

FLIGHT PERIOD: early June/middle July, in a single brood, in the 
most part of Siberia; in South Ural and the southern Far East 
two broods develop, late May/late June and July/August, 
respectively. In Primorye the second brood butterflies are 
maller and more numerous. 

PREIMAGINAL PHASES: Foodplants: Lathyrus vernus, less frequently 
other Fabaceae: Vicia, Caragana. Eggs: greenish, covered with 
hairs and white specks, laid on the surface or tips of the 
foodplant leaves. The larvae before hibernation, which occurs 
among the leaf fall, live in leaf-made shelters. Mature larva: 
greenish, brownish-grey or yellowish-brown with a whitish-green 
"saddle" on the back and spiny projections on the 1st, 2nd, 3rd, 
5th and 11th segments. Pupa: short, ground-yellow or yellowish 
with mate golden bloom, with two spots on the head and numerous 
ones on the thorax; wing cases margined by brown line. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 20-27 mm. On fore wing 
upperside there is an even row of small white spots along outer 
margin and two large triangular spots in cell; on hind wing 
there are two well developed bands. 

GEOGRAPHICAL VARIATION: ssp. sappho inhabits S Ural and Sieria. 
Butterflies from the southern Far East are characterized by 
somewhat reduced outer band on hind wing, especially in the 
second brood. Some authors refered them to ssp. intermedia 
Pryer, 1877, described from Central China, presence of which in 
our fauna is doubtful. 


128. Neptis philyra Menetries, 1859.

TYPE LOCALITY: the Amur River: "Marienpost".

RANGE: Middle Priamurye (from the Zeya to the Goryun Rivers), 
Primorye, Central and NE China, Korea, Japan. 

HABITAT: mixed and valley broad-leaved forests. 

FLIGHT PERIOD: middle June/early August. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). 
Foodplants: Acer mono, A. japonicum, Carpinus cordata. Eggs: 
blueish with light and dark dots, laid singly on the foodplant 
leaf tips. Larva: yellowish-brown with four pairs of processes 
on the back. Pupa: contrastly coloured, with a pale-brown thorax 
and abdomen and darker wing cases with conspicuous light veins. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 27-35 mm. On fore wing 
upperside cell contains an even and narrow white stripe, there 
are no white spots in front of it at  costal margin; on hind 
wing submarginal band consists of relatively large white spots. 
Similar species: N. philyroides. 


129. Neptis philyroides Staudinger, 1887.

TYPE LOCALITY: Primorye and the Amur.

RANGE: Priamurye (from the Amursko-Zeyskoe Plateau to the Goryun 
Rivers), the southern ranges of the mountains of Bureya, 
Primorye, NE China, N Korea. 

HABITAT: wood edges and open stands in mixed forests, montane 
oak woods, bushy mountain slopes. 

FLIGHT PERIOD: middle June/middle August. 

PREIMAGINAL PHASES: Foodplants: Corylus manshurica (Kurentzov, 1970). 

PECULIAR TRAITS OF IMAGO: F.w.l.: 28-35 mm. On fore wing 
upperside cell contains an even white stripe, there are two 
small white spots in front of its distal end at costal margin; 
on hind wing submarginal band consists of relatively large white 
spots. Similar species: N. philyra. 


130. Neptis speyeri Staudinger, 1887.

TYPE LOCALITY: Primorye, the Ussuri River.

RANGE: Primorye (northwards along the coast to the Kievka River 
and southwords along the Ussury River to Khabarovsk) and the 
adjacent regions of NE China. In N Primorye this is a rare 
species 

HABITAT: river valleys, clearings and edges in montane mixed forests. 

FLIGHT PERIOD: late June/middle August. 

PREIMAGINAL PHASES: studied in Primorye by M.M. and M.A. 
Omel'ko. Foodplant: Corylus heterophylla. Eggs: sphaerical with 
a complicated knobby sculpture, laid singly at leaf margins. 
Larvae, 2-3 mm long, hatch after 7-10 days. A young larva eats 
the egg horion and then constructs a shelter. For this purpose 
it makes two long cuts from leaf margins to the central (or 
lateral) vein and cuts slightly the latter; then it makes two 
more cuttings on either side of the almost cut-off apical part 
of the leaf, fasten its edges with silk and so forms a shelter, 
which soon fades. The larva eats edges of its shelter for 3-4 
days and moults in it on 9-13th day. Then the larva eats its 
shelter and constructs a new one, this repeats also on 2nd and 
3rd moults so that the larva moves towards the leaf base. In 
August there often rests only a leaf petoile with the shelter on 
it. In September, before hibernation, the larva spins the 
shelter and the petiole with a silk and fasten it to the twig. 
Before hibernation the larva is about 5 mm long, brown with a 
dark head and with processes on both ends of the body. After 
hibernation the larva is at first little active and feeds on 
faded parts of the shelter, then on young buds, but most time it 
spends in the shelter. After hibernation the larva becomes 
light-ochre-coloured with light dashes on sides, that makes it 
similar either to a faded shelter or young pinkish-violet buds. 
In middle May the larva abandons the shelter for leaf upperside, 
where keeps to brown spots, common on leaves of C. heterophylla, 
on which it is hardly noticeable. The last 5th moults occurs on 
leaf upperside in late May. 2-3 days before pupation the larva 
moves to leaf underside, where pupates. Pupa: generally 
nacreous, 19-20 mm, its stage lasts for 13-14 days. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 24-30 mm. On fore wing 
upperside white stripe in cell incised at its fore margin, white 
spots in front of its distal end are absent; on hind wing 
submarginal band obscure, being a diffuse streak disappearing to 
costal edge. 

ETYMOLOGY: Adolf Speyer - a German entomologist of XIX century.


131. Neptis rivularis (Scopoli, 1763)
(= lucilla Denis et Schiffermuller, 1775). 

TYPE LOCALITY: Austria: Carniola.

RANGE: The non-tropical Eurasia northwards to the middle taiga 
belt, the Sakhalin, the Kuriiles, Japan. In 1994 the species was reported for 
the Polar Ural (A. Tatarinov). 

HABITAT: river and brook valleys, bushy mountain slopes, wood 
edges and openings, in the mountains goes as high as 1700 m 
altitude. 

FLIGHT PERIOD: two broods in the forest-steppe zone, in June and 
August; one brood in the forest zone and in the mountains, in 
late Jun/late July. 

PREIMAGINAL PHASES: studied in Altai (Y. Korshunov, P. 
Gorbunov). 
Foodplants: Spiraea, such as S. salicifolia, S. hypericifolia, 
S. crenata, S. aquilegifolia, less frequently Filipendula 
ulmaria, Cotoneaster. Eggs are laid singly on leaf tips on upper 
branches of a foodplant bush. The larva makes cuts from either 
margins, of the leaf 10-15 mm from its tip, to the central vein, 
fasten the edges and so forms a shelter. The larva hibernate the 
3rd instar in such a shelter, having span it with silk to fasten 
to the twig. Mature larva: brownish-grey with a yellowish back, 
lateral stripe on either side, and slanting streaks on segments 
4-11; segments 2, 3, 5, and 11 bear paired processes. Pupation 
occurs on thin branches of the foodplant. Pupa (according to 
observation of O. Kosterin in SE Zabaikal'ye): with two 
broadening in the midle part of the body so that wing case 
margins are much protruding, light-brown with a darker 
reticulate ornament, with a dark brand in the middle of wing 
case and several pairs of dark dots on the back; there is a sarp 
crest along the back, dark in the fore part of the body and 
white with brown rim further; abdomen with darker lengthise 
stripes along ventral side and either lateral side and dark 
slanting streaks on abdominal segment upperside. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 20-29 mm. On fore wing 
upperside cell usually contain either one white spot at its 
distal end or a number of small spots; on hind wing submasrginal 
band absent or there are only traces of it, medial band is 3-5 
mm wide in males; on hind wing underside there is a light brand 
along fore margin; black dots absent. 

GEOGRAPHICAL VARIATION: Ssp. coenobita Goeze, 1779, close to the 
nominotypical and having widest white pattern, inhabits Ural, 
West Siberian Lowland and the Kuznetskoe upland; in Altai, 
Sayans and eastwards ssp. magnata Heyne in Ruhl, 1895 ranges, 
with a reduced white pattern on wing upperside but well 
expressed white submarginal lunules on hind wing underside; SE 
Zabaikalye, Priamurye, Primorye, the Sakhalin and Kuriles are 
inhabited by : ssp. bergmanni Bryk, 1942, resembling coenobita 
with a wide white pattern. The status of Neptis kusnetzovi 
Kurentzov, 1949, described by two males by two males from a 
spruce/fir forest of the Middle Sikhote-Alin' Mts. and combining 
the characters of N, rivulars and N. pryeri, is not clear. 


132. Neptis pryeri Butler, 1871

Type locality: Japan.

RANGE: Priamurye (from the Byreya to the Goryun Rivers), 
Primorye, Central and NE China, Korea, Japan. 

HABITAT: wood edges and openings in birch/oak, broad-leaved and 
mixed, mostly montane, forests. 

FLIGHT PERIOD: late June/early August. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). 
Foodplants: Spiraea. Eggs: greyish, covered with hairs, laid 
singly on leaf margins and tips. Larva: dark, resembles 
N.rivularis but with longer processes on the back; hibernates at 
the third instar. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 21-30 mm. On fore wing 
upperside white stripe in cell split into four spots; hind wing 
underside has 8-10 large black spots at base. 

GEOGRAPHICAL VARIATION: For Primorye and Priamurye ssp. andetria 
Fruhstorfer, 1909 (=kusnetzovi Kurentzov, 1949) is reported, 
which differs from the nominotypical one by a smaller size, 
reduction of submarginal band on hind wing upperside and darker 
ground colour o hind wing underside. 

ETYMOLOGY: H. Pryer - an English lepidopterologist of XIX 
century, an author of works on Japanese fauna. 


133. Neptis alwina Bremer et Grey, 1852.

TYPE LOCALITY: China.

RANGE: Priamurye (at Kabarovsk, Nikolaevsk-na-Amure), Primorye, 
Central and NE China, Korea, Japan. 

HABITAT: orchards with fruit trees, valley broad-leaved and pine/apricot 
forests. 

FLIGHT PERIOD: July/late August.

PREIMAGINAL PHASES: Foodplants: Prunus, Armeniaca, besides, 
Spiraea and Malus were also reported. Eggs: blueish-green, 
covered with small hairs and white knobs; laid singly on 
foodplant leaf margins. The larva overwinters at the third 
instar near a hibernating bud, then it constructs a shelter. 
Mature larva: greenish with a brown ornament and four pairs of 
processes on the back. Pupa: pale-brown, suspended on a 
foodplant branch. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-41 mm. On fore wing 
upperside white stripe in cell fractured; at fore wing apex 
there is a white spot in males and a streak in females; on hind 
wing submarginal band consists of relatively large white spots. 


134. Neptis thisbe Menetries, 1859.

TYPE LOCALITY: The mountains of Bureya.

RANGE: Middle Priamurye, the southern ranges of the mountains of 
Bureya, Primorye, Central and NE China, Korea. 

HABITAT: montane oak woods, other types of broad-leaved and 
mixed forests. The butterflies keep to wood edges, open stand 
of trees, but occur under the wood canopy as well.  

FLIGHT PERIOD: middle June/August.

PREIMAGINAL PHASES: Foodplant in Priamurye: Querqus mongolica 
(Graeser 1888). 

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-43 mm. Spots and bands on 
wing upperside orange-yellow; on hind wing underside, at the 
base, at distal end of an elongate blueish basal spot there is 
one relatively large oval brand of the same colour. Similar 
species: N. tchetverikovi, N. themis. 

GEOGRAPHICAL VARIATION: ssp. ussuriensis Kurentzov, 1970, described 
from Primorye, most probably is a synonym for the nominotypical one. 


135. Neptis tschetverikovi Kurentzov, 1936
(= yunnana auct. non Oberthur, 1906).

TYPE LOCALITY: Primorye: the Ussuri headwaters.

RANGE: E Zabaikalye (the Shilka and Argun' basins), Priamurye, the 
mountains of Bureya, the Sikhote-Alin', NE China, Korea. 

HABITAT: open woods, wood edges in birch, broad-leaves, mixed 
and dark-needle forests, in the mountains up to 1000 m above sea 
level. 

FLIGHT PERIOD: prolonged, middle June/early September.

PECULIAR TRAITS OF IMAGO: F.w.l.: 28-37 mm. Spots and bands on 
wing upperside are orange-yellow; on hind wing underside at the 
distal end of a short lengthwise blueish basal spot there are 
two or three small darker brands. Similar species: N. thisbe, N. 
themis. 

ETYMOLOGY: Sergey Sergeevich Chetverikov (1880-1959) - a prominent Soviet 
geneticist and entomologist, he made a large collection of Lepidoptera and 
transfered it to Zoological Museum (St-Petersburg). 


136. Neptis themis Leech, 1892.

TYPE LOCALITY: W China: Siaolu.

RANGE: Priamurye (between the Zeya and Goryun Rivers), South 
Primorye, NE, S and Central China, Korea. 

HABITAT: polydominant broad-leaved and mixed forests. The butterflies usually 
occur together with N. thisbe but substantially less frequently. 

FLIGHT PERIOD: early July/middle August. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 27-35 mm. Spots and bands on 
wing upperside orange-yellow; on hind wing underside there is a 
lengthwise elongate blueish basal spot without brands at its 
distal end. Similar species: N. thisbe, N.tchetverikovi. 

GEOGRAPHICAL VARIATION: Primorye is inhabited by the subspecies ilos 
Fruhstorfer, 1909. 


GENUS ALDANIA Moore, [1896].
Type species: Diadema raddei Bremer, 1861.

  An East-Asian genus with two species.


137. Aldania raddei (Bremer, 1861).

TYPE LOCALITY: the mountains of Bureya.

RANGE: Middle and Lower Priamurye (between the Zeya and Goryun 
Rivers), the southern ranges of the mountains of Bureya, 
Primorye, the adjacent regions of NE China and N Korea. 

HABITAT: edges and openings in valley or, less frequently, montane 
polydominant broad-leaved and mixed forests. 

FLIGHT PERIOD: middle June/middle August in Primorye, in Middle 
Priamurye late May/early August. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 33-42 mm. Both swing sides 
light- or dark-ash-grey with black veins. 

ETYMOLOGY: Gustav Ivanovich Radde (1801-1859) - a well-known Russian 
naturalist, zoologist, and explorer of the Far East. 



SUBFAMILIA NYMPHALINAE Swainson, 1827.

    Our species are of an intermediate or relatively large size 
(f.w.l.: 16-34 mm), with wings  brightly and variously coloured, 
red, fulvous, dark-brown and cherry colours predominating. 
Sexual dimorphysm is weakly expressed. 
    The larvae have 6-7 lengthwise rows of spines; the pupae 
bear pairs of sharp projections on the head and on the back of 
the abdominal segments and a large prominence on the back of the 
thorax. The imagines hibernate (except for Vanessa cardui), in 
many species they are capable of substantial migrations 
(sometimes transoceanic), therefore they are widely distributed, 
their populations are not considerably isolated from each other, 
and geographic variatioon is weak. The abundance of individuals 
in a given locality fluctuates substantially from year to year. 


GENUS POLYGONIA Hubner, [1819].
Type species: Papilio c-album Linnaeus, 1758.

  Mostly a Holarctic genus with 15 species.


138. Polygonia c-album (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The non-tropical Eurasia northwards to the forest-tundra 
belt, the Sakhalin, the Kuriles, Japan. 

HABITAT: various forest biotopes, river valleys, bogs, 
settlements, waste lands; in the north of East Siberia and the 
Far East - the riparian floodland poplar woods. Imagines migrate 
actively, occuring in any open landscape. 

FLIGHT PERIOD: in the most part of Siberia in one brood, from 
late July to autumn and, after hibernation, until June (or, in 
the northern range, July). In southern localities the second 
brood is possible, the butterflies of which being more lightly 
coloured (f. hutchinsoni Robs. - in South Ural; f. lunigera 
Butl. - in Primorye). In July and early August in such 
localities one can sometimes observe the butterflies of both 
seasonal forms flying together, although in various ratios. In 
spring and autumn only the darker form can be found. It is 
thought (Moucha, 1979) that in Europe some larvae hatching from 
the eggs laid by overwintered buterflies develop rapidly to give 
rise to lighter second brood buterflies, while others result in 
darker butterflies appearing later; the lighter butterflies give 
offspring which develops also rapidly to result in the same 
darker individuals in autumn. Details of brood dynamics in 
Siberia is waiting an investigation. 

PREIMAGINAL PHASES: Foodplants: mostly arboreal plants of 
different families, among which reported are: for W Europe: 
Humulus lupulus, Grossularia rectinata, Cerasus; for the taiga 
zone: Urtica dioica, U. urens, Salix caprea, Ribes rubrum, 
Rubus; for the Middle Volga Basin: Ulmus glabra, Corylus 
allavena; for W Altai: Lonicera; for Primorye: Ulmus propinqua. 
Eggs: green, oval with ten light lengthwise ribs, laid singly on 
the foodplant leaves. Young larva: mottled, with yellow spots 
and blueish, white, and black dots on a dark background, and 
with seven rows of branched spines. It keeps to leaf underside 
or draws leaf edges together with silken threads to make a 
shelter. Last instar larva: bicoloured: the first five segments 
yellow, the rest ones white, either side with two reddish 
lengthwise stripes; spines light, branched. Pupa: reddish-grey 
or pale-brown with three pairs of  brilliant spots on a strongly 
bent back; head and thorax with prominences. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 21-29 mm. Wing upperside 
fulvous or brick-red with dark spots; outer margin with blunt 
teeth of different sizes; wing underside greyish-black (spring 
form) or yellowish-brown (summer form), with a white C-shaped 
mark in the centre.             

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges on 
the most part of Eurasia (the form kultukensis Kleinschmidt, 
1929, described from Baikal, is its synonym);  ssp. hamigera 
Butler, 1877 (= ssp. sachalinensis Matsumura, 1915) is reported 
for the southern Far East, the Sakhalin and Kuriles; the 
butterflies from Priamurye and Primorye differs from western 
ones by a larger size and a widened black pattern in a summer 
form, those from the Sakhalin and S Kuriles approach to Japanese 
ones by longer wing teeth and a more mottly ground colour of 
wing underside in a hibernating form. 


139. Polygonia c-aureum (Linnaeus, 1767). 

TYPE LOCALITY: Primorye: the Suchan River.

RANGE: Priamurye (from the Amur-Zeya Plateau to the Goryun 
River), the southern ranges of the mountains of Bureya, 
Primorye, NE and Southern China, Korea, Japan. 

HABITAT: different woody biotopes, orchards, parks, settlements, 
waste lands, dry meadows. 

FLIGHT PERIOD: in one or two broods in July-October, and, after 
hibernation, until middle May. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al, 1982). 
Foodplants: Cannabis sativa, Humulus japonicus. Eggs: green with 
tiny white dots and lengthwise striae; laid singly on foodplant 
leaves or buds. Larva: lives in a rolled leaf; in the last 
instar it is dark-brown with yellowish streaks and seven rows of 
orange branched spines. Pupa: brown with darker streaks along 
wing cases and sides of abdomen; on dorsal side it bears sharp 
projections and small light knobs. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 22-29 mm. Wing upperside 
fulvous with dark spots; outer margin has sharp teeth of 
different sizes; wing underside black (spring form) or ochre-
coloured (summer form) with a golden L- or C-shaped mark in the 
centre. 


     GENUS RODDIA Korshunov, 1995.

     Original description:
"Type species: Papilio l-album Esper, 1780.
   A monotypic Holarctic genus. The imago fore wing length is 
26-32 mm. The wing shape and pattern resemble those of both 
Polygonia and Nymphalis genera. The wing outer margin is 
toothed. The wing upperside is reddish-fulvous with rather large 
black spots and a white spot at the costal margin of each wing. 
The wing bases are covered with dense soft hairs. The wing 
underside is brown, the postdiscoidal area being lighter, with 
dark strokes or reddish-brown. 
   The genus is named in the honour of a forester and 
entomologist Eugeniy Georgievich Rodd (1871-1933), an explorer 
of Altai and the Upper Priobye." 


140. Roddia l-album (Esper, 1780)
(= vau-album (Denis et Schiffermuller, 1775) - nomen nudum 
(Kocak, 1981)). 

TYPE LOCALITY: Central Europe.

RANGE: The temperate Eurasia northwards to the middle taiga 
zone, the Sakhalin, Kuriles, Japan, the temperate N America; in 
its eastern range the species becomes in average more abundant. 

HABITAT: various forest biotopes, in the mountains rises to the 
subalpine belt. 

FLIGHT PERIOD: middle July/Spetember and, after hibernation, 
until middle May. 

PREIMAGINAL PHASES: Foodplants: Salix, Populus tremula are 
reported for Siberia; Ulmus propinqua and Betula mandshurica  
for the southern Far East (Kurentzov, 1970), for various parts 
of the rnge aso reported are Alnus, Tilia, Acer, Humulus, Rumex, 
Fraxinus, Morus, Spiraea, Rosa, Sorbus, Rubus. Eggs: at first 
yellowish-green, later become blue; laid by 35-45 in twig-
ringing batches. Young larva: dark with black spots and a row of 
white spots along the back, after each moult larva becomes more 
blueish and its spines - more light-coloured. Young larvae live 
gregariously on nests made out of silk-spun leaves, mature 
larvae live solitarily, they usually placed on a leaf edge in J-
like posture Mature larva: ground colour varies from black to 
blueish-grey; body covered with yellow or reddish dots; its 
ventral side is fulvous-red or brown; a double yellow line goes 
along the back and a wide yellow stripe, split by segment 
divisions, along either side; head with yellowish dots and two 
yellow strokes; spines branched, yellowish with black tips; 
spiracles reddish or reddish-yellow; head is speckled with 
yellowish dots and bears two yellowish spinules. Pupa: 
yellowish-ochre coloured, with pairs of silvery spots on the 
metathorax and the second abdominal segment and with pointed 
prominences on the head, thorax and abdomen; suspended on the 
trunks and branches. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 27-32 mm. On upperside of all 
wings there is a white spot at costal margin; hind wing 
underside with a white L-shaped mark in the centre. 


GENUS NYMPHALIS Kluk, 1802.
Type species: Papilio polychloros Linnaeus, 1758.

  Mainly a Holarctic genus with seven species.


141. Nymphalis polychloros (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: Europe, N Africa, Anterior Asia, the Pamiro-Alai Mts., NE 
Kazakhstan (reported by V. Dubatolov for Pavlodar), South Ural, 
reported for S Siberia (for Altai by Elwes (1899) and Tomsk by 
M. Ruzskii (1937), probably erroneously, since the species was 
not recorded there in recent decades. 

HABITAT: deciduous forests, settlements. Imagines migrate 
actively. 

FLIGHT PERIOD: in a single brood from late June to autumn and, 
after hibernation, until May. 

PREIMAGINAL PHASES: studied in Europe. Foodplants: Salix, 
Populus nigra, P. tremula, Ulmus, less frequently arboreal 
Rosaceae, such as Crataegus, Malus, Pyrus, Cerasus, Padus. Eggs: 
hemisphaerical, flattened beneath, with 10 vertical ribs, 
castane-coloured; laid by 20-60 on a foodplant branch about 2 m 
above the ground and higher. First instar larva: blackish-grey, 
set by dense hairs; after the first moult yellow spines appear. 
Mature larva: brownish-grey or greyish-blue, with mate-yellow 
lengthwise stripes on back and sides; spines, arranged into 
seven rows, are of the same colour, thin white hairs being 
noticeable between them. Pupa: reddish or brownish, usually with 
silvery and golden areas on the back; head projections are 
sharply pointed. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 25-31 mm. Wing upperside 
brick-red with black spots; hind wing upperside with brownish-
fulvous basal area;  legs black. Similar species: N. 
xanthomelas. 


142. Nymphalis xanthomelas (Esper, 1781).

TYPE LOCALITY: Central Europe.

RANGE: The temperate and subtropical Eurasia, the Sakhalin, the 
Kuriles, Japan, more common the in eastern part of the range. 

HABITAT: various forest biotopes, bogs. Imagines migrate  
actively occuring in diverse landscapes, for instance, in 
steppes or even arctic tundras. Their  abundance fluctuates much 
from year to year,   mass proliferations being recorded in the 
taiga zone of  Middle and Eastern Siberia and the Far East.  

FLIGHT PERIOD: from July to autumn and in spring after hibernation. 

PREIMAGINAL PHASES: Foodplants: mainly Salix species, besides, 
Lonicera and Cotoneaster were reported for W Altai, Urtica - for 
Krasnoyarsk, Populus, Betulae, Alnus - for Europe. Females are 
fertilized before hibernation. Eggs: laid in spring by 100-1500 
on thin foodplant branches. Young larvae live gregariously on 
nests made of silk-spun leaves. Mature larva (according to 
obsevation by P. Gorbunov in W Altai): dark with white or 
yellowish specks forming wide lengthwise stripes and light thin 
hairs; there are six rows of black spines, two upper rows 
containing largest spines; spines of the lowest row on segments 
4-11 rising from fulvous spots; thoracic legs flvous. Being 
disturbed, the larva rises the fore part of the body and become 
immovable to resemble a twig. Pupa: pale-brown or greyish, often 
with a greyish farinaceous bloom, without silvery spots; it 
bears two rows of small spines on dorsal side of abdominal 
segments and two sharp projections on the head and one on the 
thorax. 

PECULIAR TRAITS OF IMAGO:F.w.l.: 25-32 mm. Wing upperside brick-
red with black spots; fore wing upperside with a reddish-brown 
basal area and apically with white spot at costal margin. Legs 
yellowiish-grey. Similar species: N. polychloros. 

GEOGRAPHICAL VARIATION: The taxon sachalinensis Matsumura, 1925, 
was  described from the Sakhalin, ssp. japonica Stichel, 1902, 
reported for the Kuriles. 


143. Nymphalis antiopa (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia, the Sakhalin, the Kuriles, Japan,  
N America. A migrating species having been observed as north  
as the southern tundras of the Yamal Peninsula, in NE Asia does 
not reach polar regions, absent from Kamchatka. 

HABITAT: various forest biotopes. 

FLIGHT PERIOD: from middle June to autumn and, after 
hibernation, until June or, locally in the North, until July. 

PREIMAGINAL PHASES: Foodplants: Betula verrucosa, B. manshurica, 
B. dahurica, Salix, besides, from Europe (Niculescu, 1965) and N 
America reported are Populus, Alnus, Urtica, Humulus lupulus, 
Ulmus, Acer, Tilia, Fraxinus, Spiraea, Rosa, Sorbus, Rubus. 
Eggs: at first ochre-yellow, later become reddish-brown; laid by 
dense rings, containing up to 100 eggs, around the foodplant 
twigs. The larvae live gregariously until the last instar. 
Mature larva: dark with numerous white dots and a row of red or 
reddish-brown spots on the back of segment 3-10 and with six 
rows of black spines; ventral prolegs reddish-brown. In Siberia 
the larvae develop usualy during July. Pupa: greyish or ochre-
coloured, resembles N. xanthomelas but teeth-like projections on 
the head and abdomen are more long and more sharp. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 28-37 mm. Wing underside 
black, upperside dark-brown with a cherry tint, both with yellow 
or whitish marginal band; on upperside of both wing there is a 
row of blue spots distally of it. 

GEOGRAPHICAL VARIATION: A little variable species; small 
butterflies from Yakutia were described under the name borealis 
Wnukowsky, 1927; those from the Sakhalin and Kuriles - as asopos 
Fruhstorfer, 1909. 



GENUS VANESSA Fabricius, 1807.
Type species: Papilio atalanta Linnaeus, 1758.

  The genus includes 9 species and has a world-wide distribution. 


144. Vanessa atalanta (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: North and Central America, N Africa, Europe, Anterior 
Asia. In South and Middle Ural, the south of West Siberia to the 
Altai piedmonts and the Yenisei River known by occasional 
findings. In 40ths these butterflies appeared in mass quantities 
in the Omsk Region. 

HABITAT AND FLIGHT PERIOD: imagines migrate actively occuring in 
various open landscapes, usually in the second half of summer, 
rarely in early spring after hibernation. 

PREIMAGINAL PHASES: Foodplants: Urtica, also reported are 
Humulus, Parietaria (Urticaceae), Carduus, Cirsium, Helichrysum 
(Asteraceae), Salix. Eggs: greenish, almost spherical, with ten 
lengthwise ribs; laid singly on the foodplant leaves. Larva: 
black, red-brown, or yellow-green, with yellowish lateral 
stripes, yellow speckles, and rows of light spines rising from 
reddish warts; lives in a rolled and silk-fastened leaf. Pupa: 
grey or brown, with paired silvery spots on the back. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 26-32 mm. Wing upperside dark-
brown or black; fore wing with a slightly bent red band of even 
width throughout, going from the middle of costal margin to anal 
angle. 


145. Vanessa indica (Herbst, 1794).

TYPE LOCALITY: India.

RANGE: South Siberia east of the Pribaikalye (migrating 
specimens), the southern Far East, Kamchatka (migrating 
specimens), the Sakhalin, the Kuriles, Mongolia, China, Korea, 
Japan, India; known also from the Canary Islands and the 
Madeira. The imagines migrate actively. 

HABITAT: river valleys, vasting lands, parks, settlements, wood 
edges, other open iotopes, in Kamchatka - riparian poplar woods. 

FLIGHT PERIOD: July-September and, after hibernation, in May. 

PREIMAGINAL PHASES: Foodplants: Urtica urens, less frequently 
Populus maximoviczii are known in the southern Far East 
(Kurentzov, 1970), Urtica cannabifolia in SE Zabaikalye (O. 
Kosterin, V. Dubatolov). Eggs: blueish-green with light 
lengthwise striae; laid singly or in small batches on buds or 
young leaves of the foodplant. Mature larva: brownish-black with 
yellowish marking and branched spines longer than in V. cardui. 
It lives and pupates in a sack-like nest made out of silk-spun 
leaf. Pupa: light- brown with pale farinaceous bloom. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 25-34 mm. Wing upperside dark-
brown or black; fore wing with an irregular red band with broken 
inner margin, going from middle costal margin to anal angle. 


146. Vanessa cardui (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: Eurasia, Africa, Australia, N America, many islands, 
including the Sakhalin and Kuriles. Eastwards of Ural occur 
frequently in the temperate zone, but some migrating imagines 
reach tundras where they, however, can not establish the 
progeny. 

FLIGHT PERIOD: Due to such a migrating activity the flight span 
and time of egg-laying and development of larvae much vary in 
different regions and in different years. Imagines, scarce or 
abundant, can be found from May to autumn. 

PREIMAGINAL PHASES: Foodplants (Scott, 1986; other authors): 
Asteraceae: Carduus, Cirsium, Arctium, Centaurea, Serratula, 
Xanthium, Echinops, Tanacetum, Achillea, Helianthus, Senecio; 
rarely Urticaceae (Urtica), Plantaginaceae (Plantago), Poaceae 
(Zea mays), Polygonaceae (Rumex), Eleagnaceae (Rhanmus), 
Malvaceae (Althea, Malva), Fabaceae (Lupinus, Medicago, 
Trifolium), Boraginaceae (Lappula), Lamiaceae (Mentha, Salvia, 
Stachys), or Rosaceae (Fragaria, Prunus), and some plants of the 
families Apiaceae, Brassicaceae, Rutaceae. The cases are known 
when the larvae of this species brought about a considerable 
damage to melons, pulses, tomato, mint, and other cultivated 
plants. Following traits were reported from Europe: Eggs: 
greenish, sphaerical with 16 lengthwise ribs; laid singly on the 
foodplant leaves. Mature larva: grey or black with a double 
yellow stripe on the back and a streak of the same colour on 
either side, with yellowish specks, short hairs and six rows of 
whitish branched spines; head greyish-black. Pupa: light- brown 
with paired golden prominences on the back, more elongate than 
in V.atalanta; its stage lasts for about fortnight. In our 
climate hibernation usually occurs at the pupa stage. The second 
last instar larva was collected at Novosibirsk on Lappula sp. by 
O. Berezina. The larva is dark-grey with a complicated ornament 
and 9 rows of spines; those of 5 upper rows pinkish-red with 
yellow tips, those of four lower rows (two rows on either side) 
reddish-yellow (the difference in spine colour is a very vidiv 
character), with branches yellowish basally and black apically. 
On 2nd and 3rd thoracic segments spines of a medial back row 
absent; 1st segment lacks spines. Each segment behind a spine 
ring has two narrow yellow "belts" margined with narrow 
interrupted black lines, there are also black dots between the 
belts, they fuse between each other on thoracic segments. A wide 
double interrupted yellow stripe goes along the back, it is 
divided by a narrow black line; besides, on either side 
lengthwise yellow lines go through 2ns and 3rd spine rows 
(counting from beneath). Below the lower line the colour is 
evenly light grey. Spines of five upper rows spring from black 
semirings, on which lengthwise yellow streaks are present just 
in front of the spines of 3rd and 4th rows (from beneath). 
Spiracles black, light rimmed. Head black, with dense light 
hairs. All legs yellowish.  


PECULIAR TRAITS OF IMAGO: F.w.l.: 25-34 mm. Wing upperside pale-
orange with separate black spots; fore wing apically has five 
white spots of different sizes. 

GEOGRAPHICAL VARIATION: little expressed, ssp. japonica Stichel, 
1909 was reported for the Sakhalin and S Kuriles. 


GENUS AGLAIS Dalman, 1816.
Type species: Papilio urticae Linnaeus, 1758.

  A Holarctic genus with 4 species.

147. Aglais urticae (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia, the Sakhalin, the Kuriles, Japan. 

HABITAT: river valleys, and also man-affected biotopes, 
settlements. Following the man and the nettle this species 
deeply penetrates into taiga regioins reaching the Polar Circle. 
In the mountains these butterflies, probably migrants, occur 
above the tree-line, usually on screes and rock crests. 

FLIGHT PERIOD: in spring, summer, and autumn, i.e. almost 
contiguously during the warm season; in the North and mountains 
- in one brood, in the southern range - in two broods. 

PREIMAGINAL PHASES: Foodplants: Urtica dioica, U. urens, U. 
angustifolia, rarely (observation in Europe) Humulus lupulus, 
Cannabis sativa, Ribes, Fragaria vesca. Eggs: greenish, 
cylondrical, laid by 100-200 on a foodplant leaf. The larvae are 
gregarious until the last instar; the young ones are dark with 
greyish spots and black spines. Mature larva: black with a 
double yellow-green lengthwise stripe on the back and on either 
side and with seven rows of light branched spines. Pupa: 
angular, varying in colour from light-yellow to dark-brown, with 
a golden bloom; usually found on walls or fences. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 20-27 mm. Wing upperside brick-red with 
separate black and yellow spots; hind wing basal half black. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges in 
Siberia and the Far East; butterflies from  subpolar regioins 
and NE Asia, have a somewhat widened blak pattern that 
charaterises the race polaris Staudinger, 1871. The Sakhalin and 
the South Kuriles are inhabited by ssp. connexa Butler, 1881, 
described from Japan, probably being a separate species; it 
dffers from continental butterflies by a wavy black band across 
fore wing, formed by fusion of two discal spots. 



GENUS INACHIS Hubner, [1819].
Type species: Papilio io Linnaeus, 1758.

  A monotypical Palearctic genus.


148. Inachis io (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia, northwards to the northern taiga 
zone, the Sakhalin, the Kuriles, Japan. 

HABITAT: river valleys, various open and woody man-affected 
biotopes, settlements. 

FLIGHT PERIOD: from middle July to autumn and, after 
hibernation, until late June. 

PREIMAGNAL PHASES: Foodplants: Urtica dioica, U. angustifolia, 
rarely Humulus lupulus. Eggs: greenish with 9 lengthwise ribs, 
resembling a gooseberries in shape, laid in large batches by 
100-400, usually on the foodplant leaf underside. The larvae 
until the last instar live gregariously on the nests made of 
silk-spun leaves. Mature larva: black with numerous white dots 
and six rows of black branched spines. Pupa: varying in 
colouration from yellow to brown, with golden bloom and spots; 
sometimes hibernates. 

PECULIAR TRAITS OF IMAGO:F.w.l.: 25-30 mm. Wing upperside 
cherry-red; each wing has sepatare yellow and black spots and, 
apically, a large "peacock eye" complex ocellus. 

GEOGRAPHICAL VARIATION: butterflies from the southern Far East, 
the Sakhalin, and the South Kuriles, are attributed to ssp. 
geischa Stichel, 1908, described from Japan. 



GENUS KANISKA Moore, [1899].
Type species: Papilio canace Linnaeus, 1763.

A monotypical genus


146. Kaniska canace (Linnaeus, 1763).

TYPE LOCALITY: E China.

RANGE: South Primorye (north to the Ussuri River lower reaches 
at the Khrkhtsirskii Nature Reserve), Central, South, and North-
East China, Korea, Japan. 

HABITAT and FLIGHT PERIOD: the imagines occur rarely in May, 
August, and September in broad-leaved and mixed forests. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). 
Foodplants: Smilax oldhami, Lilium tigrinum, Streptopus 
amplexifolius, and other Liliaceae. Eggs: greenish with light 
lengthwise streaks, laid singly on shaded foodplant leaf 
upperside. The larvae live in rolled leaves. Mature larva: dark 
with reddish-brown ornament and six rows of yellowish-white 
spines with black branches. Pupa: brown with golden spots, 
dorsal side dark-brown;  abdomen with reticulate ornament; 
thorax with sharp projection; suspended underneath a foodplant 
leaf. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 25-32 mm. Wing upperside is 
dark-brown with a blueish band going through both wings.

GEOGRAPHICAL VARATION: Primorye is inhabited by ssp. charonides 
Stichel, 1908. 



SUBFAMILIA ARASCHNIINAE Butler, 1859.

GENUS ARASCHNIA Hubner, [1819].
Type species: Papilio levana Linnaeus, 1758.

  These butterflies exhibit a well-known example of great 
seasonal dimorphysm. Having close sizes (f.w.l.: 15-22 mm), the 
imagines of the spring and summer broods are strikingly 
different in wing colouration: wing upperside in the former is 
fulvous with dark spots; while in the latter it is black with 
white bands. Wing underside in both broods is reddish-brown with 
a network of yellowish stria along and crossing the veins and 
with lilac spots at the outer margin. 
  Eggs: green, ribed almost spherical, laid in 3-6 adjacent 
columns of 4-20 eggs each on the foodplant leaf underside. The 
young larvae live gregariously until the last instar; 
hibernation occurs in the pupal stage. The pupae bear paired 
pointed projections on the head and the dorsal side of the 
abdominal segments; those of the summer brood hibernate. 
   A Palaearctic genus with six species.


150. Araschnia levana (Linnaeus, 1758).

TYPE LOCALITY: Germany.

RANGE: The temperate Eurasia northwards to the forest-tundra 
belt, the Sakhalin, Japan. 

HABITAT: river valleys, banks of water bodies, edges and 
openings in light woods, turf bogs, settlements, gardens, 
wasting lands; in the mountains penetrates along brook valleys 
to the tree-line.  

FLIGHT PERIOD: in southern lowlands in May/June and July/August; 
the  summer brood often is less abundant. In taiga and mountains 
regions the spring brood flies in June/July while the summer 
brood is as a rule missing. 

PREIMAGINAL PHASES: Foodplants: Urtica, rarely Humulus lupunus. 
Young larva: black, covered with tiny whitish dots and with 
light marking along the back. Mature larva: black, reddish- or 
light-brown with seven rows of black, blueish, or yellowish 
branched spines; head also bears two long dark spines; yellow-
brown strokes or dots can be expressed over the body. Pupa: 
brown with a marble ornament and dark spots at wing cases. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-22 mm. In male genitalia 
caudal processus of valva is tooth-shaped (Table  ). Spring 
brood (f. levana Oschenheimer, 1807): fore wing upperside 
apically with two fused white spots and a row of white dots on a 
dark background. Summer brood (f. prorsa L.): rows of red spots 
on wing upperside absent or weakly expressed, white bands 
narrower than in A. burejana. Individuals of the third brood may 
appear which have an appearance transitory between the two 
former broods but closer to prorsa (f. porima Ochsenheimer, 
1807). Similar species: A. burejana. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges over 
the most part  of the territory considered; the southern Far 
East, the Sakhalin and the South Kuriles are inhabited by ssp. 
wladimiri Kardakov, 1928 (with the summer form borgesi Kardakov, 
1928), differing by somewhat lighter colouration of wing 
upperside grond colour in the spring form and a wider white band 
in the summer form. 


151. Araschnia burejana Bremer, 1861.

TYPE LOCALITY: Priamurye: the mountains of Bureya. 

RANGE: Middle and Lower Priamurye, Primorye, the Sakhalin, the 
South Kuriles, Central and NE China, Korea, Japan. 

HABITATS: meadows in broad-leaved and mixed forests; in the 
mountains reaches the dark-needle taiga belt. 

FLIGHT PERIOD: middle May/middle June and middle July/late 
August, in average this species flies a fortnight later than A. 
levana. In the mountains a single brood flies in June/July. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). 
Foodplants: Urtica. Mature larva: dark with orange spots at 
spiracles and black spines longer than in A.levana. Pupa: dark-
brown with lighter areas on the sides. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-22 mm. In male genitalia 
caudal processus of valva large, club-shaped and barbed (Table  
). Spring brood (f. burejana): fore wing upperside apically with 
a yellow spot at outer margin fused with fulvous ground colour 
in its lower part. Summer brood (f. fallax Janson, 1882): rows 
of elongate red spots are well expressed on wing upperside, one 
on fore wing and two on hind one. Specimens with traits 
transitory between those of the two broods are known. Similar 
species: A. levana. 

GEOGRAPHICAL VARIATION: In Priamurye, Primorye and the Sakhalin 
ssp. burejana ranges; a Japanese ssp. strigosa Butler, 1866 is 
reported for the S Kuriles; later ssp. kurilicola Bryk, 1942 was 
described from the Kunashir, differing from continental 
butterflies by a more developed black patter. 



SUBFAMILIA MELITAEINAE

  Butterflies of intermediate size predominantly. Wing upperside 
varies in ground colour from light-grey to brick-red, with 
reticulate or mottled dark ornament. For hind wing underside two 
reddish or orange transversal bands on a white or yellowish 
background are characteristic: in discoidal region (a discoidal 
band) and in postdiscoidal zone (a postdiscoidal band). Hind 
wing has no transversal veins. 
  Eggs are mostly sphaerical or pear-shaped, apically with a 
funnel surrounded with vertical ribs; laid in batches on the 
foodplants. Young larvae live gregariously on silken nests, 
after hibernation - solitarily. They bear conical protuberances 
called false spines, by 9-11 on each segnment from the first to 
the seventh and by 2-4 on each of the rest segments. Pupae are 
usually light-coloured with dark markings; their head is blunt, 
without "horns". 
  The subfamily includes about 400 species, the majority of 
which belong to the fauna of Central and South America. 


GENUS EUPHYDRYAS Scudder, 1872.
Type species: Papilio phaeton Drury, [1773].

  F.w.l.: 16-23 mm. The wing upperside ground colour is light 
with a reddish-orange sumbarginal band and two spots of the same 
colour in the cell; the dark ornament is reticulate. 
  A Holarctic genus with 15 species.


152. Euphydryas maturna (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia, except for the Far East, eastwards 
to Central Yakutia and E Zabaikalye, in the North lovcally 
reaches the forest-tundra zone. A local species. 

HABITAT: mesophilous forest meadows, wood edges, banks of water 
bodies, in the mountains of South Siberia rises up to 1400 m 
altitude. Butterflies feed on the flowers of Polgonum bistorta, 
Veronica, Viburnum opulus etc., congregate on wet ground in hot 
weather, males occupy perches most often being prominent 
branches of the bushes and chase out other individuals.  

FLIGHT PERIOD: early June/late July, depending on the locality. 

PREIMAGINAL PHASES: Foodplants: Veronica septentrionalis 
reported for Polar Ural, V. longifolia for Novosibirsk Region, 
Viburnum opulus, Lonicera, Spiraea crenata, Cotneaster 
melanocarpa, Salix, Caragana frutex, Artemisia - for W Altai (P. 
Gorbunov), besides, for Europe reported are Thalicrum, Populus 
tremula, Viola, Syrigna, Digitalis, Scabiosa, Plantago. Eggs: 
pale-green, ribbed apically; laid on foodplant leaf underside. 
In Novosibirsk Region (Y. Korshunov) young larvae hatched in 
erly August, at the first instar they live gregariously on silk-
spun leaves of herbaceous plants and overwinter on the nests 
made out of leaves. According to observations in Central Europe, 
after hibernation they feed on trees and bushes, but in 
Novosibirsk Region (O. Kosterin) they were observed to continue 
feeding on Veronica longifolia. Mature larva: black with yellow 
marking concentrating into a wide yellow back stripe and a 
similar stripe on either side; false spines black, hairy. Pupa: 
whitish with small black and orange spots. 

PECULIAR TRAITS OF IMAGO: F.w.l.:  17-24 mm. Wing upperside 
ground colour whitish; on wing underside fulvous colour 
predominates over white and yelowish; outer border of hind wing 
underside fulvous. Similar species: E. intermedia, E. iduna. 

GEOGRAPHICAL VARIATION: In Siberia there widely ranges ssp. 
staudingeri Wnukowsky, 1929 (= uralensis Staudinger, 1852), 
characterizing by a white ground colour in wing central; 
butterflies from Polar Ural by a wide pulvous band approach the 
nomnotypical subspecies. 


153. Euphydryas intermedia (Menetries, 1859).

TYPE LOCALITY: "Khotoum" [the Vilyui River].

RANGE: The Alps, Middle and South Ural, the middle and southern 
taiga zone and North Baraba on the West Siberian Lowland, 
mountain regions of South Siberia and the Far East, the 
Sakhalin, Mongolia, NE China, N Korea. A local species. 

HABITAT: bushy wood edges, openings, river and brook valleys, 
subalpine parklands, stone pine and alder dwarf woods; in the 
mountains of South Siberia is recorded as high as 2200 m above 
sea level. Feeding of imagines was observed on the flowers of 
Anthriscus sylvestris, Senecio nemorensis, Spiraea beauverdiana, 
and other plants with large inflorescences. 

FLIGHT PERIOD: middle June/late July.

PREIMAGINAL PHASES: Foodplants: Veronica reported from the Upper 
Ob' basin; Lonicera maakci - from the southern Far East, beyond 
Asian Russia also Thalictrum, Salicaceae. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-23 mm. Wing upperside 
ground colour varies from light yellowish-orange to brick-red; 
on hind wing postdiscoidal band often contains minute black 
dots; outer margin on hind wing underside fulvous. Similar 
species: E. maturna. 

GEOGRAPHICAL VARIATION: ssp. altaica Seitz, 1908 ranges in Ural, 
West Siberia, and Altai Mts.; itermedia inhaits Yakutia and, 
probably, also Tuva, the Sayans, Pribaikalye, Zabaikalye, and 
the upland Stanovoe Nagorye, in the latter regions specimens 
predominate with ochre-coloured spots in central part of wings, 
known as mongolica Staudinger,  1892; butterflies from the 
southern Far East difer from Siberian ones by even brick-red 
wing upperside ground colour and to some extent reduced dark 
pattern; similar butterflies from the Sakhalin were descred as 
ssp. konumensis Matsumura, 1927. 


154. Euphydryas iduna (Dalman, 1816).

TYPE LOCALITY: Sweden.

RANGE: The very North of Europe (Fennoskandia, the record 
closest to the considered therritory is the surroundings of the 
Ukhta town in Komi Republic), the Alps, the Caucasus, the 
Putorana Plateau, the mountauins of East and South Siberia, of 
the northern Far East (on the Ochot coast is lowered to coastal 
tundras), and of Mongolia. A local species. 

HABITAT: in northern lowlands: mossy (Sphagnum) larch woods, 
bogs and thickets of the dwarf birch; in the mountains: 
detrituous and the dwafr birch montane tundras, alpine and 
subalpine meadows, along river banks in the upper part of the 
forest belt. 

FLIGHT PERIOD: middle June/middle July. 

PREIMAGINAL PHASES: studied in Scandinavia (Henriksen, Kreutzer,
1982). Foodplants: Veronica alpina, V. fruticans, Plantago,
Vaccinium. A biennial species. Eggs: yellowish-green, conical,
with a funnel-shaped concavity on apex and numerous fine ribs;
laid in small groups on foodplant leaves. Larva at the 2nd-3rd
instars: black with numerous yelowish-white markings concentrating
at segment joints, prolegs pale-brown, spiracles ringed by light colour.
Larvae live in groups in a small web. Pupa: pale-ochre-grey with black
latitudinal spots on wing cases, black spots on thorax and black lines
on back and sides divided by segment joints.

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-24 mm. On hind wing 
underside the light-grey or white colour predominates over the 
fulvous and brown; on wing upperside white spots in the centre 
are larger than in E. iduna; black spots in postdiscoidal band 
of hind wing as a rule absent (their traces can be present in 
eastern specimens). Similar species: E. maturna. 

GEOGRAPHICAL VARIATION: butterflies from N Siberia are very 
close to the nominotypical suspecies; the mountains of South 
Siberia are inhabited by ssp. sajana Higgins, 1950, differing by 
somewhat paler wing patern. 


155. Euphydryas aurinia (Rottemburg, 1775).

TYPE LOCALITY: Paris.

RANGE: S Europe, S Ural and S Zauralye, the mountains of S 
Siberia east to Kentei Mts., E Kazakhstan and Mongolia; the 
Prilenskoe Plateau; Ural and Yakutia rare. 

HABITAT: steppes and steppefied meadows in intermontane hollows 
and south-exposed slopes. In South Siberia, from Altai to 
Kentei, beside of the typical form there occurs a peculiar f. 
banghaasi Seitz, 1908, which differs from the former by paler 
colour and substantially smaller size and is confined to 
highlands, inhabiting mostly marchy dwarf-birch thickets at 
1600-2800 m above sea level, sometimes considered as a distinct 
species (Lukhtanov, Lukhtanov, 1994). Only this highland form is 
present in Zabaikalye. 

FLIGHT PERIOD: late May/middle July, depending on the locality. 

PREIMAGINAL PHASES: studied in Europe (Niculescu, 1965;
Henriksen, Kreutzer, 1982; Ebert, 1991, other authors).
Foodplants: Succisa pratensis, Knautia arvense
(Dipsacaceae), also  reported Plantago, Digitalis, Veronica,
Theucrium, Geranium, Lonicera, Centaurea scabiosa. In W ALtai (P.
Gorbunov) the larvae were found on Spiraea crenata, Cotoneaster
melanocarpa, Caragana frutex, Viburnum opulus, Lonicera altaica,
Artemisia. Eggs: yellow, truncated conical shaped, with 12-16
lengthwise ribs, laid in batches by 30 and more on  foodplant
leaves at the ground. The larvae live gregariously on a nest
until the second hibernation. Larva: black with small white dots
which concentrate into wide lengthwise bands on the sides; false
spines black; spiracles white-rimmed; prolegs pale-brown. Pupa:
stout, white or light-yellow with two lengthwise black streaks on very convex
wing cases, head black, leg cases yellowish-orange with black spots, each
abdominal segment with a black streak interrupted by orange-
yellow spots on rounded warts.

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-23 mm. Wing upperside 
ground colour varies from pale-yellow to brick-red; submarginal 
band on hind wing usually orange-red; on hind wing underside 
outer border white, postdiscoidal band contains six large black 
dots seen also on the upperside. 

GEOGRAPHICAL VARIATION: The species exhibit great geographical 
variation. Ssp. sareptensis Staudinger, 187, differing from 
thenominotypical by somewhat narrower and very contrasted 
ditinct pattern, ranges in S. Ural; ssp. siberica Staudinger, 
1861 (pro desfontanesi Eversmann, 1851, nec desfontanesi 
Boisduval, 1832), with the type locality Irkutsk, ranges in the 
mountains of S Siberia (ecept for Zabaikalye, differing from 
sareptensis by a more een colouration that makes it approcahing 
the type subspecies; a close subspecies  laeta Christoph, 1893 
was described from the Prilenskoe Plateau in Central Yakutia. 
Investigations by V.V.Dubatolov revealed that butterflies of 
steppen Zabaikalye and the Far East belong to a separate 
species, which is given below. 


156. Euphydryas davidi Oberthur, 1881
(= aurinia auct.).

TYPE LOCALITY: N China: Peking.

RANGE: S and E Zabaikalye, Upper and Middle Priamurye, Primorye, 
E Mongolia, NE China, Korea. 

HABITAT: dry meadows in valley broad-leaved and mixed forests. 
In the Bol'shoi Bukukun River valley (S Chita Region) the 
imagines flied together with those of E. aurinia (V. Dubatolov). 

FLIGHT PERIOD: June/middle July. 

PREIMAGINAL PHASES: studied in SE Zabaikalye by V. Dubatolov. 
Foodplant: Veronica incana. Larva: black with back and 
spiracular lines consisting of white spots of different size and 
shape, lateral stripes split by narrow black winding lines; 
spiracles black with wide white rims; false spines black; hairs 
black, above prolegs brown.   

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-24 mm. On the hind wing 
underside the postdiscoidal band contains six small black dots 
in large light spots, only 2-3 dots being seen on upperside; 
wing upperside ground colour pale-orange, outer margin wide, 
yellow-orange, that on hind wing underside contains six small 
black dots on large pale spots, on hind wing upperside only 2-4 
black dots are seen. In male genitalia the arms of the uncus is 
substantially longer than in the similar species E. aurinia 
(Higgins, 1950). 

GEOGRAPHICAL VARIATION: In Middle Priamurye and Primorye there 
ranges ssp. tjutjujensis Higgins,1950, differing from 
Zabaikalian butterflies by somewhat widened black pattern 
elements on wing upperside. 

ETYMOLOGY: Armand David - a French missioner in China, who 
collected butterflies for K. Oberthur in the southern part of 
the Great Khingan at the end of XIX century. 



GENUS MELITAEA Fabricius, 1807.
Type species: Papilio cinxia Linnaeus, 1758.

  F.w.l.: 12-31 mm. Wing uppersude ground colour varies from 
pale-ash-grey to brick red, dark pattern reticulate or spotty; 
on fore wing there are no postdiscoidal band, spots in 
cell do not differ in colour from background. 
  Mainly a Palearctic genus including not less than 50 species.


157. Melitaea athalia (Rottemburg,  1775).

TYPE LOCALITY: France: the environs of Paris.

RANGE: The temperate Eurasia, in West Siberia northwards to the 
middle taiga belt; eastwards of the Sayans the species is known 
by scarce records from Central Uakutia (the settlement 
Khaptagai), the Stanovoe Nagorye upland (the settlement 
Vasil'evskii), the Magadan Region (the Kolyma River headwaters), 
Kamchatka (the settlement Kozyrevsk), Lower Priamurye (the 
Goryun River), S Primorya (the Suchan River). 

HABITAT: forest, floodland and highland meadows, meadow steppes, 
forest clearing; in the mountains goes up to an altitude of 2300 
m (SE Altai). 

FLIGHT PERIOD: prolonged, early June/early August.

PREIMAGINAL PHASES: studied in Europe and W Siberia. Foodplants: 
Plantago (Plantaginaceae), Melampyrum, Veronica, Digitalis  
(Scrophulariaceae), Valeriana (Valerianaceae), Centaurea, 
Chrysanthemum, Hieraceum, Tanacetum. Eggs: yellowish-white with 
24-26 ribs at apex, laid in batches by several dozens on 
foodplant leaf underside. Young larvae live gregariously and eat 
leaf mesophyl leaving veins and upper epidermis intact, they 
hibernate in a silken nest, at the last instar become solitary. 
Mature larva: dark with a lighter ventral side and transversal 
rows of white and blueish dots; false spines fleshy, yellow or 
brownish, with dark hairs. Pupa: light-grey with black spots of 
different shape and small orange specks on sides; suspended on 
leaf underside. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 14-20 mm. Wing upperside 
ochre-orange with black reticulate markings; hind wing underside 
ground colour white; postdiscoidal band contains diffuse ochre-
orange spots between the veins. Male genitalia: caudal process 
of valva with a long straight tooth (see Fig. ). Similar 
species: M. ambigua, M. britomartis, M. aurelia, M. menetriesi, 

GEOGRAPHICAL VARIATION: butterflies from Ural and W Siberia are 
close to ssp. athalia; those from the mountains of S Siberia 
differ by on average more developed black pattern and were 
describe as ssp. reticulata Higgins, 1955 - from Central Altai 
and ssp. tinica Fruhstorfer, 1910 from the surroundings of 
Irkutsk. Specimens from Yakutia, Magadan Region, Kamchatka and 
the Stanovoe Nagorye upland are very small (with f.w.l. 14-18 
mm) and have a pale wing upperside ground colour; ssp. asiae 
Verity, 1940 was described from Primorye, in general it 
resembles the nominotypical one. 


158. Melitaea ambigua Menetries, 1859.

TYPE LOCALITY: the Amur Region: "Djai" (the surroundings of 
Blagoveshchensk).

RANGE: The East Sayan, Pribaikalie, Zabaikalie, Priamurye 
(downstream to the Goryun River), Primorye, the Sakhalin, E 
Mongolia, NE China, Korea, Japan. 

HABITAT: various meadow and meadow steppe patches. 

FLIGHT PERIOD: middle June/late July. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al, 1982). 
Foodplants: Scrophulariaceae, Cirsium and Artemisia 
(Asteraceae) were also reported. Larva: dark with orange 
streaks; set with hairs; false spines white and fleshy. 
Pupa: light, almost white, covered with small black spots; 
suspended usually on a foodplant stem. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-24 mm. Dark reticulate 
pattern on wing upperside much narrowed in postdiscal area; hind 
wing underside ground colour, as different from M. athalia, with 
dictinct yellowish tint; differs from M. athalia by uncul apes 
in male genitallia (Fig. ). Similar species: V. athalia, M. 
britomartis. 

GEOGRAPHICAL VARIATION: ssp. ambigua (?= bathilda Fruhstorfert, 
1919) is described from Priamurye; from more westerly parts of 
the range there were described the taxa sayaskalpina Verity, 
1940 (E Sayan) and kenteana Seitz, 1909 (Zabaikalye);in Primorye and the 
southern Sakhalin there ranges ssp. niphona Butler, 1878 (= 
mandshurica Fixsen, 1887; = sachalinensis Matsumura. 1925) 
differing gy a larger size and more intense suffusion of yellow 
scales on wing underside.


159. Melitaea britomartis Assmann, 1848.

TYPE LOCALITY: Poland: the town Vrozlav, the Odra River.

RANGE: The temperate Eurasia, in Siberia northwards to the southern 
taiga belt. 

HABITAT: mostly mesophilous and damp meadows of various kinds; 
in the mountains rises as high as 2300 m altitude (Central 
Altai). The butterflies actively visit flowers, they are often 
seen on wet ground. 

FLIGHT PERIOD: prolonged, June/August. 

PREIMAGINAL PHASES: studied in Europe (Henricksen, Kreutser,
1982 and others). Foodplants: Plantago (Plantaginaceae), Veronica, Linaria,
Melampyrum (Scrophulariaceae), Tanacetum (Asteraceae). Eggs:
yellow, barrel-shaped with numerous fine ribs and a small apical funnel;
laid by 30 and more on foodplant leaves. Before hibernation the larvae
live gregareously ona web from which they move for feeding leaving a
silken thread. After hibernation they start living solitarily in
individual webs. Larva: black but looks greenish or greyish due
to numerous light markings (larger and more numerous than in M.
athalia) on back and sides; set with black hairs; false
spines light-yellow with white apices. Pupa: brown with
paler stripes, a darker back stripes and wing cases, abdomen with
yellow spots and pale segment joints.

PECULIAR TRAITS OF IMAGO: F.w.l.: 13-19 mm; wing uppreside 
brownish-orange with a black reticulate pattern; the hind 
wing underside ground colour is yellowish. Male genitalia: 
uncus consists of two teeth; valva is not stretched, nearly 
rounded, its caudal processus stout, but no more than thrice 
as short as valva, with the upper teeth elongate (Table...). 
Similar species:  M.  aurelia,  M. menetriesi, M. athalia, 
reliable identification is possible only by male genitalia. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges in 
Ural and the south of West Siberia. Ouuth Siberian taxa 
seminigra Seitz, 1909 (the Sayans), frigidaltaica Verity, 1940 
(Altai), imitans Verity, 1930 (Zabaikalie), and others, were 
synonymized by L.G.Higgins (1944) with the oldest taxon 
amurensis Staudinger, 1892, described from Priamurye; the 
subspecies latefascia Fixsen, 1883 (= coreae Verity, 1940) was 
also reported for Primorye, it is characterized by larger size 
and somewhat reduction of dark pattern in wing centre. 


160. Melitaea menetriesi Caradja, 1895
(= centralasae Wnukowsky, 1929).

TYPE LOCALITY: Kamchatka. 

RANGE: The south-eastern part of the West Sibarian Lowland (the 
upper Ob'River basin), the mountains of South and East Siberia, 
the mountains of Bureya, Kamchatka, Mongolia. 

HABITAT: in the Novosibirsk region: mesophilous herbaceous 
meadows on river terraces; in the mountains of South Sberia: 
montane, predominantly steppefied, medows; in Kamchatka: long-
forb meadows; in the mountains rises to the tree-line. 

FLIGHT PERIOD: early June/early August, depending on locality. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 14-19 mm. Wing upperside 
ochre-orange or brownish; black marking sometimes to some extent 
reduced; spots of ground colour within postdiscoidal row wider 
than others.  Male genitalia: the valve substantally elongate 
(Tabl....). Simlar species: M. britomartis, M. aurelia, M. 
athalia; for a reliable identification examinatioin of the male 
genitalia is necessary. 

GEOGRAPHICAL VARIATION: At first this species was described 
under the name orientalis Menetries, 1859, this preoccupied name 
was later replaced by menetriesi Caradja, 1895. In 1892 the 
species mongolica Staudinger was described from the Kentei, 
however, this name was also replaced by centralasiae Wnukowsky, 
1929. Thus, ssp. centralasiae Wnukowsky, 1929 ranges 
in West and South Siberia; the nominotypical subspecies is 
distributed in Kamchatka (butterflies from the southern 
Ochot coast and the mountains of Bureya were also included into 
the type series, but the consubspecificity of the entire series 
is problematic) and differ from the previous ssp. by hind wing 
underside, namely, presence of white spots along entire outer 
margin and enlarged black spots in postdiscal area. Similar 
butterflies were collected also in the Magadan Region 
(settlement Madaun); ssp. kolymskia Higgins, 1955, was described 
on the materials from the surroundings of the settlements 
Zhigansk and Srednekolymsk,it is characterized by a small size 
and enlarged white spots on hind wing underside. 


161. Melitaea rebeli Wnukowsky, 1929.

TYPE LOCALITY: SE [Russian] Altai, the Chuya River.

RANGE: A little-known species found in North-East Altai [its Russian 
part implied] (the surroundings of the village Onguday, the stows 
Tyurguno and Tete in the vicinity of the village Kuray, the 
Saylyugem mountain range) and NE Mongolia (the Huvs-Nur and 
Bayan-Ulegey aimaks); reported for the East Sayan (the Tunkinskie 
Belki mountain range). 

HABITAT: larch parkland, highland mountains and detrituous 
ridges at 2000-2900 m above sea level. 

FLIGHT PERIOD: middle June/middle July. 

PECULIAR TRAITS OF IMAGO: A little known taxon, species 
independence of which needs comfirmation. By extermal features 
and male genitalia most resembles M. menetriesi centralasiae. 
Among main diagnostic features the shape of aedeagus, straight 
and stout, sharply ending with a steep "keel", used to be 
mentioned (Higgins, 1955).

ETYMOLOGY: G. Rebel - a German lepidopterologist, the co-author 
of O. Staudinger. 


162. Melitaea aurelia Nickerl, 1850.

TYPE LOCALITY: Czechia: Bohemia.

RANGE: Europe (except for the North), Anterior Asia, South Ural, 
the south of West Siberia (east to the Karasuk District of the 
Novosibirsk region). 

HABITAT: meadow and meadow-steppe patches among kolki (birch 
groves), in pine fortests, along wind-break strips, at water 
bodies, on mountain slopes. 

FLIGHT PERIOD: prolonged, early June/late July. 

PREIMAGINAL PHASES: studied in Europe. Foodplants: Plantago 
(Plantaginaceae), Veronica, Melampyrum, Digitalis 
(Scrophulariaceae), Tanacetum (Asteraceae). Young larvae live 
gregariously, later - solitarily. Mature larva: darker than that 
of M. athalia,  with very small white dots, set with short 
hairs; false spines brownish or reddish with white tips. Pupa: 
yellowish-white with brownish and orange dots of various size 
and shape. 

PECULIAR TRAITS OF IMAGO: F.w.l: 14-18 mm. Wing upperside ochre-
orange with dark-brown reticulate ornament; hind wing ground 
colour ochre-yellow. Male genitalia: uncus not expressed 
(Table...). Similar species: M. britomartis, M. menetriesi, M. 
athalia. 


163. Melitaea plotina Bremer, 1861.

TYPE LOCALITY: Priamurye: the mountains of Bureya.

RANGE: The Novosibirsk Region east of the Ob' River (the valleys 
of the Shadrikha Rivulet and of the right tributary of the Malyy 
Elbash rivulet; between the settlements Salair and Suenga), 
Kemerovo Region (the village Ust'-Travyanka, the town Targai), 
Altaiskii Krai (the village Soldatovo), the East Sayan (the 
Irkut River), Zabaikalie (the Chikoy river, Kyakhta), Priamurye, 
Primorye, NE China, Korea. A local species. 

HABITAT: wet floodland meadows (in the Novosibirsk region - with 
the reed (Phragmites communis)), "mari" - bogged up open larch 
woodland. 

FLIGHT PERIOD: July. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 13-17 mm. Wing upperside 
ochre-orange with black reticulate ornament; on hind wing 
underside ground colour much darkened and contains separate 
yellowish spots, oval in postdiscal area, semiround in 
submarginal area. 

GEOGRAPHICAL VARIATION: weakly stidied. Ssp pacifica Verity, 
1932, has been described from S Zabaikalye, ssp. ussuriae 
Verity, 1932 - from S. Primorye. 

 
164. Melitaea diamina (Lang, 1789).

TYPE LOCALITY: Germany: Augsburg.

RANGE: The temperate Eurasia, locally as north as the northern 
taiga. 

HABITAT: damp forest and forest-steppe meadows, mostly in river 
valleys; in the mountains rises up to the tree-line. 

FLIGHT PERIOD: early June/middle July. 

PREIMAGINAL PHASES: studied in Europe (Niculescu, 1965 and 
others). Foodplants: Plantago (Plantaginaceae), Veronica, 
Melampyrum (Scrophulariaceae), Valeriana, Patrinia 
(Valerianaceae), Polygonum bistorta (Polygonaceae), Asteraceae. 
Eggs: pale-yellow, with 24 vertical ribs; laid in batches by 
5-20 or 200-300 on foodplant leaf underside. Young larvae live 
gregariously on a silken nest; they hibernate in the fourth 
instar inside dry stems or anong dead leaves. Mature larva: 
dark-grey with darker streaks on the back, covered with minute 
blueish dots and hairs; false spines reddish or yellowish; 
head with two blue spots. Pupa: greyish-white or light-blueish-
green with large black brands on wing cases and transversal 
black streaks, interrupted by orange-yellow spots, on abdominal 
segments. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-23 mm; wing upperside 
ochre-brown with rows of light-reddish brown or ochre-yellow 
spots, that is a dark pattern extremely pronounced, as compared 
with other species, and predominates over a lighter colour; hind 
wing underside ground colour is whitish or silvery-white, five 
cells of a fulvous-brown postdiscoidal band contain black dots. 
Similar species: V. protomedia. 

GEOGRAPHICAL VARIATION: Ural and the forest zone of Siberia are 
inhabited by ssp. hebe Borkhausen, 1793, differing from the 
nominotypical one by even more developed dark pattern and a 
whitish colour of spots in wing center in females: the mountains 
of S Siberia east to Pribaikalye - by ssp. erycina Lederer, 
1853, described from W Altai and being characterized by the most 
development of dark colour on wing upperside, with contrasted 
small spots, yellowish in males and whitish in females; larger 
butterflies from SE Zabaikalie, Priamurye, and Primorye, with 
light spots in wing center enlarged, are known as ssp. 
erycinides Staudinger, 1892.


165. Melitaea protomedia Menetries, 1859
(= regama Fruhstorfer, 1915)

TYPE LOCALITY: "the Amur" at the Ussuri River mouth.

RANGE: Middle Priamurye (the surroundings of Khabarovsk), the 
southern and western Primorye, NE China, Korea, Japan. 

HABITAT: damp meadows in river valleys, open places on the 
slopes of mountains and coastal terraces. 

FLIGHT PERIOD: early July/late August. 

PREIMAGINAL PHASES: Probable foodplant: Veronica (Kuretzov, 
1970). 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-22 mm. Wing upperside in 
both sexes ochre-brown  with black reticulate ornament; on hind 
wing underside there is a fulvois-brown sumbarginal band with 4-
5 black dots in cells; this band is bordered at both sides with 
ochre-yellow spots, although in some females they are silvery-
white (f. argentea Fixsen, 1887). This species differs from the 
similar M. diamina by a lighter wing upperside colouration and 
by the shape of the processus on the valva in male genitalia 
(Table....). 


163. Melitaea baicalensis Bremer, 1861 (= arcesia Bremer, 1861).

TYPE LOCALITY: North Pribaikalye.

RANGE: The mountains of South and East Siberia (the Vitim River, 
the Suntar-Khayata mountain range), the Magadan Region, 
Priamurye (the Tukuringra mountain range, surroundings of 
Komsomol'sk-na-Amure), the Sikhote-Alin' Mts. (the Sokolovka 
River), Mongolia, North and Central China. A local species. 

HABITAT: montane steppes, steppefied, forest and alpine meadows; 
Altai - only in the mountains at evelations of 1800-3000 m. On 
the Suntar-Khayata range the species was collected on pebble 
banks of the Khadyta River headwaters at 500-700 m above sea 
level. 

FLIGHT PERIOD: middle June/late July. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 14-21 mm. Wing upperside 
ochre-orange with rows of small black spots; on hind wing 
underside there is an ochre-orange marginal streak, 
postdiscoidal band contains seven round spots of the same 
colour. Male genitalia: caudal processus of valva is like a 
large tooth. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges in 
Pribaikalie, Zabaikalie and the mountains of the Amur basin; 
ssp. minor Elwes, 1899 (?= carmana Fruhstorfer, 1915) is 
known from Altai, Tuva, and the Sayan, it differs by a small 
size and a paler colouration; similar butterflies were found out 
in E akutia and highlands of Zabaikalye (the mountain Golets 
Sokhondo). 

SYSTEMATIC NOTE: The taxon baicalensis was described by O. 
Bremer (1861) by a single male from the Radde's collection from 
North Pribaikalye as a species. A page below in the same paper 
Melitaea arcesia was described by a collection from Dahuria. 
Conspecificity of these taxa, supposed by many 
lepidopterologists (O. Staudinger, L. Higgins) leads to the use 
of the name baicalensis as a valid name for this species (see 
also Sviridov, 1981a), although for a final decision a study of 
type materials at Zoological Institute, St.Petersburg, is 
necessary. Noteworthy that butterflies resembling to Bremer's 
picture of baicalensis were recently described from highlands of 
the Khamar-Daban mountain range (S Pribaikalye) as Melitaea 
arcesia dabanica Barantshikov, 1979, among diagnostic features 
mentioned is "absence of spines on valva processes in male 
genitalia" (Baranchikov, 1979). We didn't studied yet the 
holotype of this taxon. 


superspecies didyma

  F.w.l.: 16-25 mm. Wing upperside is ochre-orange or brick-red 
in males, but in females the fore or both wings can be greyish; 
black pattern consists of separate spots, in females they may be 
fused; on hind wing underside there is a contiguous orange 
postdiscoidal band.  Male genitalia: ventral margin of valva 
dentate. 
  The constituent species are very similar, differing by details 
of colouration and on chromosomal and molecular (isozyme 
polymorphism) levels.


167. Melitaea (didyma) didyma (Esper, 1779).

TYPE LOCALITY: Bavaria.

RANGE: S Europe, N Africa, Anterior and Central Asia, Kazakhstan, 
S Ural, the southern West Siberia, North Altai. 

HABITAT: meadow-steppe patches at kolki [birch groves] edges, 
ravines, long fallow lands, in the mountaims - on barren rocky 
southern slopes and ridges. In Altai the butterflies were 
observed to feed on Goniolimon speciosm, Phlomis tuberosa, 
Leonurus sibirica, Dracocephalum nutans, Trifolium. 

FLIGHT PERIOD: prolonged, middle May/middle July, locally the 
second brood apperas in July/August, these imagines are 
noticeably smaller and paler. 

PREIMAGINAL PHASES: Foodplants: Trifolium, Plantago, Valeriana, 
Veronica, Verbascum, Linaria, Stachys, Phlomis, Linum, Scabiosa, 
Carduus, Cenrtaurea, Artemisia. In West Siberia (the Novosibirsk 
Region as well as W Altai), the larvae are found almost 
exclusively on Phlomis tuberosum, rarely on Veronica incana. 
Eggs: greenish-yellow, pear-shaped with 24 ribs at the apex. 
Young larvae live gregariously on a silken nest, in which they 
also hibernate, usually at the third instar. Mature larva 
(according to observations of P. Gorbunov in W Altai): lives 
solitarily; it is white with numerous black markings with 9 (on 
abdomen) rows of whitish false spines; head and bases of spines 
of 2nd and 4th row from beneath are orange; legs whitish. Pupa 
Pupa: white with black and orange marks, on wing cases the fused 
into two transversal stripes; suspended underneath large leaves, 
in the mountains - on stones; its stage lasts for 10-15 days. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-25 mm. Wing upperside 
ochre-orange or ochre-red,  in females never lacks reddish 
colour; as different from M. latonigena, on hind wing underside 
submarginal orange band bordered with separate black strokes 
between veins; as different from M. didymoides, black spots on 
wing underside larger, as different from M. fascelis, antennal 
club is fulvous beneath. 

GEOGRAPHICAL VARIATION: In the area considered there ranges  
ssp. neera Fischer de Waldheim,  1840 (= uralca Bryk, 
1940), described from the Volga Basin. 


168. Melitaea (didyma) latonigena Eversmann, 1847.

TYPE LOCALITY: Zabaikalie: the Kentei Mts..

RANGE: The mountains of South Siberia and Mongolia, East Siberia 
north-east to the Nera River basin. 

HABITAT: meadows steppes and steppe meadows on south-exposed 
slopes, river terraces, and intermontane hollows; found at sands 
in the Selenga River valley. 

FLIGHT PERIOD: middle June/August, depending on the locality.

PREIMAGINAL PHASES: Pupa: whitish with black and reddish specks 
of different sizes; suspended on grasses. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-24 mm. In males 
wing upperside ochre-orange or ochre-red, in females 
usually greyish with pronounced black ornament; hind wing underside 
ground colour whitish, on hind wing underside submarginal orange band 
bordered with more or less contiguous black lines. Similar 
species: M. didyma, M. didymoides, M. sutchana. A difference in 
chromosome number were shown between this and the most close 
species, M. didyma (Lukhtanov, Kuznetsova, 1989).

GEOGRAPHICAL VARIATION: ssp. latonigena inhabits Pribaikalye and 
Zabaikalye, westwards in the mountains of S Siberia there range 
two smaller subspecies: ssp. altaica Grum-Grshimailo, 1893, 
Altai, and atrata Higgins, 1935, the Sayans; arid regions of 
Yakutia are inhabited by most small subspecies polaris Grum-
Grshimailo, 1899, with a black outer border on wing upperside 
narrower than in South Siberian forms. 


169. Melitaea (didyma) didymoides Eversmann, 1847.

TYPE LOCALITY: Zabaikalie: Kyakhta.

RANGE: Tuva (the Malyi Yenisei River), the East Sayan (the 
village Mondy), S Pribaikalie, S and E Zabaikalie, Upper 
Priamurye (the Zeya River basin), S Primorye. E Mongolia, NE 
China. A local species. 

HABITAT: meadow steppes and steppefied meadows; in South 
Primorye also coastal meadows. 

FLIGHT PERIOD: in one or two broods in early June/late August, 
depending on the locality. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-24 mm. Wing upperside 
ochre-red in males, ochre-orange or light grey in females; wings 
narrower than in M. didyma and M. latonigena, dark pattern in 
central part of wings reduces, on hind wing underside 
submarginal orange band bordered with separate black strokes 
between veins. Male genitalia: teeth on harpe weakly expressed 
(See Table. ). 

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges 
Tuva, Pribaikalye, Zabaikalye, and Priamurye; ssp. latonia Grum-
Grshimailo, 1891, described from S Manzhuria, inhabits the 
Khanka Plain in Primorye, it differs by a more bright 
colouration of wing upperside with more developed black spots. 


170. Melitaea sutschana  Staudinger, 1892.

TYPE LOCALITY: Primorye: Suchan.

RANGE: E Zabaikalie, Priamurye (as downstream as the Goryun River), 
Primorye, the Sakhalin (Yuzhnosakhalinsk), NE China, Korea. 

HABITAT: meadows and open stands in broad-leaved and mixed 
forests, bush thickets. 

FLIGHT PERIOD: early June/late August, in two broods. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-24 mm. Wing upperside 
ochre-red with black spots in males, ochre-orange or light-grey 
with reticulate dark pattern in females; Male genitalia: valva 
longer than in M. didymoides, ventral margin of harpe without 
teeth at apex. 

GEOGRAPHICAL VARIATION: weakly expressed. Butterflies from Lower 
Priamurye differ from typical ones by a smaller size and 
somewhat more developed dark pattern. Small (f.w.l. 18-20 mm) 
and very peculiar butterflies from the Sakhalin Island are 
described as Meltaea sutschana graeseri P. Gorbunov, sbsp.n. 
     Original description:
   "   HOLOTYPE. A male. F.w.l. 20 mm. The wing upperside ground 
colour is ochre-orange, more faded than in the typical 
butterfliws. Among the black marking a row of spots on the 
middle of the wings, these spots are isolated from each other 
and more lengthwise elongate than in sutchana. On the fore wing 
upperside there are only four (also lengthwise elongate) spots 
at the fore margin. The hind wing underside ground colour is 
white with a yellow tint; the ochre-orange bands are wide, 
occupying about a half of the wing area. The light space between 
the external and the internal bands does not exceeds the width 
of the former (about 3 mm). In this space the black spots are 
close to the internal band, while in sutchana they are closer to 
the external one. The antennae and their clubs are fulvous. The 
genitalia structure is as in M. sutchana.
   MATERIALS: The holotype: a male - 2.07.1994, the Sakhalin, the 
environs of the city Yuzhnosakhalinsk (A.M. Basarukin). The 
paratype: a male - 2.07.1994, the same locality.
   We include only two specimens from the Sakhalin into the type 
series, although a male of a similar appearance was found also 
in our collections from the Lower Amurland (the Komsomol'skii 
Reserve, Bichi, 23.06.1988), and two more such males - in the 
ISEA collection from the southern Transbaikalia (Sokhondinskii 
Reserve, the Agutsa River middle flow, 9.08.1987). These facts 
rise a supposition of a species rank of the taxon being 
described, that need further materials for corroboration." 
     It should be added that the subspecies is named in the honour 
of Ludwig Karl Fridrich Graeser (1840-1913), a German 
lepidopterologist who studied butterflies on the Amur River in 1881-
1885.


172. Melitaea romanovi Grum-Grshimailo, 1891.

TYPE LOCALITY: "Mudshik" - a right tributary of the Huwang Ho river.

RANGE: S Zabaikalie (the basins of the Selenga and Onon Rivers,  
Lakes Bol'shoy Chindat, Zun-Ttorei,Barun-Torei), Mongolia, 
Central China (the province Gansu). A local species. 

HABITAT: steppefied meadows and steppes, locally reaches alpine 
meadows. 

FLIGHT PERIOD: late June/late July. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-19 mm. Wings elongate, 
their upperside orange with separate black spots; hind wing 
underside ground colour white, in discoidal area there present 
isolated orange spots which do not form a band. 
 
GEOGRAPHICAL VARIATION: Zabaikalie is inhabited by ssp. puella 
Higgins, 1941. 

ETYMOLOGY: Nikolay Mikhaylovich Romanov (1859-1919) - a grand 
duke, a grandson of Nikolay the First, an owner of a large 
butterfly collection and the editor of "Memores sur les 
Lepidopteres".


172. Melitaea fascelis (Esper, 1784)
(= trivia Denis et Shiffermuller, 1775).

TYPE LOCALITY: Povolzhye: the town Sarepta.

RANGE: The steppe and forest-steppe zones from West 
Europe to Lake Hubsugul (Mongolia). A local species. There are 
reliable records only for the western Ural piedmonts; the species 
was reported also for Altai (the village Kamlak, the valleys of 
the Koksa, Inya, Bukhhtarma Rivers, the surroundings of 
Ust'-Kamenogorsk in East Kazakhstan). 

HABITAT: steppefied, mostly valley meadows, steppefied southern 
slopes. 

FLIGHT PERIOD: in S Ural: middle June/late July, in on brood. 

PREIMAGINAL PHASES: studied in Europe (Niculescu, 1965). 
Foodplants: Verbascum thapsus, V. nigrum, Pedicularis, Teucrium 
are known. Larva: light-blueish-grey with blueish dots and a 
dark back stripe; with red-brown head and lengthwise streaks 
above spiracles; false spines light-yellow with white tips. 
Pupa: blueish-grey with dark specks. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-19 mm. Wing upperside 
brick-red or ochre-orange with black spots, much widened at 
outer margin; on fore wing underside black spots at the outer 
margin are lunular in shape; on hind wing underside orange spots 
of postdiscoidal band without black dots; antennal club black 
beneath. Similar species: M. uvarovi, M. didyma. 

GEOGRAPHICAL VARIATION: A very variable species, imagines of 
different populations and broods substantially differing in 
size, the degree of dark suffusion, and details of colouration. 
The nominotypical subspecies inhabits western piedmonts of S 
Ural. Yu. P. Korshunov (1977) described a peculiar female of M. 
fascelis from the surroundings of Lake Hubsugul in Mongolia. We 
suggest the name Melitaea fascelis singularia Korshunov, sbsp.n. 
for such butterflies. Translation of the cited description, 
which should be considered as the original description of the 
subspecies, is follwoing: 
  "*Melitaea trivia Denis et Schiffermuller, 1775, ?sbsp.n.
   Hubs[ugul Aimak]: Lake Hubsugul, 26 VI (D. [V.Ch. 
Dorogostaiskii], 1 female.
   From the typical form differs by light and mottled pattern, 
somewhat elongate wings. Generally the habitus resemble that of 
M. t. ignasiti Sagarra, 1926. Probably, belong to a new 
subspecies.
   Female. The palpi are set with yellow hairs. The fore wing 
lengthis 18 mm. The wing upperside is pale-fulvous, mottled 
since black dots are small and well contrasted. On the hind wing 
underside the black strokes are narrow, do not contact 
completely to the fulvous band. The middle field is light-
yellow, not differing in colour from the basal and outer parts. 
There are two rows of four irregular black spots (dots) going from 
the fore margin to the cell in the middle field. The genitalia 
are close to those in the typical form."


173. Melitaea uvarovi P. Gorbunov, 1995.

     Original description:
  "   The species is being stated by five males collected by S.V. 
Shutov in the valley of the Ural River, together with such 
spring steppen species as Zerynthhia polyxena, Triphysa phryne, 
Proterebia phygea [an error, should be "Proterebia afra" - Yu. 
P.], Thersamonia thersamon etc. From a close species M. 
fascelis, the closest population of which is known in 15 km NE 
of the place of collection of the new species (12 km S of the 
station Kuvandyk) it differs by a great number of external 
characters, and also early flight period (over-wintering in a 
phase of pupa is probable). It is not excluded that the taxon 
deing described will turn out to be conspecific to a Central 
Asian species M. robertsi Butler, having a certain similarity 
with it in appearance and life history. But now, taking into 
account the Ural populations to be so remote, we describe it in 
a species rank.
   MALE: F.w.l. 15.2-16.4 mm (15.4 mm in the holotype). The 
wings are somewhat narrower than in M. fascelis. The fore wing 
ground colourt is ochre-orange-red, noticeably darker than in a 
close Central Asian taxon M. robertsi catapelia Staudinger, 
1886. The black spots on the wing upperside are smaller and more 
clear-cut than in M. fascelis. The black marginal band is about 
1 mm wide. A row of black submarginal lunules goes along it on 
both wings, they may contact to the marginal band but do not 
form, together with it, an united marginal blackeinig, found in 
M. f. fascelis. There is a small white spot on the fore wing 
fore margin (3 mm off the apex). The basal blackeinig on the 
hind wing does not extend above the vein Cu1. The fore wing 
underside is ochre-orange with a whitish spot at the apex and 
rather small isolated black spots. The hind wing underside 
ground colour is white, without a yellowish tint peculiar to M. 
fascelis. The pattern is ain in M. fascelis but the black spots 
at the outer margin are narrower. The fringe is white, more 
clear and long than in M. fascelis, with black patches at the 
vein ends. In the genitalia structure the differences from M. 
fascelis (see Appendix) are rather weak. 
   MATERIALS: The holotype: a male - 6.05.1982, the Orenburg 
Region, the village Donskoe, a valley steppefied meadow (S.V. 
Shutov). Paratypes: 4 males - 3-6.05.1982, the same locality.
   The species is dedicated to the memory of Uvarov, Boris 
Petrovich (1888-1920), a honoured member of the All-Union 
Entomologiocal Society, who studied the fauna of Lepidoptera of 
NW Kazakhstan from a young age."


174. Melitaea phoebe (Goeze, 1779).

TYPE LOCALITY: Central Europe.

RANGE: The steppe and forest-steppe zones from the Mediterranian Sea 
to the Prilenskoe Plateau, Priamurye, Primorye, and NE China. 

HABITAT: meadow patches at the edges of pine forests and birch 
groves, in river valleys, on montane slopes, in dry larch open 
woodland. 

FLIGHT PERIOD: mostly middle June/middle July in one brood, in 
South Ural (the Orenburg Region) - late May/late August in two 
broods. 

PREIMAGINAL PHASES: Foodplants: Rhaponticum uniflorum, 
Saussuraea, Centaurea, Cirsium, Serratula centauroides 
(Asteraceae), Plantago (Plantaginaceae). Preimaginal phases are 
described for SE Zabaikalye by O. Kosterin as follows. Mature 
larva: white with fine black reticulate ornament, so that looks 
grey; this ornament fuses into a black line along the back and a 
more diffuse line on either side (between 2nd and 3rd row 
of false spines from beneath); a white stripe (without ornament) 
goes through 2nd rom beneath false spine row. False spines 
fulvous with  white apices and black branches; on abdomen they 
are arranged in 11 lengthwise rows, the lowest one (jus 
beneath prolegs) is formed by double false spines springing from 
the same point, on 1st and 2nd abdominal segments there are 
additional false spines position of which coincides with that 
of ventral prolegs on other segments; on 2nd and 3rd thoracic 
segments there are 10 false spine rows (those of the medial row 
on the back being missing); on 1st thoracic segments there are 
also 10 false spines: a "collar" of six small closely set ones 
and, on either side, two larger false spines below. Thoracic 
legs and ventral prolegs yellowish-grey; head greyish-black, set 
with tiny black hairs. Pupa: head, thorax, wing and leg cases 
black with white spots; on fore wing cases there are following white 
markings: at base, in cell, at anal angle, a slanting 
stripe going from middle costal margin to middle outer margin, 
and two rows of dots along outer margin; veins yellow; visible 
hind wing cases with a long elongate white spot. Abdominal 
segments bicoloured: black in fore part and white with black 
dots in hind part, division going through five spines with 
orange apices; ventral side of abdomen with two dark lengthwise 
stripes. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-26 mm. Wing upperside 
ground colour yellowish-orange or yellowish-brown in males, from 
yellowish-browb to pale-yellow in females, black spots partly 
fused into bands; on hind wing underside postdiscal band 
contains large round ochre-red spots between veins. As different 
from a similar species M. scotosia, in male genitalia valva apev 
has 2-3 rather long variously directed teeth. 

GEOGRAPHICAL VARIATION: ssp. ornata Christoph, 1893 from Ural 
have two rows of lighter spots on wing upperside and so look 
mottled; ssp. saturata Staudinger, 1892, widely ranging in 
SIberia, has more even wing ground colour and larger black 
spots, butterflies from humid montane regions being much darker 
than those from arid regions of W Altai, Tuva, and S 
Transbaikalia. Rather small dark butterflies from Central Yakuia 
were described as ssp. tungusa Herz, 1898; light butterflies 
from S Zabaikalye - as ssp. changaica Seitz, 1908. 


175. Melitaea scotosia Butler, 1878.

TYPE LOCALITY: Japan.

RANGE: Primorye. NE China, Korea, Japan. 

HABITAT: dry meadows on montane slopes and in river valleys, 
open montane oak woods. A rare species. 

FLIGHT PERIOD: middle June/late July. 

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). 
Foodplants: Serratula coronata, Saussurea pulchella and S. 
maximowiczii, Synurus deltoides are known. Eggs: yellow, ribbed, 
in batches by several dozens. Larvae live on silken shelters by 
5-40 individuals and hibernate at the fifth instar inside the 
stems of the foodplant or among withered leaves. Mature larva: 
dark, speckled with numerous white dots, with a wide yellowish-
brown back stripe; false spines are brown or yellowish-brown, 
covered by tiny white hairs. Pupa: white with black dots on 
abdomen and thorax; wing cases partly rimmed with a brown margin 
and confine not-fused black spots. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 26-31 mm. Wing upperside 
orange with a dark pattern somewhat reduced in postdiscal area; 
on the hind wing underside the postdiscoidal band usually 
contains six large pale-orange or black spots; submarginal band 
being narrower and paler than in M. phoebe. Male genitalia: 
valva apex with 6-7 short stout teeth. 

GEOGRAPHICAL VARIATION: On the continent this species is 
represented by ssp. butleri Higgins, 1940, differing from the 
nominotypical one by a substantial reduction of black pattern 
on wing upperside, especially in females. 


170. Melitaea cinxia (Linnaeus, 1758).

TYPE LOCALITY: Sweden: Uppsala.

RANGE: N Africa, Europe (except for the North), Anterior and 
Central Asia, N and E Kazakhstan, the southern Ural and Siberia 
NE to the Prilenskoe Plateau, the southern Far East (solitary 
findings).

HABITAT: meadow patches in various forests (kolki [birch 
groves], bory [pine woods], etc.), along wind-break strips, on 
long-fallow lands, on mountain slopes, in steppes, meadow 
steppes. The imagines feed on the flowers of Geranium 
sylvestris, Veronica, Dracocephalum, Plantago, Trifolium, 
Achillea etc. 

FLIGHT PERIOD: prolonged, late May/late July. 

PREIMAGINAL PHASES: studied in Europe (Henriksen, Kreutzer, 1982
and others) Foodplants: Viola, Veronica, Plantago, Hieracium,
Centaurea, Achillea. Eggs: yellowish-white, barrel-shaped,
ribbed in upper part, with a deep apical funnel, in batches on
leaf underside. Young larvae live gregariously, hibernate at 2nd
or 3rd instar in silken nests, then live solitarily. Mature larva:
dark with a lighter ventral side; head and ventral prolegs reddish;
each segment ringed by more than two rows of blueish-white dots
and a row of cream-white, brown or black false spines. Pupa: whitish-grey
with darker to almost black but lght-rimmed wing cases, brownish back,
orange warts and black dots on abdomen, sometimes looks lighter due to
a wax bloom.

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-22 mm. Wing upperside is 
ochre-orange in males, of the same colour or greyish in females, 
black spots partly fused into bansd; on hind wing underside 
postdiscoidal band contains 5-6 black dots, as different from a 
similar species M. arduinna, orange anal border absent. 

GEOGRAPHICAL VARIATION: Ural and W Siberia are inhabited by the 
nominotypical subspecies; ssp. tschujaca Seitz, 1908, described 
from Altai, rangess in the mountains of South Siberia, it is 
characterized by a more dull wing upperside ground colour and a 
widened dark pattern, that is especially characteristic for 
butterflies of Central Yakutia. 


177. Melitaea arduinna (Esper, 1784).

TYPE LOCALITY: Povolzhye: Kamyshin.

RANGE: Mostly mountainous regions from Greece through Caucasus, 
S Ural, Kazakhstan and  Central Asia to Altai (the villages 
Manzherok and Cherga). A local species. 

HABITAT: meadow steppes and herb meadows on south-exposed slopes 
and in river valleys. 

FLIGHT PERIOD: in South Ural prolonged, late May/middle July. 

PREIMAGINAL PHASES: A probable foodplant: Centaurea. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-23 mm. Wing upperside 
ochre-red or ochre-orange, black spots are partly fused into 
bands; on hind wing underside postdiscoidal band contains 
usually 5 large black dots, there is an orange anal border. 
Similar species: M. cinxia. 

GEOGRAPHICAL VARIATION: Butterflies from S Ural and Altai, 
exhibiting a great individual variation, are close to ssp. 
arduinna (= uralensis Eversmann, 1844); in montane-forest 
regions a red form with widened black pattern often 
predominates. 


SUBFAMILIA ARGYNNINAE Duponchel, [1835]

  The butterflies of intermediate ar relatively large size 
(F.w.l.: 14-40 mm). Wing upperside ochre-orange, less frequently 
yellowish, or greyish, with a pattern consisting of rows of 
black spots, sometimes fused into a network. Hind wing underside 
colouration  is usually a characteristic of the species. 
  Eggs are hemisphaeric or thimble-shaped. Larvae bear six 
lengthwise rows of branched spines. They feed on Viola, less 
frequently on Polygonaceae, Vacciniaceae and some others; as a 
rule they hibernate. Pupae have usually a pair of horns on head 
and paired acute prominence on dorsal side. Imagines actively 
visit various flowers. 


GENUS ARGYNNIS Fabricius, 1807
Type species: Papilio paphia Linnaeus, 1758.

  Relatively large butterflies: f.w.l. in our species: 25-40 mm. 
Wing upperside as a rule ochre-orange or ochre-red with black 
spots; fire wing outer margin straight or slightly concave; 
discoidal band absent on hind wing underside. 
  This genus, in a broad sense, includes about 55 specieswhich 
are often considered within a number of related genera. 


- Argynnis pandora (Denis et Schiffermuller, 1775).

TYPE LOCALITY: Austria.

RANGE: S Europe, N Africa, Anterior, Central and Southern Asia, 
NW, S and E Kazakhstan. Quite a common species in the adjacent to Russia 
regions of West Altai (Lake Markakol'), may be found out in SW 
Altaiskii Krai Region.

HABITAT: meadow/steppe patches in woods, on montane slopes. 

FLIGHT PERIOD: late June/middle August.

PREIMAGINAL PHASES: studied in Europe. Foodplants: Viola (V. 
tricolor and others). Eggs: yellow, pear-shaped, with 22-24 
lengthwise ribs. Larvae hibernate at early instars. Mature 
larva: purple-brown with three lengthwise light lines on back, 
slanting light dashes laterally of them, and large blueish-black 
crescent-shaped spot at fore margins of each segment, sometimes 
fused into a lengthwise band; spines short and sharp, yellowish 
or brown with dark setae; head black. Pupa: generally from 
greyish-brown to greenish-grey with metallic brands on back, 
with three dark streaks on wing cases. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 32-40 mm. Fore wing underside 
has a large brick-red area; hind wing underside greenish. 


178. Argynnis sagana Doubledau, 1847.

TYPE LOCALITY: N. China.

RANGE: The south-west of the West Siberian Lowland (Suzunskii 
Bor pine forest in the Novosibisrk; surroundings of Tomsk),the 
Salairskiy Kryazh elevation (at villages Suenga, Novososedovo in 
the Novosibirsk Region; many points in the Kemerovo Region); the 
Gornaya Shoria elevation (Kemerovo Region), the Kuznetskii 
Alatau Mts., NE Altai, the Sayans (the village Sutyagi, the 
Sisim River), the southern Pribaikalye (Irkutsk, the town 
Babushkin), Zabaikalye, the southern Far East, NE China, Korea, 
Japan. This species becomes very local westwards of Priamurye. 

HABITAT: in the Far East: mixed and broad-leaved forests; in the 
mountains the species inhabits valleys and clearings in 
spruce/fir forests and reaches the tree-line. In Siberia it 
occurs on long-forb meadows in piedmont forests, including the 
relict linden (Tilia sibirica) forests. Imagines were observed 
to feed on Cirsium heterophyllum, Phlomis tuberosa, Origanum 
vulgare; in cloudy weather the butterfly keep to the crowns of 
birches, asps and bird cherries.

PREIMAGINAL PHASES: studied in the Gornaya Shoria elevation by 
O. Kosterin and O. Berezina. Foodplant: Viola uniflora. Mature 
larva: brownish-black with a lighter dark-brown ventral side; 
abdomen with six rows of brownish-yellow spines with short black 
branches and black tips. Thoracic segments dorsally bear a pair 
of spines, those on 1st segments being 1.5 times longer and with 
blunt tips; besides, two pairs of lateral spines spring from 1/2 and 
2/3 segment joints. There are several warts set with black setae 
on sides of thoracic segments and bunches of black setae above 
prolegs. Pupa: golden-ochre coloured with a darker brownish-
grey reticulate ornament which, as becoming more dense, forms 
two lengthwise stripes along sides of abdomen and a double dark 
stripe its ventral side. There are two lengthwise strokes on the 
same colour on wing cases: in the middle and along anal margin. 
Five pair of short spines on dorsal side of thorax are rainbow-
glittering.

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-41 mm. In males basal half 
of hind wing underside ochre-coloured, outer one with a purple 
tint, border between them without white spots. The colouration 
of the females is unique for the subfamily: wing upperside 
black with a blue-green flash and large white spots and bands; 
underside brownish-green with the same white pattern. 

GEOGRAPHICAL VARIATION: The species was at first described by a 
male from China. Four years later A. Nordmann described a female 
of this species, originating from the Irkutsk vicinity, under 
the name paulina Nordmann, 1851. In the upper Ob' River Basin , 
on the Kuznetskoe Upland, NE Altai, and in the Sayan and adjacent 
territories there ranges ssp. relicta Korshunov, 1984, characterized 
by the largest black spots in males and smallest white spots in 
females; from more easterly regions the taxa paulina Nordmann, 
1851 and nordmanni Korshunov, 1984 (Zabaikalye, Priamurye) were 
described; butterflies from Primorye are usually attributed to 
the Japanese ssp. liana Fruhstorfer, 1907. 


179. Argynnis zenobia Leech, 1890.

TYPE LOCALITY: N China.

RANGE: S Primorye, NE China. A local species. 

HABITAT: open places with lime-stone rocks and screes in mixed 
and deciduous forests at elevations below 700 m. The imagines 
were seen to feed on the flowers Dracocephalum multicolor and 
Sorbaria sorbifolia. The males exhibit well expressed 
territorial behaviour, while the females spend much time on the 
flowers. 

FLIGHT PERIOD: middle July/middle September. 

PREIMAGINAL PHASES: studied by M.M. and M.A. Omel'ko (1978). 
Foodplants: Viola variegata. Eggs: ribbed, light-yellow, laid by 
1-3 on petioles or undersde of the foodplant leaf or on dry 
litter or gravel at the foodplant. The larva hatches on 13th-
15th day; it is dark with lengthwise rows of long greyish hairs. 
Itkeep to the leaf underside or petiole and eats the leaf 
margins; having been disturbed it rolls into a ring and falls 
into the litter. After the first moult in the age of 9-11 days 
there appear six lengthwice rows of black and fulvous spines. 
The larva hibernates at the third instar in dry rolled fallen 
leaves near foodplants; after hibernation it feeds on their 
leaves and flowers. The larva at the last (seventh!) instar is 
55-60 mm long, brown-black with a wide (2-3 mm) orange stripe 
along the back and six rows of black spines emerging from 
reddish warts. Pupation takes place on stones, twigs or barren 
tree roots. Pupa: brownish-black or brown (depending on 
illumination), its stage lasts for 25-29 days. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-39 mm in males, 36-44 mm in 
females; wing upperside ground colour fulvous in males, blueish-
grey in females; hind wing underside greenish with a network of 
silver cross-bands. 

GEOGRAPHICAL VARIATION: In the Russian territory there  ranges 
ssp. penelope Staudinger, 1891. 


180. Argynnis anadiomene Cajetan et Rudolf Felder, 1862.

TYPE LOCALITY: China.

RANGE: Priamurye (from the Zeya to the Gorin rivers), Primorye, 
NE and Central China, Korea, Japan.  

HABITAT: valleys, meadows, and clearings in montane mixed forests. 

FLIGHT PERIOD: prolonged, middle June/August.

PREIMAGINAL PHASES: studied in Japan (Fukuda et all, 1982). 
Foodplants: Viola spp. Eggs: yellowish-white with rows of spots; 
laid on various substrate near the foodplant. Larvae hatch 
usually in autumn and hibernate at the second instar. Mature 
larva: brown with an orange back line and rows of brown spines. 
Pupa: brown with greenish-ochre bloom, it has two rows of 
projections on abdomen, a conspicuous prominence on the thorax 
and two spines on head. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 34-38 mm in males; 37-42 mm in 
females. Fore wing outer margin distinctly concave; hind wing 
underside is greenish-ochre with a white spot at fore margin. 

GEOGRAPHICAL VARIATION: ssp. ella Bremer, 1864 (= obliterata 
Kardakov, 1928), described from the surroundings of Vladivostok, 
ranges in the southern Primorye. 


181. Argynnis paphia (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia, locally as northwards as the 
forest-tundra zone, the Sakhalin, the South Kuriles, Japan. 

HABITAT: meadows and open tree stand in various type forests, in 
the mountains locally reaches the tree-line. The imagines 
usually feed on Asteraceae and Apiaceae; they are capable of 
substantial migrations. 

FLIGHT PERIOD: prolonged, late June/early September. 

PREIMAGINAL PHASES: Foodplants: Viola, in Europe also Rubus 
(Rosaceae). Eggs: pear-shaped with 20-22 vertical ribs, at first 
yellow, later become brownish; laid singly, mostly of tree 
trunks and branches near the foodplant. Young larva: blueish-
grey with two white streaks along back and yellowish-brown 
spinules; it hibernates in the leaf fall soon after hatching. 
Mature larva: about 45 mm long; brown with two yellow lines on 
back and two ones on either side, besides, there are dark dashes 
and black and yellow dots on sides; spiracles ringed with 
yellow; spines long, brownish-red on back and black on sides; 
prothorax bears two much longer narrow processes; head black 
with white specks. Pupa: brownish-grey, pale- or dark-brown, 
with dark spots and golden dots and two obtuse prominences on 
dorsal side. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 27-36 mm. Hind wing 
underside is greenish with wide transversal silvery-white 
stripes. Females of a light-grey, with green flash, colouration 
(f. valesina Esper, 1793) are frequent. 

GEOGRAPHICAL VARIATION: The most part of the range is occupied 
with the nominotypical subspecies paphia. Ssp. neopaphia Fruhstorfer, 
1907 (=sachalinensis Matsumura, 1925) is known from Priamurye, 
Primorye, and the Sakhalin, it differs by a somewhat enlarged 
black pattern and the absence of f. valesina; ssp. geisha 
Hemming, 1934 (pro paphioides Butler, 1879) was reported for 
South Kuriles; besides, the form virescens Nakahara, 1926  has  
been described from the southern Sakhalin. 


182. Argynnis laodice (Pallas, 1771).

TYPE LOCALITY: S Russia.

RANGE: Central and Eastern Europe, Middle and South Ural,  the 
Kurgan Region, and, after a large gap, the southern Far East, 
Sakhalin, South Kuriles, NE China, Korea, Japan. 

HABITAT: meadow patches in broad-leaved, mixed, and, sometimes, 
dark-needle coniferous forests, also peat-bogs. 

FLIGHT PERIOD: from middle June to autumn.

PREIMAGINAL PHASES: studied in Europe (Schwarz, 1948 and others) 
and Japan (Fukuda et al., 1982). Foodplants: Viola (V. canina, 
V. palustris, and others). Eggs: conical with 17 lengthwise 
ribs, light yellow, later become purple. Hibernation occurs at 
the stage of the egg or just hatched larva. Mature larva: 
reddish-grey or brownish with two narrow interrupted cream-white 
or yellowish lines on back and a row of oval black spots on 
either side; spines are red and branched. Pupa: glossy, brownish 
or dark-brown, with narrow black or yellowish lines; most 
frequently suspended on dry vertical stems near the ground. . 

PECULIAR TRAITS OF IMAGO: F.w.l.: 26-35 mm. Fore wing has a 
straight outer margin and, in males, dark andraconial stripes 
along veins 2 and 3. Basal and outer halves of hind 
wing underside are of different colours, being separated by 
elongate white spots, the former is greenish-ochre, while the 
latter is pinkish-brown with a lilac tint; in females there is 
usually a small white apical spot on fore wing upperside. 

GEOGRAPHICAL VARIATION: Ssp. laodice Pallas, 1771 inhabits the 
European part of Russia,  Southern and Middle Ural. The 
butterflies from the Far East have in general larger black spots 
on wing upperside and are more variable individually and 
geographically, the following subspecies being reported: ssp. 
ussurica Kardakov, 1928: for forest region of Priamurye and 
Primorye; ssp. fletcheri Watkins, 1924: the Lowland of Lake 
Chanka; ssp. produota Matsumura, 1929: the Furughelm and Popov 
islands; ssp. ferruginea Watkins, 1924: the Sakhalin; ssp. 
japonica Menetries, 1857: the S Kuriles. 


183. Argynnis ruslana (Motschulsky, 1866).

TYPE LOCALITY: Primorye.

RANGE: Middle Priamurye, Primorye, the southern Sakhalin, the South 
Kuriles, NE China, Korea, Japan. 

HABITAT: meadows and clearings in broad-leaved and mixed 
forests, on open moderate elevations. The imagines occur 
together with A. laodice, but substantially less frequent, they 
often feed on Sorbaria sorbifolia. 

FLIGHT PERIOD: July/middle August.

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982); 
they resemble those of A. laodice. Foodplants: Viola. Eggs: conical, 
laid singly or in small batches on dead leaves or lichens. The 
larvae start feeding in early spring, they usually hide among 
withered leaves on the ground. Mature larva: greyish-brown with 
two narrow yellowish-white streaks on back and dark spots 
laterallly of them; spines brown, covered with whitish 
hairs. Pupa: brownish with marking, which is extremely dense on 
wing cases, and golden spots on thorax; the latter bears 
a prominence, on abdomen there are two rows of projections. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 30-38 mm. Fore wing has a 
concave outer margin and, in males, dark andraconial stripes 
along veins 1, 2 and 3. Hind wing underside have a transversal 
stripe composed of silvery spots which divides internal 
greenosh-yellow and outer pinkish-brown wing halves. 

GEOGRAPHICAL VARIATION: The Kurile islands are inhabited by the 
Japanese subspecies lysippe Janson, 1877, differing from the 
nominotypical one by an enlarged black pattern on wing 
upperside. 


184. Argynnis nerippe Cajetan et Rudolf Felder, 1862.

TYPE LOCALITY: China.

RANGE: southern regions of Primorye (the Gamova Peninsula, the 
Popova island, the settlement Khasan, the Chanka Lowland), 
North-East, Central, and South China, Korea, reported for 
Khabarovsk (Nebaikin, 1985). A local species. 

HABITAT: forb/Calamagrostis dry meadows and oak open woods on 
barren low elevations. 

FLIGHT PERIOD: late July/middle September.

PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). 
Foodplants: Viola (in Primorye Viola mandshurica). Eggs are laid 
by 1-3 on withered leaves, stems and gravel near the foodplants; 
the larvae hatch after 20-27 days and hibernate on the empty egg 
shells. Mature larva: grey or blueish-grey with a yellow streak 
along back and rows of brownish spines. Pupa: ochre- or dark-
brown. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 31-42 mm. Wing upperside 
light-ochre coloured, black spots of postdiscal row reduced to 
dots or entirely; hind wingh underside pattern consists of 
rounded silver spots on dark-ochre background. 

GEOGRAPHICAL VARIATION: Primorye is inhabited by the subspecies 
coreana, 1882. 


185. Argynnis niobe (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: Europe, Anterior and Central Asia, E Kazakhstan, the 
south of Ural, Siberia (northwards in Siberia to the southern taiga 
zone), and the Far East, Mongolia, N China. East of Zabaikalye 
is known by few records only. 

HABITAT: various types of meadows; in the mountains of S  
Siberia reaches the altitude of 2200 m. The imagines often occur 
together with A. adippe but usually much less frequently. They 
feed on various Fabaceae, Scrophulariaceae, Asteraceae, Apiaceae 
and other plants.

FLIGHT PERIOD: late June/late August.

PREIMAGINAL PHASES: studied in Europe (Henriksen, Kreutzer, 1982
and others). Foodplants: Viola, reported also Plantago and
Onobrychis. Eggs: conical with 14-15 lengthwise and numerous
transversal ribs, glossy, yellwoish- or reddish-yellow; laid by
1-3 on or near the foodplant; as a rule hibernate. Mature larva:
reddish-brown, with a square black spot on the back of each
segment, which is divided by a white back line and bordered
with a white spot on each side; spines light-grey, rose or
purple, covered with black hairs; a pair of spines just behind
head longer than other; head yellowish-brown. Pupa: reddish-
brown or brown-green with metallically glittering spots on spiny
prominences on dorsal side; suspended in hidden places on stems,
bark or branches.

PECULIAR TRAITS OF IMAGO: F.w.l.: 23-33 mm; in males andraconial 
stripes (sex bransd) along  veins on the fore wing weakly 
expressed; pattern of hind wing underside consists of the 
postdiscoidal row of ocelli and large silvery spots. In Ural and 
W Siberia a form with these spots being of the same colour as 
the background (f.eris Meigen) comprises about 70% of 
individuals. Similar species: A.adippe. 

GEOGRAPHICAL VARIATION: The butterflies of Ural and W Siberia 
are close to the nominotypical subspecies; ssp. changaica Reuss, 
1922, described for Zabaikalie and differing by a larger size, 
lighter ground colour and fine black pattern, ranges in the 
mountains of S Siberia; ssp. voraxides Reuss, 1921, approaching 
the nominotypical one, inhabits S Primorye. 


186. Argynnis adippe (Linnaeus, 1767).

TYPE LOCALITY: Austria.

RANGE: The non-tropical Eurasia northwards to the middle taiga 
zone, the Sakhalin, S Kuriles, Japan. 

HABITAT: various types of meadows, in the mountains reaches the 
tree-line. Imagines keep to flowering herbs, such as 
Sanguisorba, Inula, Crepis, Dianthus, Origanum, Campanula etc.  

HABITAT: middle June/late August, depending on the locaity. 

PREIMAGINAL PHASES: Foodplants: Viola, besides, Polygonum was 
also reported. Eggs: pear-shaped, yellwoish or greenish, later 
become reddish, with 18 longitudinal light-spotted ribs. 
Hibernation - at the stage of egg or first-instar larva. Mature 
larva, according to description by O. Kosterin from SE Chita 
Region: in general dark brown with six row of spines; on the 
thoracic segments the spines of the lowest row are absent and 
replaced with light lengthwise streaks, while the spines of the 
2nd row are absent on segment 1 (which have two spines only) and 
shifted to the hind margins of segments 2 and 3. The dorsal side 
is dark-brown with a fine reticulate darker ornament. Along the 
back there is a light-grey stripe interrupted on each segment 
between the pair of upper spines, in front of these spines the 
line divides a pair of large black irregularly quadrangular 
spots. The segment joints on the bakc are ash-grey, lighter than 
the ground colour. On either side a black stripe, light bordered 
above, goes through the bases of the spines of the 2nd row.  The 
spines, head and sides are light brown with tiny lighter dots. 
On the head there are dark stripes along the seams forming a 
double triangular chevrone and dark spots at the eyes. Pupa: 
usually suspended on a grass leave base in a silken net with 
large irregular cells. In SE Chita Region it was described by O. 
Kosterin as evenly reddish-brown with two rows of glittering 
knobs of which those on the mesothorax and 1st and 2nd abdominal 
segments are largest; besides, these two abdominal segments bear 
an additional pair of small glittering knobs laterally of the 
knobs of the main pair. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 25-36 mm; in males androconial 
stripes along veins 2 and 3 are conspicuous; pattern of hind 
wing underside is paler than in A. niobe; hind wing underside 
resemble that of A. niobe. In Ural and W. Siberia a form is 
common with th silvery spots replaced with those only slightly 
differing from the ground colour (f. cleodoxa Ochsenheimer, 
1807). In the Baikal region and eastwards a form predominates 
with an intensive greenish suffusion of the hind wing underside. 
Similar species: A.niobe, A. aglaja. 

GEOGRAPHICAL VARIATION: The following taxa has been described: 
zarewna Fruhstorfer, 1912: Irkutsk; vorax Butler, 1871: the 
Amur; xanthodippe Fixsen, 1887: the Ussuri; satakei Nakahara, 
1926: the Sakhalin; doii Matsumura, 1928: the S.Kuriles. The 
taxon pallescens Butler, 1873, described from Japan, is 
considered by Korean authors as a separate species and reported 
also for Kentei, the Ussury, and China, 


187. Argynnis aglaja (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The non-tropical Eurasia northwards to the northern 
border of the middle taiga zone, in the Ob' basin recorded up to 
the polar regions, the Sakhalin, the Kuriles, Japan. 

HABITAT: various meadows, including highland ones. The imagines 
are capable of substantial migrations. 

FLIGHT PERIOD: in the southern range is prolonged, late 
June/early September, in the middle taiga zone and highlands - middle 
July/late August. 

PREIMAGINAL PHASES: Foodplants: Viola, from Europe were also 
reported Polygonum bistorta, Vicia tenuifolia. Eggs: truncated 
conical with 26-28 lengthwise ribs, at first yellwoish-white, 
later become reddish-brown; laid by 1-3 underside the leaves or 
on stems of the foodplant or, less frequently, on various 
substrate near it. Larvae hibernate at early instars. Mature 
larva: up to 45 mm; brownish-black with tiny white dots, double 
interrupted white line along back, slanting light streaks at its 
sides, bright-red spots at spiracles of each abdominal segment, 
and a yellow line on eitheer side just above legs and prolegs; 
head and branched spines are dark. Pupa: brown with black areas 
on thorax; wing cases prominent, they exhibit a marbled pattern; 
abdomen strongly bent. The pupa is suspended on leaf underside. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 23-36 mm;  pattern of hind 
wing underside consists of large silver spots on a muddy-green 
ground colour; no ocell present in postdiscoidal area. 

GEOGRAPHICAL VARIATION: ssp. borealis Strand, 1901 tanges in 
Ural and the most part of Siberia; ssp. kenteana Stichel was 
reported for Zabaikalye; ssp. fortuna Janson, 1877 inhabits 
Priamurye and Primorye; Southern Primorye is inhabited with the 
most large ssp. graeseri Kardakov, 1928; the Sakhalin 
butterflies are known under the name matsumurai Nakahara, 1926 
(=otoniana Matsumura, 1928); for the South Kuriles ssp. basalis 
Matsumura, 1928 (= chishimensis Matsumura, 1928) was reported. 


GENUS NEOBRENTHIS Kocak, 1980 (= Brenthis Hubner, [1819]).
T.s.: Papilio hecate [Denis et Schiffermuller], 1775.

  F.w.l.: 15-26 mm. Wing upperside fulvous with dark spots. Hind 
wing underside has a yellowish band in discoidal area composed 
of angular spots; postdiscoidal area contains five, rarely six, 
spots. A Palearctic genus with three species. 


188. Neobrentis ino (Rottemburg, 1775).

TYPE LOCALITY: Germany.

RANGE: The non-tropical Eurasia, locally as north as the subpolar 
regions, the Sakhalin, the Kuriles, Japan. 

HABITAT: meadows of various kinds; in the mountains rises up to 
the tree-line. 

FLIGHT PERIOD: in the southern areas prolonged, middle June/middle 
August, in Kamchatka and the Magadan Region - in the second half 
of the summer.

PREIMAGINAL PHASES: Foodplants: Filipendula ulmaria, Sanguisorba 
officinalis, Rubus saxatilis, and other Rosaceae, and also 
Viola. Eggs: light-yellow, pear-shaped, with 12 longitudinal 
ribs; laid singly on leaf underside; may hibernate. The larva 
feeds only at night; it usually hibernate at early instars. In 
the Novosibirsk Region the larvae were found only on Filipendula 
ulmaria. Mature larva (O. Kosterin): light-brown with a 
complicated lengthwise ornament and a double white dorsal stripe 
bordered with a dark-brown line with a very fine white rim 
beneath. Two more dark-brown lines with fine white rims go 
through either side: just beneath the upper spine row and 
through the bases of the 2nd spine row. Besides, a white stripe 
goes through the bases of the lowest spine row, beneath it the 
ground colour becomes dark-brown. The spines are reddish-yellow 
with black spines. The head is of the same colour, with small 
black specks on the upper part, a dark horseshoe-shaped spot on 
the forheed and dark spots at the eyes; the head bears two small 
knobs and is set with black bristles. Pupa: yellowish-grey with 
a reticulate ornament, a brownish-grey darkening on the ventral 
side of the abdomen and shadows of the same colour on the sides 
of the abdominal segments, and two slanting brands on the wing 
cases, the latters  are dark-rimmed along the anal margin. There 
are two rows of glittering golden pointed knobs along the back. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-25 mm (in ssp amurensis - 
to 24 mm). As different from B. daphne, in the butterflies from 
Ural, South Siberia, Priamurye, Primorye, and the Sakhalin the  
black spots at the outer margin of the hind wing upperside are 
fused into a contiguous band; as different from B. hecate, the 
pattern of the hind wing underside consists of brownish lines 
and spots. Similar species: N. daphne. 

GEOGRAPHICAL VARIATION: The species is very variable 
geographically, the following taxa having been described from 
the territory considered: vitimensis Wnukowsky, 1929: (pro 
borealis Seitz, 1908): Ural, West and East Siberia; paidicus 
Fruhstorfer, 1907: the Altai Mts.; adalberti Fruhstorfer, 1916 
(pro Sibirica Seitz, 1908): the Sayans; limpida Krulikovsky, 
1909 (pro clara Staudinger, 1892): Zabaikalie; amurensis 
Staudinger, 1887: Priamurye and North Primorye; tarnis 
Fruhstorfer, 1919: the Ussuri River basin; magadanica 
Kurenzov,1970: the Magadan Region; siopelus Fruhstorfer, 1907: 
Kamchatka; parvimarginalis Nakahara, 1926 (pro karafutonis 
Matsumura, 1925): the Sakhalin; tigroides Fruhstorfer,1907: the 
South Kuriles. 


189. Neobrenthis daphne (Bergstrasser, 1780).

TYPE LOCALITY: Central Europe.

RANGE: The forest-steppe zone of Eurasia, the Sakhalin, the Kuriles, 
Japan. A local species. 

HABITAT: dry meadows in deciduous and mixed, mostly valley, 
woods. 

FLIGHT PERIOD: in South Ural inJune and early July; in Altai and 
easterly: late June/early August. 

PREIMAGINAL PHASES: Foodplants: Viola, Rubus idaeus, Rubus 
caesius, Filipendula, Sanguisorba officinalis. Eggs: conical 
with 14-16 lengthwise ribs, at first yellowish, later become 
reddish-purple; laid singly on the foodplant leaf underside. 
Mature larva: dark-brown with two yellowish streaks along back 
and three orange-yellow ones on either side; spines ochre-yellow 
with black tips; head yellowish-brown with brown spots 
and two short spinules. Pupa: ochre- or yellowish-grey with dark 
marbled bordering of wing cases; its strongly humped back 
has golden lunules on abdominal segments. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 21-29 mm. Black spots at outer 
margin of hind wing upperside are not fused into a band and may 
only be connected with black strokes; pattern of hind wing 
underside consists of brownish lines and spots, butterflies from 
Zabaikalye, Priamurye and Primorye have a lilac submarginal 
band. Similar species: N. ino. 

GEOGRAPHICAL VARIATION: South and Middle Ural and Altai are 
inhabited by the nominotypical subspecies; Pribaikalye, 
Zabaikalye, Priamurye, Primorye - by ssp. ochroleuca 
Fruhstorfer, 1907; the Sakhalin and Kunashir - by iwatensis 
Okano, 1943. 


190. Brenthis hecate (Denis et Schiffermuller, 1775).

TYPE LOCALITY: Austria.

RANGE: S Europe, Anterior and Central Asia, Kazakhstan, S Ural, 
steppen regions of W Siberia, Altai, the West Sayan. 

HABITAT: the patches of herb/grass steppes and steppefied meadows 
at pine wood edges, ravines, rock outcrops, dry barren mountain 
slopes. Feeding of imagines was observed on Trifolium, Goniolimon 
speciosum, Heracleum dissectum, Viburnum opulus.

FLIGHT PERIOD: June in Ural, in S Siberia locally to the end of 
July. 

PREIMAGINAL PHASES: studied in Europe (Niculescu, 1965). 
Foodplants: Filipendula vulgaris, reported also Onobrychis and 
Viola. Eggs: pear-shaped with 9-10 lengthwise ribs. Larva: 
brown. Pupa: chestnut-brown with paired golden spots on back, 
its phase lasts for forthnight. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-25 mm; hind wing underside 
colouration contrasted due to black lines and spots; submarginal 
area with a characteristic row of transversal black dashes. 
Similar species: N. ino.

GEOGRAPHICAL VARIATION: The butterflies of S Ural, the West 
Siberian Lowland, N and W Altai approach to the nominotypical 
subspecies; based on the old collections by H. Elwes from Altai 
(the villages Onguday, Kuray, the Chuyskaya Step') ssp. warren 
Kudrna, 1974 has been described. 



GENUS RATHORA Moore, 1900 (=ISSORIA Hubner, 1819)
T.s.: Papilio lathonia Linnaeus, 1758.

  F.w.l. of our species: 16-26 mm. Wing upperside ochre-orange 
with separate black spots. On hind wing underside the central 
band consists of spots with rounded angles; the largest of them, 
as well as the basal spots and spots at the outer margin, are 
silver; postdiscoidal area contains a row of seven dark spots 
with light pupils. 
  A world-wide distributed genus with 12 species.


191. Rathora lathonia (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: Europe, N Africa, the western half of Asia eastwards to the 
Upper Yenisei River basin, W Mongolia and the Himalaya; in 
Siberia does not penetrate northwards of the southern taiga 
belt. 

HABITAT: dry meadows, fields, vasting lands, in the 
mountains of S Siberia reaches the tree-line. The males 
often rest on plots of barren ground, most frequently on 
roads, and chase out any passing butterfly. 

FLIGHT PERIOD: early May/late August in two broods. The 
maximum abundance is usually observed in July, but active 
migration of imagines results in varying developmental 
schedule in different sites and different years. The second 
brood imagines often overwinter and fly early in spring. 

PREIMAGINAL PHASES: Foodplants: Viola, in the fields most 
frequently connected with a ruderal V. arvensis; Onobrychis 
arenaria, Anchusa, and Rubus were also reported. Eggs: 
greyish-white, obtuse conical with 20-22 lengthwise and also 
numerous irregular transversal ribs; laid singly on 
the foodplant leaves. In Siberia the larvae were observed in 
May and July. Mature larva: up to 40 mm in length; greyish-
black with tiny white dots, with two brown lines on either 
side and a white or yellowish stripe along 
back usually split into separate triangular spots with black or brown 
segment joints; spines short, brick-red or black, emerging from warts of 
the same colour; head cordate, brown or 
yellowish-brown with a dark face, without spines. Pupa: 
brownish or ash-grey with a white spot at wing case apex, a 
white transversal band in the middle of the body, and a 
yellowish lengthwise streak and golden spots on back. 
Hibernation occurs at any phase (including imago), most 
frequently in larval or egg one. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-26 mm. Fore wing outer 
margin straight or slightly concave; silver spots underside 
hind wing large, occupying a great area. 


192. Rathora eugenia (Eversmann, 1847).

TYPE LOCALITY: the Irkutsk Province.

RANGE: Polar and Subpolar Ural, the Putorana Plateau, the mountains 
of South and East Siberia, Kamchatka, the mountains of Mongolia, 
N China, Tibet. A local species. 

HABITAT: meadow, dwarf-birch, or stony tundras, alpine 
meadows, in the mountains of South Siberia at altitudes of 
1600-2900 m; in Polar Ural - mostly meadows in river valleys 
surrounded by larch-spruce open woodland. 

FLIGHT PERIOD: early June/July in moderate altitudes, middle 
July/late August in highlands and in the North. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-21 mm. Fore wing outer 
margin slightly conwex; on hind wing underside among silver spots 
the central one is prominent as being very elongate.

GEOGRAPHICAL VARIATION: In Pribaikalye and E Siberia there 
ranges ssp. eugenia Eversmann, 1847 (= vega Christoph, 1889, 
described from Yakutia). Butterflies from Altai and the 
Sayans, known as montana Bang-Haas, 1906 have a somewhat 
paler ground colour of hind wing underside; butterflies from 
Polar Ural have a similar appearance. Butterflies from S 
Zabaikalye (the mountain Golets Sokhondo) differ by on 
average larger size and a pale-grey wing upperside ground 
colour in females. Such dark females predominate also in 
populations of other arid regions of S Siberia and Mongolia. 


GENUS CLOSSIANA Reuss, 1920
T.s..: Papilio selene [Denis et Schiffermuller], 1775.

   F.w.l.: 14-32 mm. Wing upperside vary from ochre-coloured 
to brick-red, with black spots; postdiscoidal area of both 
wings has a row of roundish spots; on hind wing underside 
this row contains 6-7 spots. A peculiar element of  the hind 
wing underside pattern is a discoidal band which is lighter 
than the ground-colour. Having similar appearance, the 
species can be identified mostly by the hind wing underside. 
Further in the text only its pattern is characterized. 
  About 35 species of this Holarctic genus inhabit different 
tundra, meadow and marshy landscapes of the non-tropical 
Eurasia and North America. The formation centre of this 
genus is thought to be in the monutainous regions of NE 
Asia, which retains the maximal species diversity until 
present. 


193. Clossiana eunomia (Esper, 1799)
(= aphirape Hubner, 1799).

TYPE LOCALITY: Koenigsberg (at present Kaliningrad).

RANGE: The temperate Eurasia northwards to the forest-tundra, the 
Sakhalin, North America. A local species. 

HABITAT: floodland grass fens, peat meadows, alpine meadows, 
meadow and wet dwarf-birch tundras, open peat-moss bogged up 
woodland, raised bogs (including the "ryams" of the forest-
steppe), rarely pine woods. In the mountains this species 
occurs as high as 2600 m above sea level (SE Altai). The 
butterflies were observed to feed on the flowers of 
Polygonum, Hedysarum, Scorzonera radiata, Veronica, Ledum 
etc. 

FLIGHT PERIOD: mostly middle June/July, in the 
highlands and forest-tundra zone - in July. 

PREIMAGINAL PHASES: Foodplants: in S Ural and Siberia 
Polygonum bistorta, P. viviparum (Polygonaceae), Vaccinium 
uliginosum (Vacciniaceae),  Trollius europaeus 
(Ranunculaceae), from other regions also Viola palustris 
(Violaceae), Thalictrum, Caltha (Ranunculaceae), 
Pentaphylloides fruticosa (Rosaceae), Andromeda polyfolia 
(Ericaceae) are known. Eggs: conical with numerous ribs and 
an apical dimple, at first cream-white, later fulvous-brown; 
laid in batches up to 20 ones underside foodplant leaves. 
Larva: brown but looks yellowish or silvery-grey due to 
numerous  lighter markings on  back and sides, there is a 
yellow line on back and either side; head brown; spines 
orange, covered with minute hairs. It hibernates in the 
second instar. Pupa: pale-brown with dark spots and brown 
streaks on wing cases. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-23 mm. Discal band 
composed of angular silvery-white or ochreous spots;  
postdiscal area contain six small black-rimmed spots of the 
same colour. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies 
penetrates eastwards to South and Middle Zauralye 
[Transuralia]. The mountains of South Sivberia are inhabited 
by ssp. acidalia Boeber, 1809 (= asiatica Staudinger, 1901) 
described from Pribaikalye and differing from the 
nominotypical one by smaller size and a more fine black 
pattern. In the forest-tundra and taiga zones of Asia there 
widely ranges ssp. ossianus Herbst, 1800 differing by a 
darker  ground colour of both wing sides and a silvery 
colour of discal band on hind wing underside in both sexes. 


194. Clossiana selene (Denis et Shiffermuller, 1775).

TYPE LOCALITY: Vienna.

RANGE: The temperate Eurasia northwards to the forest-tundra, the 
Sakhalin, the Kuriles, Japan (Hokkaido), the temperate North 
America. 

HABITAT: forest and forest-steppe meadows, in the North also 
meadow and dwarf-birch tundras; in the mountains goes up to the 
tree-line. 

FLIGHT PERIOD: in the southern range - in two broods, in June 
and August, respectively, in the taiga zone and in the mountains 
- a single brood flies in July. 

PREIMAGINAL PHASES: Foodplants: Viola, less frequently Fragaria, 
in the north also Vaccinium uliginosum. Eggs: thimble-shaped 
with 16-18 longitudinal ribs, at first pale-yellow, later become 
greenish; laid singly or in small clusters on upperside of 
foodplant leaves. Mature larva: reddish-brown with light dots, 
double whitish streak along back and three yellowish lines on 
either side; spines ochre-yellow, reddish, or brown, with black 
hairs; those of a pair behind head much longer. In the southern 
range a part of larvae develop faster and give rise to a not 
abundant second brood, the rest of the larvae hibernate and 
pupates in May. Pupa: brown with yellowish-brown wing cases and 
four silver spots on its sides; suspended on a stem or petiole. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-21 mm. Discal band is 
composed of angular yellow spots and contains three silver 
spots, the other spots are yellow; there are seven silver spots 
at outer margin accompanied internally with six dark dots. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies ranges in 
the forest and forest-steppe zones of Siberia, butterflies from 
the mountains of Bureya, Priamurye and also Kamchatka, which 
have a somewhat more fine black pattern, are attributed to the 
same subspecies. Polar and Subpolar Ural and the forest-tundras 
of Siberia are inhabited by ssp. thalia Hubner, 1790 (= obscurum 
Sedykh, 1977), differing by a smaller size and a darker ground 
colour. In Primorye there ranges ssp. dilutior Fixsen, 1887 
differing by a narrower and paler wings and a weakly expressed 
basal suffusion. For South Primorye the taxon perryi Butler, 
1882 has been also recorded, which is considered as an 
independent species by some authors (Watkins). Ssp. chibiana 
Matsumura, 1927 was described from the Kunashir, it is also 
reported for the Sakhalin. 


195. Clossiana selenis (Eversmann, 1837).

TYPE LOCALITY: Middle and South Ural.

RANGE: The forest-steppe and forest zones from Povolzhye to 
the Far East; the mountains of South Siberia, the Sakhalin, 
Japan. The northern border needs in clarification, judging by 
few findings in the Lower Priobye (the town Labytnangi), the 
northern East Siberia (the Suntar-Khayata mountain rangem the 
Nera River, the mountains Madaunskie Gol'tsy), West Chukotka 
(the settlement Bilibino) it lies in the forest-tundra zone. 
Westwards of Altai this species becomes rare. 

HABITAT: in the southern Far East: dry meadows, steppefied 
slopes, waste lands; in more westerly range: forest and forest-
steppe meadows, open larch and pine woods; bogs in the southern 
and middle tayga belts, raised peat bogs (ryams) in the forest-
steppe; in the mountains sometimes reaches the tree-line. 

FLIGHT PERIOD: in Ural and Siberia, as a rule, in one brood 
in middle June/middle July, locally until middle August; in 
Primorye - in two broods in late May/June and August/September.

PREIMAGINAL PHASES: Foodplants: Viola is known from the southern 
range. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-23 mm. Discoidal band 
is composed of yellowish spots, less angular than in C. 
selenis, three of which can have a certain silver flash;
silver or white marginal spots absent. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies is known 
from Middle and South Ural. In South Siberia 
there ranges ssp. sibirica  Ershov,1870 (=festiva Krulikovsky, 
1923, with type locality E Siberia; = meinhardi Sheljuzhko, 
1929, with tyoe locality Tomsk) differing by a darker ground 
colour of outer part of hind wing underside; specimens from 
montane tundras of the Madaunskie Gol'tsy (the Magadan Region) 
were described as ssp. kononovi Kurenzov, 1970. This subspecies 
is characterized withan even reddish-brown ground colour of hind 
eing underside and weakly expressed postdiscal lilac lunules. 
From the northern Sakhalin ssp. onorensis Matsumura, 1925 is 
described, which is close to sibirica; Korea is inhabited with 
ssp. chpsensis Matsumura, 1927, which can be also found in the 
southern Primorye.


196. Clossiana euphrosyne (Linnaeus, 1758).

TYPE LOCALITY: Sweden.

RANGE: The temperate Eurasia north (in the Ob' River basin) to 
the forest-tundra, the Sakhalin, Japan (Hokkaido). 

HABITAT: forest, forest-steppe, and highland meadows, in the 
mountains of South Siberia reaches an elevation of 2500 m. 

FLIGHT PERIOD: in the forest-steppe zone in late May/June, in 
the North and in highlands - in July; in South Ural a second 
brood can occurs in August. 

PREIMAGINAL PHASES: Foodplants: Viola (in the Novosibirsk 
Region Viola hirta), less frequently Rubus and Fragaria 
(Rosaceae), in Finland (Seppanen, 1970) Vaccinium oxycoccum 
and Ledum palustre (Ericaceae s.l.) are known; Asteraceae 
and Hypericaceae were also reported. Eggs: thimble-shaped with 18-
20 lengthwise ribs, at first greenish-yellow, later become 
brown; laid singly on foodplant leaves. The larvae 
hibernate after 2nd-3th moult. Mature larva, according to 
observations by O. Kosterin and O. Berezina in the 
Novosibirsk Region: black with six rows of spines, those of 
subdorlas row bright-yellow, of suprastigmal and 
substigmal rows reddish-brown, spine apices and branches 
black. On either side of each segment there are several grey 
spots of different sides between suprastigmal and substigmal 
spines, which form a diffuse lighter lateral stripe. Spines 
on prothorax don't differ from others in sie. Pupa: grey 
with small dark specks and transversal dark strokes, forming 
double brackets on dorsal side of each segment; there is a 
dark stroke in the middle of wing cases.

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-23 mm. Discal band 
consists of angular ochre-yellow spots, but the central spot 
is silver and conspicuous; dark spots of postdiscal row lack 
light pupils. Similar species: C. oscarus, C. iphigenia. 

GEOGRAPHICAL VARIATION: weakly expressed. The taxa nephele 
Herrich-Schaffer (E Europe), orientisvivax Verity, 1932 (the 
E Sayan) and orphanus Fruhstorfer, 
1907 (Primorye) are quite close to the nominotypical 
subspecies, although in populations of forest-steppe 
regions larger light coloured butterflies with less 
expressed black pattern predominate. Somewhat more light 
coloured butterflies are found also in Kamchatka from where 
ssp. kamtschadalus Seitz, 1908 was described.


197. Clossiana iphigenia (Graeser, 1888).

TYPE LOCALITY: "the Amur downstream of Khabarovsk".

RANGE: Lower Priamurye, the eastern ridges of the mountains 
of Bureya, the Sikhote-Alin' (southwards to the Terney Bay), 
the Sakhalin, the South Kuriles, Japan, NE China. 

HABITAT: damp meadows in dark-needle and mixed forests, 
coastal meadows with the bush alder. 

FLIGHT PERIOD: middle June/middle August.

PREIMAGINAL PHASES: studied in Japan (Fukuda et al, 1982). 
Foodplants: Viola selkirkii is known from the continent, V. 
grypoceras - from the Sakhalin. Eggs: greenish with white 
ribs; laid singly on foodplant leaves or near it on 
faded leaves. The larvae feed at daytime. Mature larva: 
dark-brown with rows of rosy spines. Pupa: slender, with a 
sharp prominence on thorax, brown, covered with tiny 
lighter knobs, abdomen lighter; wing cases with a marble 
pattern. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 20-25 mm. Discal 
band consists of angular yellow spots, central spot 
silver; postdiscal spots contain silver dots. Similar 
species: C. euphrosyne, C. oscarus.

GEOGRAPHICAL VARIATION: The Sakhalin and Kunashir are 
inhabited by butterflies, described as ssp. sachalinensis 
Masumura, 1925, which little differ from the nominotypiacl 
subspecies ranging on the continent.


198. Clossiana oscarus (Eversmann, 1844).

TYPE LOCALITY: Altai.

RANGE: The temperate Eurasia to the east of the Priobye [the 
Ob' River basin), the Sakhalin; locally penetrates far 
northwards, to the town Surgut in the West and to Magadan 
in the East. Reported for Middle Ural (V. and A. Lukhtanov) 
but we have no data from there. A local species. 

HABITAT: forest meadows of lowlands and mountains, 
open peat-moss larch woodland ["mar's"]; in the Koni Peninsula 
(the  Magadan  region)  these  butterflies  were  observed  on 
meadows among sparse bushes of dwarf pine (Pinum  pumila)  and 
bush alder (Dushekia fruticosa). Feeding of imagines was 
observed on Polygonum bistorta, Dracocephalum ruyschiana, 
etc.

FLIGHT PERIOD: late May/July; in the Magadan Region middle 
July/middle August.

PREIMAGINAL PHASES: Foodplants: Viola. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 19-26 mm. Hind wing 
underside ground colour brick-red; discal band composed of 
yellowish spots without brilliancy; there are seven 
semicircular silver spots at outer margin. 

GEOGRAPHICAL VARIATION: Siberia, the mountains of Bureya 
and the lower Amur basin are inhabited by ssp. oscarus. 
Ssp. oscaroides Menetries, 1859, described from Yakutia, is 
close to it, differing by somewhat larger black spots on 
wing upperside. Middle Priamurye and Primorye are inhabited 
by a larger and more brightly coloured ssp. australis 
Graeser, 1888. For the Sakhalin ssp. sachalinensis Matsumura 
has been reported, this name needs replacement as a homonym.


199. Clossiana angarensis (Ershov, 1870).

TYPE LOCALITY: "the Irkutsk Province".

RANGE: The forest-tundra and forest zones from the 
Arkhangelsk Region to the Magadan regions and Chukotka (the 
Anadyr' River) (yet not found in Kamchatka); the adjacent 
mountains, the Sakhalin; found also within the forest-steppe 
of West Siberia: in North Baraba at the village Kayly, in 
the surroundings of Novosibirsk. In the southern part of its 
range this is a local species. 

HABITAT: damp forest meadows, dwarf birch thickets, raised 
bogs, open bogged up larch woods, pine woods; in the 
mountains reaches the tree-line. In the Krasnoyarsk environs 
the imagines were observed to feed on the flowers of 
Scutellaria galericulata, Galium menotispermum, Comarum 
palustre, Pulmonaria officinalis. 

FLIGHT PERIOD: in the southern taiga belt late June/middle 
August, in the forest-tundra in July.

PREIMAGINAL PHASES: not studied. Probable foodplant on Polar 
Ural: Polygonum bistorta.

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-26 mm. Discal band 
consists of angular ochre-coloured spots, three of which may 
be silvered; postdiscoidal area is brownish with lilac spots 
and six large black spots, the postdiscoidal spots 
proximally with adjacent diffuse lunular lilac-coloured 
spots; outer margin with seven elongate silver spots. 

GEOGRAPHICAL VARIATION: ssp. angarensis ranges in 
Pribaikalye, Zabaikalye and ?Priamurye. The butteflies from 
highlands (1300-2000 m above sea level) of the Sayans were 
described as ssp. alticola Sushkin et Tshetverikov,1907 
differing by a smal size, narrow wings, widened black 
pattern, especially in basal part, and narrow discal band on 
hind wing underside. By materials of Yakutia ssp. herzi 
Wnukowsky, 1927 was stated, rather close to alticola, it 
ranges in Central and East Siberia. In Ural and West Siberia 
ssp. samkoi Sheljuzhko, 1931 (with the type locality 
Tobol'sk) ranges. From the Sakhalin ssp. miakei Matsumura, 
1919 was described, differing by a somewhat more pale 
upperside ground colour  and large black spots on wing basal 
half while hose on outer half are somewhat reduced. Similar 
specimens are known from Lower Priamurye. In the Kedrovaya 
Pad' Nature Reservation the Korean subspecies hacutozana 
Matsumura, 1927 was recorded, earlier considered as an 
independent species (Kurenzov, 1970). 


200. Clossiana thore (Hubner, 1806).

TYPE LOCALITY: the Alps of Tirol.

RANGE: The forest-tundra and forest zones from the 
Arkhangelsk Region to the Magadan regions and Kamchatka (the 
Anadyr' River) (yet not found in Chukotka); the adjacent 
mountains, the Sakhalin; found also within the forest-steppe 
of West Siberia: in North Baraba at the village Kayly, in 
the surroundings of Novosibirsk. In the southern part of its 
range this is a local species. 

HABITAT: damp meadows, open tree stands, mostly in 
coniferous forests, dwarf birch thickets, raised bogs, open 
boggy larch woods; in the mountains reaches the tree-line. 
Feeding of imagines was observed on Polygonum bistorta, 
Veronica longifolia, Ligularia sibirica, Senecio praticola, 
Crepis sibiiirica, Dactylorhiza fuchsii. 

FLIGHT PERIOD: middle June/August, depending on the locality. 

PREIMAGINAL PHASES: studied in Finland (Pekkarinen, 1977) 
and the Ob' River basin (Y. Korshunov). Foodplants: Viola 
altaica, V. biflora, V. repens, V. selkirkii, V. riviniana, 
V. canina, V. palustris are known. Eggs: greenish with white 
grainy bloom, thimble-shaped with longitudinal ribs; laid 
singly on leaves of foodplant or near it on withered grass. 
Young larva: green with white dots along back and light 
spinules; it feeds at day-time; after the first moult 
becomes brownish-grey. In Finland two hibernations were 
observed, at 2nd and 4th instar. Mature larva: brown with an 
interrupted yellow streak along back and a contiguous one 
along eitgher side, with dark-brown spots on sides; spines 
yellowish or pinkish with black setae. Pupa: yellowish-brown 
or greyish with lighter brands, with conspicuous paired 
projections on head,thorax, and abdomen; leg and antennae 
cases black. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-23 mm. Hind wing 
underside ground colour ochre-brownish; discal band consists 
of angular ochre-coloured spots; the postdiscoidal area has 
a lilac tint at fore and anal margins and diffuse dark-brown 
spots; silver, white or distinct black spots absent. 

GEOGRAPHICAL VARIATION: For Siberia ssp. borealis 
Staudinger, 1861 used to be reported, which was described 
from Lapland. However, in Ural, the Lower and Middle Priobye 
butterflies range with much enlarged black pattern and oval, 
extended along veins, postdiscal spots. We attribute them to 
ssp. transuralensis Sheljuzhko, 1931. In the mountains of 
South Siberia there ranges ssp. hypercala Fruhstorfer, 1907, 
stated from Altai, with smaller black spots, especially on 
fore wing underside ahere they are alvays separate from each 
other. For the souhtern Far East  ssp. hyperusia 
Fruhstorfer, 1907 (= meridionalis Kardakov, 1928) was 
reported, which is characterized by a clear-cut black 
pattern, weakly expressed basal suffusion on hind wing 
upperside. Butterflies from the Sakhalin were described 
under the name karafutonis Matsumuro, 1919. 


201. Clossiana frigga (Becklin in Thunberg, 1791).

TYPE LOCALITY: Lapland.

RANGE: The tundra, forest-tundra and northern forest zones of 
Eurasia, the adjacent mountain countries southwards to the 
Tarbagatai, Mongolia, and the Sikhote-Alin', the northern 
Sakhalin, the northern half of North America. A local 
species.

HABITAT: raised bogs, peat-moss 
open woodland, tundras of bushy, fruticulose, or Dryas/lichen 
types, in the mountains of South Siberia mostly marshy areas with 
dwarf birch and bush willows in brook headwaters, on plateaux, 
saddles, flattened ridges at altitudes of 1900-3000 m. The 
imagines were observed to feed on Ledum palustre, Lagotis 
integrifolia, Polygonum viviparum, Pedicularis. 

FLIGHT PERIOD: in different sites from middle June (the middle 
taiga zone) to August. 

PREIMAGINAL PHASES: studied in Scandinavia (Henriksen,
Kreutzer, 1982). Foodplants: Rubus chamaemorus, from Alaska
Dryas integrifolia is also known, and also Salix and Betula
are known. A biennial species. Eggs: pale salmon-coloured,
cone-shaped, ribbed. Larva after first hibernation: dark
olive-green with six rows of black branchy spines, set with
fine light-brown dots and dense hairs, head black with two
spines; ventral prolegs brown. Mature larva after
second hibernation: brown with wide lateral lines composed of
small pale-ochre spots. Pupa: obtuse, brown with darker wing
cases and dorsal side, light lateral streaks, dark-brown spots
on dorsal sides of segments, and pairs of pale spinules on abdomen.

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-24 mm. Basal area of 
hind wing underside reddish-brown,  postdiscal area is 
lighter, reddish-violet or ochre-brownish; discal band 
consists of roundish yellowish or brownish spots, two of 
which,  central and that at fore margin, are much lighter 
and usually silvered. 

GEOGRAPHICAL VARIATION: This species is very variable 
geographically. The nominotypical subspecies ranges in the zonal 
tundras, forest tundras and taiga zones of Eurasia; in the 
mountains of South Siberia ssp.  alpestris Elwes, 1899 ranges, 
differing by a smaller size, paler wing upperside ground colour 
and narrower discal band on hind wing underside. For thw 
southern Far East ssp. maritima Kardakov, 1928 was reported, 
which is characterized by an orange-yellow ground colour of wing 
upperside, a paler colouration of basal half of hind wing 
underside and rahter a wide discal band. - in the southern Far 
East. 


202. Clossiana improba (Butler, 1877).

TYPE LOCALITY: the Arctic America.

RANGE: The zonal tundras of Eurasia and North America. 

HABITAT: subfrutex tundras (with Empetrum, Dryas, Cassiope, 
etc.) on flat elevations. The imagines keep low to the 
ground (and so are hardly noticeable) or congregate on 
meadow patches on southern slopes of river terraces and 
ravines, where they actively visit the flowers of Trientalis 
europaea, Taraxacum, Crepis nigrescens, Polygonum viviparum. 

FLIGHT PERIOD: July/early August.

PREIMAGINAL PHASES: studied in Scandinavia (Henriksen,
Kreutzer, 1982) and North America (Scott, 1986).  Probable
Polygonum viviparum, often growing under low willow shrubs, that
may explain reports on Salix herbacea as a probable foodplant in
Finland. Eggs: thimble-shaped, smooth with
vague ribs and an apical dimple, at first cream-white, later become
orange-yellow and fulvous-brown; laid singly on foodplants and
other neighbouring plants. A biennial species. The larva
hibernates at 1st and 4th instar. In the first year the
larva is  dark-brown with black back stripes, covered with
dense hairs; spines dark-brown; head black with two spines and
brown spots above eyes. Later cream-white lateral stripes
appear. Pupation takes place between two silk-fastened leaves. Pupa:
with brown spots and fulvous-brownlines along the back;
abdonimal segments 5-7 with numerous grey triangles
accompanied with two black lunules.

PECULIAR TRAITS OF IMAGO: F.w.l.: 
14-17 mm. Hind wing underside resembles much that of C. 
frigga but is more heavily suffused with dark scales.


203. Clossiana titania (Esper, 1793) (= amathusia auct., 
nec Esper, 1784).

TYPE LOCALITY: Sardinia.

RANGE: Mostly mountain regions of the temperate Eurasia, 
including Ural (from South to Subpolar), the southern half 
of Siberia NE to SW Yakutia (the Nyuya River). By few 
records is known from S Zabaykalye (the mountain Golets 
Sokhondo), the Lower Priamurye and the North Sakhalin 
(Kurentzov, 1970). A local species. 

HABITAT: forest and higland meadows, open larch woodland, 
stony and dwarf birch montate tundras; in Lower Priamurye 
and the Sakhalin - peat moss bogs. In West Siberia feeding 
of imagines was observed on Polygonum bistorta, Scabiosa 
ochroleuca, Myosotis caespitosa, Veronica longifolia, Inula, 
Leucanthemum, Crepis sibirica.

FLIGHT PERIOD: late June/middle August, depending on the 
locality.

PREIMAGINAL PHASES: Foodplants: Polygonum bistorta reported 
from Middle Ural (Y.N. Baranchikov), Filipendula ulmaria and 
Trollius asiaticus from the Krasnoyarsk environs (Y.P. 
Korshunov), Viola and Vaccinium uliginosum from Europe. Eggs: 
thimble-shaped, ribbed, brownish with a white suffusion; laid 
singly underside leaves or on twigs of foodplants. The larva 
hibernates usually at the third instar. Mature larva: 
greyish-brown, ash-grey, or dark-brown, with black lateral 
and back stripes interrupted by the segment joints;  
spines yellow or dark-yellow; black head bears longer and 
lighter spines. Pupa: brownish- or yellowish-grey with a 
black pattern on back and wing cases; abdominal knobs black; 
the pupal stage lasts for about fortnight. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 19-24 mm. Discoidall band 
consists of angular ochre-coloured spots with veins suffused 
with brown, postdiscoidal area contains a row of lunular lilac 
spots distally of dark ocelli; outer margin without white or 
silvery spots. 

GEOGRAPHICAL VARIATION: From Middle and South Ural the East 
European subspecies rossica Hemming, 1934 (= bivina Fruhstorfer, 
1908) is known. North Ural, West Siberia ane easterly ssp. 
staudingeri Wnukowsky, 1929, ranges, which differ by an enlarged 
black pattern, especially in the butterflies from Subpolar and 
North Ural, probably deserving separation into a special 
subspecies. For the Sakhalin ssp. miyakei Matsumura, 1919 has 
been described, probably, the same subspecies ranges in the 
lower Amur River basin. 


204. Clossiana chariclea (Schneider, 1794).

TYPE LOCALITY: Lapland.

RANGE: Polar regions of Eurasia, Kamchatka, Alaska, North 
Canada. 

HABITAT: in the Yamal Peninsula: riparian Carex/subfrutex 
asociations among willow bushes within the subzone of typical 
tundras; in the Taymyr Peninsula - in the same biotopes and, 
besides, in polygonal tundras; in Chukotka and Kamchatka prefers 
in montane tundras of various types. 

FLIGHT PERIOD: July/middle August, depending on the 
peculiarities of the year.

PREIMAGINAL PHASES: Probable foodplants: Cassiope tetragona is 
reported from Lapland, Salix arctica and S. reticulata from the 
Arctic America, Dryas integrifolia from Greenland. Eggs: light-
yellow, with lengthwise ribs and tiny hairs. Larva and pupa are 
not known. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-19 mm. Discal band consists 
of angular ochre spots separated by brown-suffused veins, three 
of the spots, rarely all, are whitish; at outer margin there are 
seven elongate silver spots. 

GEOGRAPHICAL VARIATION: The most part of Siberia is inhabited by 
the nominotypical subspecies; Chukotka and Kamchatka - by ssp. butleri 
W.H. Edwards, 1883, differing by a more saturated ground colour 
of wing upperside and somewhat reduced dark postdiscal spots. 
Some authors consider it as an independent species.


205. Clossiana dia (Linnaeus, 1767).

TYPE LOCALITY: Austria.

RANGE: The forest-steppe and southern forest zones of Eurasia,  
east of Zabaikalye is known by few findings from West Yakutia 
and the Magadan Region.

HABITAT: forest and forest-steppe meadows, in the mountains also 
larch parklands, alpine meadows and dwarf birch tundras up to 
2500 m altitude (Central Altai). 

FLIGHT PERIOD: late May/late June in the southern forest zone, 
in the mountains - to middle July; in the forest-steppe zone 
flies in two broods from May-June to August. 

PREIMAGINAL PHASES: Foodplants: Viola, also reported Rubus 
idaeus, Prunella vulgaris. Eggs: pale-green, truncated conical, 
with 20 lengthwise ribs, dimpled. Larva: up to 30 mm in length; 
dark- or blueish-grey with dark white-margined stripe along  
back, on either side of each segment there is a yellow spot 
ringed with wide black margin; spiracular line and   head 
reddish-brown; spines yellowish, lighter on sides, they emerge 
from reddish-yellow warts; frons is marked with  a  black 
triangle. Pupa: chestnut-brown with golden spots.  

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-18 mm. Hind wing underside 
ground colour dark-brown with a distinct violet tint; spots of 
discal band roundish, three of them distinctly bright-silvered, 
cell with a light black-rimmed spot. 

GEOGRAPHICAL VARIATION: weakly expressed. Small butterlies from 
the Altai highlands were described as ssp. alpina Elwes, 1899, 
which is pertained to the mountains of South Siberia. 


206. Clossiana freija (Becklin in Thunberg, 1791).

TYPE LOCALITY: Sweden.

RANGE: North Europe, tundra, forest-tundra and forest zones of 
Asia; the adjacent mountain countries southwards to Mongolia, 
Japan (Hokkaido), the northern half of North America; found on 
the Sakhalin (the Tymovskii district, collected by S. 
Pervukhin).  

HABITAT: open pine or larch woods with lichen or peat-moss 
ground layer, raised bogs, damp Eriophorum meadows,  thickets of 
willow bushes, bushy and subfrutex tundras; in the mountains of 
South Siberia the dwarf birch tundras of gentle northern slopes, 
marshy areas, damp forest openings, at 1000-2500 m altitudes. 
Butterflies often rest on wet moss, observed to feed on 
Polygonum bistorta, Ledum palustre, Dryas octopetala. 

FLIGHT PERIOD: in different sites from May (the southern 
taiga) to late July (the extreme North and high elevations), in 
one brood. 

PREIMAGINAL PHASES: Foodplants: in Siberia: Rubus chamaemorus,
Vaccinium uliginosum, V. vitis-idaeus, Empetrum nigrum; in
Scandinavia: Arctostaphilus uva-ursi, A. alpina; North America:
Dryas integrifolia; Japan: Rhododendron aureum, Sieversia. Eggs:
yellowish, later become orange; laid singly underneath leaves of
foodplants or neighbouring plants. Larva at the first instar
(Henricksen, Kreutzer, 1982): brownish-black, covered with long
pale-brown hairs, which after the first moult are replaced by
low branched red-brown hairy spines. Hibernation occurs at the
fourth instar. Pupa: stout, purple-brown with a bent abdomen
bearing two rows of small knobs; wing cases outlined with black;
suspended beneath foodplant leaves close to the ground.

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-21 mm. Hind wing underside 
mottled in a peculiar way: discal band outlined with dentate 
black lines, it consists of acute-angular spots stretched out 
along veins, 2-3 of spots white, others suffused with brown 
scales. 

GEOGRAPHICAL VARIATION: The nominotypical subspecies is known  
from North Europe, Ural and the West-Siberian Lowland; Altai, 
the Sayans, S Pribaikalye, S Zabaikalye, and Yakutia are 
inhabited by ssp. jakutensis Wnukowsky, 1927 (= calais 
Fruhstorfer, 1916, type lolality: Tuva) stated for Central  
Yakutia, it differs with a much more light (light-ochre) 
wingupperside ground colour, more fine black pattern, darker 
ground colour of basal part and reduced black postdiscal spots 
on hind wing underside. 


207. Clossiana polaris (Boisduval, 1828).

TYPE LOCALITY: Nordkap (North Norvegia)

RANGE: Polar regions of Eurasia (including the Wrangel island) 
and North America. 

HABITAT: subfrutex, Dryas or Cassiope, and stony tundras on hill 
tops and slopes, on plateaux and river terraces  in mountainous 
regions. In the surroundings of the settlement Tareya in the 
Taymyr Peninsula the imagines were observed to feed on Lagotis 
minor and Dryas octopetala. 

FLIGHT PERIOD: depending on the locality, from July to 
middle August. 

PREIMAGINAL PHASES. The natural history of the species is poorly 
investigated, notwithstanding that this was the first butterfly 
species the biology of which was attempted to be studied in 
Siberia. Lewaski (1887) discovered mature larvae as long ago as 
in the end of the previous century, in the Shaitan mountains in 
the Taymyr Peninsula, on sunny slopes on 10th of May. They, 
according to the author's note, fed on "mosses". In the middle 
of June they pupate and the butterflies hatched after 20-22 
days. By recent observations: foodplants: Dryas octopetala is 
known from Polar Ural, Cassiope tetragona and Vaccinium 
uliginosum from Lapland. Eggs: greenish-yellow with fine 
lengthwise ribs. A biennial species hibernating just after 
hatching from the egg and in the last instar. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-23 mm. Basal part of hind 
wing underside reddish-brown, outer part  ochre-fulvous. 
Discoidal band composed of angular spots, strongly dusted with 
brown scales, inconspicuous on ground colour; outer margin with 
seven roundish white spots. In male genitalia the apex of costal 
processus of valva club-shaped. Similar species: C. distincta, 
C. erda. 


208. Clossiana erda (Christoph, 1893).

TYPE LOCALITY: Yakutia.

RANGE: E Siberia from the Putorana Plateau (the Tanlakh River) 
to West Chukotka (the settlement Bilibino), south to the Lena 
River sources, Zabaikalye (the Yablonovyy mountain range), and 
Lower Priamurye (Komsomol'sk-na-Amure). A local species. 

HABITAT: meadows and rocks in the valleys of mountain rivers and 
brooks, open south-exposed slopes, pebble banks, open larch 
woods, montane tundras. 

FLIGHT PERIOD: middle June/middle July.

PREIMAGINAL PHASES: not studied. On the Suntar-Khayata mountains  
oviposition was observed by V. Dubatolov on Vaccinium vitis-
idea.

PECULIAR TRAITS OF IMAGO: F.w.l.: 21-26 mm. As different from 
similar species C. polaris, C. alberta, C. astarte and C. 
tritonia, wing ground colour more bright, ochre-orange, black 
spots being smaller and more distinct; pattern of hind wing 
underside also more bright and mottled; in male genitalia the 
apex of costal processus of valva club- or shortened foot-
shaped. 

GEOGRAPHICAL VARIATION: poorly studied. The taxon dulkeiti 
Kurenzov, 1970, described as a species by a male and female 
from the Omsukchanskii mountain range in the Magadan Region, 
most probably in fact belongs to this species. 


209. Clossiana alberta (Edwards, 1890).

TYPE LOCALITY: Canada: the Alberta Province.

RANGE: This characteristic species for North American montane 
tundras has been found, together with Clossiana astarte, in the 
tundras of Chukotka (the Provideniya Bay, Goryachie Klyuchi [Hot 
Sources]) and to the west of the peninsula, on the Anyuyskiy 
mountain range. 

HABITAT: stony tundras. The imagines keep to rocky crests and 
hill tops in tundras. 

FLIGHT PERIOD: July/early August.

PREIMAGINAL PHASES: Foodplants: Dryas octopetala (Kurentzov, 
1970), from North America Vaccinium was also reported. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-21 mm. Wing upperside 
ground colour is ochre, more dull than in C. erda, black basal 
suffusion beingless developed than in C. polaris and C. astarte. 
Discal band on hind wing underside is somewhat wider than in C. 
astarte. In male genitalia thr apex of  costal processus of 
valva club-shaped, that of caudal processus has two teeth 
(Table...). 

GEOGRAPHICAL VARIATION: The subspecies kurentzovi Wyatt, 1961 
was described from Chukotka, differing by a more developed black 
pattern on wing upperside, a darker groud colour of basal half 
on hind wing underside and a wider discal band. 


210. Clossiana tritonia (Boeber, 1812)

TYPE LOCALITY: Pribaikalye.

RANGE: Pribaikalie,  Zabaikalye, Priamurye,the mountains of 
Bureya, E Mongolia. A local species. 

HABITAT: rock outcrops within the forest belt, steppefied 
slopes. Feeding of imagines was observed on Thymus and Spiraea.

FLIGHT PERIOD: eary June/middle July.

PREIMAGINAL PHASES: Foodplants: Saxifraga bronchialis is known 
(Graeser, 1888). 

PECULIAR TRAITS OF IMAGO: F.w.l: 24-30 mm. Hind wing underside 
ground colour ochre-orange, discal band evenly ochre- or 
whitish-coloured, with a slight suffusion of dark scales. As 
different from C. astarte and C. matveevi,  general pattern is 
more bright, dark basal suffusion on hind wing upperside is less 
expressed. As different from C. erda, in male genitalia the apex 
of costal processus of valva is foot-shaped, caudal processus is 
short, with 2-3 apical teeth (see Table ...). 

GEOGRAPHICAL VARIATION: Ssp. tritonia is known from Pribaikalye. 
Butterflies from the Malakhanskii mountain range in Zabaikalye 
were described as elatus Staudinger, 1892; those from Upper 
Priamurye as amphilochus Menetries, 1857. Comparison of 
specimens from N Pribaikalye and Priamurye (the Komsomol'skii 
Nature Reservation) showed great differences not only in the 
wing pattern (the specimens from Priamurye have on hind wing 
underside a more evenly coloured ochre-yellow discal band and a 
more rounded white marginal spots isolated from the wing margins 
with black lunules) but also in the length of the caudal 
processus of the valva in the male genitalia. Probably, they are 
different species, more extensive materials being necessary to 
solve this problem. 


211. Clossiana astarte (Doubleday at Hewiston, 1847).

TYPE LOCALITY:

RANGE: Polar Ural (the railway station 110th km), the 
Putorana Plateau, the Prilenskoe [pertained to the Lena River] 
Plateau, the mountains of East Siberia south to N Pribaikalye 
(the Khamar-Daban mountain range), Chukotka, the northern Far 
East, reported for Kamchatka (Sedykh, 1979) and the Central 
Sikhote Alin' Mts. (Kurentzov, 1971); Alaska. The western and 
southern borders need in clarification. 

HABITAT: montane tundras, rock outcrops in dry open larch 
forests. 

FLIGHT PERIOD: early June/middle July in the Prilenskoe Plateau, 
late June/late July in mountainous regions. 

PREIMAGINAL PHASES: On the Suntar-Khayata mountain range 
oviposition was observed by V.V.Dubattolov on dry peduncles of 
Saxifraga spinulosa, in Chukotka the butterflies keep to the 
bunches of Saxifraga punctata. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 17-26 mm. Hind wing underside 
ground colour varies from ochre-orange to reddish-brown. As 
different from C. tritonia, general pattern is more dull, dark 
basal suffusion on hind wing upperside is more expressed; on 
hind wing underside a discal band is strongly suffused with dark 
scales, in submarginal area there are a row of transversally 
elongate white spots united with wing margin with white strokes. 
As different from C. alberta, C. erda, C. polaris, and similarly 
to C. tritonia tritonia, in male genitalia the apex of costal 
processus of valva is foot-shaped, caudal processus, as 
different from C. matveevi, has 2-3 apical teeth. 

GEOGRAPHICAL VARIATION: A complex taxon probably embracing 
several distinct species. The following subspecies are known: 
the Putorana Plateau; 
ssp. dubatolovi Korshunov, 1987 was descrived the Khamar-Daban 
mountain range (the surroundings of the Cherskiy peak); ssp. 
machati Korshunov,  1987 from Polar Ural (between 100th and 112th km 
of the railway Seyda - Labytnangi). Chukotka is inhabited by the 
smallest ssp.  
tschukotkensis Wyatt, 1961, characterizing by in general 
darker of wing upperside, ssp. distincta Gibson, 1920 is known 
from Alaska. Below a subspecies from the Prilenskoe Plateau is 
described.

     Clossiana astarte ershovi Korshunov et P. Gorbunov, 1995 

     Original description:
   "...Butterflies from the Prilenskoe [near the Lena River] 
Plateau differ by relatively large size (f.w.l. is 22-24 mm in 
males, 23-26 mm in females) and the pattern of the hind wing 
underside. 
   Clossiana astarte ershovi Korshunov et P. Gorbunov, sbsp.n.
   HOLOTYPE: A male. F.w.l. 22 mm. The wing upperside is fulvous 
with an intensive black pattern; the hind wing is substantially 
blackened in the basal half. The wing underside is mottled, a 
row of round black spots at the outer margin and their light 
rims are especially conspicuous. On the hind wing underside the 
outer half ground colour is reddish-brown; the discal band on 
the hind wing is more mottled than in other subspecies, it 
contain contrasted patches of whitish and brown colour.
   MATERIALS: The Holotype : a male - 19.06.1968, the settlement 
Khaptagai on the Lena River (Yu. Ammosov). Paratypes: 2 males - 
23.06 and 11.071987, the settlement Khaptagai, the road to Tammu 
(Yu. Ammosov); 2 females - 20.07.1962, Yakutsk, the botanical 
garden (Alexandrova); a male - 7.06.1979, the same locality; a 
ma;e - 8.06.1985, the same locality (V. Dubatolov), a female - 
23.06.1992, the same locality (P. Gorbunov).
     The subspecies is dedicated to the memory of Ershov, 
Nokolai Grigoryevich (1837-1896), one of the first real members 
of the Russian Entomological Society, the author of numerous 
works and the first catalogue of the Siberian [an error - must be 
Russian - Yu.K.] butterflies."


212. Clossiana matveevi P. Gorbunov et Korshunov, 1995.

     Original description:
  "MALE: 22.6-23.8 (in holotype 22.8 mm). The  wing  upperside  
is ochre-orange, more faded than in C. tritonia and C. erda and 
close to that in C. astarte. The black pattern in general 
corresponds to that in C. tritonia but is noriceably fine; a 
more narrw black margin without widening at the veins, as is 
characteristic for C. tritonia, is among significant 
differences. On the fore wing underside the pattern is the same 
as on the upperside, but the black spots are more narrow and two 
black parallel lines replace the margin. The hind wing underside 
ground colour varies from muddy-orange to muddy brick-red; the 
ground-ochre or greyish discoidal band is relatively narrow, 
2.2-2.5 mm wide at the vein M1 (in other taxa of the tritonia-
group it is not less wide than 2.5 mm), its both sides being 
bor dered with conspicuous black streaks, and locally it is much 
suffused by dark scales, which entirely hide it at the anal 
margin. The postdiscoidal area contains a row of seven well-
expressed whitish spots. Outside of it there is a row of six 
black spots and then the area of ochre-red colour 2.0-2.5 mm 
wide. The black bracked-shaped submarginal spots are distally 
margined with white spots streched out across the veins; outside 
of them there is a double marginal line. In the male genitalia 
the caudal processus of the valva ends with a single large 
tooth, not with a row of small teeth as in C. astarte and C. 
tritonia, the costal processus is large, foot-shaped (for the 
figure see Appendix). 
   MATERIALS: the holotype: a male, Altai, the Kurayskiy mountain 
range. the settlement Aktash, 2700 m above sea level, a dry 
scree, 8th July 1990 (E.A. [must be E.M. - O.K.] Matveev leg.). 
Paratypes: two males one female, the same locality 16th July 1992;  
4 males - the Kuraiskiy mountain range, the Yarly-Ayry river 
headwaers, 3000 m altitude, 7-12th July 1992 (Yu. Prokofyev). 
   The taxon is named by the surname of Evgenii Aleksandrovich 
[an error - must be EMvgenii Mikhailovich] Matveev, a Moscow 
naturalist, a butterfly lover, who first collected these 
butterflies. Although it is not excluded that a butterfly of this 
very species (identified by A.A. Meingard as C. amphilochus 
Menetries) was caught by V.V. Sapozhnikov on 6th June 1905 in the 
Tsagan-Kol River headwaters in Mongolia." 


GENUS BOLORIA Moore, 1900.
T.s.: Papilio pales [Denis et Schiffermuller], 1775.

  F.w.l.: 15-21 mm. The fore wing is pointed apically. The 
hind wing underside ground colour varies from ochre to brick-red; 
the central band consists of angular ochre or darker spots; its 
part within the cell is silver; the spots at the outer margin 
are also silver. Species have rather a similar appearance 
accompanied with a great individual variation. Identification is 
difficult without examining the male genitalia and taking into 
account the locality where an individual has been taken from. 
  A Holarctic genus with about 10 species. If to follow the 
species attribution according to the classical work by B. Warren 
(1944) the butterflies from the Asian Russia should be regarded 
within either of two complex "species", namely, Boloria napaea 
and B. pales, which we consider as species groups (superspecies).


superspecies napaea

  On the hind wing underside the discal band is as a rule ochre-
coloured, weakly suffused with red scales. The wing upperside 
ground colour in females is as a rule much darker than in males, 
in some cases to entire elemination of fulvous tint in B. 
alaskensis. In the male genitalia the length of the dentate part 
of the outer edge of the apical part of the harpe is not less 
than a half of the total harpe length (Table ...). 
  These butterflies keep to meadow and tundrous biotopes of the 
alpine zone of mountains and the northern tundral zone. 


223. Boloria (napaea) altaica (Grum-Grshimailo, 1893).

TYPE LOCALITY: West Altai: the Kurchumskiy mountain range: Kara-Baga.

RANGE: The mountains of South Siberia from Altai to Kentei, the 
mountains of East Siberia (the Cherskogo and 
Verkhoyanskiymountain ranges). 

HABITAT: in Altai: meadows in 
highlands and in the upper part of the forest belt, meadow and 
dwarf-birch montane tundras at elevations of 1600-2900 m above 
sea level. The imagines feed on the flowers of Sajania monstrosa, 
Polygonum bistorta, P. viviparum, P. nitens, Solidago davurica, 
Senecio turczaninovi, Scorzonera radiata, Allium lineare etc. 

FLIGHT PERIOD: prolonged, late June/late August.

PREIMAGINAL PHASES: Probable foodplant: Polygonum bistorta. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 18-23 mm. The wing upperside 
ground colour in females is darker than in males but always 
reddish. The hind wing underside usually is ochre-coloured, 
including a little contrasted central band. 

GEOGRAPHICAL VARIATION: Ssp. altaica ranges in Altai, the 
Sayans, the mountains of Tuva. From montane tundras of the 
Verkhoyanskiy mountain range (the Kokchin river) the subspecies 
vinokurovi Dubatolov, 1992, has been described. A subspecies 
from the elevation of Gornaya Shoria (the Kemerovo Region) is 
described below. 
     Boloria altaica pustagi Korshunov et Ivonin, 1995

     Original description:
   "By the materials from Gornaya Shoria [elevation] the 
subspecies Boloria altaica pustagi Korshunov et Ivonin, sbsp.n. 
is being described, differing from altaica first of all by a 
darker (brick-red) ground colour.
   HOLOTYPE: a male. F.w.l. 21 mm. The wing upperside is brick-
red with the black pattern enlarged, as compared with the 
subspecies altaica. The black spots along the outer margin are 
not fused into a streak. On the hind wing underside the discal 
band is bright, well contrasted to the ground brick-red 
background; the black round spots in the postdiscal area are 
contrasted, some of them contain white nucleoli. 
   In females the upperside is lighter, yellowish, the underside 
is bright, as in males.
   MATERIAL: The holotype: a male - 13.07.1992, Gornaya Shoria, 
the environs of the settlement Sheregesh, the mountain Pustag, 
1000-1300 m (V. Ivonin). Paratypes: 7 males, 3 females - 
13.07.1992, the same locality."


214. Boloria (napaea) frigidalis Warren, 1944.

TYPE LOCALITY: Altai (the Chuya River) and Mongolia (the Arsayn-Gol 
mountain range ["Arasan"]).

RANGE: West, Central and South-East Altai, the Sayans, the 
mountains of Tuva and the western half of Mongolia. A local 
species. 

HABITAT: alpine meadows, dwarf birch and detrituous tundras at 
altitudes within 2300-2900 m. The imagines are usually found 
among those of B. (napaea) altaica but are much less frequent. 

FLIGHT PERIOD: July/middle August.

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-18 mm. The hind wing 
underside ground colour is dark-reddish-brown, with a purple 
tint; the central band is ochre-coloured. 


215. Boloria (napaea) alaskensis Holland, 1900.

TYPE LOCALITY: Alaska.

RANGE: The polar regions of Eurasia and the western North 
America; the mountains of North Ural, Central and East Siberia, 
Kamchatka, the mountains of Bureya, Lower Priamurye. Details of 
distribution in the southern East Siberia and Far East are not 
studied.

HABITAT: lowland and montane tundras of various types, meadows 
in river valleys. In the Yamal Peninsula the imagines congregate 
on rich in herbs meadow patches at river valleys or ravines and 
actively visit the flowers of Pedicularis, Hieracium, Crepis 
nigrescens, Polygonum, Veronica. 

FLIGHT PERIOD: July/early August.

PREIMAGINAL PHASES: studied in Lapland (Henricksen, Kreutzer, 
1982). Foodplants: Polygonum bistorta, Viola biflora is known. 
Eggs: golden-orange, conical, with a rounded top and 10 
lengthwise ribs. Larva: brown-grey with a double yellow line 
along the back and black spots laterally of it at the fore 
margin of each segment. The spines are pinkish-coloured. In the 
extreme North and high altitudes the larva hivernates twice. 
Pupa: yellowish-brown with darker margins of the wing cases; 
yellow back and light spots on the sides, sometimes much 
darkened. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-22 mm. The wings are more 
short and angular in shape than in B.altaica. The wing upperside 
in females is sometimes without traces of red colour. The hind 
wing underside ground colour is brick-red; the discoidal band is 
ochre-coloured and contrasted to the background. 

GEOGRAPHICAL VARIATION: insufficiently studied. Ssp. sedykhi 
Crosson du Cormier, 1977; has been described from Polare Ural; 
ssp. nikolaewsky Heydemann, 1920 - from the lower Amur (and the 
middle Lena?). A detailed investigation is necessary of the 
peculiar butterflies from NE Siberia. 


superspecies pales

  On hind wing underside the discal band can be strongly 
suffused with reddish scales. Sexual dimorphism is usually weak; 
the wings of males and females are of the same ochre-orange or 
ochre-red ground colour. In the male genitalia the length of the 
dentate part of the outer edge of the apical part of the harpe 
is less than a half of the total harpe length. 
  The butterflies tend to raised bogs, marshy valleys, open 
forests, damp forest meadows. Relationships between the two 
following taxa are studied insufficiently. We formally draw the 
border between them by the Yenisei River. An opinion exists that 
B.banghaasi is only a subspecies of B. aquilonaris.


216. Boloria (pales) aquilonaris (Stichel, 1908).

TYPE LOCALITY: Lapland.

RANGE: Europe (except for the South), Ural, West Siberia, ? North
Altai. This species is very local in its southern range. 

HABITAT: raised and transitory bogs (including "ryams" of the 
forest-steppe zone), open peat-moss pine woods. 

FLIGHT PERIOD: late June/late July.

PREIMAGINAL PHASES: studied in Scandinavia (Henricksen,
Kreutzer, 1982). Foodplant: Oxycoccus quadripetalus, O.
palustris, in Sweden were also reported Vaccinium, Viola, for
Central Europe Polygonum. Eggs: conical with lengthwise ribs and
an apical dimple, greenish; laid singly on foodplant stems and
leaves. In Scandinavia the larvae hibernate in the moss usually
in the third instar, in Polar Ural, according to observations by
A.G. Tatarinov, in the first instar. Young larva: brown with
balck head, set with black warts bearing black hairs. Mature
larva: dark-brown with a double yellowish streak along the
back, indistinct lateral stripes of the same colour and small
whitish-yellow spots on the sides; small yellow branched spines
emerge from the orange warts. Pupa: obtuse, brown sometimes with
sparse pale spots on the back; abdomen with dark brown
transversal spots; the wing cases with dark streaks and light
margin.

PECULIAR TRAITS OF IMAGO: F.w.l.: 15-20 mm. The hind wing 
underside ground colour is brick-red or ochre-brown; the ochre-
coloured central band is strongly suffused with red scales and 
so is not contrasted to the ground colour. 


217. Boloria (pales) banghaasi Seitz, 1909.

TYPE LOCALITY: Mongolia: Kentei.

RANGE: The mountains of South and East Siberia and the northern Far 
East, Mongolia. A local species. 

HABITAT: meadow patches at the tree-line, rarely in the forest 
belt on cowberry or bog bilberry heaths and bogged up dwarf 
birch or bush alder thickets; in SW Kamchatca - the coastal 
peat-moss/cranberry bogs. The imagines were observed to feed on 
Comarum palustre, Pentaphilloides fruticosa, Senecio tundricola. 

FLIGHT PERIOD: July/August. 

PECULIAR TRAITS OF IMAGO: F.w.l.: 16-20 mm. The hind wing underside ground 
colour is brick-red or ochre-brown; the ochre-coloured central 
band is scarcely suffused with red scales, as different from B. 
acquilonaris. 

GEOGRAPHICAL VARIATION: On the Sakhallin the subspecies neopales 
Nakahara, 1922 is known. To this taxon we attribute, yet 
conventionally, the butterflies collected in Central Altai on 
the Katunskiy (the headwaters of the Katun', Belaya Berel' and 
Yazovaya Rivers, on damp valley meadows with Pentaphylloides 
fruticosa bushes at 1500-1800 m above sea level) and Chikhacheva 
(the Yustyd river) mountain ranges They seem to belong to a 
distinct subspecies or even species. 

ETYMOLOGY: Andreas Bang-Haas (1846-1925) and his son Otto 
(1882-1948) - well known German lepidopterologists.

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