NOTE: the text corresponds to (Korshunov & Gorbunov, 1995) with additions and corrections (Korshunov, 1996) incorporated
Special thank to Dr. Cris Guppy (Quesnel, Canada) who kindly corrected many spelling and other lingustic errors.
FAMILIA PIERIDAE Duponchel,1835]
The butterflies of intermediate size. The wing ground colour is usually white, yellow or orange, with black or greenish markings. The head rounded. All the legs are equally developed and used for walking.
Eggs: elongate, ribbed longitudinally, 0.7-1.5 mm in length. The larvae are mostly green, often with markings and stripes, covered by sparse short and fine hairs, feed predominantly on Brassicaceae and Fabaceae. The pupae have the head pointed; they are attached head upwards or horizontally by tail hooks and a cincture, or a belt of silk threads around the middle. This family embraces all main pests among butterflies.
The world fauna includes about 1000 species, in Asian Russia
40 species are known.
GENUS LEPTIDEA Billberg, 1820.
Type species: Papilio sinapis Linnaeus, 1758.
F.w.l.: 18-27 mm. The wings are elongate, white; the body is long and very thin. As a rule two broods develop, the imagines of the summer brood being larger and with less expressed suffusion. The flight of imagines is weak and slow.
Eggs: spindle-shaped with twelve ribs, at first white, later of a cream colour, laid singly on the young stems and leaves of foodplants. The larvae of our species are very similar, they are thin, green, with a darker back line and a yellow line on either side at the spiracles. The pupae are sharply pointed on either end and bear a conspicuous projection on the thorax, which make them triangle-shaped; yellow or green, with more or less expressed two-coloured, yellowish and reddish-brown lateral streaks. They are attached with a cincture head upwards or downwards.
A Palearctic genus with five species.
71. Leptidea sinapis (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: Morocco, Europe, Anterior and Central Asia, Kazakhstan, the southern halves of Ural and Siberia to Priangarye, in West and Middle Siberia northwards to the Middle Priobye.
HABITAT: forest edges, glades, openings, brook valleys, meadows, in the mountains (Altai) rises up to 1500 m altitude; on the most part of the range flies together with L. morsei in the same habitats. The imagines often visit the flowers of Lathyrus, Trifolium, Ranunculus, Cerastium pauciflorum (Korshunov, 1969)
FLIGHT PERIOD: in two broods, almost permanently from late April to late August.
PREIMAGINAL PHASES: studied in Europe (Eckstain, 1913; Lorkovic, 1947; Ebert, 1991). Foodplants: Lathyrus, Vicia, Trifolium, Medicago, Astragalus, Lotus, Coronilla, and other herbaceous Fabaceae. Eggs: cylindrical, ribbed, yellowish-white or light- green. Larva: green with a dark back line and yellow spiracular lines. Pupa ochre-yellow or green, with a reddish streak on either side.
PECULIAR TRAITS OF IMAGO F.w.l.: 18-23 mm. The fore wing apex is rounded; the hind wing underside is yellowish, the greyish pattern in the spring brood imagines (f. lathyri Hübner, 1823) is distinct but the lengthwise strokes between the veins are absent, in the imagines of the summer brood (f. deniensis Boisduval, 1840) it consists of two vague stripes or is absent entirely. Similar species: L. morsei.
72. Leptidea morsei (Fenton in Butler, 1881).
TYPE LOCALITY: Japan: Hokkaido.
RANGE: The temperate Eurasia westwards to the Balkans and the lowland Austria, northwards in Siberia to the northern border of the middle taiga zone; the Sakhalin, Japan.
HABITAT: forest edges, glades, open woodland, brook banks, meadows; in the mountains rises up to 1800 m above sea level (N. Altai). The imagines were observed to feed on the flowers of Pulmonaria, Geranium, Iris, various Fabaceae.
FLIGHT PERIOD: early May/June and July/August, in two broods.
PREIMAGINAL PHASES: Foodplants: Vicia cracca, V. amoena, Lathyrus, other herbaceous Fabaceae. Eggs: according to Lorcovic (1947) differ from those of L. sinapis by somewhat greater size. Larva: as compared with that of L. sinapis, has a lighter colouration, a wider back line and more conspicuous yellowish spiracular lines. Pupa: green with yellowish or reddish line on either side, it differs from that of L. sinapis by a longer head prominence and a straight profile of the dorsal side.
PECULIAR TRAITS OF IMAGO F.w.l.: 19-25 mm.; as different from L. sinapis, the fore wing is slightly pointed apically. The hind wing underside is white with a distinct greyish pattern. In the spring brood butterflies (f. croatica) on hind wing underside at outer margin there are distinct dark intervein lengthwise strokes; in the summer brood the pattern consists of two distinct greyish stripes going across the veins, one of which, as different from L. amurensis, crosses the cell between the veins M1 and M2. Similar species: L. amurensis, L. sinapis.
GEOGRAPHICAL VARIATION: Ural and, most probably, eastwards throughout the continent there ranges the subspecies major Grund, 1905 described from Yugoslavia. Besides, the taxon morseides Verity, 1911 was described from the Sayans and micromorsei Verity, 1947 from Priamurye.
73. Leptidea amurensis (Ménétriés, 1859).
TYPE LOCALITY: Priamurye.
South Siberia east of Verkhnee Priobyethe upper Ob' basin], Central Yakutia, NE Kazakhstan, Mongolia, NE China, Priamurye, Primorye, Korea, Japan.
HABITAT: meadows and bushes in river valleys, dry meadows, open woodland, birch forest-steppe.
FLIGHT PERIOD: in two broods: May/Middle June and July/late August.
PREIMAGINAL PHASES: studied in Japan (Fukuda et al. 1982): Foodplants: Vicia is known. The preimaginal stages resemble those of L. morsei. Pupa: that of summer generation light green, hibernating pupa of the spring brood - pale brown.
PECULIAR TRAITS OF IMAGO F.w.l.: 21-27 mm. The fore wing apex is more acutely pointed than in L. morsei and is somewhat stretched out. In the spring brood the hind wing underside is pale ochre- coloured with a greyish pattern and the lengthwise white beam going throughout the wing, not being interrupted with a greyish spot, in the summer brood the underside is entirely white or has two obscure greyish transversal bands, which, as different from a similar species L. morsei, do not enter the space between the veins M1 and M2.
GEOGRAPHICAL VARIATION: The taxa emisinapis Verity, 1911 (Pribaikalye) and vibiloides Verity, 1911 (Priamurye) are close to the nominotypical subspecies. Ssp. vernalis Graeser, 1892, was described from Priamurye. The subspecies jacutia P. Gorbunov et Korshunov, 1995 was described from Yakutia.
"Butterflies from Central Yakutia differ by a noticeable yellowish tint on the wing upperside and more intensive, than in other subspecies, ochre suffusion on the underside of the hind wing and on the most part, at the apex and the fore margin, of the fore wing. By these characters we state a subspecies Leptidea amurensis jacutia P. Gorbunov et Korshunov, sbsp.n.
MATERIALS: Holotype: a male - 2.06.1985, Yakutsk, the botanical garden (V. Dubatolov). Paratypes: a male - 21.06.1962, the same locality; 2 males - 28.05.1985, the same locality; a male - 2.06.1985 - the same locality."
SUBFAMILIA PIERINAE Duponchel,1835]
TRIBUS PIERINI Duponchel,1835]
GENUS APORIA Hübner,1819].
Type species: Papilio crataegi Linnaeus, 1758.
F.w.l.: 28-33 mm. The wings are white or yellowish, in our species without black spots but with conspicuous black veins.
A Palearctic genus with 12 species.
74. Aporia crataegi (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: The non-tropical Eurasia, except for deserts and arctic tundras.
HABITAT: forest meadows, other woody and open landscapes, bogs, gardens. In Siberia and the Far east this species repeatedly (every 4-5 years) exhibits mass proliferation, and almost every year this is the most abundant butterfly in the forest-steppe zone, in E Europe the last mass proliferation was observed in 30ths. The imagines migrate actively, some specimens penetrating into montane and lowland tundras. Copulation as a rule occurs immediately after the imagines have hatched from the pupae, with the wings yet not spread. According to our observations, active contacts between the mature males and females, which superficially resemble mating, are not in fact copulations, but rather an abortive form of mating behaviour. These butterflies as a rule congregate in large numbers on wet ground at pools, brooks, lakes etc., males predominating in these congregations. The imagines feed on various flowers the pollen of which often colours brightly their hind wings.
FLIGHT PERIOD: early June/middle July, sometimes to early August.
PREIMAGINAL PHASES: The main foodplants: mostly Padus avium, and also Sorbus aucuparia, S. sibirica, Spiraea media - in forest and forest-steppe zone of Ural and West Siberia (our observations); Padus avium, Prunus spinosa, Amigdalus nana, Rosa, Malus - in the steppen zone of Preduralye (Kuznetsov, Martynova, 1954); Vaccinium uliginosum in Yakutia, Zabaikalye, and Priamurye (Kurenzov, 1970; Sviridov, 1981a); Armeniaca sibirica - in SE Transbaikalia (V. Dubatolov); Crataegus maximowiczii, C. pinnatifida, Malus manshuricus in Primorye (Kuranzov, 1970). The various species of the genera Spiraea, Crataegus, Cotoneaster, Malus, Pyrus, Cerasus, Amygdalus, Filipendula, and also Quercus and Juglans are reported as the foodplants from other parts of the range. Eggs: ribbed, yellow- orange, in batches by 20-200 most frequently on the leaf underside of foodplants. The young larvae are gregarious, in August they construct a nest by spinning several leaves or twigs with silk, in which they overwinter. After hibernation they at first live on the nests and feed on buds, later they keep together in groups by several individuals and eat leaves and inflorescences, but after the last molt live solitarily. Mature larva: brown or dark-ash-grey with blueish-grey ventral side, on either side with two wide reddish-brown lengthwise stripes split into fragments by segment divisions; covered by long light hairs. Pupa: whit, yellowish or greenish, with numerous black spots, among which a pair of round spots on the back of the thorax is largest, the dorsal rib of the thorax is also black; the head bears a bright-orange-yellow process with a black base and a pair of black-rimmed spots of the same colour, a orange spot presents also on either side of abdominal segment margins. The pupae with a white ground colour have larger spots than those of yellowish or greenish colour. The pupae are found on trunks and branches, often by numerous groups.
PECULIAR TRAITS OF IMAGO F.w.l.: 23-36 mm. The wings are milky- white with black veins, in females often with transparent areas; yellow scales are absent from the hind wings.
GEOGRAPHICAL VARIATION: weakly expressed in N Asia. The taxa meinhardi Krulikowsky, 1903 (W Siberia), sibirica Verity, 1911 (Tomsk), asiatica Meinhard, 1915 (again Tomsk), sajana Verity, 1911 (E Sayan), banghaasi Bryk, 1921 (Priamurye), ussurica Kardakov, 1928 (Primorye), and sachalinensis Matsumura, 1925 (Sakhalin) most probably should be attributed to the nominotypical subspecies. A Japanese subspecies adherbal Fruhstorfer, 1910 was reported for the South Kuriles.
75. Aporia hippia (Bremer, 1861).
TYPE LOCALITY: Priamurye: the basins of Zeya, Bureya, and Ussuri Rivers.
RANGE: Zabaikalye, Priamurye, Primorye, E Mongolia, NE China, Korea, Japan.
HABITAT: deciduous and mixed forests, gardens, mostly in river valleys.
FLIGHT PERIOD: late June/middle August.
PREIMAGINAL PHASES: studied in Japan (Shirozu, Hara, 1969; Fukuda et al., 1982): Foodplants: Berberis sinensis, B. amurensis is known. Eggs: elongate, yellow with lengthwise lines, laid in batches by 50-200, usually by several layers, on leaf underside of the foodplant. The larvae live gregariously on loose silken nests, hibernate at the third or fourth instar. Mature larva: dark-brown with lengthwise darker lines and spots on the back and sides, set with long light hairs; the back of the first, second, and the last segment are reddish; the head is black. Pupa: cream-white or yellow with numerous black markings on the thorax and abdomen, with black lines margining the antennal cases and joints of thoracic segments.
PECULIAR TRAITS OF IMAGO F.w.l.: 27-37 mm. The wing underside ground colour is yellowish, with a bright yellow spot at the base; the veins are accompanied with broad dark suffusion.
GEOGRAPHICAL VARIATION: is weakly expressed. The taxon occidentalis O. Bang-Haas, 1927, was described for the Malakhanskii mountain range in W Transbaikalia, which is very close to ssp. hippia, known from Primorye and Priamurye.
GENUS PIERIS Schrank, 1801.
Type species: Papilio brassicae Linnaeus, 1758.
F.w.l.: 17-34 mm. The wing upperside is white, less frequently yellowish; the discoidal spot on the fore wing is absent; the sexual dimorphism is expressed through appearing or enlarging the black spots in the postdiscoidal area in females. The larvae are generally green, they live on Brassicaceae. The pupae are light-coloured with black markings; with the head pointed and with angular projections on the dorsal side: one on the thorax and two on the first abdominal segment; those of the autumn/spring brood hibernate.
The genus includes about 30 species, mostly inhabiting the arid mountains of the subtropical Asia.
77. Pieris brassicae (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: Europe, N. Africa, Anterior and Central Asia. Kazakhstan, China, North, Central and South Ural, the south of Siberia westwards to Pribaikalye. The species is actively expanding. In W. Siberia P. brassicae became abundant only in the latest couple of decades (Y.P. Korshunov). Recently it has established in S. Primorye and since the autumn 1993 is abundant there. Recently was found out also at Khabarovsk. In 70s years the species invaded Chile where became a pest.
HABITAT: meadows, fields, waste lands, settlements; in the mountains goes as high as 1700 m above sea level (N. Altai); a serious pest.
FLIGHT PERIOD: May/September in 2-3 broods.
PREIMAGINAL PHASES: Foodplants: Brassicaceae, predominantly cultivated: Brassica, Raphanus, Sinapis, Lepidium, Rorippa, etc., and also Reseda lutea (Resedaceae). Eggs: conical with 14 ribs, yellow, in clusters by 20-100 on the foodplant. The young larvae are gregarious, eat the mesophyl of the young leaves from their underside, not perforating the upperside epidermis. Mature larva: blueish-green with yellowish ventral side and yellow back stripe, sprinkled with black spots of different sizes and set with short light hairs; the ventral side is yellowish; lives solitarily. Pupa: yellowish-green or white with small black and orange dots and spots, attached to tree trunks, fences, and other rather high places; hibernates. The pupae from Primorye differ by those from Europe and Siberia by presence of thorn- like projections on the sides (V. Dubatolov, pers. comm.)
PECUILAR TRAITS OF IMAGO F.w.l.: 25-34 mm. Fore wing has a black area at the apex, extended along the fore and outer margins, females have also two round black spot in the postdiscoidal area. The hind wing underside is yellowish with a light even suffusion of black scales.
SIMILAR SPECIES: Pieris rapae
78. Pieris rapae (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: Eurasia, N. Africa, N. America, Australia. In Ural and West Siberia reaches the polar regions but is not found in the north of East Siberia and the Far East.
HABITAT: steppes, meadows, fields, wasting lands, settlements, roads and clearings in woods.
FLIGHT PERIOD: in the southern areas from May to September in 2- 3 broods, in the North in a single brood in early summer.
PREIMAGINAL PHASES: Foodplants: Brassicaceae: Alliaria, Arabis, Barbarea, Brassica, Cardamine, Descurainia, Erysimum, Hesperis, Raphanus, Turritis; and also Reseda lutea (Resedaceae). Eggs: pear-shaped, with 12 ribs, yellowish-orange, usually laid singly on leaf underside. Larva: velvety, with fine pubescence, mate- green with a yellowish back line and two stripes or rows of spots of the same colour along either side; the head is brownish-green, the ventral side is yellowish-green. It develops rapidly, pupates after the fourth molt, usually on leaf under surface. Pupa: yellowish- or greyish-green or pale brown, depending on the substrate colour, with yellowish streaks and black spots on the wing cases, back, and sides of abdominal segments, it has prominences on the head and thorax. Its stage lasts for 7-10 days, or the pupa hibernates.
PECULIAR TRAITS OF IMAGO F.w.l.: 20-30 mm. Dark apical spot on fore wing is slightly extended along the costal margin and does not reach the middle of the outer edge; the hind wing underside is evenly pale-yellow or whitish.
GEOGRAPHICAL VARIATION: is little studied. The butterflies from Ural and the majority of Siberia are in general close to the subspecies rapae (? = viluensis Rober, 1907, described from the Vilyui River). The specimens from Transbaikalia and more eastern regions are characterized by a widened dark pattern on the fore wing upperside in females, a suffusion by dark scales in the internal half of this wing being well expressed, they were reported under the names: kenteana [Staudinger] in Rober, 1907 (Kentei), orientalis Oberthur, 1880 (Primorye), metra Stephens, 1827 (the Sakhalin), crucivora Boisduval, 1836 (the Sakhalin and Kuriles) but all they are insufficiently studied, the differences being very weak. Numerous individual variations are known for this species.
79. Pieris canidia (Sparrman, 1768).
TYPE LOCALITY: China.
RANGE: The southern half of Asia, including Central Asia, NE. China, Korea, Japan; recently was discovered by E.A. Belyaev in South Primorye at the village Ryazanovka.
FLIGHT PERIOD: in N. Korea in the second half of July.
PREIMAGINAL PHASES: Foodplants: Brassicaceae. Larva resembles that of P. rapae. The pupa has a sharp projection on either side of the first abdominal segment.
PECULIAR TRAITS OF IMAGO F.w.l.: 24-28 mm. The apical spot strongly extends along the fore and, especially, outer margin of the fore wing upperside; on the fore wing upperside there is one round black spot in male and two ones in female; the hind wing underside ground colour is white.
79. superspecies napi
F.w.l. of the first (spring) brood imagines usually 18-24 mm; the apical dark spot on the fore wing is narrow, stretched along the costal margin, frequently replaced by a dark suffusion of the vein endings; the hind wing underside with a heavy dark suffusion along the veins. F.w.l. of the second (summer) brood imagines is up to 31 mm; the hind wing underside has traces of a dark suffusion along the veins or is clear white.
The systematic situation within the group is very complicated due to a high level of individual variation. In the work by Eitschberger (1983) a new attempt has been made to split Pieris napi L. s.l. into a number of separate species, seven of which being reported for the territory considered. However, special study, especially with regard of the preimaginal stages, are missing in those regions, so the conclusions still remain to be the matter of future.
80. Pieris (napi) napi (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: Europe, Anterior and Central Asia, Kazakhstan, Ural, West Siberia, the mountains of South Siberia east to ?Pribaikalye.
HABITAT: meadows of various kinds, fields, settlements.
FLIGHT PERIOD: usually in two broods in May/June and July/August.
PREIMAGINAL PHASES: Foodplants: various Brassicaceae (Alliaria, Arabis, Armoracia, Barbarea, Brassica, Cardamine, Descurainia, Draba, Erysimum, Lepidium, Raphanus, Sisymbrium, Thlaspi etc.) and also Reseda lutea (Resedaceae). Eggs: bottle-shaped, with 14 ribs, yellowish, laid singly on the leaves or stems of the foodplant. Larva: velvet-green with yellow interrupted streak on either side, sprinkled with black dots and pale warts; its development lasts for 2-3 weeks. Pupa: green with black dots, more slender and angular than in P. rapae. F.w.l.: 19-26 mm.
PECULIAR TRAITS OF IMAGO F.w.l.: 17-26 mm. As compared with P. (napi) bryoniae, the basal darkening on the wings is weaker, the hind wing underside ground colour is of a noticeable yellowish tint, in females the wing upperside has no yellowish tint and a weaker vein suffusion.
GEOGRAPHICAL VARIATION: The nominotypical subspecies napi inhabits Ural and West Siberia. In the mountains of South Siberia from the West Tannu-Ola mountain range to the Selenga River basin in Buryatia a subspecies euorientis Verity, 1908 ranges. It has been described from the East Sayans and is considered by U. Eitschberger (1983) as an independent species.
81. Pieris (napi) bryoniae (Hübner, 1791).
TYPE LOCALITY: Switzerland: "Gebirge bei Gent".
RANGE: Distributed mostly in the mountain regions of the northern half of Eurasia, Reported for the mountains of South, Central, and East Siberia (Eitschberger, 1983). The range is to be studied.
FLIGHT PERIOD: June/July in a single brood.
PREIMAGINAL PHASES: studied in Lapland (Eitschberger, Strohle, 1990) Eggs are green spindle-shaped, with 17 ribs. A young larva is light-green, covered with white hairs. Mature larva is dark- green, with a whitish-green ventral side, covered with tiny black dots. The spiracles are brown with wide yellow rims. Pupa: green, with yellow keel-like projections, or ochre-coloured with black spots of small or varying size.
PECILUAR TRAITS OF IMAGO F.w.l.: 18-25 mm. As compared with P. (napi) napi, the dark basal suffusion is more expressed, in females the wing upperside usually has a strong yellowish tint, the dark suffusion along the veins being heavier.
GEOGRAPHICAL VARIATION: The subspecies bryonides Scheljuzhko, 1910 ranges in the mountains of South Siberia; the subspecies vitimensis Verity, 1911 is described from West and Central Yakutia; schintimeisteri Eitschberger, 1983 - from the Aldan upland and Verkhoyanskiy mountain range; sheljuzhkoi Eitschberger, 1983 - from the Magadan Region (the Kolyma basin and city Magadan); peculiar butterflies from Kamchatka was described as ssp. kamtschadalis Rober, 1907. Probably it is P. bryoniae to which a little-known taxon sichotensis Kurenzov, 1941 from the Sikhote-Alin' Mts. should be attributed.
SYSTEMATIC NOTES: By external and biological features this species much resembles P.(napi) napi. In Central Europe differences were found out between these taxa, on the molecular level (isozyme variation) (Gieger, 1978). However, during population studies in the Alps other authors (Benninger et al., 1993) revealed fertility of the hybrids, that indicated rather a subspecies rank. In Siberia such studies are missing.
82. Pieris (napi) dulcinea (Butler, 1882).
TYPE LOCALITY: N. Korea.
RANGE: Priamurye, Primorye, the Sakhalin, the South Kuriles, NE. China, Korea. There are also reports on its occurrence further westwards to the Kodar mountain range in NE. Zabaikalye (a male in the ISEA collection collected by Savin, expedition headed by Starikovskii, on 7.08.1986) and northwards to the Suntar-Khayata mountain range in S. Yakutia.
HABITAT: meadow patches, wood edges, bush thickets, fields, settlements.
FLIGHT PERIOD: in two broods in middle April/middle May and middle June/late August.
PREIMAGINAL PHASES: Foodplants: Arabis hirsuta was reported for Primorye (Kurenzov, 1970).
PECULIAR TRAITS OF IMAGO As different from P. melete, on the fore wing upperside the cell has no dark suffusion. Spring brood imago: F.w.l. 19-25 mm; the dark postdiscoidal spots on the fore wing underside are of close size (in females) or absent (in males); the hind wing underside has no yellowish tint and a yellowish spot at the wing base usually is not seen. The summer brood imago: F.w.l.: 24-30 mm; on the fore wing upperside the apical dark spot is triangular or only vein endings are dark suffused, in the postdiscoidal area there is usually one spot in males and two ones in females; the hind wing underside is white. By many features this taxon is close to P. (napi) napi.
GEOGRAPHICAL VARIATION: The subspecies dulcinea is distributed in the continental part of the range. The subspecies saghalensis Nakahara, 1926 was described from the Sakhalin, which differ by on average smaller size of the butterflies of the summer brood and reduction of the dark patter on the fore wing upperside. Besides, a taxon tomariana Matsumura, 1928, described from the Kunashir Island and considered as a distinct species by Eitschberger (1983) is close to the spring brood of P. dulcinea. A Japanese taxon nesis Fruhstorfer, 1909 was also reported for the Sakhalin and South Kuriles, which is also given a species rank by Eitschberger (1983)..
83. Pieris (napi) melete Ménétriés, 1858.
TYPE LOCALITY: Japan.
Range: Priamurye (known from the city Blagoveshchensk to the Gorin River), Primorye, NE. China, Korea, Japan.
FLIGHT PERIOD: in two broods from middle May to early September.
PREIMAGINAL PHASES: Foodplants: Brassicaceae, Arabis hirsuta was reported for Primorye (Kurenzov, 1970), a number of other species - for Japan.
PECULIAR TRAITS OF IMAGO As different from similar species of the napi group, the lower vein of the cell on the hind wing underside is heavily suffused with dark scales. Spring brood imago: F.w.l. 21-27 mm; the hind wing underside is light-yellow with the veins suffused, a distinct yellowish spot usually presenting at the base; on the fore wing underside the upper dark postdiscoidal spot is much smaller than the lower one, the traces of which are usually seen even in males. The summer brood imago: F.w.l.: 26-31 mm; on the fore wing upperside the apical dark spot is large and usually quadrangular, in the postdiscoidal area two spots are present in males, in females they are very large; the wing underside is white.
GEOGRAPHICAL VARIATION: The continental part of the range is inhabited by the subspecies orientis Oberthur, 1881, described from the Askold Island and also considered by U. Eitschberger as a distinct species.
GENUS PONTIA Fabricius, 1807.
Type species: Papilio daplidice Linnaeus, 1758.
F.w.l.: 17-27 mm; the wing upperside is white; the fore wing has a black discoidal spot and separate spots at the apex; in females the hind wing has dark spots; the hind wing underside is yellowish-green or dull green with white spots; in the cell there is one white spot isolated from the others. Hibernation occurs at the pupal stage.
The genus includes six species ranging in Eurasia and Africa.
84. Pontia edusa (Fabricius, 1777) (= daplidice auct. non Linnaeus, 1758).
TYPE LOCALITY: Central Europe.
RANGE: The non-tropical Eurasia, in Siberia penetrates as north as the middle taiga zone, locally to the northern taiga zone..
HABITAT: dry meadows, steppes, waste lots, long fallows, fields, settlements.
FLIGHT PERIOD: early May/September in 2-3 broods.
PREIMAGINAL PHASES: Foodplants: Brassicaceae: Alyssum, Arabis, Berteroa, Erysimum, Sisymbrium, Thlaspi, Turritis; Resedaceae: Reseda lutea, also reported are some Fabaceae: Vicia, Lathyrus, Pisum, Trifolium. Eggs: cylindrical with rounded apex and 14 ribs, at first yellow, later become orange-yellow; laid singly on the leaf margin of the foodplant shoots. According to observations at Novosibirsk (Y. Korshunov) and in Middle Ural (P. Gorbunov), the mature larva is blueish-green with two pale- yellow stripes on either side, the head and body bears black warts and fine pubescence, black on the back and whitish on the ventral side; the head is greyish or light-green with black dots and has two yellow spots above the eyes. The larvae live by families or solitarily. Pupa: the abdomen is coloured alike the larva: green with a brownish or greyish tint, tiny black dots, and yellowish-white streaks; the wing cases are evenly green with lengthwise rows of very fine black dots; the head apex and dorsal crests are brownish. The hibernating pupae are smaller than the summer ones.
PECULIAR TRAITS OF IMAGO F.w.l.: 15-20 mm in a spring form; 22- 30 mm in a summer form. The hind wing underside is yellowish- green with separate white spots of different sizes, that in the cell being roundish.
SYSTEMATIC NOTES: For a long time this species was known as Pontia daplidice L. However, the isozyme analysis has revealed that P. daplidice ranges in S Europe, N Africa and the Near East, while the temperate zone of Eurasia is inhabited by P. edusa F., which differs from the former species not only on the molecular level but also by the genitalia structure (Geiger et al., 1988).
GEOGRAPHICAL VARIATION: the nominotypical subspecies ranges in Ural and Siberia, ssp. orientalis Kardakov, 1928 was reported from Zabaikalye, Priamurye, Primorye, and the Sakhalin.
[84.1]. Pontia daplidice (Linnaeus, 1758)
TYPE LOCALITY: Africa and S Europe.
RANGE: N Africa, S Europe, S Asia, in Russia is reported for S Primorye: the Kedrovaya Pad' Nature Reserve (Keskula, 1990) and Golubinyi Utes in the Gamov Peninsula (V. Dubatolov).
HABITAT: Various meadow-steppe associations.
FLIGHT PERIOD: February/March and later in two and more broods in the main range, the findings in the south of Primorye happened in the middle of the summer.
PECULIAR TRAITS OF IMAGO: F.w.l.: 18-24 mm; the spring brood is smaller and its wing underside green ground colour is darker. The main difference concerns main genitalia: in daplidice the valva is rounded, in edusa is angular.
85. Pontia chloridice (Hübner, 1808).
TYPE LOCALITY: the Balkans.
RANGE: The steppe and forest-steppe zones of Eurasia from Greece to Bol'shoy Khingan and Korea, the mountains of Central Asia, reported for the west of USA.
HABITAT: steppes of various kind, steppen and forest-steppen meadows in open woods, birch groves, montane slopes.
FLIGHT PERIOD: in two broods in middle May/late June and middle July/September.
PREIMAGINAL PHASES: Foodplants: various Brassicaceae, in Novosibirsk Region observed on Descurainia sophia (Korshunov, 1981), in Preduralye - on Sisymbrium junceum (Bartel, 1914). According to observations from N. America, the eggs are yellow, later become orange; laid singly on various parts of the foodplant. Larva: blueish-green or greenish-white, with small black warts, each bearing a black chaeta with a white tip, and rather long greenish hairs; the segment joints are orange or yellow; a light-green or yellowish stripe goes along the back and a wide stripe of the same colour in either side; the head is light-green with black dots. As different from P. edusa, the back stripe is free from the black warts. In a larva ready to pupation the lateral stripes become inconspicuous. Pupa: brownish-green with a light streak along the back and cream wing cases; the first and second abdominal segments are greyish- white.
PECULIAR TRAITS OF IMAGO F.w.l.: 17-22 mm. On the hind wing underside green colour extending along the veins forms two full waving bands; the white spot in the cell is elongate.
GENUS PARAPIERIS de Niceville, 1897 (= Synchloe auct.)
Type species: Papilio callidice Hübner (1799-1800)
86. Pontia callidice (Hübner, 1800).
TYPE LOCALITY: the Swiss Alps.
RANGE: Polar and mountain regions of the non-tropical Eurasia, the north and west of N America. A local species. In 20-30th years of this century this montane species was first found in the West Siberian Lowland in the upper Ob' basin (at Kamen'-na- Obi and Barnaul). In 1991-1993 it appeared to be quite common on the lowland in the forest-steppe and southern steppe zones from South and Middle Ural to the Ob' River and was also found in lowland steppes of S Zabaikalye.
HABITAT: montane and lowland tundras, shingle banks, floodland meadows of rivers and brooks, steppefied south-exposed slopes; in the forest steppe zone found on pastures, wasting and long fallow lands at settlements, i.e. the habitats of Pontia edusa.
FLIGHT PERIOD: middle June/August; in lowlands two broods develop, flying in middle May/late June and middle July/middle August.
PREIMAGINAL PHASES: Foodplants: Brassicaceae; in Middle Ural found on Erysimum cheirantoides (P. Gorbunov), in Pribaikalye on Erysimum sp. and Brassica napus (V. Ivonin), in SE Transbaikalia the butterflies keep to the riparian thickets of Dimorphostemon pectinatus (O. Kosterin, O. Berezina); from other parts of the range other Brassicaceae were reported, namely Arabis, Barbarea, Brassica, Descurainia, Draba, Erysimum, Lepidium, Sisymbrium, Thlaspi; and also Reseda lutea (Resedaceae) and Orostachys (Crassulaceae). Larva (according to observations of V. Ivonin in Pribaikalye): 35-40 mm long; greenish or blueish-green with lengthwise black and orange streaks; the body is covered by black warts and short black hairs; the head is grey with numerous black dots and a symmetrical V-like line. Pupa: 20-21 mm; muddy- grey with sparse black dots.
PECULIAR TRAITS OF IMAGO: F.w.l.: 20-26 mm. On the hind wing underside the greyish- or brownish-green ground colour is cleft between the veins by narrow triangular white spots.
GEOGRAPHICAL VARIATION: North Eurasia is occupied by the nominotypical subspecies. The butterflies from the southern and steppen regions usually have a lighter ground colour of the hind wing underside and somewhat narrower dark pattern.
SUBFAMILY ANTHOCHARINAE Tutt, 1896.
F.w.l.: 16-26 mm. The wing upperside is white with a dark pattern and, sometimes, with orange areas on the fore wing; the hind wing underside is greenish with light spots of different sizes, one of which going from the cell to the fore margin. In our conditions only a single brood develops. The larvae feed on Brassicaceae; hibernation occurs at the pupal stage.
GENUS ZEGRIS Boisduval, 1836.
Type species: Papilio eupheme Esper,1805].
A West-Palearctic genus with two species.
87. Zegris eupheme (Esper,1805]).
TYPE LOCALITY: Crimea: surroundings of the city Sevastopol.
RANGE: NW Africa, Spain, SE Europe, S Ural (the surroundings of the cities Orsk and Orenburg), N and E Kazakhstan (in S Altai south of the Narymskii and Kurchumskii Mt. Ranges). A local species.
HABITAT: steppen areas, mostly with chalk outcrops.
FLIGHT PERIOD: late April/early June.
PREIMAGINAL PHASES: studied in S Europe (Higgins, Riley, 1980). Foodplant: only Hirrschfeldia incana is known. Larva: short, stout, set with rather long hairs. Pupa: stout; in a frail cocoon.
PECULIAR TRAITS OF IMAGO F.w.l.: 20-26 mm. The fore wing apex is stretched out but not pointed, with an orange spot on a black background. The hind wing underside is greenish-yellow with large roundish yellowish-white spots.
GEOGRAPHICAL VARIATION: S Ural is inhabited by ssp. eupheme.
GENUS MICROZEGRIS Alpheraky, 1913.
Type species: Pontia pyrothoe Eversmann, 1832.
A monotypical West-Palearctic genus.
[87.1]. Microzegris pyrothoe (Eversmann, 1832).
TYPE LOCALITY: NW Kazakhstan: the valley of the Ural River: Indersk.
RANGE: Nizhnee Povolzhyethe lower Volga basin], South Preduralyethe western piedmont plain of Ural], Kazakhstan, Central Asia. This species has also been reported by H.J. Elwes (1899), by collections of Rueckbeil, for SW Altai, the village Kenderlik ("south of the Irtysh River"), according the collection by Ruckbeil, but so far has not been found within the territory of Russia. A rare species over its entire range.
HABITAT: in the Volga valley the imagines were observed near bank bushes (Kumakov, Korshunov, 1979).
FLIGHT PERIOD: April and May.
PECULIAR TRAITS OF IMAGO F.w.l.: 16-20 mm. The fore wing apex is black with an orange spot; the hind wing underside is greenish- yellow with several large transversely elongated white spots.
GENUS PARAMIDEA Kusnezov, 1929.
Type species: Anthocharis scolymus Butler,1866].
A monotypical genus.
88. Paramidea scolymus (Butler,1866]).
TYPE LOCALITY: Japan.
RANGE: Primorye northwards to Khabarovsk along the Ussuri valley, West, Central and North-East China, Korea, Japan.
HABITAT: forest edges, roads, floodland meadows in broad-leaved and mixed forests, waste lands at settlements, coastal meadows. A common species.
FLIGHT PERIOD: late April/early June.
PREIMAGINAL PHASES: studied in Japan (Shirozu,, Hara, 1960; Fukuda et al., 1982). Foodplants: Barbarea, Arabis, Cardamine, Draba, and other Brassicaceae; in Primorye was observed on Draba sp. and Descurainia sophia (Beljaev et al., 1989). Eggs: oval, at first white, later orange, laid singly on the pedicules and fruits of the foodplant. A freshly hatched larva is yellow with a black head. Larva: green with numerous black dots and white spiracular line, with a diffuse upper edge, on either side, covered with short hairs, it feeds on the seeds, fruits, buds and stems of the foodplant. Pupa: brown, with a longer head projection than in Anthocharis cardamines and distinct black spots on the dorsal side, hibernates.
PECULIAR TRAITS OF IMAGO F.w.l.: 17-21 mm, the fore wing apex is stretched out and pointed, with a small orange spot in males.
GEOGRAPHICAL VARIATION: The butterflies from Primorye refer to ssp. mandschurica Ballow, 1930 and differ from Japanese ones by smaller size, reduction of the basal suffusion on the wing upperside, and somewhat smaller and paler orange spot at the fore wing apex in males.
GENUS ANTHOCHARIS Boisduval, Rambur et Graslin, June, 1833].
Type species: Papilio cardamines Linnaeus, 1758.
A Holarctic genus with ten species.
89. Anthocharis cardamines (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: The non-tropical Eurasia northwards (in Siberia) to the forest-tundra, the Sakhalin, Hokkaido.
HABITAT: river and brook banks, wood edges, glades, various meadows (including highland ones), wasting lands, fields, settlements. In Priamurye flies in forests with participation of larch but not in broad-leaved ones (A. Streltsov)
FLIGHT PERIOD: May/middle July, in the subpolar and alpine regions - late June/late July.
PREIMAGINAL PHASES: studied in Europe (Eckstein, 1913 etc.). Foodplants: Cardamine, Alliaria, Arabis, Barbarea, Brassica, Erysimum, Hesperis, Sinapis, Sisymbrium, Thlaspi, Turritis and other Brassicaceae, Reseda lutea. Eggs: conical, with 14 ribs, at first yellowish-white, later orange, laid singly on the flowers, pedicules and young fruits of the foodplant. Larva: thin, blueish-green, covered by fine wrinkles and black dots, on either side with white a stripe at dark spiracles. It feeds on the flowers petals and fruits of the foodplant. Pupa: green, yellowish or brown, with much extended and pointed head; there are a white streak with tiny red dots on either side; hibernates.
PECULIAR TRAITS OF IMAGO F.w.l.: 17-25 mm. The wings are of a common shape; the hind wing underside green with a white marble pattern; the fore wing apex has black marginal spots extended along the outer margin; in males the distal parts of the fore wing is brightly orange.
GEOGRAPHICAL VARIATION: This species displays little geographic variation, which is much less than individual. Nevertheless, the following taxa has been described for our territory: progressa Sovinskyi, 1905 (Irkutsk), sajana Verity, 1908 (E Sayan), sibirica Haanshus, 1920 (W Siberia); septentrionalis Wnukovsky, 1927 (Yakutsk); koreana Matsumura, 1937 (Korea), kobayashii Matsumura, 1925 (Sakhalin); all they are very close to the nominotypical subspecies.
GENUS EUCHLOE Hübner,1819].
Type species: Papilio ausonia Hübner, var. esperi Kirby, 1871.
F.w.l.: 17-25 mm. The wings are white, in females often suffused with black and, sometimes, yellow scales; the fore wing apex has a wide black area containing white spots; the orange spots are absent. The butterflies are very variable individually and geographically.
A Holarctic genus with nine close species.
F.w.l.: 17-25 mm. The hind wing underside ground colour is yellowish- or dull-green with a marble pattern of white spots of more or less rounded shape and various size. A group complicated with respect of taxonomy, includes about five species. The taxonomy within the genus was problematic until recently because of use of the name simplonia Freyer, 1829, which in fact is a synonym for ausonia, while the butterflies from the Simplon Pass in the Alps have been described under the name simplonia Boisduval, 1828, that is now fixed by W. Back (1990, Atalanta, Bd.21, pp. 187-205).
90. Euchloe (ausonia) ausonia (Hübner, 1799) (= simplonia Freyer, 1929).
TYPE LOCALITY: North Italy.
RANGE: S Europe, N Africa, Anterior and Central Asia, South and Middle Ural, Central, West and South-East Altai Mts.
HABITAT: in Ural: meadow areas in various forests and birch groves, along wind strips; Rather a rare species. In Altai: forest and highland meadows at 1600-2300 m above sea level, most frequently at brook banks.
FLIGHT PERIOD: in S Ural: middle May/August. In Middle Ural the butterflies of a small spring form fly in June, while soon in July larger butterflies of a summer form appear, with a greater yellow suffusion on the hind wing underside. It is not excluded they are in fact different species. In Altai: middle June/late July depending on the locality, in one brood.
PREIMAGINAL PHASES: studied in Europe (Back, 1990; other authors). Foodplants: Barbarea, Sinapis alba, Sisymbrium, Turritis, and other Brassicaceae. Eggs: spine-shaped, pale- brown. Mature larva: 30-35 mm in length, green or ochre-coloured with two wide stripes along either side, sometimes little expressed, and a violet stripe along the back, speckled with rather large dark dots, the spiracles are white with dark rims. Pupa: with the head sharply pointed, pale brown or ochre- coloured with a dark line along the back and dark streaks along the anal margin of the wing cases.
PECULIAR TRAITS OF IMAGO F.w.l.: 18-25 mm. As different from E. naina, the fore wing costal margin is white with black marks; white spots on the hind wing underside are larger; sexual dimorphism is weakly expressed.
GEOGRAPHICAL VARIATION: Ural is inhabited by the subspecies volgensis Krulikovsky, 1907 described from Saratov by large summer individuals. The butterflies of a small spring "form" were described under the name uralensis Bartel, 1902. Both forms differ from the nominotypical subspecies, from Central Europe, with an enlarged black discal spot on the fore wing. Altai is inhabited by ssp. dubatolovi Korshunov, 1995; the original description is follows:
"... The butterflies from Altai, described below, differ from the western subspecies by a faded muddy-green hind wing underside ground colour and smaller white spots. By these and other traits they look like a peculiar transition (especially in the population of the Chuya River valley) to E. naina. In fact the butterflies from Chuya valley were not mentioned in the description, they belong to E. naina , that was stated below]).
HOLOTYPE: a male. F.w.l: 22 mm. The wing upperside is white; the fore wing apex is suffused with dark scales, the suffusion containing three white spots; the dark discoidal spot is crescent-shaped. The fore wing underside is white, the dark discoidal spot with a contrasted light discoidal vein; the apex is suffused with dark and green scales and has the same white spots as on the upperside; the costal margin with short black striae. The hind wing underside is greyish-green with white spots of different sizes, mainly with sharp edges.
ALLOTYPE: a female. F.w.l.: 23 mm. The wing upperside is white; the dark discoidal spot is large, concave on its sides, and, as different from E. n. naina [an error - must be E. ausonia ausonia - Yu.K.], is not fused along suffused veins] with the apical suffusion, which contains three elongate white spots.
MATERIALS: the holotype: a male, Altai, Katunskiy mt. range, 15 km SW of village Katanda, interfluve of the Kuragan and Kucherla rivers, alpine meadow, 2300 m, 13th July 1983 (V.V. Dubatolov leg.); the allotype: a female, the same locality, 14th July 1983 (V.V. Dubatolov leg.); paratypes: Altai, Ust'-Kan district, Karlinskiy Pass, 15th July 1970 (Kosinykh leg.); a male - the same locality as for the holotype, 12th July 1983 (V.V. Dubatolov leg.); 1 male 1 female - Katunskiy mt. range, the Katun' river headwaters at the Kapchal river mouth, 1700 m, 9th July 1987 (O. Kosterin leg.); a male - the Kapchal valley, 2300 m, 10th July 1987 (O. Kosterin leg.); 2 males 1 female - the Katun' headwaters, stow Altyn-Bulak, an old felled land, 1800 m, 13th July 1987 (O. Kosterin leg.); a male - Katunskiy mt. range, lower reaches of the Koksu river, 1600 m, 2th July 1988 (O. Kosterin leg.); a male - left bank of the Koksu river, long-forb meadow on wood opening, 1900 m, 12th July 1988 (O. Kosterin leg.); 1 male 2 females - the Argem (Direntay) river (a tributary of the Koksu) headwaters, an alpine meadow, 2200 m, 21st July 1988 (O. Kosterin leg.); a male, Kholzun mt. range, watershed of the Tigirek and Khamir river headwaters, 2300 m, 17th July 1983 (V.A. Lukhtanov leg.).
The subspecies is named after the surname of Vladimir Vladimirovichmust be Viktorovich instead of Vladimirovich - Yu.K.], an entomologist of ISEA (Novosibirsk) who collected a part of the type series of the new subspecies. "
91. Euchloe (ausonia) naina (V. Kozhantshikov, 1923) (= simplonia auct, non Boisduval,
1828; belia auct. non Cramer, 1782).
TYPE LOCALITY: the West Sayan: Lake Buyba.
RANGE: The North of Middle Siberia, Yakutia, the Magadan region, Kamchatka, the mountains of Bureya (the Ezop mountain range), SE Altai, the Sayans, including North Mongolia, the mountains of E. Kazakhstan.
HABITAT: in the mountains of South Siberia mostly highland meadows, in Altai - forest (rarely), subalpine (most frequently), and alpine meadows at altitudes of 1600-2300 m, observed also on barren south slopes at 1600 m above sea level; in the North - tundras, open larch or Betula ermani (or B. lanata) woods, steppefied south-exposed slopes, pebble banks, at altitudes of 200-1600 m.
FLIGHT PERIOD: middle June/middle July. On the Suntar-Khayata mountain range (Yakutia) a last instar larva was found by L.I. Popova on Descurainia sophioides (Brassicaceae), which pupated on 25th of July, 1985, the pupa hibernated, and the imago hatched on 16th of May, 1986.
PECULIAR TRAITS OF IMAGO 17-24 mm. The hind wing underside is dull green with more rounded white spots than in the previous species; the green colour predominates. In females the wings are suffused with yellow or black scales, often rather heavily, the black pattern on the fore wing is enlarged.
GEOGRAPHICAL VAIATION: The subspecies naina ranges in SE Altai, the Sayans and the mountains of Pribaikalye. In Yakutia, Magadan Region, and Kamchatka ssp. jacutia Back, 1990 ranges, which differs by on average smaller size and whitish tint of the majority of females. The butterflies from Taymyr and the Putorana Plateau were described as ssp. kuznezovi Korshunov, 1995. Original description was as follows:
"...Below we describe the butterflies from the Yenisei River low reaches, Taymyr, and the Putorana Plateau. They are closer to the subspecies naina than to jacutia, but differ from the former by in average more narrow wings and a darker hind wing underside ground colour.
HOLOTYPE: a male. F.w.l: 22 mm. The wing upperside is white; the fore wing costal margin from the base to the discoidal vein is suffused with black scales, further to the apex there are alternating black and white patches along the margin; the discoidal vein is accompanied by a figured dark spot which is to some extent more rounded than in E. naina naina. At the apex there are a dark stripe 2-3 mm wide which is parallel to the outer margin, and dark spots at the vein endings at the margin; the space between the stripe and the margin being generally white while in E. naina naina it is almost fully suffused. On the fore wing underside the dark discoidal spot is larger than in E. n. naina. The hind wing underside is almost exactly identical to that of E. n. naina, but the veins are not so conspicuous.
ALLOTYPE: a female. F.w.l.: 23 mm. On the wing upperside the white ground colour is almost displaced by a dark suffusion, the white areas remaining only in the center and at the edges of the wing. The hind wing underside ground colour is greyish-green without a dark suffusion; the veins are not contrasted.
MATERIALS: the holotype - the Yenisey lower reaches, the Irbo-Keta river middle flow, 18th July 1982 (D.M. Pupavkin leg); allotype - the same locality, 21st July 1982; paratypes: 5 males 3 females - the Yenisey lower reaches, the Rybnaya river middle flow, 12th-21st July 1982 a male - the Yenisey lower reaches, the Togbym river middle flow, 12th July 1982; 14 males 11 females - the Yenisey lower reaches, the Irbo-Keta river middle flow, 18th July 1982 (D.M. Pupavkin leg); a male - Taymyr, the Ary-Mas forest, July 1983 (Polovinkina leg.)
The subspecies being describing was earlier reported (Korshunov, 1985) as E. ausonia arctica, as it was designated in the labels by Nikolay Yakovlevich Kusnezov (1873-1948), a well- known Russian lepidopterologist and zoogeographist of the first half of this century. ".
92. Euchloe creusa (Doubleday and Hewitson, 1847).
TYPE LOCALITY: USA: Nevada.
RANGE: The mountains of South, Middle, and East Siberia, Chukotka, Kamchatka, the mountains of Bureya, Priamurye (the Blagoveshchensk environs, the Khekhtsir mountain range), Primorye (the Sasskii District), Mongolia, the north-west of North America.
HABITAT: in South Siberia: meadows and glades in montane forests, stone steppes on slopes and in intermontane hollows, rocks; in the North - open larch woods, steppefied south- exposed slopes, montane tundras; in Primorye these butterflies were found in an oak/aspen replacement forest flying together with Paramidea scolymus.
FLIGHT PERIOD: mostly June, locally (C Altai, Kamchatka) up to middle July. depending on the locality; early May in Primorye.
PREIMAGINAL PHASES: studied in N America (Scott, 1986). Foodplants: Draba lanceolata, Arabis glabra. Eggs are laid singly on the foodplant; the larvae feed on flowers and fruits. A probably foodplant in Primorye is Cardamine fritida, the flowers of which are regularly visited by the imagines.
PECULIAR TRAITS OF IMAGO F.w.l.: 15-21 mm. The hind wing underside is dull green with numerous white spots, angular and irregular in shape, which are stretched transversely. Similar species: E. naina, E. ausonia.
GEOGRAPHICAL VARIATION: The butterflies from Altai, which differ from other forms by a yellowish-green ground colour of the hind wing underside and so representing a clear-cut subspecies, were described as a "race" emiorientalis Verity, 1911 of Euchloe belia orientalis Bremer, 1864. The subspecies orientalis Bremer, 1864 ranges in the Sayans, Zabaikalye, the Stanovoe Nagorye upland, E Mongolia, ?the mountains of Bureya; ssp. kurentzovi Beljaev, 1986 in North Siberia and Kamchatka; ssp. nemoralis Beljaev, 1986, was described from the Spassk district of Primorye. According to Beljaev (1986); in ssp. kurentzovi the discal spot is relatively short and narrow, in ssp. nemoralis and orientalis it is longer and bended proximally, in males of ssp. nemoralis it is very narrow, in females and in ssp. orientalis is wide; besides, in ssp. kurentzovi the black basal suffusion is much more expressed.
**[There were two versions of Subfamily Coliadinae in the translation which was downloaded. I have assumed that the first version is the translation of Korshunov and Gorbunov (1995), and the second version the revision by Korshunov (1996). I have amalgamated the two versions, with the 1996 revisions indicated through strikeout deletes and underlined additions]**
SUBFAMILIA COLIADINAE Swainson, 1827.
TRIBUS CALLIDRYINI Kirby, 1896.
GENUS GONEPTERYX [Leech],1815].
Type species: Papilio rhamni Linnaeus, 1758.
F.w.l.: 25-35 mm. The wing upperside is yellow in males and whitish in females; the dark outer border is absent; the outer margin of each wing has a tooth. The pupae have a pointed head, a blunt prominence on the back of the thorax and a very large one - on the ventral side at the wing cases; they are usually attached to leaf underside. The imagines hibernate.
An Eurasian genus with seven species.
93. Gonepteryx maxima (Butler, 1885) (= rhamni auct.).
TYPE LOCALITY: Japan.
RANGE: Middle and Low Priamurye (from the city Blagoveshchensk to the village Troitskoe), Primorye, NE China, Korea, Japan.
HABITAT: dry open woods, bush thickets, wood edges.
FLIGHT PERIOD: late July/October and, after hibernation, middle May/June.
PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). Foodplants: in Primorye Rhamnus ussuriensis (Graeser, 1888). Eggs: cigar-shaped, greenish-yellow; laid singly on the young leaves of the foodplant in May/June. Larva: green with a white spiracular line. Pupa: light-green with several brown spots.
PECULIAR TRAITS OF IMAGO: F.w.l.: 30-34 mm; the fore and hind wings are of the same colour, yellow in males and light-yellow in females. The medial spots are relatively large, about 2-2.5 mm in diameter, of orange-red colour. Similar species: G. aspasia.
GEOGRAPHICAL VARIATION: On the continent there occurs the subspecies amurensis Graeser, 1888.
94. Gonepteryx rhamni (Linnaeus, 1758).
TYPE LOCALITY: Sweden.
RANGE: Europe. N Africa, the temperate Eurasia, including the Tian- Shan, eastwards to S Zabaikalye (Kyakhta) and northwards to the middle taiga zone.
HABITAT: forest edges and openings, glades, river and brook valleys in deciduous forests, birch and pine groves in the forest-steppe, woody mountain slopes, settlements. The imagines feed on various flowers (Corydalis bracteata, Pulmonaria mollis, Veronica longifolia, Pedicularis resupinatum, Arctium tomentosum, Carduus crispus, Centaurea scabiosa, Crepis sibirica), in springs - often on inflorescences of willows and sticky birch buds.
FLIGHT PERIOD: The butterflies appear from late June to middle August, depending on peculiarities of the season, the early emerged butterflies are capable of summer hibernation until autumn. Imagines overwinter in various shelters and appear amongst the earliest butterflies on thaws in March/April.
PREIMAGINAL PHASES: Foodplants: Rhamnus catharia, Frangula alnus, and the bird cherry (Padus avium) was also reported (Migranov, 1991; and others). Eggs: conical, yellowish or greenish, laid singly or by 2-6 on the young leaves of the foodplant. Larva (according to observations of I. Porchinskii): mat- green with uncontrasted whitish stripe (only its lower margin is well seen) along the green spiracles. Pupa: green with a light yellow lengthwise stripe on either side; the head is green. Numerous tiny black spinules, looking as black dots, cover the body, each spinule produces a drop of a liquid of a golden colour; these tiny drops make the larva a beautiful golden shine at sunlight. The larva moves extremely slowly, being threatened it rises the fore part of the body and bends it back and produce a drop of a liquid from the mouth. The pupa is usually attached head-downwards on the trunks, branches or stems of bushes and herbs.
PECULIAR TRAITS OF IMAGO: F.w.l.: 26-31 mm. The wings are yellow in males, greenish-white in females; medial spots is small, about 1-1.5 mm in diameter, of pale-orange colour.
95. Gonepteryx aspasia (Ménétriés, 1859).
TYPE LOCALITY: the Amur.
RANGE: Priamurye (downstream to the Gorin River), Primorye, Central and North-East China, Korea, Japan.
HABITAT: meadowy openings in valley and montane deciduous woods.
FLIGHT PERIOD: late July/October and, after hibernation, April/middle May.
PREIMAGINAL PHASES: studied in Japan (Fukuda et al., 1982). Foodplants: in Priamurye Rhamnus dahurica (Graeser, 1888). Eggs: at first white, later orange or red, laid on twigs of the foodplant. The larva resembles to that of G. maxima. Pupa differs from that of G. maxima by a smaller projection on the ventral side.
PECULIAR TRAITS OF IMAGO: F.w.l.: 26-30 mm. In males the fore wing upperside is bright yellow, the hind wing upperside is light yellow; the wings of females are greenish- yellow; the medial spots are small, 1-1.5 mm in diameter. Similar species: G. maxima.
TRIBUS COLIADINI Swainson, 1827.
GENUS COLIAS Fabricius in Illiger, 1807.
Type species: Papilio hyale Linnaeus, 1758.
F.w.l.: 19-34 mm. The wing outline is even, without pointed projections. The wing ground colour varies from white to orange- red or brownish-green. On the fore and, usually, hind wing there is a dark outer border often containing light spots. On the discoidal vein of the fore wing there is a black medial spot, while on that of hind wing - a light medial spot. The imagines often display an individual polymorphism but rather a weak geographical variation. Flight of these butterflies is strong and impetuous; some species are capable of substantial migrations.
The larvae of the majority of species are connected trophically with legumes. The pupae have a pointed head, a blunt prominence on the thorax back, and a larger one - on the ventral side at the wing cases. Hibernation occurs at the larval stage.
The genus includes about 70 species, the majority of which are connected with the mountains of C. Asia, N and S America.
96. Colias palaeno (Linnaeus, 1758).
TYPE LOCALITY: Sweden: Uppsala.
RANGE: The tundra, forest-tundra, and forest zones of Eurasia, the adjacent mountains, the Sakhalin, Japan, NE of North America. In the southern parts of the range the species is local and rare.
HABITAT: in the northern range: various types of tundras, forest meadows, open coniferous forests, bogs; in the mountains: the forest belt up to 1800 m above sea level; in the forest-steppe zone: raised and transitory bogs (the so- called "ryams" and "mar's"). On the Koni Peninsula (the Magadan Region) feeding of imagines was observed on Scorzonera radiata and Pentaphylloides fruticosa (Kosterin, 1994b).
FLIGHT PERIOD: mostly late June/late July, in the polar and mountain regions - until middle August.
PREIMAGINAL PHASES: studied in Europe (Eckstein, 1913; Ebert, 1991, etc.) and the Far East (Graeser, 1888). Foodplants: Vaccinium uliginosum and V. myrtillus. Eggs: elongate, with 20 ribs, yellowish, later become red; laid singly or in small batches on the foodplant leaves. Young larva: greenish-brown with a black head. The young larvae eat the leaf mesophyl and leave veins, hibernate at the third instar in leaf fall. Last instar larva: green with bright- yellow lateral streaks outlined above with black lines; the body is covered with small warts. The larva just after hibernation eats the buds, later - the leaves. It usually rests on the central vein of the leaf; being disturbed it rolls itself into a ring. Pupa: greenish-yellow with darker wing cases and a yellowish streak along either side of the abdomen; its stage lasts for about 15 days.
PECULIAR TRAITS OF IMAGO: F.w.l.: 20-28 mm. In males the wing upperside is bright or pale yellow, rarely yellowish-white (f. cretacea Schilde); in females it is white, sometimes yellow (f. illgneri Ruhl.). In both sexes the black outer border is as a rule without light spots, sometimes with lengthwise light strokes between the veins in females. The medial spot on the fore wing is absent or present as an elongated stroke. Individual variation is substantial.
GEOGRAPHICAL VARIATION: into a great extent is masked by an individual variation, most greatly expressed in the forest- tundra zone. Ural, the forest-tundra and forest zones of West Siberia are inhabited by ssp. palaeno. East of the Enisei River the subspecies orientalis Staudinger, 1892 (=? arctica Verity, 1908; = sachalinensis Matsumura, 1919; = karaganica Kurenzov, 1970) is widely distributed, differing from the nominotypical one by in general smaller size, a bright lemon-yellow ground colour in males and a wider dark border. The butterflies from the typical and arctic tundras of Siberia cam be attributed to ssp. lapponica Staudinger, 1871, described from the northern Lapland. In these populations lighter males with a narrower border are predominating. A Korean subspecies sugitani Esaki was reported for the southern Skihote-Alin' Mountains. The butterflies from Chukotka were described as Colias palaeno gomojunovae Koshunov, 1996, with the following original description:
"...It was Kurenzov (1970) who first noticed that Chukotka is inhabited by a peculiar variety of palaeno but he thought that it is identical to the form of Alaska, designed by him as schippewa (lapsus calami - Yu.K.) instead of chippewa Edw. However, the males of chippewa resemble our specimens only by the wing upperside, but the butterflies are much more yellow [above] and the marginal band is wider and is not cut through by [light] veins; the wing underside is more greenish, the discal spots are absent or hardly noticeable only on the fore wings. In the females from Alaska [I have studied 7 males and 3 females from there] black discal spots are seen above and beneath, the ground colour is yellowish. The females from Chukotka have a yellowish-white ground colour and no discal spots. There is also a difference in the genitalia, that was mentioned by Kurenzov (1970). By these characters we describe a new subspecies Colias palaeno gomojunovae Koshunov, sbsp.n.
HOLOTYPE: a male. F.w.l.20 mm. The wing upperside is yellowish green, the marginal spots on the fore wing are wider than on the hind wing, the colour of the marginal band is brownish, the band is not cut through by light veins. There is no discal spot on the fore wing upperside, but it is slightly seen on the underside of both wings. The hind wing underside is yellowish- green, with scattered grey scales. The fore wing underside is of the same colour, but grey scales present only at the costal margin. On the hind wing underside the silvery spot large, its diameter approximately equals to the length of elongate (and also silvery) spot on the transversal vein.
ALLOTYPE: a female. F.w.l. 21 mm. The wing upperside is yellowish-white. The marginal band at the apex is relatively wide, it is tapering to the anal angle, forming a kind of a triangle. Light spots present only in its wide part. On the hind wind there is only a narrow marginal streak at the fore angle. The discal spots are missing. The hind wing underside is muddy- green, a silvery spot is smaller than in the male, there is an additional tiny round spot above it. On the fore wing upperside there is only traces of the discal spot, the apex is of the same colour as the hind wing, the rest part of the wing is light.
In the male paratypes the marginal band is distinctly cut through by light veins only in 3 [of 7] specimens, which are worn out, in fresh specimens it is darker and is covered with silvery scales becoming worn out later.
MATERIALS: The holotype: a male - 30.07.1968, the Magadan Region, The Bilibino District (N. Gomoyunova); the allotype: a female - 12.07.1968, the left bank of the Khoroshaya River, a tributary of the Molondzha River, herbs/dwarf birches and willows/cloudberry (N. Gomojunova). Paratypes: 2 males - 3 and 15.07.1967, the settlement Markovo on the Anadyr' River (A.G. Mirzaeva); 2 males 1 female - 12 and 13.07.1968, the Khoroshaya River left bank; a male - 19.07.1969, the Molondzha River floodland; 2 males 2 females - 10 and 14.07.1969, the Magadan Region, the Kegali River, a larch parkland with the dwarf birches and willows; 2 males - 25.07 1 and 2.08.1986, West Chukotka, [the settlement] Ust'-Chaun, 100 km south of Pevek (V. Dubatolov).
The subspecies is dedicated to the memory of Nina Petrovna Gomojunova (1933-1973) - an entomologist and parasitologist of the Biological Institute of Siberian Division of the Academy of Sciences, who collected in Chukotka following the nomadic reindeer breeders. She tragically perished in an aircraft catastrophe."
97. Colias erate (Esper, 1804).
TYPE LOCALITY: the Volga basin: Sarepta.
RANGE: The arid and semiarid regions of the temperate and subtropical Eurasia, in Siberia northwards to the southern forest-steppe, in the Far East - to the so-called sub-taiga, i.e. the relatively narrow belt of contiguous deciduous forests to the south of the coniferous taiga; the South Sakhalin, the South Kuriles, Japan.
HABITAT: montane and lowland steppes, desertefied steppes, steppefied meadows, waste lands, fields, settlements, dry open pine and larch forests. This species is rather rare in Siberia.
FLIGHT PERIOD: in S Ural as well as in S Primorye in 2-3 broods, May/September, depending on the cite.
PREIMAGINAL PHASES: Foodplants: Fabaceae, such as Melilotus officinalis, Glycirrhiza korshinskyi reported for S Ural (Bartel, 1914; Migranov, 1991), Trifolium, Vicia, Glycine reported for the southern Far East (Kurenzov, 1970); sometimes this species becomes a pest of the soybean. Eggs: spindle-shaped with distinct ribs, greenish-white or yellowish, later become orange and, just before hatching of the larvae, violet-brown. Young larva: muddy yellowish- green, densely covered with hairs, with a black head set with small warts bearing whitish hairs. Larva, according to observations in Japan (Fukuda et a., 1982): greenish-white with a yellow spiracular line and orange and black dots at the hind margin of each segment, covered with tiny hairs. In Europe the larva was described (Mansell, 1987) as velvety green, covered with small warts and hairs, the spiracular line is white with a yellowish upper rim accompanied with reddish strokes, below the line a ground colour become lighter and black spots of an irregular shape present on each segment, those on segments 4-10 being smaller than on the others; the spiracles are white, dark-rimmed; the head is light-green. Pupa: light-green, sometimes with a yellowish streak on either side; black dots scattered over the body concentrate at the margins of wing cases. Hibernation - either at larval or pupal stage.
PECULIAR TRAITS OF IMAGO: F.w.l.: 22-30 mm. The wing upperside is brightly-yellow in males, of the same colour or white (f. albina Biener) in females; orange males and females (f. chrisodona Kindermann) are known from S Ural, which resemble C. croceus. As different from C. hyale, the dark border of the fore wing reaches the anal edge and (except for males of ssp. erate) contains isolated spots of the ground colour. The border on the hind wing is very narrow or split into separate spots. The medial spot on fore wing is round. The specimens from the mountains of S Siberia can be distinguished reliably from C. hyale only by the structure of the male genitalia, in which the valva is noticeably widened at the middle (see Fig...).
GEOGRAPHICAL VARIATION: The nominotypical subspecies is rather common locally in S. Ural, Kazakhstan and the Altai piedmonts but hardly penetrates into the West Siberian Lowland. Further eastwards a group of subspecies ranges which are sometimes considered as separate species C. polyographus and is characterized by a presence of light spots on the fore wing dark border in both sexes. The subspecies polyographus Motchulsky, 1860 proper ranges in Zabaikalye, the southern Far East and the Sakhalin. The ssp. naucratis Fruhstorfer, 1909 is known by few findings from S Pribaikalye (the settlement Taezhnyi) and Prisayanye (the environs of Krasnoyarsk), which is more common in Mongolia; it differs from polyographus by a smaller size, by reduction of dark border at the fore wing anal angle and on the hind wing, that makes this species to resemble C. hyale. The butterflies from the S Kuriles are known under the name tomarias Bryk, 1942 (= tokotana Bryk, 1942) and differ from the continental ones by a paler ground colour and a bleached border; the first brood specimens from the Sakhalin are similar to them.
98. Colias hyale (Linnaeus, 1758).
TYPE LOCALITY: S England.
RANGE: The temperate Eurasia eastwards to the Prilenskoe Plateau, the Stanovoe Nagorye upland and E Transbaikalia, in the North locally reaches the polar regions (the Lower Priobye: Labytnangi), in the mountains rises up to the tree-line.
HABITAT: meadows of various types, fields, fallow and waste lands, settlements. The imagines feed on various flowers: Trifolium, Medicago, Melilotus, Chamaecystis, Vicia, Compositae, and many others.
FLIGHT PERIOD: in southern parts of the range in 2-3 broods in middle May/September, in the North - in a single brood in late June/early July.
PREIMAGINAL PHASES: studied in Europe (Eckstein, 1913; Ebert, 1991; other authors) and Ural (Chislov, 1977). Foodplants: Fabaceae, such as Vicia, Coronilla, Chamaecystis, Lotus, and, especially, Melilotus alba, M. officinalis, Trifolium pratense, and Medicago falcata (the alfalfa). Eggs: spindle-shaped with 26-28 ribs, whitish or yellowish, later become dark-reddish with a yellow top; laid by 1-2 on leaf underside of stems; the larvae appear after a week. The larva of spring/summer brood hibernates in the second instar inside a silk-spun rolled leaf or freely, in or under the litter. The larva feeds for more than 30 days before hibernation and 20-23 days after it. Mature larva: dark-green with a darker dorsal side, yellow lateral lines and pink spiracles. Pupa: grey or greenish, with yellow lateral streaks; there are six black dots on either wing case and three ones on the ventral side of each segment; usually on bush twigs or stems of large herbs.
PECULIAR TRAITS OF IMAGO: F.w.l.: 20-30 mm. In males the wing upperside is pale-yellow, rarely white (f. albescens Metschler, 1922) or yellowish-orange (f. junior Geest, 1905), in males - whitish, rarely yellow (f. inversa Alpheraky). In both sexes the outer border on the fore wing contains large light spots, the last of them at the anal angle being fused with the light wing area. As different from a similar species C. erate, in male genitalia the valva is not widened at the middle (see Fig...).
GEOGRAPHICAL VARIATION: is poorly known in Siberia. The nominotypical subspecies inhabits Ural and South Siberia; the taxon altaica Verity, 1911 (type locality - the Chuya River valley) is very close to it; from Yakutia (the Vilyuy River) ssp. palidis Fruhstorfer, 1910 was described, differing by a paler ground colour in males and reduction of the submarginal dark spots on the wing underside; similar butterflies are known from Pribaikalye, from where (Irkutsk) the taxon irkutskana Staudinger, 1929 was described.
SYSTEMATIC NOTES: There are no reliable reports of the South- European species Colias alfacariensis Ribbe, 1905 (= C. australis Verity, 1911) from Siberia, however, in forest-steppe areas from S Ural to Pribaikalye larger and brighter specimens with a well expressed border on the hind wings (external characters of C. alfacariensis) are found. Moreover the taxon saissanica Reissinger, 1989, was recently described from Lake Zaissan, which lies between the Altai and Tarbagatai Mts. (East Kazakhstan) as a subspecies of C. alfacariensis. Further studied are needed, including examination of the preimaginal phases by which these species differ reliably. A mature larva of C. alfacariensis has two yellow stripes on either side, each accompanied with a row of adjacent quadrangular black spots situated on segment hind margins, the spots at the lower stripe being smaller (Henriksen, Kreutzer, 1892).
Besides, it should be noted that both names, Colias alfacariensis and C. australis, are in fact junior synonyms for Colias sareptensis Staudinger, 1871! Thus name was proposed for a presumed hybrid between C. hyale and C. erate, but later S. Alpheraky (1876, in the Proceedings of Russian Entomological Society, V.8: 153-154) changed its status and consider it as "a persistent variety usual in S and SE Russia". He wrote that males of sareptensis are large than those of a typical hyale, have more intense yellow colour, the black border is much wider on both wings; the females are also larger, yellowish-white, the black border reaches the anal angle of the fore wings, as in females erate ab. pallida; the wings in both sexes are much more elongate than in C. hyale. This form flew together with C. erate and C. hyale. So, a nomenclature change is needed: Colias sareptensis Staudinger, 1871 (= alfacariensis Ribbe, 1905; = australis Verity, 1911).
99. Colias mongola Alpheraky, 1897.
TYPE LOCALITY: Mongolia: Urga (at present Ulan-Bator)
RANGE: The mountains of Mongolia, West China, Central and SE [Russian] Altai Mts. (known from the Kurayskiy, Severochuyskiy, Saylyugem, and Katunskiy mountain ranges), Tuva (Lukhtanov, Likhtanov, 1994), S Pribaikalye (Baranchikov, 1979). A local species.
HABITAT: short-forb alpine meadows at brook sources, detritus ridges with sparse tundra vegetation, at altitudes of 2400-2900 m.
PREIMAGINAL PHASES: not studied. Probable foodplant in Altai: Oxytropis oligantha (Kosterin, 1994a).
PECULIAR TRAITS OF IMAGO: F.w.l.: 19-23 mm. The wing upperside is muddy-brownish-green; the dark outer border is wide, containing a row of elongate yellowish spots; on the hind wing upperside there is a whitish stroke between the medial spot and the wing base.
100. Colias tyche Bober, 1812. (= melinos Eversmann, 1847)
TYPE LOCALITY: Pribaikalye.
RANGE: The polar regions of Eurasia and Alaska, the mountains of the temperate Asia.
HABITAT: meadow and steppe patches on southern slopes, river terraces, in the extreme northern regions (the Yamal Peninsula) - river terrace meadows, tundrous slopes with willow thickets; in the Upper Priamurye - raised bogs with larch, open oak woods on dry slopes (Sviridov, 1981a), in the mountains - forb subalpine meadows, dwarf birch thickets up to 2400 m above sea level (Central Altai).
FLIGHT PERIOD: depending on locality, from late May (Zabaikalye) to August (the Taymyr Peninsula).
PREIMAGINAL PHASES: studied in Scandinavia (Henriksen, Kreutzer, 1983). Foodplants: Astragalus alpinus and Vaccinium are known from Scandinavia, Oxytropis nigrescens is reported from Taymyr (Korshunov et al., 1985), Caragana sp. - from Altai (V. Barkhatov). Eggs: pale-yellow, ribbed, barrel-shaped, laid singly on the stems and leaves of the foodplant. Larva: green with two yellow lengthwise stripes on the back and a red line beneath them and a white spiracular line on either side; the body is covered with sparse hairs up to 0.8 mm long. It hibernates, sometimes repeatedly. Pupa: green, yellowish-green, or straw- coloured, with a slightly darker back.
PECULIAR TRAITS OF IMAGO: F.w.l.: 18-28 mm. The wing upperside is whitish, with yellowish or greenish tint; on the fore wing the outer border is pale and contains a row of large light spots, they often fuse to split the border into two parallel ones. Similar species: C. nastes.
GEOGRAPHICAL VARIATION: is mostly masked by an individual variation. The subspecies werdandi Setterschtadt, 1840, occupying the extreme North of Europe, Polar Ural and North Siberia to the Taymyr Peninsula, further eastwards, to the Magadan Region, ssp. herzi Staudinger, 1901 ranges. Besides, two taxa were described from this area: zemblida Verity, 1911 from the Novaya Zemlya Isles and, by one male from the Okhota River, relicta Kurenzov, 1970. (According to the opinion of J. Troubridge, it is identical to the Canadian thula Hovanitz, 1915, which is considered to be a colour morph of Colias boothii Curtis, 1875 which, in turn, is considered by some authors as a hybrid between C. hecla and C. nastes). The subspecies tyche (= melinos Eversmann, 1847) ranges in Pribaikalye and the East Sayan, it differs from more northern butterflies by in general lighter ground colour of the wing upperside. The taxa montana Verity, 1911 (Altai Mts.), vitimensis Austaut, 1899 (the Stanovoe Nagorye upland); deckerti Verity, 1909 (Zabaikalye); chryseis Verity, 1911 (the Upper Priamurye) are very close to tyche.
101. Colias nastes Boisduval, 1832.
TYPE LOCALITY: the Labrador Peninsula.
RANGE: The north of East Siberia, of the Far East (including Kamchatka), and of North America. A local species.
HABITAT: shingle banks, steppefied south-exposed bank slopes, alpinotype meadows and montane tundras.
FLIGHT PERIOD: July/middle August.
PREIMAGINAL PHASES: studied in North America (Scott, 1986). Foodplants: Astragalus, Oxytropis, Hedysarum, and also Salix arctica. A biennial species. Larva: dark-green with small black dots, two yellowish streaks along the back and a white line on either side; hibernates twice, in early and the last instars.
PECULIAR TRAITS OF IMAGO: F.w.l.: 18-26 mm. The wing upperside is more strongly suffused with dark and greenish scales than in C. tyche, the light spots on the border are smaller and do not fuse to the ground colour. The fore wing underside is ash-grey, the hind wing underside is yellowish-green, with a silvery discal spot ringed with pink or reddish rim.
GEOGRAPHICAL VARIATION: In the Suntar-Khayata mountain range and the basins of Yana and Indigirka Rivers the subspecies jacutica Kurenzov, 1970 (= jacuttica Ferris, 1985; = jacuticola Weiss et Mracek, 1989) is distributed, which is characterized by a smoky-brown wing upperside with a wide border with contrasted white spots on both fore and hind wings; from Chukotka and the Kolyma basin the subspecies sibirica Kurenzov, 1970, has been described, the name of which was preoccupied by Colias aurora sibirica Lederer, 1853, so, this name was replaced by us with the name dezhnevi Korshunov, 1995 (etymology: Semen Dezhnev - a Russian kazak who first sailed round the Chukotka Peninsula). In this subspecies the wing upperside is pale-yellowish-green in males and pale-yellow in females, on the fore wing the outer border is split by a row of large light spots into two parallel fragments.
102. Colias chrysotheme (Esper, 1781).
TYPE LOCALITY: Hungary: Cremnitz.
RANGE: The steppen and forest-steppen zones from Central Europe to East Zabaikalye, the mountains of East Kazakhstan, South Siberia, and Mongolia.
HABITAT: lowland and montane steppefied meadows up to the altitude of 2200 m, less frequently - fields and long-fallow lands. The imagines were observed to feed on Aster alpinus, Dracocephalum nutans, Brassica, Trifolium pratense.
FLIGHT PERIOD: in South Ural and the forest-steppe zone of West Siberia in two broods in May and July/August, in Altai and further eastwards - in a single brood in middle June/late July.
PREIMAGINAL PHASES: studied in Central Europe (Niculescu, 1963; other authors) Foodplants: Astragalus and Vicia are known. Eggs: cylindrical with pointed apex; at first white, later become yellowish. The young larva is muddy-green with a dark head. Mature larva: green with numerous small lengthwise-directed reddish strokes and a white lateral line on either side (which is more distinct in the fourth instar); the ventral prolegs are greenish-yellow. The larva feeds at night. Pupa: greenish-yellow with dark markings on the edges of the projections and the wing cases.
PECULIAR TRAITS OF IMAGO: F.w.l.: 21-29 mm. The wing upperside usually is yellowish-orange or orange in both sexes. The yellow males and females (f. shugurovi Krulikowsky, 1903) and white females (f. hurlei Aigner-Abafi, 1902) are known from Ural. The outer border in males is 3-4 mm wide, with conspicuous yellowish veins throughout its length; in females it is 4-5 mm wide, with a row yellow spots, which is incomplete on the fore wing. In the male the androconial spot is absent. Similar species: C. thisoa, C. croceus.
GEOGRAPHICAL VARIATION: In the southern Ural the nominotypical subspecies is distributed, South Siberia, from Zauralye to S Zabaikalye is inhabited by ssp. andre Hemming, 1933 (instead of sibirica Grum-Grzhimailo, described from the environs of Krasnoyarsk) differing from the nominotypical subspecies by a pale orange-yellow ground colour and a wider border. Very peculiar butterflies of the Altai Mountains, the Sayans, and the mountains of Tuva, differing from lowland subspecies by a more even and bright colouration of the wing upperside, narrower light streaks along the veins in the dark marginal band in males and darker and more contrasted colouration of females, which tend to resemble C. thisoa by these characters, were described as Colias chrysotheme elena P. Gorbunov, 1995.
Original description is as follows:
" MALE: F.w.l. 24.4-27.9 mm (26.2 mm in the holotype). The wing upperside is light-orange with a narrow yellowish stripe along the forewing fore margin and an area of the same colour at the anal margin of the hind wing. A slight suffusion with dark scales is noticeable on the hind wing. The marginal band on the fore wing in its middle part is 3.5-5 mm wide, usually all over its length it is cut through with narrow light streaks along the veins, which most often don't reach the fringe. The maximum width of the marginal band on the hind wing is 2.4-4 mm; light venal streaks on it are even narrower than on the fore wing, in some specimens they are hardly seen at the internal margin of the band. Underside the ground colour of the hind wing is yellow, that in the central area of the fore wing is orange- yellow, the pattern is as in the subspecies andre. The genitalia are alike those in C. chrysotheme andre and C. thisoa [the statement concerning thisoa is evidently of no relevance - Yu.K].
FEMALE: F.w.l. 25.5-28.7 mm (27.6 mm in the allotype). The fore wing upperside is bright orange-red (while in the subspecies andre they are yellowish-orange or orange-yellow) with a dark suffusion in the basal area and at the fore margin. The marginal band on the fore wing is blackish-brown, 4.8-6.0 mm wide in its narrowest part, with very contrasted 6-7 yellow spots. The black discal spot is large, usually of a triangular shape. The hind wing upperside is muddy-orange (much darker than in the females of andre) with an orange discal spot about 2 mm in diameter. On the hind wing the marginal band contains two yellow spots at the apex, four other yellow spots interrupt the band fusing with the ground colour at its internal margin. The discal spot is oval-shaped, 3.0-3.6 mm long. The wing upperside ground colour is greenish-yellow, but orange in the central area of the fore wing. There is a row of 3-5 black spots along the outer margin of the wore wing. The discal spot on the fore wing underside is somewhat larger than on the upperside and contains a light dot. On the hind wing underside just above an oval silvery discal spot there is an additional small silvery spot.
MATERIALS: The holotype: a male - 29.06.1993, SE Altai, the village Aktash (V. Barkhatov). The allotype: a female - 26.07.1982, Tuva, Choon-Khem (O. Fileva). Paratypes: 3 males - 2-3.07.1971 , Tuva, the settlement Shuurmak, a steppe (S. Nikolaev); a female - 25.06.1971, the same locality; a female - 16.07.1980, Altai Mts., the Seminskii Pass (Yu. Korshunov); 2 males - 25.06.1992, SE altai, the village Aktash (V. Barkhatov); 14 males 1 female - 27-29.06.1993, the same locality."
103. Colias thisoa Ménétriés, 1832.
TYPE LOCALITY: Daghestan, the mountain Shakhdag.
RANGE: The mountain regions of Asia Minor, the Caucasus, the Elburs, Central Asia, East Kazakhstan, Central and SE Altai Mts., Tuva (the Moren River), the East Sayan (the Mondy village environs). A local species.
HABITAT: meadowy steppe areas on slopes and plateau at altitudes of 400-2400 m, together with C. chrysotheme. The feeding of imago was observed on the flowers of Dracocephalum nutans.
FLIGHT PERIOD: middle June/late July.
PREIMAGINAL PHASES: Foodplants: Astragalus and Oxytropis are known from the Caucasus (Nekrutenko, 1990). Hibernation - at the larval stage.
PECULIAR TRAITS OF IMAGO: F.w.l.: 22-28 mm. The wing upperside is orange in males and orange-red in females; in males the outer border is 2-3 mm wide with weakly dentate inner margin, on which the veins can be contrasted by orange scales only at the apex; in females it is 3-4 mm wide and contains a row, usually incomplete (sometimes absent), of rather small yellow spots, the hind wing upperside in females is strongly suffused with dark scales. The androconial spot in males is absent.
GEOGRAPHICAL VARIATION: the South Siberian butterflies in general resemble those from Central Asia, from where the subspecies aeolides Grum-Grshimailo, 1890 (type locality: Karategin, Zaalai) and urumtsiensis Verity, 1909 (type locality: Urumqi) have been described. The butterflies from the highlands of Central and SE Altai, as different from those from the piedmont steppes of W Altai, usually are smaller and have a narrower border.
104. Colias hyperborea Grum-Grshimailo, 1899.
TYPE LOCALITY: the mouth of the Vilyuy River, the Yana River (at Verkhoyansk and the mouth of the Adycha River), the Kolyma valley.
RANGE: The northern Middle Siberia, East Siberia, W Chukotka, the East Sayan, Pribaikalye, Zabaikalye.
HABITAT: steppe-like south-exposed slopes, edges of larch woods, bank meadows, shingle banks, in the Ochot coast of the Koni Peninsula the species was restricted to south-exposed loam bluffs of the coastal terrace with sparse mesophilous vegetation with numerous flowers (Kosterin, 1993b).
FLIGHT PERIOD: middle June/August.
PREIMAGINAL PHASES: not studied. On the Suntar-Khayata mountain range oviposition was observed by V.V. Dubatolov on Astragalus alpinus.
PECULIAR TRAITS OF IMAGO: F.w.l.: 22-27 mm. The wing upperside is orange in both sexes. The outer border is uninterrupted in males and contains a row, usually full, of yellow spots in females. As different from similar species C. viluensis and C. hecla, in males there is a pink androconial area at the very base of the hind wing costal margin.
GEOGRAPHICAL VARIATION: poorly studied. From the town Srednekolymsk environs the taxon viluensoides Verity, 1909 was described.
SYSTEMATIC NOTES: Judging by the presence of the androconial spot, V.V. Dubatolov (personal communication) stated the taxa tuncuna Austaut, 1912 (type locality: the Tunkinskie Goltsy mountain range within the East Sayan) and kurnakovi Kurenzov,1970 (described by a single specimen from the Omsukchanskiy mountain range in the Magadan Region as a subspecies of C. hecla) to belong to this species.
105. Colias viluensis Ménétriés, 1859.
TYPE LOCALITY: E Siberia, the Vilyuy River.
RANGE: the Putorana Plateau, Yakutia, the Magadan Region, Pribaikalye, Zabaikalye, the mountains of Bureya, NE Mongolia, ?Chukotka, ?Alaska.
HABITAT: highland and valley meadows, less frequently open larch stands, montane tundras.
FLIGHT PERIOD: in Zabaikalye and the Upper Priamurye middle June/middle July. In the northern range the imagines can fly together with C. hyperborea, but appear 7-10 days later, usually in late June.
PECULIAR TRAITS OF IMAGO: F.w.l.: 22-27 mm. The wing upperside is bright-orange or orange-yellow (f. sulphureaflava Sheljuzhko, 1918) in males and bright-orange, less frequently white (f. alba Verity, 1909), in females. The outer border in males is uninterrupted, about 4 mm wide, in females it usually contain a full row of yellow or white spots; as different from C. hyperborea, the androconial spot in males is absent.
GEOGRAPHICAL VARIATION: The nominotypical subspecies inhabits the northern part of the range, the subspecies dahurica Austaut, 1899 ranges in Pribaikalye and Zabaikalye.
SYSTEMATIC NOTES: From the Olenyok and Yana River basins the species C. aquilonaris Grum-Grshimailo, 1899 was described. According to the original diagnosis (Grum-Grshimailo, 1899) the main specific feature of this species separating it from C. hecla consists in dark muddy-yellow colouration of the wing underside and the wings being wider and of a somewhat different shape; the outer border is also wider, especially at the fore wing apex. According to a personal communication of V.V. Dubatolov, the type of this taxon, preserved in the collection of the Zoological Institute in St. Petersburg, is conspecific to C. viluiense.
106. Colias hecla Lefebvre, 1836.
TYPE LOCALITY: Greenland.
RANGE: The polar regions of Eurasia and North America.
HABITAT: lowland dry herbage-grass tundras, on flat elevations in marshy areas, meadows in river and brook valleys, at willow thickets; these butterflies were found in the extreme northern wood Ary-Mas in the Taymyr Peninsula.
FLIGHT PERIOD: July/middle August.
PREIMAGINAL PHASES: studied in Lapland (Henriksen, Kreutzer, 1982). Foodplants: Astragalus alpinus, Trifolium repens (Fabaceae), Arctostaphylos, Vaccinium (Vacciniaceae), Dryas octopetala (Rosaceae); in N America (Scott, 1986) - also Salix arctica (Salicaceae); in the Polar Ural P. Gorbunov observed oviposition on Hedysarum sp. Eggs: yellowish or orange, barrel- shaped with about 17 ribs, laid singly or in small batches on the foodplant. Larva: green with a yellow line on either side, a dark-green spot on each segment beneath them, and a whitish- yellow spiracular line; spiracles reddish. Hibernation occurs on either larval or pupal stage. Pupa: light-green dorsally, dark- green laterally, with yellowish lateral streaks, light green wing cases, abdomen may be olive-brown.
PECULIAR TRAITS OF IMAGO: F.w.l.: 18-24 mm. The wing upperside is usually orange, less frequently white in females; the outer border in males is uninterrupted, in females it contains usually a full row of yellow spots. As different from C. hyperborea, the border is in general narrower, the hind wing underside is darker as being suffused by greenish-grey scales, the nacreous discal spot has a wider dark-pinkish rim; the androconial spot is absent in males. Similar species: C. viluensis, C. hyperborea.
GEOGRAPHICAL VARIATION: The butterflies from the Polar Ural and the northern West Siberia are most close to the subspecies orientis Wnukovsky, 1929 (pro orientalis Grum-Grshimailo, 1893), known from Yakutia, the Magadan Region and Chukotka. From a Lapland subspecies sulitelma Aurivillius, 1890 they differ by in general a wider border, which in females contains smaller and more clear-cut light spots, always separated from the ground colour on the fore wings. A report of C. hecla orientalis for Altai (Rehnelt, 1983 in Nota Lepidoptera 6(4): 244) was based on both wrong labeling and wrong identification (in fact it was Colias hyperborea).
107. Colias heos (Herbst, 1792) (pro Papilio aurora Esper, 1781, nec Papilio aurora Cramer, 1780; = sibirica Lederer, 1852).
TYPE LOCALITY: Siberia: Nerchinsk.
RANGE: This local species ranges from Verkhnee Priobye (the surroundings of cities Barnaul, Tomsk, in the Novosibirsk Region at the village Kargat and the Baksa River) and the Altai Mts. through the mountains of South Siberia and Mongolia to Priamurye, Primorye and NE China.
HABITAT: various meadows, meadow steppes, in Zabaikalye and the Far East also rough southern slopes and fields. In Zabaikalye the imagines often visit flowers of Lilium pumilum, L. buschianum.
FLIGHT PERIOD: middle June/middle July, in the mountains - to late July.
PREIMAGINAL PHASES: Foodplants in Primorye: Vicia, Astragalus, and other Fabaceae, including the cultivated soybean (Glycine max) (Kurenzov, 1970). Hibernation at the larval (Priobye) or pupal (the Far East) stages. The pupa is attached to the foodplant or herbs with large stems.
PECULIAR TRAITS OF IMAGO: F.w.l.: 25-32 mm. This is a remarkably variable species. The wing upperside colouration in males is orange of various aspects, in females it can be orange, muddy- orange, sometimes almost black (f. nigra Tshugunov, 1914), white (f. chloe Eversmann, 1847), or white with a slight dark (f. obscurissima Verity, 1911) or orange (f. decolorata Staudinger, 1897) suffusion, sometimes salad-green with an orange spot (f. semenovi Standel, 1960). In any case in both sexes the veins are to some extent suffused with dark scales. In females the outer border contains a row of yellow or white spots, which is incomplete on the fore wing. The males have an orange androconial area at the hind wing base upperside.
108. Colias myrmidone (Esper, 1781).
TYPE LOCALITY: Hungary: Turnau.
RANGE: Central and Eastern Europe, Middle and South Ural, the south of the Kurgan and Tyumen' Regions.
HABITAT: forest and forest-steppe meadows, edges of birch or pine woods. The imagines feed mostly on the flowers of Fabaceae plants.
FLIGHT PERIOD: in South Ural in two broods in June and August, respectively; in Middle Ural usually a single brood in middle June/middle July.
PREIMAGINAL PHASES: Foodplants: Chamaecytisus ruthenicus is known. Eggs: green. Larva: green with a dark back line and a light-green line on either side, speckled with dark dots. Pupa: green with a yellowish lateral stripe on either side and tiny dark dots on the ventral side of the abdomen; it is usually attached to a large stem.
PECULIAR TRAITS OF IMAGO: F.w.l.: 23-28 mm. The wing upperside is orange with noticeable violet bloom in males and orange, less frequently white (f. alba Staudinger, 1871) or yellow (f. flavescens Garbowski, 1892) in females. In males the outer border is 3-4 mm wide with the veins of the same colour, in females it usually contains a row of seven yellow spots. The males have a light-orange androconial area at the hind wing base upperside. Similar species: C. croceus.
GEOGRAPHICAL VARIATION: Ural and Zauralye are inhabited by the subspecies ermak Grum-Grshimailo, 1890, differing from the nominotypical one by a somewhat larger size and a brighter ground colour..
109. Colias crocea (Geoffroy in Fourcroy, 1785).
TYPE LOCALITY: France: Paris.
RANGE: Europe, N Africa, Anterior Asia. The species is capable of long migrations. In Middle and South Ural and the Kurgan region the imagines are observable rarely and not every year. HABITAT: meadows, fields, steppefied areas, settlements.
FLIGHT PERIOD: late May/September, in two broods.
PREIMAGINAL PHASES: studied in Europe. Foodplants: Onobrychis, Astragalus, Medicago, Cytisus, Coronilla, Chamaecytisus, Lotus, Trifolium, Vicia, and other Fabaceae. Eggs: spindle-shaped with 24-26 ribs, yellowish, laid singly on the foodplant. Larva: green, set with small warts and short hairs, on either side with a yellow spiracular stripe, with a white rim above, containing lengthwise red dashes and. Pupa: green with a lighter ventral side and yellow streak on either side of abdomen, shorter than in C. hyale.
PECULIAR TRAITS OF IMAGO: F.w.l.: 24-27 mm. In males and females wing upperside is yellowish-orange. White females (f. helice Hübner, 1803) and yellow males (f. erateformis Nickerl) are known. Outer border in male about 5 mm in width, with veins suffused with yellow scales, in females it usually contains 3-4 yellow spots. The males have a light-orange androconial area at the hind wing base upperside. Similar species: C. myrmidone.
110. Colias fieldi Ménétriés, 1855.
TYPE LOCALITY: the Himalaya.
RANGE: Central Asia, NE China. In South Primorye the only male was collected by A.V. Tsvetaev on 6th of September 1966 in the Kedrovaya Pad' Nature Reservation.
PECULIAR TRAITS OF IMAGO: F.w.l.: 27-30 mm. The wing upperside is bright yellowish-orange; the border is clear-cut in males, about 5 mm wide, and diffuse in females, much widened at the fore wing apex, on the hind wing it is stretched along the costa; the medial spot on the fore wing upperside is large, almost round but slightly stretched to the fore margin, the corresponding spot on the underside containing a nacreous stroke; the hind wing upperside has a light-orange quadrangular spot in the center, clear-cut in males and obscure in females; the hind wing underside is greenish-yellow with an oval nacreous medial spot.
GEOGRAPHICAL VARIATION: The Far East is inhabited by ssp. chinensis Verity, 1909.
ETYMOLOGY: Genrikh Andreevich Field (1822-1875), a member of the Russian Entomological Society, a co-author of N.G. Ershov of the catalogue of Lepidoptera of the Russian Empire (1870).