MASAYA YAGO
Biosystematics Laboratory, Graduate School of Social and Cultural Studies, Kyushu University
COMPARATIVE MORPHOLOGY AND IDENTIFICATION OF THE SUBGENUS KULUA, WITH DESCRIPTION OF A NEW SPECIES FROM VIETNAM (LEPIDOPTERA, LYCAENIDAE, HELIOPHORUS)
M. Yago, 2002. Comparative morphology and identification of the subgenus Kulua, with de- scription of a new species (Lepidoptera: Lycaenidae: Heliophorus). – Tijdschrift voor Ento- mologie 145: 145-171, figs. 1-80. [ISSN 0040-7496]. Published 1 December 2002. A new lycaenid species, Heliophorus (Kulua) smaragdinus Yago & Monastyrskii sp. n., is de- scribed from Central Vietnam. It belongs to the tamu-group of the subgenus Kulua Zhdanko. The male and female genitalia of Kulua species are described and illustrated, and a key to species of this subgenus is proposed. Correspondence: M. Yago, JSPS Research Fellow, Biosystematics Laboratory, Graduate School of Social and Cultural Studies, Kyushu-Univ., 4-2-1 Ropponmatsu, Chuo-ku, Fukuoka, 810- 8560 Japan. E-mail: [email protected] Key-words. – Heliophorus; genitalia; Kulua; Lycaenidae; Lycaeninae; new species; systematic position; Vietnam.
The genus Heliophorus Geyer, 1832, in the subfam- oval or spherical; those of the saphir-group are 1) jux- ily Lycaeninae (s. str.), presently comprises 20 species ta produced into a pair of extraordinarily elongate occurring in tropical to temperate zones of the process and 2) E3 marking on underside of hindwing Palearctic and Oriental Regions. Yago et al. (2000) re- extremely enlarged; those of the yunnani-group are 1) cently updated the classification of Heliophorus, and wings rounded with evenly curved termen, 2) hind- divided the genus into two subgenera, Heliophorus s. wing lacking tail, 3) M1 on underside of wings absent, str. and Kulua Zhdanko, 1995 as well as an unnamed 4) underside of wings lacking black markings of E1, E2 group for H. kiana (Grose-Smith, 1889). In particu- and E3, 5) space between E1 and E3 extremely widened, lar, they considered the following character states as 6) male genital valva with slightly subbasal area bor- autapomorhies for the clade composed of Kulua: 1) dered by internal ridge and 7) male genital juxta with M1 on underside of wings arranged in a continuous a pair of deep invaginations and a pair of long process- band, 2) E3 on underside of hindwing shifted inward- es. The subgenus Kulua characterized by the above ly in cells 4 and 5, 3) female genital plate with large features comprises 14 species occurring in Kashmir lamellate antevaginal process more or less bifurcate, 4) across Himalayan region (including Nepal, Sikkim, Female genital plate with bank-like swelling along Bhutan and Assam) to Myanmar, Thailand, Laos, posterior margin, 5) female ductus bursae short, etc. Vietnam and China. In addition, Yago et al. recognized the following three The male genitalia of most species of the subgenus species groups in Kulua: tamu-group, saphir-group Kulua were briefly described with simple illustrations and yunnani-group. According to them, the presumed by Riley (1929) and Huang (1999). Here I redescribe autapomorphies of the tamu-group are 1) hindwing and illustrate in more detail the male genitalia of all inner margin distinctly longer than costa, 2) hindwing species of Kulua, except for H. brilliantinus Huang, tail short and broad, 3) M1 on underside of wings 1999 and H. pulcher Chou, 1994 which could not be arranged in a conspicuous continuous band, 4) female examined. Female genitalia are only illustrated for H. genital plate very wide anteriorly with wide lamellate tamu (Kollar, 1844) (Zhdanko 1995), and H. brahma flange of antevaginal process, 5) female ductus bursae Moore, 1858, H. saphir (Blanchard, 1871) and H. extremely short and 6) male genital valva tending to be yunnani D’Abrera, 1993 (Yago et al. 2000). General-
145 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
1 2
3 4
Figs. 1-4. Heliophorus smaragdinus. – 1, Holotype, male, upperside; 2, ditto, underside; 3, paratype, female, upperside; 4, dit- to, underside. ly, identification of female species of Kulua is very dif- after careful examination of the general appearance ficult because of their very similar wing markings. I and the male and female genitalia of this material, I have found distinct differences among these species reached the conclusion that it is undescribed. This dis- mainly in the lamella antevaginalis of the female gen- covery may be worthy of note because Heliophorus italia as described below. Based on detailed analysis of species are well clarified in the Lycaenidae because of wing markings and genitalia, I provide a key to both their striking metallic wing markings in males, and males and females for the species of Kulua. show high endemism in Central Vietnam (Anony- Five species of Heliophorus have been recorded pre- mous 1994, Collins et al. 1991). In this paper we also viously from Vietnam: H. epicles (Godart, 1824), H. describe a new species of the subgenus Kulua and dis- ila (de Nicéville, 1896), H. kohimensis (Tytler, 1912) cuss its phylogenetic position and classification. and H. brahma (Fruhstorfer 1912, Metaye 1957, Eliot 1963, Yago et al. 2000). In late March 1998, one of MATERIALS AND METHODS authors of the new taxon, A. L. Monastyrskii partici- pated in a biological survey in Vietnam, in cooperation Material with the BirdLife International Vietnam Programme. Specimens examined in this work are dried adults. During this survey he collected some specimens of a Their data and depositions are mentioned under the species of Heliophorus in Ngoc Linh Nature Reserve, descriptions of each species. They were from the fol- located in Kon Tum province, Central Vietnam. This lowing collections: The Natural History Museum, species is similar to H. tamu, H. eventa Fruhstorfer, London (BMNH), the Kunming Institute of Zoology, 1918, H. hybrida (Tytler, 1912), H. brahma, H. pul- China (KIZC), the South China Agricultural Universi- cher and the nominotypical subspecies of H. androcles ty (SCAU), the Museum of Nature and Human Activ- (Westwood, 1852) in having the metallic green ap- ities, Hyogo (MNHAH), the Kitakyushu Natural His- pearance on the upperside of both wings. However, tory Museum (KNHM), the Kurashiki Museum of
146 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
Fig. 5. Habitat of Heliophorus smaragdinus, Ngoc Linh (1,700 m alt.), Kon Tum Province, Central Vietnam.
Natural History (KMNH), the Biosystematics Labora- stereoscopic microscope with magnifications up to tory, Kyushu University (BLKU) and from the follow- ϫ126. For detailed microstructure, a Leica MZ12 ing private collections: A. L. Monastyrskii (ALMPC), stereoscopic microscope with magnification up to M. Saito (MSPC), Y. Watanabe (YWPC). Particularly, ϫ400 was used. many Chinese and Japanese entomologists, whose Terminology of the male genitalia follows Shirôzu names are mentioned in the acknowledgements of (1960), except for the substitution of brachium by this paper, donated specimens to BLKU. Recent mate- falx, and that of the female genitalia by Shirôzu & Ya- rial was collected under the permissions of the Forest mamoto (1956) except for the term corpus bursae. In Protection Department of Ministry of Agriculture wing markings, we adopt the system proposed by and Rural Development of Vietnam, the Department Schwanwitsch (1949). However, as to E3 of the un- of Hotel & Tourism of Myanmar and the National derside, we included not only the outer black lines but Democratic Association of Kachin, or in cooperation also the inner white and black lines in E3. The cells 1’ with the BirdLife International Vietnam Programme, and 1 in his system were treated as cells 1a and 1b+c, the Vietnam Russian Tropical Center, the Forest In- respectively. ventory and Planning Institute in Vietnam and the Hanoi Agricultural University. TAXONOMIC PART Methods Checklist For the observation of adult genital structures, api- cal parts of the abdomen were placed in a 5% KOH so- Heliophorus Geyer, 1832 lution at about 50° for 3-5 hours depending on the subgenus Kulua Zhdanko, 1995 size of material, then neutralized in 2% acetic acid. Af- ter this treatment, they were washed with distilled wa- Heliophorus tamu-group ter and placed in 80% ethanol for dissection and obser- tamu (Kollar, 1844) vation. Chlorazol black E and Delafield’s hematoxylin brahma Moore, 1858 were used to stain the membranes. Structures were ex- eventa Fruhstorfer, 1918 amined and illustrated under an Olympus SZ-60 androcles (Westwood, 1852)
147 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
hybrida (Tytler, 1912) male genitalia, valva rather large and broad; juxta pulcher Chou, 1994 short, hardly produced anteriorly. In female gen- moorei (Hewitson, 1865) italia, lamella antevaginalis not Y-shaped ...... 4 gloria Huang, 1999 4. Discal lines (M1) of underside straight, prominent oda (Hewitson, 1865) (especially on forewing), dark and often covered bakeri Evans, 1927 with reddish scales in cells 2 and 3 (sometimes also smaragdinus Yago & Monastyrskii sp. n. in cell 4) of forewing. Discal area on underside of forewing with a distinct and extensive orange H. saphir-group (flushed orange) patch. In male genitalia, tegu- saphir (Blanchard, 1871) men with a pair of long lateral processes (fig. 52); saphirioides Murayama, 1992 valva expanded and truncate on posterior margin brilliantinus Huang, 1999 (fig. 52). In female genitalia, lamella antevaginalis narrowly and longitudinally swollen on midven- H. yunnani-group tral area of basal portion, produced posteriorly yunnani D’Abrera, 1993 into broad, short process with marginal serration (fig. 56) ...... oda – Discal lines (M1) of underside absent, or if faintly Key to species of the subgenus Kulua indicated then consisting of a row of fine arched The following key based on wings and genitalia can lines most readily seen in cells 2 and 3 of forewing. distinguish the species of the subgenus Kulua of the Discal area on underside of forewing without a dis- genus Heliophorus. Some species were based on the tinct and extensive orange (flushed orange) patch. published descriptions and illustrations of wings and In male genitalia, tegumen with a pair of lateral male genitalia of Riley (1929), D’Abrera (1986, 1993), processes greatly reduced (fig. 55); valva rounded Chou (1994), Huang (1999) and Yago et al. (2000). on posterior margin (fig. 55). In female genitalia, Females of H. hybrida, H. sahirioides and H. brillianti- lamella antevaginalis small and slender, produced nus are unknown, and thus not included in this key. posteriorly on apical 1/3 into a pair of triangular processes (fig. 58) ...... bakeri 1. Upperside of forewing without elliptical orange 5. Upperside of both wings metallic bronze red, green discal patch (males) ...... 2 (including golden green, bronze green and silvery – Upperside of forewing with elliptical orange dis- green) or with at least basal greenish scales ...... 6 cal patch (females) ...... 17 – Upperside of both wings metallic bluish (includ- 2. Upperside of wings non-metallic, deep blue or pur- ing silvery blue, purplish-blue brilliant blue or plish-blue and rather silky. Hindwing lacking tail at deep blue) ...... 12 vein 2, at most only a tooth-like projection...... 3 6. Metallic area on upperside of forewing represent- – Upperside of wings with at least basal metallic ed by basally frosted (powdered) green scales ...7 scaling, generally for the most part some shade of – Metallic area on upperside of forewing not repre- various color (e. g. deep blue, purplish-blue, sil- sented by basally frosted (powdered) scales and very blue, green, golden green, bronze, or cop- evenly occupying at least basal half of wing ...... 8 pery). Hindwing with a tail at vein 2 ...... 5 7. Metallic scales usually sparsely frosted, occasional- 3. Black submarginal bar (E31) of cell 1b+c on un- ly a few greenish ones visible. Orange submargin- derside of forewing very obscure or absent. On al lunules on upperside of hindwing extending up underside of hindwing submarginal border even- to cell 5 (sometimes extending to cell 3). Wing ly pale pinkish, marginal spots (E2) almost absent. small, 12.0-17.0 mm in forewing length. In male Outer margin of hindwing smoothly rounded genitalia, tegumen with a pair of lateral processes and evenly curved. In male genitalia, valva long, greatly reduced (fig. 29); basal process of costa of narrow and ending in small dorsally projecting valva with short apically pointed projection on keel (fig. 75); juxta strongly produced anteriorly posterior margin (fig. 28) ...... eventa ( into a pair of large coeca (fig. 75). In female gen- – Metallic scales usually rather densely frosted. Or- italia, lamella antevaginalis plate-like but slender, ange submarginal lunules on upperside of hind- Y-shaped (fig. 79) ...... yunnani wing restricted to cells 1a, 1b+c and 2, and some- – Black submarginal bar (E31) of cell 1b+c on un- times also slightly distinguishable in cell 3. Wing derside of forewing distinctly present. On under- large, 16.5-19 mm in forewing length. In male side of hindwing submarginal border reddish and genitalia, tegumen with a pair of long lateral suffused with white scales, marginal spots (E2) processes (fig. 19); basal process of costa without rather prominent, especially in cell 2. Outer mar- short apically pointed projection on posterior gin of hindwing dentate at tip of each vein. In margin (fig. 18) ...... tamu (
148 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
8. Orange submarginal lunules on upperside of distal portion; perivesical area of phallus sur- hindwing at least extending up to vein 6 ...... 9 rounded by weak cornutus ...... 15 – Orange submarginal lunules on upperside of 13. Upperside metallic deep blue with broad black hindwing not reaching vein 6 ...... 12 marginal border (summer form), or dim light blue 9. Upperside metallic gold or golden coppery, shift- with narrow black marginal border (spring form). ing in color from fiery red to greenish bronze with Discal lines (M1) of underside almost absent, or direction of light. Upperside of hindwing with represented by a row of fine lines. Submarginal very broad reddish lunules which are noticeably border on underside of hindwing red (summer wider in cells 4 or 5 than elsewhere. In male geni- form) or pinkish (spring form). White lunules on talia tegumen without lateral processes (fig. 22) .. underside of hindwing very large. Wing tail very ...... brahma ( long (summer form) or short (spring form). In – Upperside metallic brassy green, not shifting in male genitalia, juxta very long and almost as long color from fiery red to greenish bronze with di- as valva (fig. 62); costa of valva without long basal rection of light. Upperside of hindwing with process (figs. 61, 62)...... saphir ( rather narrow reddish lunules which are not no- – Upperside metallic blue with slight greenish tint. ticeably wider in cells 4 or 5 than elsewhere. Discal lines (M1) of underside rather prominent Tegumen with short pointed lateral processes and arranged in a conspicuous continuous band (fig. 39) ...... hybrida ( or a row of distinct lines. Submarginal border on 10. Metallic area on upperside of forewing occupying underside of hindwing reddish. White lunules on about basal half, and on upperside of hindwing underside of hindwing rather small. Wing tail confined to posterior half, so hardly extending to rather short and broad. Juxta much shorter than cell 5 and anterior half of discoidal cell ...... valva (figs. 66, 71); costa of valva with long sub- ...... smaragdinus sp. n. ( basal process (figs. 65, 66, 69, 70) ...... 14 – Metallic area on upperside of forewing extending 14. Upperside of forewing with metallic area extend- distinctly beyond basal half, and on upperside of ing hardly beyond end of cell. Orange submar- hindwing extending beyond posterior half, so dis- ginal lunules on upperside of hindwing up to vein tinctly extending up to cell 5 and anterior half of 3. In male genitalia, distal portion of valva nar- discoidal cell ...... 11 row and small (fig. 66) ...... saphirioides ( 11. Upperside metallic golden green. Orange sub- – Upperside of forewing with metallic area extend- marginal lunules on upperside of hindwing re- ing well beyond end of cell. Orange submarginal stricted to cells 1b+c and 2 and almost connected lunules on upperside of hindwing restricted be- with metallic area. Underside discal lines very ob- low vein 3. Distal portion of valva larger and scure (Chou 1994) ...... pulcher (1) broader (fig. 69) ...... brilliantinus ( – Upperside metallic silvery green or metallic green 15. Upperside of both wings metallic blue with pur- with slight blue tint. Orange submarginal lunules plish tint. Upperside of forewing with metallic on upperside of hindwing usually extending to area extending well beyond veins 6, 7+9 and 10. cells 1b+c and 2, and faintly in cell 3 (it may ex- In male genitalia, valva narrow and produced api- tend beyond cell 3 in dry-season form), and dis- cally into small dorsal projection (fig. 45) ...... tinctly separated from metallic area. Underside ...... gloria ( discal lines prominent and dark ...... – Upperside of both wings metallic deep blue (bril- ...... androcles androcles ( liant blue) or silvery blue. Upperside of forewing 12. Hindwing rounded, its inner margin almost as with metallic area extending hardly beyond veins long as costa. In male genitalia, lateral process of 6, 7+9 and 10. Valva rounded, rather oval and not tegumen short, thick and terminating distinctly produced into small dorsal projection ...... 16 anterior to distal portion of socius; inner margin 16. Upperside of both wings metallic silvery blue. In of valva smooth on distal portion; perivesical area male genitalia, dorsal inner margin of valva with- of phallus surrounded by strong cornutus ...... 13 out a short apically pointed projection at middle, – Hindwing elongated, its inner margin much basal process of costa wide, plate-like and not longer than costa. In male genitalia, lateral sharply pointed at apex (fig. 31)...... process of tegumen long, rather slender and ter- ...... androcles androcles ( minating slightly anterior or posterior to distal – Upperside metallic deep blue (brilliant blue) or portion of socius; inner margin of valva serrate on silvery blue. Dorsal inner margin of valva with a short apically pointed projection at middle, basal 1 H. pulcher was treated by Watanabe (1998) as a synonym process of costa slender and sharply pointed at of H. saphir, and the fact may be correct. However, we defer apex (fig. 41) ...... moorei ( this treatment because we did not examine the types. 17. Hindwing lacking tail at vein 2, at most only a
149 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
tooth-like projection ...... 32) – Orange submarginal lunules on upperside of – Hindwing with a tail at vein 2 ...... 18 hindwing extending up to cell 5. White lunules 18. Discal lines (M1) of underside almost absent, very on underside of hindwing appearing in various obscure, or represented by a row of fine lines ...19 degrees. In female genitalia, a pair of posterior – Discal lines (M1) of underside prominent and processes of lamella antevaginalis not inwardly arranged in a conspicuous continuous band or a produced and widely divergent from each other; row of distinct lines ...... 20 ductus seminalis arising from dorsal portion of 19. Orange submarginal lunules on upperside of base of corpus bursae ...... 23 hindwing rather narrow and extending to veins 5 23. In female genitalia, lamella antevaginalis very or 6. Submarginal border on underside of hind- narrow in basal portion, produced posteriorly wing red (summer form) or pinkish (spring form). into a pair of widely divergent lateral lobes (fig. White lunules on underside of hindwing large and 33) ...... eventa & prominent. Wing tail very long (summer form) or – In female genitalia, lamella antevaginalis wide or short (spring form). In female genitalia, lamella rather narrow in basal portion, produced into a antevaginalis produced on apical 3/4 into a pair of pair of widely divergent posterior processes .....24 long, slender, pointed processes with short spines 24. In female genitalia, lamella antevaginalis rather (fig. 77) ...... saphir & narrow in basal portion, produced posterolateral- – Orange submarginal lunules on upperside of ly into a pair of broad processes with rounded hindwing broader and distinctly extending to vein and serrate apices; midventral portion of lamella 7. Submarginal border on underside of hindwing antevaginalis strongly swollen (fig. 35). Lamella red. White lunules on underside of hindwing nar- postvaginalis with a longitudinally oval swelling rower. Wing tail rather short (Chou 1994) ...... (fig. 35) ...... androcles & ...... pulcher & – In female genitalia, lamella antevaginalis wide in 20. Orange submarginal lunules very broad, always basal portion, produced posteriorly into a pair of extending up to vein 7 or more, and noticeably triangular processes with sharply pointed apices; wider in cell 5 than elsewhere (more than 1.5 mm midventral portion of lamella antevaginalis weak- wide in cell 5) ...... brahma & ly swollen. Lamella postvaginalis with a rather – Orange submarginal lunules broad or narrow, ex- transversely oval swelling just posterior to ostium tending to veins 6 or 7 and not wider in cell 5 ...... 25 than elsewhere (less than 1.5 mm wide in cell 5). 25. In female genitalia, lamella antevaginalis very ...... 21 short and wide in basal portion, produced into a 21. Red submarginal border on underside of hind- pair of very long triangular lobes; midventral por- wing broad, about 3.0 mm in width in cell 5. In tion of lamella antevaginalis narrowly swollen female genitalia, base of lamella antevaginalis ex- (fig. 23). Lamella postvaginalis with a weak panded dorsally on both sides and connected swelling just posterior to ostium (figs. 23, 24) .... with oval swelling of lamella postvaginalis (figs...... tamu & 15, 16) ...... smaragdinus & – In female genitalia, lamella antevaginalis rather – Red submarginal border on underside of hind- long and wide on basal half, produced on apical wing narrow, just or less than 1.5 mm in width in half into a pair of triangular processes; midventral cell 5. In female genitalia, base of lamella ante- portion of lamella antevaginalis widely swollen vaginalis expanded laterally and not connected (fig. 46). Lamella postvaginalis with a strong with oval swelling of lamella postvaginalis on swelling just posterior to ostium (figs. 46, 47) ..... both sides ...... 22 ...... moorei & 22. Orange submarginal lunules on upperside of hindwing extending distinctly up to cell 5 and DESCRIPTIONS faintly (sometimes distinctly) to cell 6. White lunules on underside of hindwing always large and Heliophorus (Kulua) smaragdinus prominent. In female genitalia, a pair of posterior Yago & Monastyrskii sp. n. (figs. 1-16) processes of lamella antevaginalis rather conver- gent to each other, so posterior margin circularly Diagnosis concave at middle (fig. 48); ductus seminalis aris- Upperside of male wing metallic emerald green ing from dorsal portion slightly anterior to base of with broad black marginal border; upperside of male corpus bursae. (fig. 49) ...... gloria & hindwing with two narrow, orange submarginal lunules. In male genitalia, costa of valva produced 2Females running to couplet 3 have the same underside wing into wide triangular process at base and bearing long markings as males. pointed projection on its posterior margin.
150 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
Description blackish in cells 1a and 1b+c; outer scales brown, Male. – Head, thorax and abdomen (figs. 1, 2): whitish in cells 1b+c and 2. Similar to congeners; male fore-tarsi fused to single Underside of hindwing yellowish-ochreous in segment ending in rounded point with spines; length ground color as on forewing; M2 (subbasal dots) ap- of antenna 8.6-9.2 mm. pearing in cell 1b+c and discoidal cell as small black Male wing (figs. 1, 2). – Forewing triangular in dot, and in cell 7 as faint blackish bar; D1 discernible shape; costal margin almost straight, weakly curved as very obscure dark bar; M1 appearing as slightly ob- near apex, more or less arched near base; apex weakly scure black line irregularly arched, extending from angulated; outer margin slightly arched on anterior cell 1a to cell 7, shifted inwardly in cell 1a and out- half, almost straight on posterior half; inner margin al- wardly in cell 3, sometimes disappearing in cells 1a to most straight. Hindwing with short, broad tail at tip 1b+c; E3 represented by black-bordered white of vein 2; costal margin almost straight, distinctly lunules, arranged almost parallel to outer margin, shorter than inner margin; outer margin faintly shifted inwardly in cells 4 and 5; width of lunule in arched, slightly produced at apex of each vein but cell 1b+c about 0.2 mm; red submarginal border be- strongly so at tip of vein 6; anal angle developed. tween E2 and E3 rather broad, particularly extended Length of forewing: 16.5-17.0 mm. Length of tail: inwardly in cells 4 and 5, partially suffused with 2.0-2.5 mm. whitish scales, bordered inwardly with series of nar- Upperside of forewing metallic emerald green on row white lunules representing part of E3 and bor- basal half, with broad black outer marginal and nar- dered outwardly with series of blackish E2; width of row costal marginal borders; metallic area extending to red submarginal border (width from inside of E1 to basal 3/4 of cell 1a, basal 4/5 of cell 1b+c, basal 1/2 of inner margin of submarginal border) 1.6-2.4 mm in cell 2, basal 1/3 of cell 3, base of cells 4 and 5, and dis- cell 1a, 1.6-2.0 mm in cell 1b+c, 1.8-2.0 mm in cell coidal cell; fringe consisting of inner short blackish 2, 1.6-2.2 mm in cell 3, 2.4-2.6 mm in cell 4, 2.5-2.8 and outer long brown scales, mixed with many long mm in cell 5, 1.5-1.6 mm in cell 6, 0.6-0.8 mm in white scales at outer margin of cells 1b+c and 2. cell 7; E2 represented by semicircular blackish spot in Upperside of hindwing metallic emerald green as each cell, becoming larger in cell 2 and connected on upperside of forewing, with broad black costal, with white marginal stripes; white marginal stripes outer marginal and inner marginal borders; metallic very slender and separated by red submarginal border area extending to basal 2/3 of cell 2, basal 1/2 of cell 3, extending along each vein; E1 appearing as narrow red basal 1/5 of cell 4 and posterior 2/3 of discoidal cell, line along outsides of marginal white stripes and con- slightly present on anterior portion of cell 1b+c and nected with red submarginal border expanded along base of cell 5; orange submarginal lunules apparent in each vein; fringe as on upperside of hindwing, but in- cells 1b+c and 2, obscure in cell 3, expanded to anal ner short scales becoming reddish. angle in cell 1b+c; width of submarginal lunules 0.6- Male genitalia (figs. 6-13). – Rounded as a whole. 0.9 mm in cell 1b+c and 0.7-1.0 mm in cell 2; fringe Ring slightly inclined posteriorly. Tegumen very consisting of inner short blackish and outer long short, lateral process hardly produced. Socius long, whitish scales, the latter darker at tips of veins and to- about half as long as height of ring, covered with many ward anal angle. longish hairs above, in lateral view slender, weakly Underside of forewing yellowish-ochreous in arched, directed posteroventrally and tapered to apex, ground color; cell 1a and posterior half of cell 1b+c in dorsal view broad basally, divergent from each oth- whitish-gray; D1 (discocellular bar) recognizable as er to apical 2/3, then slightly curved inwardly at apical obscure dark line; M1 (discal line) blackish, covered 1/3, tapered apically and weakly pointed at apex. Vin- with red scales from M11b+c or M12 to M15, and ap- culum narrowed. Saccus short, slightly arched and pearing as slightly arched line extending from cells 0.3ϫ as long as height of ring. Falx slender, extending 1b+c or 2 to 7+9 or 10; E3 (submarginal bars) appear- beyond apex of socius, pointed at apex. Valva large, ing as large elliptical black bar in cell 1b+c, very nar- broad, 0.8ϫ as long as height of ring, in lateral view row in cell 2, obscure in cell 3 and also barely present slightly elliptical, more or less angulated at apex, in in cells 4 and 5, and white-bordered along inside in dorsal view very broad and evenly thick throughout cell 1b+c and faintly in cell 2; red submarginal border length, roundly arched on outer wall, widely concave between E1 (outer marginal line) and E3 present in cell on inner wall, strongly curved inwardly on apical por- 2, weak in cell 3; E2 (inner marginal line or bars) fused tion and sharply pointed at apex; dorsal inner margin with E1, narrowly lined along inside of fringe, repre- of valva produced inwardly into transtilla-like hori- sented by black in cells 1a and 1b+c and then red zontal process at base of costa, deeply concave at mid- from cell 2 to apex; fringe mainly consisting of two dle, middle to apical portions strongly serrate; basal rows of scales, inner short and outer long ones; inner process of costa wide, plate-like, triangular, with long scales reddish and slightly mixed with brownish ones, and sharply pointed projection on posterior portion;
151 Downloaded from Brill.com10/01/2021 08:24:11AM via free access
T E, 145, 2002
6 7
8
9
10
11 12 13
Figs. 6-13. Male genitalia of Heliophorus smaragdinus, scale 0.5 mm. – 6, Dorsum, dorsal view; 7, right valva, dorsal view; 8, genitalia as a whole, lateral view; 9, phallus, lateral view; 10, ditto, dorsal view; 11, juxta, lateral view; 12, ditto, dorsal view; 13, ditto, ventral view.
152 Downloaded from Brill.com10/01/2021 08:24:11AM via free access
Y: Morphology of Heliophorus (Kulua)
14
15 16
Figs. 14-16. Female genitalia of Heliophorus smaragdinus, scales: 1.0 mm (14), 0.5 mm (15, 16). – 14, Genitalia as a whole, ventral view; 15, genitalia except for corpus bursae, lateral view; 16, genital plate, ventral view. ventral inner margin of valva almost straight, slightly weakly produced and rounded; relative lengths of in- serrate on subapical portion, lower apical margin ner margin to costal margin on hindwing slightly strongly produced into narrow triangular process. shorter than in male. Length of forewing: 15.5 mm. Phallus rather short, thick, 1.5ϫ as long as height of Upperside of forewing blackish brown, with large, ring; suprazonal portion slightly shorter than subzon- elliptical, orange discal patch extending from vein 2 al portion, straight, gradually tapered on apical 1/5 to vein 10 and apical portion of discoidal cell, be- and ending in pointed apex; perivesical area on less coming obscure in cell 7+9, 3.0 mm in transverse than apical 1/3 of dorsal surface of suprazonal portion, length, 6.0 mm in longitudinal length; fringe mainly surrounded by weak cornutus; vesica with long, ser- consisting of brown scales, mixed with white scales in rate cornuti; subzonal portion in lateral view evenly cell 1b+c and slightly in cell 2. curved dorsally, coecum penis short. Juxta well devel- Upperside of hindwing blackish brown, with row oped, about 0.4ϫ as long as height of ring, in dorsal of wide orange submarginal lunules from cells 1a to 5, view moderately broad and flattened at base, slightly obscurely in cell 6; some orange scales scattered in- concave on dorsocentral region, produced into pair of wardly in cells 4 and 5; width of orange lunules 0.9 long triangular processes posteriorly; lateral portion mm in cell 1b+c, 1.0 mm in cells 2 and 3, 1.2 mm in slightly produced into flexed projection with which cell 4 and 1.0 mm in cell 5; fringe consisting of short, basal extension of costa of valva is articulated; ventro- black basal scales and long, white apical scales, the lat- median projection of juxta short. Length of male gen- ter much darker at tips of veins and toward tornus. italia: 2.1-2.2 mm (n = 3). Underside of fore- and hindwing almost as in male; Female. – Head, thorax and abdomen (figs. 3, 4): width of red submarginal border (width from inside Very similar to male; apical portion of tarsi bearing pair of E1 to inner margin of submarginal border) on of movable lateral craws; length of antenna 8.0 mm. hindwing 1.8 mm in cell 1a, 1.8 mm in cell 1b+c, 2.0 Female wing (figs. 3, 4). – wing shape similar to mm in cells 2 and 3, 2.8 mm in cell 4, 3.0 mm wide male, but outer margin of forewing evenly arched; in cell 5, 2.0 mm in cell 6, 0.6 mm in cell 7. relative lengths of costal margin to inner margin on Female genitalia (figs. 14-16). – Eighth abdominal forewing shorter than in male, so that apex more tergum large and trapezoidal, narrowed posteriorly.
153 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
17 18
19
Figs. 17-19. Male genitalia of Heliophorus tamu (after Yago et al. 2000), scale 0.5 mm. – 17, Dorsum, dorsal view; 18, right valva, dorsal view; 19, genitalia as a whole, lateral view.
Genital plate almost as long as 8th abdominal tergum. vaginalis; posterior 1/5 of lamella postvaginalis strong- Lamella antevaginalis in ventral view long and rather ly and transversely swollen, with small irregular ventro- wide on basal half, produced posterolaterally into pair median desclerotizations and bearing many shallow of rather long broad processes on apical half; broad transverse wrinkles; length of lamella postvaginalis 0.7 processes weakly and narrowly divergent from each mm. Bursa copulatrix consisting of very short ductus other, roughly serrate marginally, rather rounded api- bursae and elongate membranous corpus bursae. Duc- cally, 1.5ϫ as long as wide; midventral portion widely tus bursae thick, hemi-cylindrical, as long as wide and and strongly swollen ventrally, its base expanded dor- connected with base of projecting coecum bursae at sally on both sides and connected with oval swelling of end, in dorsal view deeply concave or furrow-like. lamella postvaginalis; in lateral view lamella antevagi- Ductus seminalis arising from dorsal portion of base of nalis very broad basally, tapered to apical portion and corpus bursae close to ductus bursae. Coecum bursae truncate at apex, in dorsal view deeply, widely and lon- very short, cylindrical, tapered to apex, sclerotized on gitudinally concave and forming furrow; total length of luminal side and covered with membrane on coelomic lamella antevaginalis about 0.6 mm. Ostium bursae side. Corpus bursae long, directed anteriorly, 5ϫ as opened just posterior to base of lamella antevaginalis long as 8th tergum, arising from dorsal surface of ante- on anterior 1/3 of genital plate. Lamella postvaginalis rior extremity of ductus bursae, evenly slender on prox- weakly sclerotized on anterior 4/5 and strongly sclero- imal 1/2, then rather cylindrically thickened from dis- tized on posterior 1/5, in ventral view with large, tal 1/2, spherically swollen on distal 1/3, of which strong, longitudinally oval swelling present just poste- diameter is 1.5ϫ as long as 8th tergum; signa absent. rior to ostium; oval swelling almost as wide as half of Papilla analis rounded apically, rather strongly sclero- basal width of lamella antevaginalis and tightly pressed tized on basal 3/5, bearing many setae of various against furrow-like dorsal concavity of lamella ante- lengths on apical 2/5; apophysis posterioris slender,
154 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
about as long as 8th tergum. Length of female geni- two as follows: 1) posterior projection of costal process talia: bursa copulatrix (from ostium to distal portion of longer and more sharply pointed at apex, 2) ventral in- corpus) 4.8 mm; ductus bursae + coecum bursae 0.2 ner margin of valva more strongly produced on apical mm; genital plate 1.2 mm. portion into narrower triangular process. Holotype. – 1(, 26.iii.1998, Ngoc Linh Natural This new species is assigned to the subgenus Kulua Reserve (1,600-1,700 m alt.), Kon Tum Province, C. on the basis of wing markings and genitalia, and has Vietnam [A. L. Monastyrskii leg.] (BMNH). six autapomorphies supporting the monophyly of the Paratypes. – 1&, 22.iii.1998 (BMNH), 1(, 22. iii. tamu-group within this subgenus and mentioned in 1998 (BLKU), 1(, 25.iii.1998 (Museum Nationale the introduction of this paper. The male genitalia of d’Histoire Naturelle, Paris), 1(, 27.iii.1998 (Y. this species has a pointed projection on the posterior Takanami collection), 2(, 28.iii.1998 (Vietnamese margin of the costal process of the valva (fig. 7) and Museum of National History), Ngoc Linh Nat. Res. greatly reduced pair of lateral processes of the tegu- (1,600-1,700 m alt.), Kon Tum Prov., C. Vietnam men (fig. 8), which are shared with H. brahma (figs. [A. L. Monastyrskii leg.]. 21, 22) and H. eventa (figs. 28, 29) of the tamu- Etymology. – The specific name, smaragdinus, is group. These character states appear to be apomor- derived from the feature of the wing markings, which phic within the group and consequently these species shine like an emerald. are considered to comprise a monophyletic clade Geographical distribution. – This species is known within the tamu-group. only from Kon Tum Prov., C. Vietnam. Biology. – At Ngoc Linh Natural Reserve, both Descriptions of the genitalia of Kulua-species sexes of this species were observed on the banks of a mountain stream at 1,600-1,700 m alt. within a river- Heliophorus tamu (Kollar, 1844) ine forest (fig. 5). This site is situated in the vicinity of (figs. 17-19, 23, 24) the summit of Ngoc Linh mountain (2,596 m). The stream was 5-10 m wide and surrounded by montane Male genitalia (figs. 17-19, after Yago et al. 2000). evergreen broadleaf forest. Broadleaf trees are domi- – Lateral processes of tegumen triangular, long, rather nant at these altitudes, and include members of the slender and usually extended slightly beyond distal Fagaceae (genera Lithocarpus and Quercus) and the portion of socius. Socii in dorsal view moderately di- Lauraceae. The banks were covered with some species vergent from each other, thick but tapered apically. of herbs, including species of Persicaria (Polygo- Falx in lateral view strongly curved throughout naceae), such as P. perfoliata (L.) H. Gross, P. caespi- length. Saccus short. Valva in lateral view very large, tosa (Blume) Nakai, P. chinensis (L.) Nakai, P. hy- broad, rounded and more or less serrate on posterior dropiper (L.) Spach (Ronse Decraene & Akeroyd margin; dorsal inner margin produced inwardly at 1988). P. chinensis (= Polygonum chinense) is a host base of costa, deeply and narrowly concave at middle, plant of H. epicles, H. ila and H. kiana (Johnston & subapical to apical portions serrate posteriorly, basal Johnston 1980, Morishita 1986, Hamano 1986, process of costa wide, triangular, with weak tooth on Igarashi & Fukuda 1997, Aoyama 1998, Shu 1999, posterior portion and angulated or often minutely Bascombe et al. 1999). In this locality, H. smaragdi- serrate at apex; ventral inner margin widely and rather nus was rather frequent from mid March to early strongly curved and strongly produced at subapical to April. They fly close to the ground among stones or apical portions into serrate triangular process. Phallus in sunlit patches for patrolling and mating. Speci- rather short; suprazonal portion slightly shorter than mens were collected between 9 to 12 a.m. on stones, subzonal portion; dorsal surface of perivesical area warmed by the sun. In some other places, the streams surrounded by weak cornutus. Juxta small, with pair were wider, and this species was observed on small of short triangular lobes produced posteriorly. Length stones in grassland near the waterside. of male genitalia: 2.0-2.3 mm. Remarks. – In general appearance, the male of the Female genitalia (figs. 23, 24). – Lamella antevagi- present new species differs from other species having nalis in ventral view very short and wide in basal por- metallic green wings by the following characters: 1) tion, produced posteriorly into pair of very long tri- metallic green scales on upperside of wings more yel- angular lobes which are strongly serrate on outer lowish (emerald green) and not basally frosted (pow- margins and sharply pointed at apices, midventral dered), 2) metallic green area restricted to about basal portion narrowly and weakly swollen, its base ex- 1/2 on forewing and to basal 2/3 and posterior 1/3 on panded laterally and not connected with oval swelling hindwing, 3) orange submarginal lunules on hindwing of lamella postvaginalis on both sides, in lateral view upperside narrow and confined from cells 1a to 3 (faint rather narrow, tapered to apical portion and pointed in cell 3). In the male genitalia, this species is similar to at apex; length of lamella antevaginalis 0.6-1.0 mm. H. brahma and H. eventa, but differs from the latter Lamella postvaginalis with weak and transversely oval
155 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
20 21
22
igs. 20-22. Male genitalia of Heliophorus brahma, scale 0.5 mm. – 20, Dorsum, dorsal view; 21, right valva, dorsal view; 22, genitalia as a whole, lateral view.
swelling just posterior to ostium; posterior portion of subapical to apical portions produced posteriorly into lamella postvaginalis small, narrowly and strongly many short spines, basal process of costa wide, trian- swollen. Ductus bursae short and small, with very gular, with short pointed projection on posterior short, protuberance-like coecum bursae anteriorly. margin; ventral inner margin slightly curved, strongly Ductus seminalis arising from dorsal portion of base and widely produced into strongly serrated, triangu- of corpus bursae close to ductus bursae. lar process at subapical to apical portions. Phallus and Specimens examined. – 13( 6&, Nepal (BLKU); juxta as in H. tamu. Length of male genitalia: 2.1-2.3 2(, Sikkim (MNHAH); 10( 1&, Kachin, N. Myan- mm. mar (BLKU & MSPC). Female genitalia (figs. 25, 26, modified after Yago et al. 2000). – Lamella antevaginalis in ventral view rather short and very wide on basal half, produced Heliophorus brahma Moore, 1858 posteriorly on apical half into pair of broad short (figs. 20-22, 25, 26) processes which are rounded and serrated marginally, Male genitalia (figs. 20-22). – Lateral process of midventral portion widely and rather strongly tegumen almost absent, only slightly produced. Socii swollen, its base as in H. tamu, in lateral view rather in dorsal view similar to those in H. tamu, but more broad, tapered to apical portion and serrated at apex; slender. Falx and saccus as in H. tamu. Valva in later- length of lamella antevaginalis 0.5-0.7 mm. Lamella al view large, broad, slightly elliptical and rounded on postvaginalis with strong and rather longitudinally posterior margin; dorsal inner margin produced in- oval swelling just posterior to ostium; posterior por- wardly at base of costa, weakly concave at middle, tion of lamella postvaginalis widely and strongly
156 Downloaded from Brill.com10/01/2021 08:24:11AM via free access
Y: Morphology of Heliophorus (Kulua)
23 25
24 26
Figs. 23-26. Female genitalia of Heliophorus (Kulua) species, scale 0.5 mm. – 23, H. tamu, ventral view; 24, ditto, lateral view; 25, H. brahma (Modified after Yago et al. 2000), ventral view; 26, ditto, lateral view.
swollen. Ductus bursae short and small as in H. tamu. inner margin similar to H. brahma, but basal process Ductus seminalis arising as in H. tamu. of costa more angulated and with more strongly Specimens examined. – 1(, Sikkim, India (MN- pointed projection on posterior margin; ventral inner HAH); 4( 1&, Naga Hills, India (MNHAH); 19( 8&, margin as in H. brahma, but subapical to apical por- Kachin, N. Myanmar (BLKU & MSPC); 6( 2&, Xam tions more narrowly produced into triangular process Neua, Laos (BLKU); 9( 1&, Lao Cai, Vietnam with rather weak serration. Phallus and juxta as in H. (KMNH, BLKU & ALMPC)); 1&, Kon Tum, Vietnam tamu. Length of male genitalia: 1.6-2.0 mm. (ALMPC); 3( 3&, Sichuan, China (SCAU & BLKU); Female genitalia (figs. 33, 34). – Lamella antevagi- 3( 2&, Yunnan, China (SCAU & BLKU). nalis in ventral view short and narrow in basal por- tion, produced posteriorly into pair of extremely di- vergent lateral lobes, occasionally bent dorsally on Heliophorus eventa Fruhstorfer, 1918 outer margins and serrated or pointed at apices, mid- (figs. 27-29, 33, 34) ventral portion narrowly and strongly swollen, its Male genitalia (figs. 27-29)–Lateral process of base as in H. tamu but rather weakly expanded later- tegumen almost absent as in H. brahma. Socii in dor- ally, in lateral view very narrow, thin, straight and ser- sal view widely divergent from each other, moderate- rated or pointed at apex; length of lamella antevagi- ly thick, gently tapered apically, convergent at apices. nalis 0.4-0.6 mm. Lamella postvaginalis with rather Falx as in H. brahma, but rather weakly curved. Sac- strong and longitudinally oval swelling just posterior cus as in H. tamu. Valva in lateral view as in H. brah- to ostium; posterior portion of lamella postvaginalis ma, but shorter and more rounded as a whole, dorsal moderately wide and strongly swollen. Ductus bursae
157 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
27 28
29
Figs. 27-29. Male genitalia of Heliophorus eventa, scale 0.5 mm. – 27, Dorsum, dorsal view; 28, right valva, dorsal view; 29, gen- italia as a whole, lateral view. as in H. tamu. Ductus seminalis arising as in H. tamu. at least Aoyama (1998), D’Abrera (1993), Leech Specimens examined. – 18( 15&, Kachin, Myan- (1892-1894), Seitz (1927), Watanabe (1998) and mar (BLKU, MSPC & YWPC); 5( 5&, Shan state, Yago et al. (2000). Thus H. viridipunctata sensu the Myanmar (BLKU); 1&, Thailand (BLKU); 9( 3&, above authors also becomes a synonym of H. eventa. Yunnan, China (KIZC, MNHAH & BLKU); 3( 3&, Sichuan, China (SCAU & MNHAH). Heliophorus androcles (Westwood, 1852) Remarks. – In appearance, this species is very simi- (figs. 30-32, 35, 36) lar to H. tamu, however, the former is easily distin- guishable from the latter on the basis of male genitalia Male genitalia (figs. 30-32). – Lateral process of in having the tegumen lacking a pair of long lateral tegumen strongly developed and terminated slightly processes (figs. 19, 29) and the valva lacking a short anterior or posterior to distal portion of socius. Socii projection on the posterior margin of the basal moderately thick. Falx as in H. tamu. Saccus slightly process (figs. 18, 28). longer than in H. tamu. Valva very similar to H. This species was described by Fruhstorfer (1918) tamu, but in lateral view slightly more elliptical, and as a subspecies of H. viridipunctata (de Nicéville, basal process of costa rather weak, more posteriorly 1890), and was recently raised by Huang (1999) to produced and without weak tooth on posterior mar- specific status. H. viridipunctata has been identified gin; ventral inner margin slightly curved at middle by Riley (1929) as a junior synonym of H. tamu. and very weakly produced into serrated triangular However H. eventa was treated as the former name by process at subapical to apical portions. Phallus and
158 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
30 31
32
Figs. 30-32. Male genitalia of Heliophorus androcles, scale 0.5 mm. – 30, Dorsum, dorsal view; 31, right valva, dorsal view; 32, genitalia as a whole, lateral view.
juxta as in H. tamu. Length of male genitalia: 2.0-2.2 Heliophorus hybrida (Tytler, 1912) mm. (figs. 37-39) Female genitalia (figs. 35, 36). – Lamella antevagi- nalis in ventral view rather short and rather narrow on Male genitalia (figs. 37-39). – Lateral process of basal half, produced posterolaterally on apical half tegumen short, about 1/2 as long as socius. Socii in into pair of broad processes which are rounded and dorsal view as in H. androcles, but more slender in serrated marginally, midventral portion rather nar- apical half. Falx rather slender, more strongly curved rowly and strongly swollen, its base not connected in middle. Saccus slightly longer than in H. androcles. with oval swelling of lamella postvaginalis on both Valva very similar to H. androcles, but basal process of sides as in H. tamu, in lateral view rather narrow, ta- costa more strongly produced posteriorly and round- pered to apical portion and truncated at apex; length ed at apex; dorsal inner margin more narrowly con- of lamella antevaginalis 0.4-0.5 mm. Lamella post- cave on subbasal to middle portions, middle to sub- vaginalis with strong, longitudinally oval, rather small apical portions more roughly and strongly serrated swelling just posterior to ostium; posterior portion of laterally, apical portion minutely serrated posteriorly. lamella postvaginalis moderately wide and strongly Phallus and juxta as in H. tamu. Length of male gen- swollen. Ductus bursae and ductus seminalis as in H. italia: 2.2 mm. tamu. Female. – Unknown. Specimens examined. – 4( 1&, Chin state, Myan- Specimens examined. – 1(, Naga Hills, Assam mar (BLKU); 7( 1&, Kachin, Myanmar (BLKU & (BMNH). YWPC); 3(, Naga Hills, India (MNHAH). Remarks. – In appearance, this species is very simi- lar to H. brahma, however, the former differs from the latter in the male genitalia on the basis of the
159 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
33 34
35 36
Figs. 33-36. Female genitalia of Heliophorus (Kulua) species, scale 0.5 mm. – 33, H. eventa, ventral view; 34, ditto, lateral view; 35, H. androcles, ventral view; 36, ditto, lateral view. tegumen with a pair of short apically pointed lateral subapical to apical portions. Phallus and juxta similar processes (figs. 22, 39) and the valva lacking a short to H. tamu. Length of male genitalia: 1.9-2.3 mm. projection on the posterior margin of the basal Female genitalia (figs. 46, 47). – Lamella antevagi- process (figs. 21, 38). nalis in ventral view rather long and wide on basal half, produced posteriorly or rarely inwardly on apical half into pair of triangular processes which are strong- Heliophorus moorei (Hewitson, 1865) ly or weakly serrated marginally and sharply pointed (figs. 40-42, 46, 47) at apices, midventral portion widely and weakly Male genitalia (figs. 40-42). – Lateral process of swollen, its base as in H. tamu, in lateral view rather tegumen as in H. tamu, but slightly shorter and al- narrow, tapered apically and pointed at apex; length ways extended slightly anterior to distal portion of so- of lamella antevaginalis nearly 0.6-0.8 mm. Lamella cius. Socii in dorsal view moderately thick, more or postvaginalis with strong, transversely oval, large less narrowly divergent from each other. Falx rather swelling just posterior to ostium; posterior portion of weakly curved throughout length. Saccus longer than lamella postvaginalis widely and weakly swollen. H. tamu. Valva in lateral view large, broad, rather el- Ductus bursae and ductus seminalis as in H. tamu. liptical and more or less produced on posterior mar- Specimens examined. – 44(21&, Nepal (BLKU); gin; dorsal inner margin produced posteriorly rather 4(, Sikkim (MNHAH); 7(1&, Bhutan (BLKU). than inwardly at base of costa, deeply concave but Remarks. – H. moorei had been regarded as a sub- with short pointed projection at middle, subapical to species of H. androcles by Riley (1929) and Evans apical portions weakly serrated, basal process of costa (1932). However, it was recently revived as species by slender and pointed at apex; ventral inner margin D’Abrera (1993) and Huang (1999). Huang (1999) weakly curved, and serrated but hardly produced at concluded that moorei consistently differed from H.
160 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
37 38
39
Figs. 37-39. Male genitalia of Heliophorus hybrida, scale 0.5 mm. – 37, dorsum, dorsal view; 38, right valva, dorsal view; 39, genitalia as a whole, lateral view. androcles in the presence of the crest of the valva (a to H. tamu, but suprazonal portion distinctly shorter small projection on dorsal inner margin of valva) in than subzonal portion. Juxta as in H. tamu. Length of the male genitalia. For this reason, I also treated moor- male genitalia: 1.9-2.1 mm. ei as a distinct species. Female genitalia (figs. 48, 49). – Lamella antevagi- nalis in ventral view rather long and wide slightly be- yond basal half, rather inwardly produced before api- Heliophorus gloria Huang, 1999 cal half into pair of short triangular processes which (figs. 43-45, 48, 49) are serrated on outer margins and more narrowly di- Male genitalia (figs. 43-45). – Lateral process of vergent from each other, midventral portion as in H. tegumen as in H. tamu, but slightly shorter, slender moorei, its base as in H. tamu, in lateral view as in H. and always terminated slightly anterior to distal por- moorei; length of lamella antevaginalis 0.5-0.6 mm. tion of socius. Socii in dorsal view moderately thick Lamella postvaginalis with strong, transversely oval and rather narrowly divergent as in H. moori, more swelling just posterior to ostium as in H. moorei; pos- strongly extended posteroventrally. Falx and saccus as terior portion of lamella postvaginalis moderately in H. moorei. Valva in lateral view rather narrow, wide and rather weakly swollen. Ductus bursae rather strongly elliptical and produced on apical portion short, with very short, protuberance-like coecum bur- into small dorsal protuberance; dorsal inner margin sae posteriorly. Ductus seminalis arising from dorsal produced posteriorly at base of costa, weakly and nar- portion slightly anterior to base of corpus bursae. rowly concave at middle, subapical to apical portions Specimens examined. – 19( 7&, Kachin, Myan- rather weakly serrated, basal process of costa long, mar (BLKU & MSPC). rather slender, acutely triangular and pointed at apex; Remarks. – Yago et al. (2000) did not treat H. glo- ventral inner margin almost straight, only slightly ria in their classification. This species should be in- arched and serrated on posterior half. Phallus similar cluded in the tamu-group. The valva of this species is
161 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
40 41
42
43 44
45
162 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
46 48
47 49
Figs. 40-42. Male genitalia of Heliophorus moorei, scale 0.5 mm. – 40, Dorsum, dorsal view; 41, right valva, dorsal view; 42, genitalia as a whole, lateral view.
Figs. 43-45. Male genitalia of Heliophorus gloria, scale 0.5 mm. – 43, Dorsum, dorsal view; 44, right valva, dorsal view; 45, genitalia as a whole, lateral view.
Figs. 46-49. Female genitalia of Heliophorus (Kulua) species, scale 0.5 mm. – 46, H. moorei, ventral view; 47, ditto, lateral view; 48, H. gloria, ventral view; 49, ditto, lateral view. narrower than in the other species of this group. pointed projection, basal process of costa long, very However the structure of the valva is most closely re- slender, style-like and sharply pointed at apex; ventral lated to the tamu-group rather than the yunnani- or inner margin narrowly and rather deeply curved, and the saphir-groups, and this species also shares the oth- strongly produced at apical portion into triangular er five autapomorphies with the tamu-group as men- process bearing minute serrations at apex. Phallus and tioned above. juxta as in H. tamu. Length of male genitalia: 2.0-2.3 mm. Female genitalia (figs. 56, 57). – Lamella antevagi- Heliophorus oda (Hewitson, 1865) nalis in ventral view strongly, narrowly and longitudi- (figs. 50-52, 56, 57) nally swollen on midventral area of basal portion, Male genitalia (figs. 50-52). – Lateral processes of produced posteriorly into very broad, short process tegumen as in H. moorei or H. gloria. Socii in dorsal which is roughly and strongly serrated marginally but view moderately thick, rather narrowly divergent as in weak on mid-posterior margin, its base much more H. moorei. Falx weakly curved throughout length. strongly expanded posterolaterally on both sides; Saccus slightly longer than that of H. tamu. Valva in midventral swollen area of lamella antevaginalis nar- lateral view rather large and broad, elliptical but ex- rowly extended anteriorly and widely bulged posteri- panded on apical portion, and ended as truncate apex orly, anterolateral portions narrowly and weakly con- with small dorsal projection; dorsal inner margin pro- cave as furrow; lamella antevaginalis in lateral view duced rather posteriorly at base of costa, weakly and broad, weakly tapered apically and serrated at apex; widely concave at middle, apical portion with short length of lamella antevaginalis 0.5-0.6 mm. Lamella
163 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
50 51
52
53 54 Figs. 50-52. Male genitalia of Heliophorus oda, scale 0.5 mm. – 50, Dorsum, dorsal view; 51, right valva, dorsal view; 52, genitalia as a whole, lateral view.
Figs. 53-55. Male genitalia of Heliophorus bakeri, scale 0.5 mm. – 53, Dorsum, dorsal 55 view; 54, right valva, dorsal view; 55, genitalia as a whole, lateral view.
Figs. 56-59. Female genitalia of Heliophorus (Kulua) species, scales: 0.5 mm (56, 57); 0.45 mm (58, 59). – 56, H. oda, ventral view; 57, ditto, lateral view; 58, H. bakeri, ventral view; 59, ditto, lateral view.
164 Downloaded from Brill.com10/01/2021 08:24:11AM via free access
Y: Morphology of Heliophorus (Kulua)
56 58
57 59
postvaginalis with strong, longitudinally oval swelling middle, subapical to apical portions smooth, basal just posterior to ostium; posterior portion of lamella process of costa rather long, triangular and pointed at postvaginalis rather widely and strongly swollen. apex; ventral inner margin weakly concave, slightly Ductus bursae rather long, with very short, protuber- serrated and weakly produced at subapical portion. ance-like coecum bursae anteriorly. Ductus seminalis Phallus as in H. tamu. Juxta shorter. Length of male arising as in H. tamu. genitalia: 1.8 mm. Specimens examined. – 6(, Jumla, Nepal (BLKU); Female genitalia (figs. 58, 59). – Lamella antevagi- 2&, Kulu, India (BMNH). nalis in ventral view small and slender on basal 2/3, Remarks. – In appearance, this species is similar to produced posteriorly on apical 1/3 into pair of sharply H. bakeri. However, it differs from the latter in hav- pointed triangular processes, midventral portion nar- ing the tegumen of the male genitalia bearing a pair of rowly swollen, its base not connected with oval long lateral processes, and the apical truncate valva swelling of lamella postvaginalis on both sides as in H. with a small dorsal projection. tamu, invaginated just anterior to both sides into weak concavities of which ventral portions bear pair of small bumps; lamella antevaginalis in lateral view rather Heliophorus bakeri Evans, 1927 broad basally, gently tapered, small and weakly (figs. 53-55, 58, 59) swollen dorsally at middle, pointed at apex; length of Male genitalia (figs. 53-55). – Lateral processes of lamella antevaginalis 0.5 mm. Lamella postvaginalis tegumen almost absent, only slightly produced. Socii with weak, small, oval swelling just posterior to os- as in H. oda, but more convergent on apical 1/3. Falx tium; posterior portion of lamella postvaginalis rather as in H. oda, but more strongly curved. Saccus similar widely and strongly swollen. Ductus bursae rather to H. oda, but in dorsal view more slender. Valva in short, with very short, protuberance-like coecum bur- lateral view rather large and broad, elliptical and sae anteriorly. Ductus seminalis arising from dorsal rounded on posterior margin; dorsal inner margin portion distinctly posterior to base of corpus bursae. produced laterally at base of costa, narrowly concave Specimens examined. – 1( 1&, Kashmir (BMNH). at subbase, produced into large, triangular process at
165 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
60 61
62
63
64 65
66
67
166 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
69
71
71 68
72
Figs. 60-63. Male genitalia of Heliophorus saphir (after Yago et al. 2000), scale 0.5 mm. – 60, Dorsum, dorsal view; 61, right valva, dorsal view; 62, genitalia as a whole, lateral view; 63, perivesical area of phallus, dorsal view.
Figs. 64-67. Male genitalia of Heliophorus saphirioides, scale 0.5 mm. – 64, Dorsum, dorsal view; 65, right valva, dorsal view; 66, genitalia as a whole, lateral view; 67, perivesical area of phallus, dorsal view.
Figs. 68-72. Male genitalia of Heliophorus brilliantinus (after Huang 1999). – 68, Tegumen, vinculum and saccus, lateral view; 69, right valva, lateral view; 70, left valva, dorsal view; 71, juxta, ventral view; 72, phallus, lateral view.
H. saphir (Blanchard, 1871) which are represented by weak invagination on ven- (figs. 60-63, 77, 78) tral inner portion; posterior portion of juxta strongly produced posteriorly into pair of very long triangular Male genitalia (figs. 60-63, after Yago et al. 2000). projections. Length of male genitalia: 2.3-2.5 mm. – Lateral process of tegumen triangular, rather short, Female genitalia (figs. 77, 78, modified after Yago thick and terminated distinctly anterior to distal por- et al. 2000). – Lamella antevaginalis in ventral view tion of socius. Socii in dorsal view narrower on com- cylindrical, rather long and narrow on basal 1/4, pro- posite area of basal portion, very narrowly divergent duced posteriorly on apical 3/4 into pair of long, slen- from each other, rather slender throughout length. der, triangular processes which bear short spines mar- Falx rather weakly curved throughout length, hooked ginally, tapered apically and sharply pointed at apices, at apex. Saccus long. Valva in lateral view elongated, midventral portion very weakly and widely swollen, narrowed, rounded on posterior margin; dorsal inner its base extended inwardly on both sides but not con- margin with small protuberance at base of costa, very nected with oval swelling of lamella postvaginalis, in weakly and broadly concave at middle, subapical to lateral view narrow, slender, tapered apically and apical portions widely swollen; ventral inner margin pointed at apex; length of lamella antevaginalis 0.9- almost straight, lacking projections. Phallus long; 1.1 mm. Lamella postvaginalis with large, posteriorly suprazonal portion distinctly longer than subzonal tapered, longitudinally oval swelling just posterior to portion; dorsal surface of perivesical area surrounded ostium; posterior portion of lamella postvaginalis nar- by strongly sclerotized cornutus with several denti- rowly and strongly swollen. Ductus bursae rather cles. Juxta very long, about as long as valva; anterior long, broad, with very short, protuberance-like co- portion of juxta produced into pair of small coeca ecum bursae anteriorly. Ductus seminalis arising
167 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
73 74
75
76
Figs. 73-76. Male genitalia of Heliophorus yunnani (After Yago et al. 2000), scale 0.5 mm. – 73, Dorsum, dorsal view; 74, right valva, dorsal view; 75, genitalia as a whole, lateral view; 76, perivesical area of phallus, dorsal view.
from dorsal portion slightly posterior to base of cor- pendently in H. yunnani D’Abrera 1993 (fig. 76). pus bursae. Thirdly, the longer transtilla (a basal process of the Specimens examined. – 6( 7&, Sichuan, China costa of the valva) is not observed in H. saphir (figs. (MNHAH & BLKU); 2( 1&, Zhejiang, China (SCAU & 61, 62). Finally, the length of the lateral process of the BLKU); 4(, Shaanxi, China (SCAU & BLKU); 2(, tegumen is variable in each species within Kulua as Hubei, China (SCAU & BLKU). mentioned above. Remarks. – Yago et al. (2000) recognized the As a result, the apparent distal projection of the val- saphir-group (including only H. saphir) as a species va, which is produced posteriorly into a broad and group within Kulua. While Huang (1999), published rounded process without marginal serration, is the nearly simultaneously, considered that H. saphir, H. only character presented by (figs. 61, 62, 65, 66, 69, saphirioides Murayama, 1992 and H. brilliantinus 70) that is synapomorphic for all three included Huang, 1999 should all be assigned to a subgroup be- species. In addition to the valva, the socii, which are cause of the following characters: 1) the more round- narrowed on a composite area of the base and not as ed wing-shape, 2) the fully developed cornutus, 3) the strongly divergent posteriorly, are also considered longer transtilla, 4) the reduced lateral process of the herein as a synapomorphy for the three species (figs. tegumen and 5) the apparent distal projection of the 60, 64). Thus the saphir-group sensu Yago et al. valva. I accept the latter (5), however the remaining (2000) should be modified based on the above new characters are not synapomorphic for the species- definition, and comprises the following three species: group. Firstly, rounded wing-shape is considered ple- H. saphir, H. saphirioides and H. brilliantinus. siomorphic in the subgenus Kulua (Yago et al. 2000). Incidentally, the developed E3 markings on the un- Secondly, the fully developed cornutus is possibly derside of the hindwing and the juxta with extraordi- apomorphic, but it is also observed perhaps inde- narily elongate paired processes, which were autapo-
168 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
77 79
78 80
Figs. 77-80. Female genitalia of Heliophorus (Kulua) species, scale 0.5 mm. – 77, H. saphir (Modified after Yago et al. 2000), ventral view; 78, ditto, lateral view; 79, H. yunnani (Modified after Yago et al. 2000), ventral view; 80, ditto, lateral view.
morphies of the saphir-group (Yago et al. 2000, fig. curved, lacking projections. Phallus and cornutus as 62), are not observed in H. saphirioides (Murayama in H. saphir. Juxta similar to H. saphir, but shorter 1992, fig. 66) and H. brilliantinus (Huang 1999, fig. and less than half of valva. Length of male genitalia: 71). Hence these two character states are considered 2.2-2.4 mm. as apomorphic for only H. saphir. Female. – Unknown. Specimens examined. – 2(, Yunnan, China (KNHM & MNHAH). Heliophorus saphirioides Murayama, 1992 Remarks. – In the male genitalia, this species is very (figs. 64-67) similar to H. brilliantinus (figs. 68-72, after Huang Male genitalia (figs. 64-67). – Tegumen, socii and 1999). It is said that the distal portion of the valva is saccus as in H. saphir. Falx similar to H. saphir, but much narrower and smaller than in the latter species. more strongly curved on distal portion. Valva in lat- However, I have examined one male identical to H. eral view broad but rather narrowed toward distal brilliantinus in general appearance, of which the valva portion and rounded on posterior margin; dorsal in- shows the same structure as that of H. saphirioides. It ner margin produced posteriorly at base of costa, is possible that the valva varies in its distal shape weakly and rather broadly concave at middle, subapi- among individuals or changes its shape with direction cal to apical portions widely swollen, basal process of of observation. Hence H. brilliantinus may be a sea- costa very long, slender, style-like and pointed at sonal form or individual variation of H. saphirioides. apex; ventral inner margin widely and very weakly
169 Downloaded from Brill.com10/01/2021 08:24:11AM via free access T E, 145, 2002
H. yunnani D’Abrera, 1993 na of Ngoc Linh Nature Reserve. I wish to express my (figs. 73-76, 79, 80) sincere thanks to Dr. A. L. Monastyrskii (Vietnam Russian Tropical Center), who is the junior author of Male genitalia (figs. 73-76, after Yago et al. 2000). the new species, for the loan of many materials and giv- – Lateral process of tegumen triangular, short, pos- ing the opportunity to describe the new species togeth- teroventrally extended slightly beyond middle of so- er with him. I express my cordial thanks to Prof. H. cius. Socii in dorsal view long and slender throughout Shima, Prof. O. Yata and Assoc. Prof. K. Araya (BLKU), length, widely divergent from each other on basal 1/3, and Assoc. Prof. K. Ogata (Institute of Tropical Agri- then gently convergent on apical 2/3. Falx evenly slen- culture, Kyushu University), for their kind guidance. I der, long and weakly curved throughout length. Sac- am much indebted to Dr. B. J. Sinclair (Zoologisches cus long. Valva in lateral view rather long, narrow, ta- Forschungsinstitut und Museum Alexander Koenig) pered from apical 3/4 to apical portion and ending in and Mr. A. Tordoff (BIVP) for his critical reading of this small dorsally projected keel; dorsal inner margin al- manuscript. Special thanks go to Dr. T. Saigusa most straight, strongly curved inwardly at apex, base (Fukuoka) for his constant guidance and Dr. A. of costa with slightly swollen flange bordered by inter- Sibatani (Kyoto) for the loan of his extensive collection nal ridge on outer surface of dorsal margin, lacking of lycaenine butterflies which is donated to MNHAH. special projection; ventral inner margin straight, Thanks are also due to Prof. A. Nakanishi (MNHAH) for minutely serrated throughout. Phallus long; suprazon- the loan of the Sibatani’s donative collection. I am al portion distinctly longer than subzonal portion; deeply grateful to Mr. K. Hatoyama (Tokyo) for his dorsal surface of perivesical area bearing short, nar- support. I am also much indebted to Dr. Y. Hirashima rowly sclerotized cornutus with several denticles near (Fukuoka) and Assist. Prof. J. Yokoyama (Department vesical opening. Juxta long, about as long as valva and of Ecology and Evolutionary Biology, Biological Insti- more flattened dorsally; anterior portion of juxta tute, Graduate School of Science, Tohoku University) strongly developed, forming pair of large coeca repre- for their advice on scientific names of some butterflies sented by broad and deep invagination on ventral in- or plants. I also wish to express our sincere thanks to ner portion; posterior portion of juxta produced pos- Dr. P. R. Ackery (BMNH), Dr. W. Ji and Dr. J. Xiong teriorly into pair of long triangular projections. (KIZC), Assoc. Prof. M. Wang (Laboratory of Insect Length of male genitalia: 2.0 mm. Ecology, SCAU), Prof. D. Yang (Department of Ento- Female genitalia (figs. 79, 80, modified after Yago mology, China Agricultural University), Dr. K. Ueda et al. 2000). – Lamella antevaginalis rather slender and (KNHM), Prof. T. Fujioka (Research Institute of Sci- Y-shaped, in ventral view long and plate-like beyond ence and Technology, Tokai University), Mr. H. basal half, produced before apical half into pair of Mikami (Ishikawa Insect Museum), Mr. R. Matsumo- slender, weakly tapered processes, midventral portion to (Osaka Museum of Natural History), Mr. Y. almost flattened, its base extended outwardly on both Okushima (KMNH), Mr. K. Shibahara (Nagasaki), Mr. sides and connected with pair of funnel-like concavi- K. Sasaki (Fukuoka), Mr. Y. Watanabe (Osaka), Mr. ties present on lateral portions of oval swelling of T. Hane (Shiga), Mr. K. Morishita (Kanagawa), Mr. Y. lamella postvaginalis; lamella antevaginalis in lateral Takanami, Mr. H. Kanmuri, Mr. T. Miyashita, Mr. Y. view very narrow, slender, weakly tapered apically; Ohno, Mr. M. Saito and Mr. H. Yui (Tokyo), and Mr. length of lamella antevaginalis 0.8 mm. Lamella post- H. Ôkido and Mr. K. Odagiri (BLKU), for valuable in- vaginalis with small, rather longitudinally oval formation or materials to compare with the new swelling just posterior to ostium; posterior portion of species. This study, contribution No. 85 from the lamella postvaginalis cordate, almost flattened, only BLKU, was supported by the Research Fellowships of weakly swollen. Ductus bursae rather long and slen- the Japan Society for the Promotion of Science for der, with short coecum bursae anteriorly. Ductus sem- Young Scientists (No. 13010131). inalis arising from dorsal portion of base of corpus bursae close to ductus bursae. REFERENCES Specimens examined. – 6( 3&, Yunnan, China (KIZC & MNHAH). Anonymous, 1994. Biodiversity action plan for Vietnam. – Government of the Socialist Republic of Vietnam and the Global Environment Facility Project VIE/91/G31, ACKNOWLEDGEMENTS Hanoi, 212 pp. Aoyama, J., 1998. Butterflies of China. – Tokai University First of all I would like to express my great thanks to Press, Tokyo, xviii+392 pp. Mr. J.C. Eames, Programme Manager of the BirdLife Bascombe, M. J., Johnston, G. & Bascombe, F. S., 1999. International Vietnam Programme (BIVP), and to the The butterflies of Hong Kong. – Academic Press, Lon- administration of the Forest Inventory and Planning don, xiii+422 pp. Institute for the opportunity to study the butterfly fau- Chou, I., 1994. Monographia Rhopalocerorum Sinensium,
170 Downloaded from Brill.com10/01/2021 08:24:11AM via free access Y: Morphology of Heliophorus (Kulua)
Monograph of Chinese Butterflies. – Henan Scientific Morishita, K., 1986. The largest copper in the world. – The and Technological Publishing House, Cheng-chou, Rhopalocerist’s Magazine Cyô-Cyô 98: 2-14. [in Japanese] 18+854 pp. [In Chinese]. Riley, N. D., 1929. Revisional notes on the genus Heliopho- Collins, N. M., Sayer, J. A. & Whitmore, T. C., 1991. The rus (Lycaenidae) with description of new forms. – Journal conservation atlas of tropical forests Asia and the Pacific. of the Bombay Natural History Society 33: 383-402, 28 – Simon & Schuster, New York, 256 pp. figs. D’Abrera, B., 1986. Butterflies of the Oriental Region, Part Ronse Decraene, L.-P. & Akeroyd, J. R., 1988. Generic lim- III Lycaenidae & Riodinidae. – Hill House, Victoria, pp. its in Polygonum and related genera (Polygonaceae) on the i-xvi+ 535-672. basis of floral characters. – Botanical Journal of the Lin- D’Abrera, B., 1993. Butterflies of the Holarctic Region, Part nean Society 98: 321-371, 31 figs. III Nymphalidae (concl.), Libytheidae, Riodinidae & Ly- Schwanwitsch, B. N., 1949. Evolution of the wing-pattern caenidae. – Hill House, Victoria, pp. i-viii+ 333-524. in the lycaenid Lepidoptera. – Proceedings of the Zoo- Eliot, J. N., 1963. The Heliophorus epicles (Godart, 1823) logical Society of London 119: 189-263. complex. – The Entomologist 96: 169-180. Seitz, A., 1927. (ed.) Gross-Schmetterlinge der Erde, 9. Die Evans, W. H, 1932. The identification of Indian butterflies, Indo-Australische Tagfalter. – Alfred Kernen, Stuttgart. 2nd edition, revised. – The Bombay Natural History So- pp. 1-1196 (text); 1-175 (pls: separately bound). ciety, Madras, 454 pp, 32 pls. Shirôzu, T., 1960. Butterflies of Formosa in colour. – Fruhstorfer, H., 1912. Uebersicht der Lycaeniden des Indo- Hoikusha, Osaka, 481 pp, 479 figs., 76 pls. [In Japanese]. Australischen Gebiets. – Berliner Entomologische Shirôzu, T. & Yamamoto, H., 1956. A generic revision and Zeitschrift 56: 197-272. the phylogeny of the tribe Theclini (Lepidoptera; Ly- Fruhstorfer, H., 1918. Revision der Lycaenidengruppe He- caenidae). – Sieboldia 1: 329-421. liophorus auf Grund der Morphologie de Klammeror- Shu, Y., 1999. Butterflies of Taiwan. 1. – Feng Huang Ku gane. – Tijdschrift voor Entomologie 61: 45-53. Bird Park, Taiwan, 344 pp. [In Chinese]. Hamano, E., 1986. Ecological encyclopedia of Taiwanese Watanabe, Y., 1998. Butterflies of China. – Tombow Pub- Butterflies. – Kodansha Ltd., Tokyo, 467 pp. lishing, Osaka, 255 pp. Huang, H., 1999. Some new butterflies from China-1 Yago, M., Saigusa, T. & Nakanishi, A., 2000. Rediscovery of (Rhopalocera). – Lambillionea 99: 642-676. Heliophorus yunnani D’Abrera and its systematic position Igarashi, S. & Fukuda, H., 1997. The life histories of Asian with intrageneric relationship in the genus Heliophorus butterflies, Vol. I. – Tokai University Press, Tokyo, 317 (Lepidoptera: Lycaenidae). – Entomological Science 3 (1): pls., pp. i-xix, 234-549. 81-100. Johnston, G. & Johnston, B., 1980. This is Hong Kong: Zhdanko, A. B., 1995. On the systematics of the genera Ly- Butterflies. – Government Information Series, Hong caena F. and Heliophorus Geyer (Lepidoptera: Ly- Kong, 244 pp. caenidae). – Entomologicheskoye Obozreniye 74 (3): Leech, J. H., 1892-1894. Butterflies from China, Japan and 652-661. [in Russian] Korea. – Private Publication, London, lviii+681 pp, 43 pls. Metaye, R., 1957. Contribution à l’étude des Lépidoptères du Viêtnam (Rhopalocera). – Annales de la Faculté des Sciences, Saigon 1957: 69-106. Murayama, S., 1992. Some new lycaenid species of Chinese Rhopalocera. – The Nature and Insects 27 (5): 37-41. [in Received: 22 February 2002 Japanese] Accepted: 24 April 2002
171 Downloaded from Brill.com10/01/2021 08:24:11AM via free access