Bull. Natl. Mus. Nat. Sci., Ser. A, 35(4), pp. 249–278, December 22, 2009The Species-group of Arge captiva (Insecta, Hymenoptera, Argidae)
Akihiko Shinohara1, Hideho Hara2 and Jung-Wook Kim3
1

Department of Zoology, National Museum of Nature and Science,
3–23–1 Hyakunin-cho, Shinjuku-ku, Tokyo, 169–0073 Japan
E-mail: shinohar@kahaku.go.jp
2
Hokkaido Forestry Research Institute, Koshunai, Bibai-shi,
Hokkaido, 079–0198 Japan
E-mail: hara.hideho@pref.hokkaido.lg.jp
3
Public Health Pest Management, DENR, Raleigh NC, USA
E-mail: jung.kim@ncdenr.gov

Abstract Arge captiva (Smith, 1874) and its allied taxa are revised and four species are recognized. All relevant type material was studied, except for that of Arge praesternalis Malaise, 1934.
Arge macrops sp. n. (Arge sanguinolenta auctt., nec Mocsáry, 1909) is described and illustrated
from the Primorskij Kraj, Korea and China. The following synonymy is established or confirmedfor Arge captiva: Arge captiva (Smith, 1874) (Hylotoma captiva Smith, 1874Hylotoma flavicollis Cameron, 1876 [syn. n.]Arge sanguinolenta Mocsáry, 1909 [syn. n.]Arge kolthoffi Forsius, 1927 [syn. n.]Arge captiva rufoscutellata Takeuchi, 1927). Arge siamana Togashi, 1988, isremoved from synonymy with A. captiva. A little known species, A. praesternalis, is treated as avalid species.
Key words : Argidae, Arge captiva group, Arge macrops, new species, new synonymy.

The sawfly genus Arge (Hymenoptera, Argidae) is represented by more than 300 described
species in the world, including about 120 species
recorded from eastern Asia (Gussakovskij, 1935;
Takeuchi, 1939; Smith, 1989, 1992; Wei et al.,2006). Among the eastern Asian representativesof the genus, there are two groups of species
whose adults are black or bluish black with reddish areas on the thorax. The species of the twogroups are similar in color pattern but greatly
different in structure. One of the groups, herein
named the rejecta group, includes A. rejecta(Kirby, 1882) and its allies. The members of therejecta group are comparatively small species
characterized by the medially distinctly carinate
supraclypeal area on the head and the pilose
outer margins of the wings. The other group,
herein named the captiva group, is represented
by A. captiva (Smith, 1874) and its allies, which
are comparatively large and robust species distinguished from the members of the rejecta groupby the roundly swollen supraclypeal area without

a median carina and the nearly glabrous outermargins of the wings. The larvae of A. rejecta are
known to feed on Rubus spp., whereas thoseof A. captiva are associated with Ulmus spp.
(Okutani, 1967).
A total of nine previously described taxa may
belong to the two groups: A. captiva (Smith,
1874), A. flavicollis (Cameron, 1876), A. rejecta(Kirby, 1882), A. vulnerata Mocsáry, 1909, A.
sanguinolenta Mocsáry, 1909, A. kolthoffi Forsius, 1927, A. captiva rufoscutellata Takeuchi,
1927, A. praesternalis Malaise, 1934, and A. siamana Togashi, 1988. Arge captiva var. watanabei
Takeuchi, 1932, is regarded as an infrasubspecific unavailable name (Blank et al., 2009) and thus
will not be discussed further in the present paper.
Taxonomy of these species is in need of revision,
because recent works differ in the interpretationof these taxa, particularly as to the placement of
A. flavicollis and A. siamana (Wei and Wen,
2000; Wei and Nie, 2003; Saini et al., 2006). Wei
and Wen (2000) synonymized A. flavicollis with

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Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

A. captiva, thus placing the species in the captiva
group, whereas Saini et al. (2006) treated them“as two separate species which are similar incolor but differ in morphology”, thus suggesting
the placement of A. flavicollis in the rejecta
group. Wei and Wen (2000) also placed A. siamana in synonymy with A. captiva, but Saini etal. (2006) mentioned, “this placement requires
closer scrutiny”. It should be noted that theseworks were not based on the examination of thetype material, and this is doubtless the main
source of confusion.
We were able to examine the type material of
all the taxa stated above, with the exception of A.
praesternalis. In the following lines, we will revise the taxa belonging to the captiva groupbased on the examination of the type material
and a series of specimens from various parts ofeastern Asia. Besides A. rejecta and A. vulnerata,
all the taxa cited above belong to the captiva
group. In conclusion, we have recognized four
species in the captiva group, namely A. captiva,
A. siamana, A. praesternalis and A. macrops sp.
n. Arge flavicollis, A. sanguinolenta, A. kolthoffi
and A. captiva rufoscutellata are synonymouswith A. captiva, which is a widespread species
showing large variations in the color pattern of
the thorax. The color variations are generally associated with geography but not clear-cut enoughto recognize subspecies in our view. Judging
from the original description, A. praesternalis
may also be a synonym of A. captiva, but wetreat it as a valid species until more information
becomes available. Arge siamana is very similar
to A. captiva in external morphology but distinguishable from it in the shape of the ovipositor. A
peculiar species characterized by the exceptionally large eyes and ocelli was previously identified
with A. sanguinolenta, but this is a misidentification and the species will be described under thename of A. macrops below.
The material used in this work is kept in thefollowing collections: BMNH – The Natural History Museum, London; CAS – California Academy of Sciences, San Francisco; CSFU – Central
South Forestry University, Changsha; HFRI –Hokkaido Forestry Research Institute, Bibai;
HNHMB – Hungarian Museum of Natural History, Budapest; HSC – H. Suda collection, Sakura;
HSLC – H.-S. Lee collection, Suwon; HU –Hokkaido University, Sapporo; HYC – H. Yoshida collection, Kobe; KU – Kobe University,
Kobe; MCZ – Museum of Comparative Zoology,
Cambridge; MNHAH – Museum of Nature andHuman Activities, Hyogo, Sanda; NRMS –Swedish Museum of Natural History, Stockholm;
NSMT – National Museum of Nature and Science, Tokyo; OMNH – Osaka Museum of Natural
History, Osaka; OPU – Osaka Prefecture University, Sakai; SDEI – Senckenberg Deutsches Entomologisches Institut, Muncheberg; SNM – Natural History Museum of Denmark, Copenhagen;
SNU – Seoul National University, Suwon (partly
on permanent loan from Kangwon National University, Chuncheon, and Gyeongsang National
University, Jinju); USNM – National Museum of
Natural History, Washington, D.C.; ZSSM –Bavarian State Collection of Zoology, Munich.
For the scientific names of the host plants, wefollowed Yonekura and Kajita (2009).
The Arge captiva GroupThe species of the captiva group are characterized as follows: Rather large and robust species(Fig. 1), length 7–12.5 mm in females and 7–11
mm in males. Black, with bluish or purplish reflection; thorax usually with large reddish areas.
Interantennal carinae (Fig. 2A, E, J) distinct,
nearly straight, subparallel or convergent below
between level of dorsal margins of antennal sockets and level of middle inner orbits, and then becoming indistinct and strongly convergent ventrally on supraclypeal area; supraclypeal area
roundly swollen, top surface nearly flattened,
without median carina, but with very shallow, inconspicuous longitudinal depression or groove
along median line enclosed by ventral parts ofusually inconspicuous V-shaped interantennal
carinae. Right mandible with inner margin distinctly notched. Cell 2Rs in forewing (Fig. 2P–R)very broad (apical length [crossvein 3r–m

Arge captiva Species-group

length] about 0.6–0.9 posterior width), not or
weakly widened towards apex, with anterior
width about as long as or shorter than posterior
width (0.7–1.0 : 1.0); crossvein 2r–m usually bent
towards base or sinuate and crossvein 3r–m nearly straight or weakly curved outwards; in bothwings, wing margin between veins Rs and Cu
nearly glabrous, setae not protruding beyond
wing margin (Fig. 2S). Sawsheath rather short,rounded, with median lobe very small and located at base in posterodorsal view (Fig. 2D, H, M),with large swelling at base in lateral view (Fig.
2I). Lance (Fig. 4) with apical crest well developed, roughly serrate and with narrow membranous areas at middle and with groups of rudimental setae at intervals along ventral margin at
middle. Lancet (Figs. 5–7) in lateral view rather
thick, with rounded apex and about 22–27 serrulae as recognized by number of marginal sensilla;
middle annuli substraight; each serrula roundedor subtriangular in outline; gap between 2nd and3rd serrulae not much larger than gaps between
other serrulae. Male genitalia (Figs. 8–10):
Gonostipes in ventral view narrowed apicallywith apical width about as long as or longer thanbasal width of harpe and with medial margin
posteriorly subtruncate or roundly convex. Harpe
gradually narrowed toward apex. Penis valve
with valviceps rather simple, narrow subtriangular or oval with elongate apex in outline in lateralview, swollen laterally at middle in dorsal view,
and with lateral carina at middle running dorsallyfrom ventral margin in lateral view.
This species-group is possibly a part of thepectoralis group defined by Smith (1989), who
included A. captiva in the pectoralis group. The

251

members of the captiva group agree with the diagnosis of the pectoralis group, except that the
gap between the 2nd and 3rd serrulae of thelancet is not larger than between others.
Extremely dark specimens of A. captiva, having no reddish areas on the thorax, look similar
to black or bluish black species with the swollen
and non-carinate supraclypeal area and theglabrous outer margins of the wings, such as A.
pullata (Zaddach, 1859) or A. enodis (Linnaeus,
1767). These two species will be distinguishedfrom A. captiva by the roundly swollen supraclypeal area without very shallow longitudinalgroove or depression at middle, the narrow andusually apically widened cell 2Rs in the forewing(ratio of apical length to posterior width is
0.6–0.9 in A. captiva group, whereas it is 0.9–1.4
in A. pullata, 0.9–1.1 (female) and 0.8–0.9(male) in A. enodis), elongate sawsheath and thevery different shape of the penis valve.
The species of the rejecta group (A. rejecta
and A. vulnerata) have a very similar color pattern to that of the captiva group, but differ in a
number of morphological features including themedially sharply carinate supraclypeal area, thedensely pilose outer margins of the wings, rather
narrow and apically widened cell 2Rs in theforewing, and the slender, apically strongly narrowed lancet. Two western Palearctic species, A.
thoracica (Spinola, 1808) and A. pleuritica(Klug, 1834), also have a similar color pattern.
These two species have much in common withthe rejecta group, sharing the same set of characters given above, though the median carina of thesupraclypeal area is rather blunt.

Key to species (A. praesternalis not included)
1. Eyes and ocelli exceptionally large and malar space very narrow (Fig. 2A–C, N); lancet with longitudinal row of minute setae at base of 1st annulus rudimentary or missing (Fig. 5); male genitalia
as in Figs. 8A–C, 9A–I. [Upper head with bluish reflection, purplish only in posterior part; interantennal carinae broadly separated and subparallel; supraclypeal area with side slopes roundly
swollen; anterior margin of clypeus broadly, rather shallowly emarginated, lateral lobes subtriangular (Fig. 2B). Lancet with about 25 serrulae as recognized by number of marginal sensilla (Fig. 5);
distance between 1st and 15th serrulae about 3.1–3.4 distance between 15th and terminal serrulae; 2nd annulus sclerotized only ventrally.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. macrops, /?

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Akihiko Shinohara, Hideho Hara and Jung-Wook Kim— Eyes, ocelli and malar space normal (Fig. 2E–G, J–L, O); lancet with longitudinal rows of minutesetae at base of first annulus present (Figs. 6–7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Upper head with purplish reflection (Fig. 2G); interantennal carinae rather narrowly separated anddistinctly convergent below (Fig. 2E); supraclypeal area with side slopes very weakly rounded; anterior margin of clypeus rather narrowly and roundly incised at middle, lateral lobes nearly truncate at apex (Fig. 2F); lancet with about 27 serrulae as recognized by number of marginal sensilla(Fig. 6A); distance between 1st and 15th serrulae about 2.5 distance between 15th and terminal
serrulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. siamana, /— Upper head with bluish, not purplish, reflection (Fig. 2L); interantennal carinae broadly separated
and subparallel or very slightly convergent below (Fig. 2J); supraclypeal area with side slopes
roundly swollen; anterior margin of clypeus broadly, rather shallowly emarginated, lateral lobes
subtriangular (Fig. 2K); lancet with about 22 serrulae as recognized by number of marginal sensilla (Figs. 6B–D, 7); distance between 1st and 15th serrulae about 3.4–6.1 distance between 15th
and terminal serrulae. [Male ganitalia as in Figs. 8D–F, 10.] . . . . . . . . . . . . . . . . . . . A. captiva, /?
Arge macrops sp. n.
(Figs. 1A–D, 2A–D, N, P, S, 3, 4A–C, 5, 8A–C, 9A–J)
Arge sanguinolenta: Kuznetzov-Ugamskij, 1927: 235;
Malaise, 1931: 152; Gussakovskij, 1935: 233, 266,
403; Zhelochovtsev and Zinovjev, 1995: 400 (misspelled as sanguinicollis); Lelej and Taeger, 2007: 943.
Nec Mocsáry, 1909.

Female (Fig. 1A–B). Length about 11–12.5
mm. Head, metathorax, legs, and abdomen entirely black with distinct bluish reflection, reflection on posterior upper and lateral parts of headpurplish; antenna black, without distinct reflection; mouth parts blackish brown to black; prothorax and mesothorax orange, with most or all
cervical sclerite, posterior part of mesepimeron,and often ventral half of mesopleuron bluishblack; wings blackish infuscated; veins and stigma blackish.
Surface generally smooth and shining; punctures on anterior part of head fine and distinct,
partly irregular but generally separated from eachother. Head and thorax covered with rather shorthairs, those on upper and lateral parts of headand on ventral part of thorax sparse; hairs generally blackish on dark areas and colorless on paleareas.
Head with exceptionally large eyes and ocelli;in dorsal view (Fig. 2C), posterior part behind
eyes very short, scarcely dilated; in lateral view,
gena very narrow. Distance between eyes 1.0

vertical diameter of eye; eye with vertical diameter 1.5–1.6 horizontal diameter. Postocellar
area scarcely convex, anterior and lateral furrows
indistinct (Fig. 2C). Ocellar area concave between ocelli. Frontal area concave in front of anterior ocellus; median fovea in large deep pit; interantennal carinae low and blunt but distinct,
subparallel, with very shallow depression withflattened bottom between them; just below levelof ventral margin of antennal sockets, carinae becoming indistinct and convergent ventrally toform inconspicuous V-shaped platform, oftenwith indistinct longitudinal depression along
midline (Fig. 2A); supraclypeal area with sideslopes roundly swollen. Malar space exceptionally short, only 0.1 width of front ocellus.
Clypeus flattened, with anterior margin broadly,rather shallowly emarginated, lateral lobes subtriangular (Fig. 2B). Antennal length 1.4–1.7
maximum width of head; flagellum hardly compressed, weakly curved basally and midapicallyand rather strongly thickened midapically (Fig.
1A–B).
In forewing, cell 2Rs with anterior width 0.8–
1.0 posterior width, and crossvein 3r-m weakly
curved outwards; in both wings, wing margin between veins Rs and Cu nearly glabrous, setae notprotruding beyond wing margin (Fig. 2S).
Abdominal terga glabrous, except for distinctly pilose lateral parts; seventh and more posterior

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253

Fig. 1. Arge macrops (A–D), A. siamana (E–F) and A. captiva (G–M), type material. — A, Paratype of A.
macrops n. sp., female, Mt. Taibaishan, China, dorsal view; B, do., ventral view; C, holotype of A. macrops n.
sp., male, dorsal view; D, do., ventral view; E, holotype of A. siamana Togashi, 1988, female, dorsal view; F,
do., ventral view; G, lectotype of Hylotoma captiva Smith, 1874, female, dorsal view; H, lectotype of Hylotoma flavicollis Cameron, 1876, female, dorsal view; I, do., ventrolateral view; J, syntype of Arge sanguinolenta Mocsáry, 1909, female, dorsal view; K, do., ventral view; L, holotype of Arge kolthoffi Forsius, 1927,
female, dorsal view; M, syntype of Arge captiva rufoscutellata Takeuchi, 1927, female, dorsal view.

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Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

Fig. 2. Arge macrops, /, paratype, Mt. Taibaishan, China (A–D, P, S), A. siamana, /, holotype (E–I, Q), A. captiva, /, Sapporo, Japan (J–L, R), A. captiva, /, Jirisan, Korea (M), A. macrops, ?, holotype (N), and A. captiva, ?, Kawasaki, Japan (O). — A, E, J, Head laterofrontal view; B, F, K, clypeus, frontal view; C, G, L,
head dorsal view; D, H, M, sawsheath, posterodorsal view; I, sawsheath, lateral view; N-O, head, frontal
view; P–R, cell 2Rs of left forewing; S, margin of forewing with vein M.

terga with sparse minute setae. Seventh sternum
with posterior margin roundly produced at middle. Sawsheath in posterodorsal view (Fig. 2D)

short subtriangular in outline, with lateral margin
nearly straight or rounded, apex very narrowly
rounded, and medially nearly truncate; dorsome-Arge captiva Species-group

Fig. 3.

255

Distribution of A. macrops based on specimens examined.

dial ridge blunt and dorsal surface weakly convex; in lateral view, ventral margin, except for
conspicuous basal convexity, roundly convex,
dorsal margin nearly straight, and apex rounded;
inner surface spinose.
Lance with several narrow membranous areasat middle (Fig. 4A–C). Lancet (Fig. 5) with about
25 serrulae as recognized by number of marginal
sensilla; distance between 1st and 15th serrulae
about 3.1–3.4 distance between 15th and terminal serrulae; dorsal margin glabrous on 1st annulus, with group of sparse setae on each of 2nd to
6th annuli, and continually setose on 7th andmore apical annuli; narrow non-annulate area
present dorsoapically; transverse rows of minutesetae present between annular plates; marginal
sensilla long; basal annular plates dorsally rough;1st annulus with small sclerotized plate ventrally
but transverse row of minute setae rudimentaryor missing; 2nd annulus sclerotized in ventral
half; serrulae roundly and very strongly convex,
rather narrow and regular in shape; each serrula
in median part of lancet with basal slope slightly
shorter than apical slope.
Male (Fig. 1C–D). Length 9–11 mm. Head,metathorax, legs, and abdomen entirely blackwith distinct bluish reflection, reflection on posterior upper and lateral parts of head often purplish; antenna black, without distinct reflection;mouth parts blackish brown to black; prothorax
and mesothorax orange, with most or all cervicalsclerite, sunken area of mesoscutal lateral lobe,
part of mesoscutellum, posterior part ofmesepimeron, and ventral half of mesopleuron
bluish black; wings blackish infuscated; veinsand stigma blackish.
Structure generally similar to female. Distance between eyes 0.8–0.9 vertical diameter ofeye; eye with vertical diameter 1.4 horizontal
diameter. Malar space linear, 0.03–0.06 widthof front ocellus. Antennal length 2.0–2.3 maximum width of head; flagellum weakly curved,
not compressed (Fig. 1C–D). Cell 2Rs in
forewing with anterior width 0.8–1.0 posterior
width. Subgenital plate (Fig. 9J) with posterior
margin very widely rounded in dorsal view andstrongly convex ventrally at base in lateral view.
Gonostipes in ventral view narrowing apicallywith apical width longer than basal width of
harpe and with medial margin posteriorly trun-

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Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

Fig. 4. Apices of lances, lateral view, A. macrops (A–C), A. siamana (D), and A. captiva (E–O). — A, Ussuriysky Zapovednik, Russian Far East; B, Ulju-gun, Korea; C, Mt. Taibaishan, China; D, holotype, Thailand;E, syntype of A. sanguinolenta; F, holotype of A. kolthoffi; G, Vladivostok, Russian Far East; H, “Kasan”,
Korea; I, Daegu, Korea; J, “WenChuanShien”, China; K, Paolai, Taiwan; L, Obihiro, Japan; M, Kurisawa,
Japan; N, Kawasaki, Japan.

cate or roundly convex (Figs. 8B, 9B, E, H).
Harpe gradually and roundly narrowed toward
apex, basally also narrowed. Penis valve withvalviceps narrow subtriangular in outline in lateral view, roundly swollen laterally at middle indorsal view, ventromedially with long obliqueand dorsally bent carina in lateral view; apex
pointed and bent inward and dorsally (Figs. 8AC, 9).
Immature stages and life history. Unknown.
Type material examined. Holotype (NSMT-IHym 41670; Fig. 1C–D): ?, “14 km west ofNakhodka, Primorskij kraj, Russia, 11. VIII.
1992, A. Saito”. Kept in NSMT. Paratypes:
RUSSIA, Primorskij kraj: 8?, “1–24. VIII. 1994,

Vladivostok, Ussuriskij, Pogranitschnyi, Primorsky Kraj, Russia, leg. J. Roloff ” (ZSSM);
1?, “14/7” “Vladivostok, Suchan, Malaise”(NRMS); 1?, “15/7” “Vladivostok, Suchan,
Malaise” (NRMS); 5?, “18/7” “Vladivostok,
Suchan, Malaise” (1? labeled “Arge sanguinolenta Mocs., Malaise det. 1933”) (NRMS); 1?,“24/7” “Vladivostok, Suchan, Malaise” (NRMS);
1?, “Vladivostok, Suchan, Malaise” “Arge sanguinolenta Mocs., Malaise det. 1930” (BMNH);
1?, “8/7” “Vladivostok, Tigrovaja, Malaise”“Arge sanguinolenta Mocs., Malaise det. 1931”(NRMS); 1?, “Jakovlevka, Distr. Spassk, Prov.
Ussuri, 17. VII. 1926, Djakonov & Filipjev”“Brit. Mus. 1939-210” “Arge sanguinolenta

Arge captiva Species-group

257

Fig. 5. Lancets, A. macrops, paratypes. — A, Ussuriysky Zapovednik, Russian Far East; B, Icheon-ri,
Gyeongsangnam-do, Korea; C, Mt. Taibaishan, China. Arrows in A-B showing rudimentary longitudinal row
of minute setae at base of 1st annulus (row of setae missing in C).

Mocs., Gussakovskij det. 1938” (BMNH); 1/,“Ussuriyskiy Zapovednik, 250 m, 33 km SE Ussuriysk, 43.37 N 132.18 E, 14. VI. 1993, A.
Taeger” (SDEI); 1?, “Anisimovka, Sukhodol
river, 43.1 N 132.5 E, 6–10. VIII. 1993, E. Groll& C. Kutzscher” (SDEI); 1?, “St. Tigrovaja, [?],
7. VII. 1926, Vasijalev” “Far East, South-Ussuri,
Railroad Station, Tigrovaja, 7. VII. 1926” “Argesanguinolenta Mocsáry, ?, redescribed”(USNM). KOREA: Gangwon-do: 1?, “Mt.

Jeombong-san, GW, 10. VIII. 1992, K. T. Park”(SNU); 4?, “2007. VII. 31, Kangwon-do,
Yangyang-gun, So-myon, Hwangi-li, coll. H. S.
Lee, N 37.5625.8 E 128.3119.7, SL256 m”(2? in HSLC; 2? in NSMT, NSMT-I-Hym
41671, 41672); 1?, “Mt. Chiak, Korea, 15. Aug.
99, Sang-Wook Park” (HSLC); 1?, “95. VII. 24,
Chuncheon, Jiamli, Kyekok, Mun Chang-Seob”(SNU); 1?, Tapdongri, 37.44.36 N 128.35.18E,
660 m, Jinbu-myeon, Pyeongchang-gun (light258

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

trap), 7. VI. 2009, S.-W. Park (NSMT, NSMT-IHym 41673); 3?, “Mureung Valley, Donghae, 3.
VIII. 2009, H.-S. Lee” (2? in HSLC; 1? in
NSMT, NSMT-I-Hym 41674). Chungcheongbukdo: 1?, “Chungnam [sic], Yeongdong, Yangsan,
Gaseon-li, 21. IV. 2002, Tripotin rec.” (USNM).
Chungcheongbuk-do/Gyeongsangbuk-do: 2?,“20. VIII. 1999, alt. — 600 m, Pilosa, Soebaeksan, Tripotin coll.” (USNM). Gyeongsangbukdo: 1?, “6. VIII. 1998, Lampe U. V., alt. — 700m, Irwol-san, Yong-yang, Tripotin coll.”(USNM). Jeollabuk-do: 1?, “Manjusanghoe,
Dalgungri, Sannae-myeon, Namwon-gun, 25–26.
VII. 1990, Ku Deok-Seo” (SNU). Gyeongsangnam-do: 1?, “Jungsan-ri, Samjang-myeon, Sancheong-gun, 12–13. VII. 1990, Yun, Ju-Ik”(SNU); 1/1?, Ulju-gun, Sangbuk-myeon,
Icheon-ri, 1989. VI. 27 (SNU). Jeju-do: 1?,“Donneko, Seoguipo City, Jeju Island, 30. VIII.
2006, Lee, H. S.” (HSLC). CHINA: Heilongjiang
Prov.: 2?, “Pinchiang, Yablonia [Yabuli],
11–12. VI. 1941, S. Asahina” (NSMT, NSMT-IHym 41675, 41676); 2?, “Laoyeling, Maoershan, Shangzhishi, 19 July 1988, Li” (CSFU).
Jilin Prov.: 2?, “Changbaishan, 2 July 1999, Meicai Wei & Haiyan Nie” (CSFU). Gansu Prov.:
2?, “Dangchuan, Tianshuishi, 7 August 1993”(CSFU). Shaanxi Prov.: 1/ (Fig. 1A–B), Kaitianguan, 2000 m, 34 00 N 107 51 E, Mt. Taibaishan,
Qinling Mts., 2. VI. 2007, A. Shinohara (NSMT,NSMT-I-Hym 41677); 1?, “Zhouzhixian, 7
August 1994, Ku Lü/at light” (CSFU). Hebei
Prov.: 1?, “Wulingshan, Xinglongxian, 27 July
1957” (CSFU). Henan Prov.: 2?, “Baiyunshan,
Songxian, 24, 27 July 2003, Yingke He”(CSFU); 1?, “Longyuwan, Luanchuanxian, 23
July 2002, Jigang Jiang” (CSFU); 1?, “Baotienman, Neixiangxian, July 1998, Wei Xiao”(CSFU).
Distribution (Fig. 3). Russia (Primorskijkraj), Korea, northern and central China.
Remarks. This is a very peculiar species,
characterized by the extremely large eyes andocelli in both sexes. It has been misidentified
with A. sanguinolenta Mocsáry, 1909, which was
described from “Sibiria or. Raddefka”. Kuznet-

zov-Ugamskij(1927),
without
studying
Mocsáry’s type material, identified his specimensfrom Ussuri region with A. sanguinolenta anddescribed the males for the first time. The males,
according to Kuznetzov-Ugamskij, are characterized by the head strongly narrowed behind theconspicuously large eyes and by the very longantennae. Malaise (1931) apparently agreed with
Kuznetzov-Ugamskij and noted about the unusually developed ocelli of the males associated with
the nocturnal activity. Gussakovskij (1935), following these authors, used the large eyes, strongly narrowed posterior part of the head, and thelong antennae of the male as the good distinguishing characters for A. sanguinolenta. An examination of the type material of A. sanguinolenta (Fig. 1J–K, female) has revealed that it has
normal-sized eyes and ocelli and is not distinguishable from the type of A. captiva.
Malaise (1931) noted that the males of thisspecies were “Nachttiere” and came to light.
Many of the male paratypes have lepidopteranscales on their bodies, suggesting that moth collectors caught them in light traps. It is interesting
that, of over 50 specimens examined, only three
are females. Malaise (1931) examined “2// undviele ??”, but the females in his collection actually belong to A. captiva. The females also havelarge eyes and ocelli and should possibly be active at night, but probably they are not active
fliers as the males or they are not attracted tolight for some reason.
Arge siamana Togashi, 1988, sp. rev.
(Figs. 1E–F, 2E–I, Q, 4D, 6A)
Arge siamana Togashi, 1988: 103; Wei and Wen, 2000: 21(misspelled as siamatica, as syn. of A. captiva).

Female (Fig. 1E–F). Length about 10 mm.
Head, metathorax, legs, and abdomen entirely
black with distinct purplish reflection; antenna
black, without distinct purplish reflection; mouthparts blackish brown; prothorax and mesothorax
orange, most of cervical sclerite, ventral half ofmesopleuron, posteromedian corner of mesoscutellum, and narrow posterior margin of

Arge captiva Species-group

mesepimeron purple black; wings blackish infuscated; veins and stigma blackish.
Surface generally smooth and shining; punctures on anterior part of head fine and distinct,
generally separated from each other. Head andthorax covered with rather short hairs, those on
upper and outer orbits and on ventral part of thorax rather sparse; hairs generally blackish on
dark areas and colorless on pale areas.
Head in dorsal view (Fig. 2G) slightly dilated
behind eyes. Distance between eyes 1.3 vertical
diameter of eye; eye with vertical diameter 1.5horizontal diameter. Postocellar area weakly convex, anterior furrow distinct but lateral furrows
indistinct (Fig. 2G). Ocellar area scarcely concave between ocelli. Frontal area raised withnearly flattened top, anteriorly slightly concave
medially; median fovea in large deep pit; interantennal carinae high and sharply defined, rather
narrowly separated and distinctly convergent
below, with roundly concave depression between
them; just below level of ventral margins of antennal sockets, carinae becoming low ventrally
and finally fused with each other at ventral end of
supraclypeal area (Fig. 2E) enclosing very shallow longitudinal depression or groove along midline; supraclypeal area with side slopes very
weakly rounded. Malar space 1.0 width offront ocellus. Clypeus flattened, with anterior
margin rather narrowly and roundly incised at
middle, lateral lobes nearly truncate at apex (Fig.
2F). Antennal length 1.6 maximum width ofhead; flagellum distinctly compressed, weakly
curved basally and midapically and rather strongly thickened midapically (Fig. 1E–F).
In forewing, cell 2Rs with anterior width 0.8
posterior width, and crossvein 3r-m nearlystraight; in both wings, wing margin between
veins Rs and Cu with very short setae, generally
not protruding beyond wing margin.
Abdominal terga glabrous, except for distinctly pilose lateral parts; seventh and more posterior
terga with sparse minute setae. Seventh sternum
with posterior margin roundly produced at middle. Sawsheath in posterodorsal view (Fig. 2H)
short subtriangular in outline, with lateral margin259

rounded, apex very narrowly rounded, and medially nearly truncate; dorsomedial ridge rather
sharp and dorsal surface nearly flattened; in lateral view (Fig. 2I), ventral margin, except for conspicuous basal convexity, roundly convex, dorsal
margin nearly straight, and apex rounded; innersurface spinose.
Lance with few very narrow membranousareas at middle. Lancet (Fig. 6A) with about 27
serrulae as recognized by number of marginal
sensilla; distance between 1st and 15th serrulae
about 2.5 distance between 15th and terminal
serrulae; dorsal margin glabrous on 1st annulus,
with group of setae on each of 2nd to 6th annuli,
and continually setose on 7th and more apical annuli; large non-annulate area present dorsoapically; transverse rows of minute setae present between annular plates; marginal sensilla long;
basal annular plates dorsally rough; 1st annulus
with transverse row of minute setae and withsmall sclerotized plate ventrally; 2nd annulus
sclerotized in ventral half and on narrow dorsal
margin; serrulae roundly and very strongly convex, rather narrow and regular in shape; each serrula in median part of lancet with basal slope
slightly shorter than apical slope.
Male. Unknown.
Immature stages and life history. Unknown.
Type material examined. Holotype of Argesiamana Togashi, 1988 (Fig. 1E–F): /, “Thailand: Chieng Mai Province, Doi Inthanon N. P.:
Huai Sai Luang 10–1100 m, 13. X. 1981, Zool.
Museum Copenhagen leg.” “Arge siamana
Togashi, 1983, det. Togashi, 1983”, “Typus”(SNM).
Distribution. Thailand.
Remarks. This species is known only by thefemale holotype. Based only on the descriptions,Wei and Wen (2000) synonymized A. siamana
with A. captiva. The characters given in the foregoing key will distinguish these two species. Togashi (1988) compared the holotype of A. siamana with the original description of A. kolthoffi
Forsius, 1927, a junior synonym of A. captiva, towhich A. siamana would run in Gussakovskij’s(1935) key. Togashi discussed four characters

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Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

Fig. 6. Lancets, type material. — A, Holotype of A. siamana; B, syntype of A. sanguinolenta; C, holotype of A.
kolthoffi; D, syntype of A. captiva rufoscutellata. Arrows in A and C showing longitudinal row of minutesetae at base of 1st annulus.

Arge captiva Species-group261(shape of the 2nd and 3rd cubital crossveins in
the forewing and the distinctness of the lateraland postocellar furrows on the head), but his interpretation of Forsius (1927) and Gussakovskij(1935) was erroneous concerning the features ofthe 2nd cubital crossvein and the postocellar furrow. Togashi (1988, p. 104) mentioned, “the 2nd
cubital crossvein in the forewing is nearlystraight” and “the postocellar furrow is rather illdefined” in A. kolthoffi. However, Forsius (1927)
and Gussakovskij (1935) never referred to the“2nd cubital crossvein” of A. kolthoffi and, regarding the postocellar furrow, Forsius (1927, p.
4) noted “bei A. kolthoffi ist der Scheitel vorn
durch eine deutliche Furche begrenzt” and Gussakovskij (1935, p. 403) noted “Scheitelplatte [ofA. kolthoffi] vorne von den Ozellen durch eine
deutliche Furche geschieden”. The alluded differences in these two characters therefore have nogrounds. A direct comparison of the holotypes of
A. siamana and A. kolthoffi (Fig. 1E–F, L) has revealed no significant differences in the four characters mentioned by Togashi (1988).

1997: 560; Nambu, 1998: 17; Togashi, 1998a: 253; Togashi, 1998b: 33; Wei and Wen, 2000: 21; Kubo, 2000:
288; Yoshida, 2002: 62; Nakamura, 2003: 253; Katayama, 2004: 48; Nagase, 2004: 1244; Naito et al., 2004:
10; Hara and Shinohara, 2005: 276; Wei et al., 2006:
513; Saini et al., 2006: 597; Yoshida, 2006: 24; Shinohara and Hara, 2007: 131.
Arge flavicollis: Konow, 1905: 19; Konow, 1908: 54; Gussakovskij, 1935: 233, 265, 403, 420; Ermolenko, 1976:
261; Okutani, 1977: 294; Chou and Naito, 1991: 86;
Saini and Thind, 1995: 107; Zhelochovtsev and Zinovjev, 1995: 400; Wei and Wen, 2000: 21 (as syn. of A.
captiva); Saini et al., 2006: 597 (as sp. rev.); Lelej andTaeger, 2007: 943.
Arge sanguinolenta Mocsáry, 1909: 4. Syn. n.
Hylotoma (“Arga” [sic]) captiva: Matsumura, 1912: 35;
Matsumura, 1930: 70 (“Arge”).
Arge kolthoffi Forsius, 1927: 3; Gussakovskij, 1935: 233,
403. Syn. n.
Arge captiva rufoscutellata Takeuchi, 1927: 381;
Takeuchi, 1932: 35; Doi, 1938: 29; Takeuchi, 1939:
405 (as syn. of A. captiva).
[Arge captiva var. watanabei Takeuchi, 1932: 35, infrasubspecific; Takeuchi, 1939: 405; Abe and Togashi,
1989: 543; Wei et al., 2006: 513; Saini et al., 2006:
597.]
Arge captiva captiva: Wei and Nie, 2003: 174.
Arge captiva flavicollis: Wei and Nie, 2003: 175.

Arge captiva (Smith, 1874)

Female (Fig. 1G–M). Length about 7–12mm. Head, cervical sclerite, ventral half of mesopleuron, posterior margin of mesepimeron,metathorax, legs, and abdomen black with dullbluish reflection; antenna black, without distinct
bluish reflection; mouth parts blackish brown;
pronotum, mesonotum, and dorsal parts of mesopleuron entirely or partly orange, or entirely
black with dull bluish reflection; wings blackish
infuscated; veins and stigma blackish.
Surface generally smooth and shining; punctures on anterior part of head fine and distinct,
partly confluent with each other. Head and thorax
covered with rather short hairs, those on upperand outer orbits and on ventral part of thorax
rather sparse; hairs generally blackish on darkareas and colorless on pale areas.
Head in dorsal view (Fig. 2L) distinctly dilated
behind eyes. Distance between eyes 1.2–1.4
vertical diameter of eye; eye with vertical diameter 1.5–1.7 horizontal diameter. Postocellar
area weakly convex, anterior and lateral furrows(Figs. 1G–M, 2J–L, M, O, R, 4E–O, 6B–D, 7,
8D–F, 9K–M, 10–13)
Hylotoma captiva Smith, 1874: 376; Kirby, 1882: 62;
Dalla Torre, 1894: 328; Marlatt, 1898: 504; Nakagawa,
1899: 202; Nakagawa, 1902: 89; Matsumura, 1931: 85(“captivum”); Matsumura, 1932: 30, 42 (“captivum”);
Shinohara and Hara, 2007: 131.
Hylotoma flavicollis Cameron, 1876: 460; Kirby, 1882:
63. Syn. n.
Arge captiva: Konow, 1905: 18; Konow, 1908: 55; Rohwer, 1910: 118; Takeuchi, 1919: 188; Takeuchi, 1932:
35; Tosawa, 1932: 84; Gussakovskij, 1935: 233, 264,
403; Watanabe, 1937: 5; Takeuchi, 1938: 83; Takeuchi,
1939: 405; Saito, 1941: 130; Takeuchi, 1949: 50;
Takeuchi, 1955: 128; Okutani, 1959: 582; Kim, 1963:
279 (misspelled as captina); Sato, 1965: 224; Okutani,
1967: 46; Sato and Tosawa, 1967: 178; Kim, 1970:
143, 717 (misspelled as captina); Zombori, 1974: 454;
Kondo and Miyake, 1976: 7; Okuno et al., 1977: 125;
Dai, 1979: 217; Okutani, 1982: 413; Yang and Bianweihui, 1983: 879; Miyoshi, 1988: 189; Abe andTogashi, 1989: 543; Smith, 1989: 125; Yuan and Ding,
1993: 640; Xiao et al., 1992: 70; Kim et al., 1994: 218;
Lee and Chung, 1997: 114; Nakamura and Enoki,262

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

usually indistinct (Fig. 2L). Ocellar area scarcely
concave between ocelli. Frontal area raised andmedially shallowly concave in front of medianocellus; median fovea in large deep pit; interantennal carinae low but sharply defined, subparallel or very slightly convergent below, rather
broadly separated, with nearly flattened or widelyrounded bottom between them; just below levelof ventral margins of antennal sockets, carinae
becoming inconspicuous and strongly convergent
ventrally and fused with each other at ventral endof supraclypeal area, often V-shaped (Fig. 2J),
enclosing shallow, often inconspicuous longitudinal depression along midline; supraclypeal areawith side slopes roundly swollen. Malar space
0.6–1.0 width of front ocellus. Clypeus flattened, with anterior margin broadly, rather shallowly emarginated, lateral lobes subtriangular(Fig. 2K). Antennal length 1.3–1.6 maximum
width of head; flagellum hardly compressed,
weakly curved basally and midapically and thickened midapically (Fig. 1G–H, J–M).
In forewing, cell 2Rs with anterior width
0.7–1.0 posterior width, and crossvein 3r–m
weakly curved outwards; in both wings, wing
margin between veins Rs and Cu with very shortsetae, generally not protruding beyond wing margin.
Abdominal terga glabrous, except for distinctly pilose lateral parts; sixth and more posterior
terga with sparse minute setae also dorsally. Seventh sternum with posterior margin roundly or
subtriangularly produced at middle. Sawsheath inposterodorsal view (Fig. 2M) short subtriangularin outline, with lateral margin rounded, apex very
narrowly rounded, and medially nearly truncate;dorsomedial ridge blunt and dorsal surface nearlyflattened; in lateral view, ventral margin, except
for conspicuous basal convexity, roundly convex,
dorsal margin nearly straight, and apex rounded;
inner surface spinose.
Lance with several narrow membranous areasat middle (Fig. 4E–N). Lancet (Figs. 6B–D, 7)
with about 22 serrulae as recognized by numberof marginal sensilla; distance between 1st and15th serrulae about 3.4–6.1 distance between

15th and terminal serrulae; dorsal margin
glabrous on 1st annulus, with group of setae oneach of 2nd to 6th annuli, and continually setose
on 7th and more apical annuli; narrow non-annulate area present dorsoapically; transverse rowsof minute setae present between annular plates;
marginal sensilla long; basal annular plates dorsally rough; 1st annulus with transverse row ofminute setae and with small sclerotized plate
ventrally; 2nd annulus fully sclerotized (e.g. Fig.
7A) or weakly sclerotized dorsally (e.g. Fig. 7B);
serrulae roundly and strongly convex in basal
half and weakly convex in apical half; each serrula with basal slope much shorter than apicalslope.
Male. Length 7–9 mm. Black with dullbluish reflection; antenna black, without distinct
bluish reflection; mouth parts blackish brown;
pronotum, cervical sclerite, mesonotum, and dorsal parts of mesopleuron often entirely or partly
orange; wings blackish infuscated; veins andstigma blackish.
Structure generally similar to female. Distance between eyes 1.2–1.3 vertical diameter ofeye; eye with vertical diameter 1.4–1.6 horizontal diameter. Malar space 0.4–0.8 width offront ocellus. Antennal length 1.7–2.2 maximum width of head; flagellum nearly straight,
not compressed. Cell 2Rs in forewing with anterior width 0.6–0.9 posterior width. Subgenital
plate with posterior margin very widely roundedor sometimes truncate in dorsal view and convex
ventrally at base in lateral view (Fig. 9K–M).
Gonostipes in ventral view narrowed apicallywith apical width about as long as basal width of
harpe and with medial margin posteriorly subtruncate (Figs. 8E, 10B, D, F). Harpe gradually
narrowed toward apex. Penis valve with valviceps narrow oval with elongate apex, with round
dorsal incision at middle in lateral view, swollen
laterally with angular top in dorsal view, and ventromedially with short lateral carina (Figs. 8D–F,
10).
Larva (Figs. 11–12). Late instar (Fig. 12)
about 23 mm long, creamy white, with head, thoracic legs and spots on body black as in Figs. 11–

Arge captiva Species-group

263

Fig. 7. Lancets, A. captiva. — A, Vladivostok, Russian Far East; B, Dalian, Liaoning, China; C, “WenChuanShien”, Sichuan, China; D, Chuul, Korea; E, Kasan, Korea; F, Daegu, Korea; G, Sapporo, Hokkaido, Japan(“holotype” of A. captiva var. watanabei Takeuchi); H, Kurisawa, Hokkaido, Japan, reared from larva on
Ulmus laciniata; I, Obihiro, Hokkaido, Japan, reared from larva on Ulmus davidiana var. japonica; J,
Kawasaki, Honshu, Japan; K, “Takasago” (Taiwan); L, Paolai, Taiwan.

12; thorax and apical abdominal segments often
slightly tinted with pale orange. Antenna conical,
rather elongate (Fig. 12B); clypeus and labrum
each with two pairs of setae; mandible with twosetae on outer surface; maxillary palp four-segmented; palpifer with two setae; labial palp
three-segmented. First to ninth abdominal seg-ments each three-annulated; black dorsal spots
on trunk more or less tuberculate; tubercles except for small ones each with one large seta; lateral lobes with some large setae; setae on trunk
missing in final instar, except for those on legs,
ventral tubercles and anal segment; prolegs on
second to sixth and tenth abdominal segments;

264

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

Fig. 8. Male genitalia, A. macrops, holotype (A–C) and A. captiva, Kawasaki-shi, Japan (D–F). — A, D, Dorsal
view; B, E, ventral view; C, F, penis valve, lateral view (left dorsal).

tenth tergum in dorsal view broadly rounded apically.
Cocoon. Length 10.5–14 mm in female,
9–10 mm in male. Pale brown. Elongate oval,
double walled; outer wall netted and inner wallparchment like.
Type material examined. Lectotype of Hylotoma captiva Smith, 1874 (Fig. 1G): /, “Type H.
T.” “B.M.TYPE, HYM. 1.71” “Hylotoma
captiva, Type Smith” “Hiogo/T 4/6” “Kb.1:
4/14” (BMNH). Lectotype of Hylotoma flavicollis Cameron, 1876 (Fig. 1H–I): /, “Type H. T.”“B.M.TYPE, HYM. 1.69” “Arge (Hylotoma)
flavicollis, Cam.” “61, 49” “Type” (BMNH).
Syntype of Arge sanguinolenta Mocsáry, 1909:
1/ (Fig. 1J–K), labeled “Sibiria or. Raddefka”“Typus, Arge sanguinolenta Mocs., 1909”(HNHMB). Holotype of Arge kolthoffi Forsius,
1927 (Fig. 1L): /, “Provins Kiangsu” “China,
Kolthoff ” “Juni” “Type” “Arge kolthoffi n. sp., /,Type, R. Forsius det.” (NRMS). Syntypes of Arge
captiva rufoscutellata Takeuchi, 1927: 1/ (Fig.
1M), labeled “17. VI. 1926, Shinshu [Jinju,
Gyeongsangnam-do, Korea], Hasegawa” “Lecto-

Arge captiva Species-group

265

Fig. 9. Male genitalia, dorsal view (A, D, G), ventral view (B, E, H), penis valves, lateral view (left dorsal) (C,F, I), subgenital plates (J–M), A. macrops (A–J) and A. captiva (K–M). — A–C, Vladivostok, Russia; D–F,
Chuncheon, Korea; G–J, “Yablonia”, China; K, “Taikyu”, Korea; L, “Dairen”, China; M, “Takasago”,
Taiwan.

266

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

Fig. 10. Male genitalia, A. captiva, dorsal view (A, C, E), ventral view (B, D, F), and penis valves, lateral view(left dorsal) (G–M). — A, B, G, Gribnoye, Russia; C, D, J, Jinju, Korea; E, F, M, Mt. Lalashan, Taiwan; H,
Dalian, China; I, Daegu, Korea; K, Nishiobihiro, Japan; L, “Takasago”, Taiwan.

Arge captiva Species-group

267

Fig. 11. Larvae, A. captiva. — A, Early instar (rearing record H, AS080804) on Ulmus davidiana var. japonica,
near Shirakaba-ko Lake, Nagano Prefecture, Honshu, August 4, 2008; B, early instar (rearing record E,
AS090702) on Ulmus davidiana var. japonica, Nopporo Forest Park, Ebetsu, Hokkaido, July 2, 2009; C, middle instar (rearing record F, HH080717H) on Ulmus davidiana var. japonica, Kikyo, Hakodate, Hokkaido,
August 2, 2008; D, middle instar, same as B, July 5, 2009; E, late instar (rearing record B, HH040810A) on
Ulmus laciniata, in Manji, Kurisawa, Hokkaido, August 11, 2004; F, late instar (rearing record G,
AS061008A) on Ulmus parvifolia, in Kamigo-cho, Yokohama-shi, Kanagawa Prefecture, Honshu, October
17, 2006. Digital images taken by Shinohara (A, B, D, F) and Hara (C, E) on the given date.

type [unpublished], Arge captiva var. rufoscutellata Takeuchi, D. R. Smith” (OPU); 1?, labeled“17. VI. 1926, Shinshu, Hasegawa” (OPU).
Other material examined. RUSSIA: Primorskij kraj: 1/, “3/7” “Vladivostok, Tigrovaja,
Malaise” (NSMT); 1/, “26/7” “Vladivostok,

Sedanka, Malaise” (NRMS); 1/, “Vinogradovka,
Ussur. kr., 30. V. 929, Kirichenko” (NRMS); 1/,“Frolovka, Suchan, Ussur. kr., 11.VII.926,
Mordvilko” (NRMS); 1?, “Gribnoye, 10km E of
Chernigovka, Prim. kray, 23–24. V. 1995, A.
Lelej” (NSMT). KOREA: Hamgyeongnam-do:

268

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

Fig. 12. Late instar larva, A. captiva (rearing record H, AS080804, fixed August 12, 2008). — A, Lateral view;B, antenna, lateral view.

2/1?, “end VII, 1933, Hokusei [Pukchong],
K. Saito” “collected in Hokusei and reared in
Suigen [Suwon], em. end of July, 1933, host:
Ulmus” (OPU). Hamgyeongbuk-do: 1/, “Shuotsu [Chuul], 16. VII. 1931, K. Sato” (NSMT).
1/, “Shuotsu, 17. VII., Y. Yano (Cor. Chi. Exp.
1937)” (OMNH). Gyeonggi-do: 1/, “Koryo[Kwangneung], 1. VI. 1924, Y. Hasegawa”(NSMT); 1/ 1?, Suigen [Suwon], 5. VII. 1922,
K. Sato (NSMT); 2/ 1?, Suigen, 10. VII. 1922,
K. Sato (NSMT); 1?, Suigen, 25. V. 1924, K.
Sato (NSMT); 1?, Suigen, 5. VI. 1924, K. Sato
(NSMT); 1?, Suigen, 5. VII. 1925, K. Sato
(NSMT); 3?, Suigen, 8. VII. 1925, K. Sato
(NSMT); 1/, Suigen, 29. V. 1929, S. Fujii(NSMT); 1/ 2?, Suigen, VII. 1925, K. Sato
(NSMT); 1/, Suigen, 4. VII. 1925, K. Sato
(NSMT); 1/, “Suwon, 6. VI. 1990, D. H. Kang”(SNU); 2/, “5. VI. 1936, Kasan [? nr. Suwon],Takeuchi” (OPU); 1/, “Nongdae [College ofAgriculture, Seoul National Univ., Suwon], 11.
IX. 95, Kim Jung-Wook” (SNU); 1/, same data
except “12. IX. 95” (SNU); 1/, “Bukhansan,
Uiryon, V. 30. 1995, Han Kyeong-Deok” (SNU);
1/, “Anyang, 31. V. 91, K.S.I.” (SNU); 1/,“Anyang Sumokwon, V. 12. 95, Kim Jung-Wook,on Hydrangea arborescens” (SNU); 1/, “Deungducheon, Soyosan, 1985. IX. 1, Park Jeom-Hui”(SNU). Gangwon-do: 1/, “Pyongchang, GW, 31.
VII. 1991, K. T. Park” (SNU); 1/, “Kangwon,
Hongcheon, Kachilbong, 1995.06.09, Lee JeongHun” (SNU); 1/, same data except “Baek JeongHui” (SNU); 1/, “Cahngchon [sic], GW, 11. VI.
1994, Y. I. Kim & J. H. Lee” (SNU); 1/, “2007.
VII. 31, Kangneung-si, Yeongok-myeon, Samsan-ri (Guhwolsaap), coll. H. S. Lee, N 37.5131
E 128.4304, alt. 186 m” (HSLC); 1/, “Chuncheon, Nam-myeon, Magag-li, Mal. trap in larch
planted forest with dense shrub layer, 24. V.–12.
VI. 2004, Tripotin Coll.” (USNM); 1/, same
locality “along Hongcheon river, alt. 70 m,
N 37 43.786 / E 127 34.589, 11. VII.–7. VIII.
2004, Tripotin rec.” (USNM); 1/, “Inje-gun,Nam-myeon, Gamdun-li, 250 m, N. 37 58.40 /E 128 07.54 on Sumac/Ampelopsis/yellow Umbelifera flowers, 5–6. VIIII [sic]. 2005, Tripotinrec.” (USNM); 1/, “Odaesan nr. Dongdaesa, 4Mal. tr. in old Korean Fir Forest, 2. vi.–2. viii,
2006, N. 37 44.31 / E 128 35.71, 800 m, Tripotinrec.” (USNM); 1/, Pugdaesa, ca. 1300 m, Mt.
Odaesan, 27. V. 2008, A. Shinohara (NSMT);
2/, “Gapyeong-gun, Seo-myeon, Magok-li,
37 42.97 N 128 35.45 E, 5 Mal. tr. in forest area,
24. VIII–31. IX. 2006, Tripotin rec.” (USNM).
Chungcheongnam-do: 1/, “Chungnam, Daejeonsi, Wadong, Mal. tr., al. forest on wild Rosa

Arge captiva Species-group

patch, 36 24.02 N / 127 25.98 E, 17. IX.–6. X.
2006, Tripotin rec.” (USNM). Chungcheongbukdo: 1?, “Seishu [Chongju], J. Yoshioka”(NSMT); 1/, “Chungbuk, Muju Valley (resort),
2–3. VIII. 2003, C. Yong rec./leg.” (USNM); 1/,“Chungnam [sic], Yeongdong-gun, Sangchonmyon, Mulhan valley, Gojadong, 4. VIII–24. IX.
2002, P. Tripotin Coll.” (USNM). Gyeongsangbuk-do: 5/, Andong, 3–6. VI. 1970, H. Yamada,K. Tobi & S. Ohkusa (KU); 8/ 10?, Taikyu[Daegu], 5. V. 1931, H. Sugiura (NSMT); 3/
2?, Taikyu, 5. V. 1931, K. Sato (NSMT); 1/ 2?,“29. V. 1936, Kinsen [?Kimchon], Takeuchi”(OPU); 1/, “Bukkokuji [Bulguksa], 7. V. 1931,
H. Sugiura” (NSMT). Jeollabuk-do: 1/, “Jirisan,
Hamyang-gun, Songjeon-li, Munsu-sa, alt. –400 m, Mal. trap, in forested area in shade,N 35 24.739 / E 127 43.818, 6–27. VI. 2004,
Tripotin Coll.” (USNM); 1/, same data except“ex 2 Mal. traps in forested area, 27. VI.–27. VII.
2004” (USNM); 1/, same data except “16.
VIII.–5. IX. 2004” (USNM); 2/, same data except “5. V.–4. VI. 2005” (USNM); 1/, same data
except “9. VII.–17. VIII. 2005” (USNM); 1/,“Jirisan, Hamyang-gun, Macheon-myon, Songjeon-li, alt. – 700 m, Mal. trap, on sm. stream,
N 35 20.930 / E 127 38.503, 27. VI.–27. VII.
2004, Tripotin rec.” (USNM); 1?, same data except “27. VII.–16. VIII. 2004” (USNM); 1/,
same data except “6. VI. 2004” (USNM); 1/,“Jirisan, Hamyang-gun, Macheon-myon, Songjeon-li, alt. – 700 m, 35 20.55 N/127 38.21
E, 27. VII. 2004, Tripotin rec.” (USNM).
Gyeongsangnam-do: 2/, “Masan, 1994.VIII.10,
Lee Hak-Yeong” (SNU); 1/, “K. N., Sancheonggun, Jungsan-ri, Sunduryu, 1981.VI. 3, Choe
Nak-sik” (SNU); 1/, “7. VI. 97, Pijindo, nr. Kojedo, Tripotin Coll.” (USNM). Jeollanam-do: 1/,“Korea, I. A. S., Suncheon, VI. 29. 1994, E. Y.
Her” (SNU). JAPAN: Hokkaido: 1/, Nayoro,
Odori-minami, 26. VI. 2009, H. Hara (NSMT);
1/, Sounkyo, 5. VIII. 1931, Kobayashi (OPU);
1/, Tenninkyo, 28. VII. 1962, K. Tsuji (KU);
4/, “Abashiri, 4. VIII. 1930, S. Fujii” “Arge captiva var. watanabei Takeuchi, det. K. Sato, ‘57”
(NSMT); 1/, Ashoro, Tokachi, 15. VII. 1964, A.

269

Nagatomi (KU); 1/, Obihiro, 18. VII. 2007, H.
Hara (NSMT); 1/ (HH040823A), Obihiro, 23.
VIII. 2004, deposited 14 eggs on Ulmus davidiana var. japonica on 23–25. VIII. in cage, H.
Hara (NSMT); 3/ (progeny of HH040823A),
Obihiro, eggs deposited on 23–25. VIII. 2004,
cocoon 21–23. IX., em. 13–15. X. 2004, host:
Ulmus davidiana var. japonica, H. Hara
(NSMT); 1?, Nishiobihiro, 20. VI. 2004, A.
Shinohara (NSMT); 1/, Memuro, Tokachi, 16.
VII. 1976, M. Suwa (HU); 1/ (HH040810A),
Manji, Kurisawa, 5 gregarious larvae on Ulmus
laciniata, 10. VIII. 2004, cocoon 12–13. VIII.,em 31. VIII. 2004, H. Hara (NSMT); 1/, Bibai,
Sorachi, 10. VI. 1996, H. Hara (NSMT); 1/,
Ichinosawa, Iwamizawa, 30. V. 2007, on Ulmus
davidiana var. japonica, H. Hara (NSMT); 1?,
Nakagoya, Tobetsu, 14. VI. 1998, H. Hara
(NSMT); 1?, “Sapporo, Matsumura, Maruyama,
26. VI. 1913” (HU); 1/, “Sapporo, Dr. Matsumura” (HU); 1/, “Maruyama, K. Igarashi, 3.
VI. 1928” (HU); 1/, “Sapporo, Y. Ota, 9. VI.
1930” (HU); 1?, Kotoni, 1. VII. 1932, H. Sugiura (NSMT); 15/ 13? [including “type series” of
A. captiva var. watanabei Takeuchi, 1932], Sapporo, 2. VII. 1932, C. Watanabe (HU, OPU); 1?,
Hokkaido, Uchida, Kotei [school campus], 5.
VII. 1932” (HU); 1/, “Sapporo, 25. VI. 1939, H.
Aoki” (OMNH); 1?, Hyakumatsuzawa, Sapporo,
20. VIII. 2001, T. Yoshida (HU); 1/, Hokkaido
Univ., Sapporo, 15. IX. 2003, H. Hara (NSMT);
1/, Hokkaido Univ., Sapporo, 26. VIII. 2004, H.
Hara (NSMT); 1/ (HH040826A), Hokkaido
Univ., Sapporo, 26. VIII. 2004, deposited 10 eggs
on leaf margin of Ulmus pumila on 26–27. VIII.
in cage, H. Hara (NSMT); 1/, Hokkaido Univ.,
Sapporo, larva coll. 12. IX. 2006, em. 25. IV.
2007, host: Ulmus pumila, H. Hara (NSMT);
23/, Hokkaido Univ. Campus, Sapporo, 43-0433N 141-20-31E, 28. VI. 2007, A. Shinohara(NSMT); 24/52?, Hokkaido Univ., Sapporo, 7.
VII. 2007, on Ulmus pumila, H. Hara (NSMT,HFRI); 1/, Botanical Garden, Hokkaido University, Sapporo 43-03-51N 141-20-34E, 28. VI.
2007, A. Shinohara (NSMT); 5/, Hokkaido
Government Office, Sapporo, 43-03-47N 141-20-

270

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

56E, 28. VI. 2007, A. Shinohara (NSMT); 1/,
Shikotsu, 24. VI. 1989, R. Inomata (MNHAH);
3/ 1? (HH080717H), Hakodate, Kikyo, eggs
coll. 17. VII. 2008, hatched 24. VII., cocoon 811. VIII., em. 31. VIII.–10. IX. 2008, host:
Ulmus davidiana var. japonica, H. Hara
(NSMT). Honshu: Tochigi Pref.: 2/, Chuzenji,
Nikko, 4. VI. 1977, A. Shinohara (NSMT); 6?
(AS080925B), Nikko, 590 m alt., 36-45-8N 13936-25E, larvae coll. 25. IX. 2008, matured 1–5.
X, em. 18–25. X. 2008, host: Ulmus davidiana
var. japonica, A. & N. Shinohara (NSMT).
Gunma Pref.: 1?, Nishikurozawa, 800 m, 36-4955N 138-57-29 E, nr. Mt. Tanigawadake, Doai,
26. VIII. 2007, A. & N. Shinohara (NSMT).
Chiba Pref.: 1/2?, Sawada, Ichikawa City, 21.
VIII. 2003, H. Suda (HSC); 1/, Moto — Kamimoto, Shisui, Inba, 8. V. 1975, H. Suda (HSC);
1/, Enokido, Yachimata City, 29. VII. 1995, H.
Suda (HSC). Tokyo Met.: 4/ 3?, Meguro, 25. V.
1934, K. Sato (NSMT); 1/, Koremasa, Fuchu,
22. VII. 1989, A. Shinohara (NSMT); 1/, Shinmachi, Oume, 18. V. 1990, H. Takahashi(NSMT); 1/, “7–18. 1930, Ulmus americana,
Komaba” (OPU); 1/, “Komaba, Tokyo, Hirayama” (OPU). Kanagawa Pref.: 2?13/(AS061008A), Kamigo-cho, Yokohama-shi, 80m
alt., 35-20-30N 139-35-02E, larvae coll. 8. X.
2006, matured 20. X.–1. XI., em. 18–29. XI.
2006, host: Ulmus parvifolia, A. & N. Shinohara(NSMT); 15 late instar larvae, same data, except
fixed 25. X. 2006 (NSMT); 30/ 61?, Kosugimachi, Kawasaki-shi, 27. IV. 1989, H. Ono(NSMT); 9/ 1?, Ebina, 20. VIII. 1957, fed on
Ulmus sp., Izumi (NSMT). Nagano Pref.: 1/,
Shimashima-dani, 29. VII. 1989, H. Takahashi(NSMT); 25 late instar larvae (AS080804), Shirakaba-ko, 1420 m alt., 36-6-22N 138-14-31E, 4.
VIII. 2008, fixed 12. VIII, host: Ulmus davidiana
var. japonica, A. & N. Shinohara (NSMT). Kyoto
Pref.: 1/7?, Kyoto, 15. V. 1927, K. Takeuchi(OPU, OMNH); 1/, Kyoto, 27. V. 1931, K.
Takeuchi (OPU); 1/, Kyoto, 21. V. 1953, K.
Takeuchi (OPU); 1/, Kose, 26. VII. 1929, K.
Takeuchi (OPU); 1?, Kyoto, 20. VII. 1924, K.
Takeuchi (OPU); 1/, “Kyoto, von Herr Ooe”

(OMNH); 1/, “Kyoto, Daimonjiyama, ‘39. 9. 3,
T. Kimura” (KU). Nara Pref.: 1?, Saki-cho, 100m, Nara-shi, 19. V. 2002, H. Yoshida (HYC).
Osaka Pref.: 1/, Yoshikawa-mura, 11. V. 1941, J.
Yoshioka (NSMT); 1?, Minoo, 11. VI. 1936, K.
Shibuya (OPU); 1/1?, “Minomo [Minoo], N.
Tosawa” (OMNH); 1/, “Minou [Minoo], 21.
VI. 1926, N. Tosawa” (OMNH); 1/, Osaka, 24.
V. 1916, K. Takeuchi (OPU); 2?, Osaka, 27. V.
1915, K. Takeuchi (OPU); 1/, “Juso, 28. IV.
1916, N. Tosawa” (OMNH); 1/, Utsubo-koen,
Nishi-ku, Osaka, 26. VI. 1991, S. Okuno(OMNH); 4?, Utsubo-koen, Nishi-ku, Osaka,
16. VIII. 1992, S. Okuno & K. Katsura(OMNH); 1?, Chishima-koen, Taisho-ku, Osaka,
30. VI. 1986, A. Ichikawa (OMNH); 1/, Chishima-koen, Taisho-ku, Osaka, 21. VI. 1994, A.
Ichikawa (OMNH); 1?, Ohama-koen, Ohamakita-machi, Sakai-shi, 12. V. 1981, K. Harusawa(OMNH); 1?, Ishibashi, 18. V. 1952, M. Koizumi (OMNH). Hyogo Pref.: 1/, “Hisayaki-mura,
Sayo-gun, Japan” “Arge captiva Sm., det.
Malaise, 1933” (NRMS); 1/, “Takarazuka, N.
Tosawa” (OMNH); 1/, “Takarazuka, 25. V.
1948, N. Tosawa” (OMNH); 1/, Nishiwaki,
Yokacho 45, 27. V. 1958, R. Inomata (MNHAH,
B1-298054); 1/, Aina, Yamada, Kita-ku, Kobeshi, 17. V. 1997, H. Yoshida (cited by Naito et al.,2004, HYC); 1/, “760718, Kasui, coll. N. H.”(KU). Tottori Pref.: 1/, Daisen, 15. VII. 1920, K.
Takeuchi (OPU). Shikoku: Kochi Pref.: 3/,
Shioyazaki, 28. VII. 1982, A. Ishida (OMNH);
3/ (AS071103A), Godai-san, Kochi-shi, ca.
100 m alt., 33-32-57N 133-34-50E, larvae coll. 3.
XI. 2007, matured 4–5. XI., em. 23–25. IV. 2008,
host: Ulmus parvifolia, A. Shinohara (NSMT);
1/, same data except em. 21. V. 2008 (NSMT); 1
late instar larva, same data, except fixed 4. XI.
(NSMT). Kyushu: Nagasaki Pref.: 1/, Tsushima,
Izuhara — Ariake-yama, 9. VI. 1941, T. Shirozu(OPU); 1/, Tsushima, Mitsushima, Ofunakoshi,
2. VI. 2002, E. & R. Matsumoto (OMNH).
CHINA: 1?, “Walker Coll., China 92–196”“8335” “British Museum” “Arge flavicollisCam., Malaise det., 1933” (NRMS). Heilongjiang Prov.: 1/, “Harbin, Manchiukuo, Aug.

Arge captiva Species-group

6. 1937, T. Touchiyama” (OPU); 1/, “Harbin,
Manchukuo, M. Weymarn.” “June 10, 1936”“Carn. Mus. Acc. 11136” “Arge captiva Smith”(USNM); 1/, “Manchuria, Ertsentientze, 62 km
E of Harbin on Chinese Eastern Railway,
15–VI–41, V. N. Alin coll.” (CAS). Jilin Prov.:
1/, “Erdao, Changbaishan, 30 June 1999, Meicai
Wei & Haiyan Nie” (CSFU). Liaoning Prov.: 1/,“On: Nire [Ulmus], Date: 1935.6.23, Place:
Yugakuzyo [Xiongyuecheng], Name Seki[Qi]” (OPU); 1/, “Yugakuzyo, S. Kariya,
1936.6.10” (OPU); 12/ 1?, “Yugakujo, 17. VI.
1917” (KU); 2/ 1?, “Dairen [Dalian], Nanmanshu, 2. VIII. 1933, Col. K. Nomura” (OPU);
1/, “Mukden [Shenyang], von E. Suenson, 2.
V. 1922” (OMNH); 1/, “Shenyang, 10 June
1999, Yuzhan Sun” (CSFU); 1?, “Shenyang, 10
June 2003, Maoling Sheng” (CSFU); 1/,“Shenyang, 26 June 2003, Maoling Sheng”(CSFU); 1/ 2?, “Shenyang, 25 July 2003,
Maoling Sheng” (CSFU); 1?, “Shenyang, 25
July 2003, Suping Sun” (CSFU); 2/, “Shenyang,
30 July 2003, Maoling Sheng & Suping Sun”
(CSFU). Neimenggu Auton. Reg.: 1/, “Yimenzhunqi, 24 June 1972” (CSFU). Ningxia HuiAuton. Reg.: 1/1?, “Shapotou, Zhongwei, April
1985, Guodong Rea” (CSFU). Gansu Prov.:
2/1?, “Yuquanguan, Tianshuishi, 24 April 2001,
Xingyu Wu” (CSFU); 1?, “Huaqiqinjiagou,
Tianshuishi, 23 May 2006, Heng Xin” (CSFU);
1/, “Xiaolongshan, Tianshuishi, 7 July 2007,
Yihai Zhong” (CSFU). Shaanxi Prov.: 1?,“Zhongnanshan, Xi’anshi, 27 May 2006, Xun
Zhu” (CSFU). Shanxi Prov.: 1?, “Shanxi Prov.,
27 July” (CSFU). Hebei Prov.: 1/, “Wulingshan,
Xinglongxian, August 1974” (CSFU); 1/ 3?,“Yunyougu, Wulingshan, 15 June 2007, ZezhanLi” (CSFU). Beijing Munic.: 1/ 1?, “Xishan,
20 June 1957, Songhe Ying & Shaohua Li”(CSFU); 1?, “Peiping, 1417” (USNM); 6/ 1?,“Peiping, Hopei, VII–VIII, 1932, G. Liu”
(MCZ). Tianjin Munic.: 1/, “Tientsin [Tianjin], June 1902, F. M. Thomson, 1904–229”(NRMS); 2/ 1?, “Tientsin, 15. 6. 06, F. M.
Thomson, 1907–200” “Arge (Hylotoma) flavicollis Cam.” (1/ in NRMS labeled “compared with271

type and found identical, Malaise, 1933”)(NRMS, NSMT); 1/, “Tientsin” “Collection Tischbein, Konow determin. 1906” “Arge flavicollis
Cam. /, Fr. W. Konow determin. 1907” (NRMS).
Sichuan Prov.: 1/, “WenChuanShien, 30 mi
NNW Kuanshien [Guanxian], VII–22. 1933”“Szechwan [Sichuan], DCGraham, 4600 ft.”
(USNM). Hubei Prov.: 1?, “Shizhishan, Wuhanshi, July 1995, Yanggan Zhu” (CSFU). Henan
Prov.: 1/, “Anyang, 3 July 1957, Xiaoquan Li”(CSFU); 1?, same data except “6 October 1957”(CSFU); 1?, same data except “22 October
1957” (CSFU); 1/, “Langyuwan, Luanchuanxian, 21 July 2004, Meicai Wei” (CSFU). Shandong Prov.: 1/, “China, Tsingtao [Qingdao],
VI–10–1926, RHLeFevre” (USNM); 1/, “Tsinan[Jinan], Shantung [Shandong], VI–10–1926,
RHLeFevre” (USNM); 1/, same locality, “A. P.
Jacot, Jun. 8, 1922” (USNM). Jiangsu Prov.: 1/,“Nanking [Nanjing], China, V. 5. 23” “Presented by ECVanDyke, Collector” “captiva, D. R.
Smith” (USNM); 1/, “Nanking, VI. 20. 1923”(USNM); 1/, Nanking, Aug. 14, 1919, H. E.
Loomis (USNM); 1/ 1?, “Nantung [Nantong], China, V–9–23” “Presented by ECVanDyke, Collector” (/ with “Arge flavicollis(Cameron) /, det. H. N. Greenbaum”) (CAS);
1/, “Penniu, China, IX. 22, 1925, Coll. C. Y.
Wong” (USNM); 1/ 1?, “Nanking, IV. 20, 23”“Presented by ECVanDyke, Collector” (CAS);1?, “Nanking, V. 1. 23” “Presented by ECVanDyke, Collector” (CAS). Shanghai Munic.:
1/, “Zô-sè [She Shan]” “Chine, Prov. Kiangsu,
Shanghai, Musee Heude” “26.6.30, O. Piel coll.”(cited by Takeuchi, 1938; OPU). Guizhou Prov.:
1/, “Fuguoshi, Fanjingshan, 28 July 2001, Mingli Chen & Ningting Huang” (CSFU). Hunan
Prov.: 1/, “Laopenggou, Mufushen, Yueyangshi,
28 May 2007, Shuaigo Nie” (CSFU); 1/, “Jiuweishan, Yongzhoushi, 23 April 2007, MeicaiWei” (CSFU). Zhejiang Prov.: 1/, “Hangchow[Hangzhou], China, V–10, 1925, Coll. T. P.
Chao” (USNM); 1/, “Hangchow, IV–5–1926, T.
P. Chao” (USNM); 1/, “Xitianmushan, Linanshi,
2 June 1990, Junhua He” (CSFU). Guangdong
Prov.: 2/ 1?, “Chebaling, Shixingxian, Shaotian-

272

Akihiko Shinohara, Hideho Hara and Jung-Wook Kimshi, 13 April 2007, Meicai Wei & Yihai Zhong”(CSFU). Fujian Prov.: 1/, “Jiangningxian”(CSFU); 1/, “Foochow [Fuzhou], C. R. Kellog
Coll.” (CAS). TAIWAN: 1?, Mt. Lala-shan,
Taoyuan Hsien, 3.VI.1986, C. C. Lo (NSMT);
1/, “Paolai, Taoyuan, Kaohsiung, 3. V. 1996, H.
Takizawa” (HU); 4/ 10?, “Takasago [Taiwan],
Go-gatsu [May]” “hachi [8, probably refers
to fig. 8 in plate XLIV in Matsumura, 1912]”“Muneaka-ruri-habachi [Japanese name for A.
captiva used by Matsumura]” (HU).
Distribution (Fig. 13). Russia (Primorskijkraj), Korea, China, Taiwan, Japan. Mongolia(Ermolenko, 1976), northern India (Saini andThind, 1995).
Host plants. Ulmus davidiana Planch. var.
japonica (Rehder) Nakai, Ulmus laciniata(Trautv.) Mayr, Ulmus parvifolia Jacq., Ulmus
pumila L. in Japan. One female examined is labeled “7–18. 1930, Ulmus americana, Komaba”,
suggesting that this specimen was reared fromthe larva feeding on a cultivated Ulmus americana L. Ulmus pumila (“Ulmus mandshurica”)
in Korea (Saito, 1941) and elm (“yu”) in China(Dai, 1979). Watanabe’s (1937) record of alder asa host plant of this species is most probably erro-Fig. 13.

neous. Yoshida and Miyashita’s (1993) record of
Zelkova needs confirmation.
Notes on rearing. The following records onrearings were made by H. Hara in Bibai, Hokkaido (A–D, F) and by A. Shinohara in Tokyo, Honshu (E, G–J).
A, HH040823A. A female collected at Obihiro, Tokachi, on August 23, 2004, deposited 14
eggs on Ulmus davidiana var. japonica on August 23–25 in the laboratory. The larvae madecocoons on September 21–23, and three females
emerged on October 13–15 of the same year.
Others died.
B, HH040810A (Fig. 11E). Five gregarious
larvae, each about 20 mm long, were found on
Ulmus laciniata in Manji, Kurisawa, on August
10, 2004. They spun cocoons on August 12–13
and one female emerged on August 31, 2004.
Others died.
C, HH060820H. A group of 33 larvae were
found on Ulmus laciniata on Mt. Sankakuyama,
Kurisawa, on August 20, 2006. They matured
and made cocoons on August 22–24. No adults
emerged.
D, HH060912. A group of larvae were collected from the leaves of Ulmus pumila in the cam-Distribution of A. captiva based on specimens examined.

Arge captiva Species-grouppus of Hokkaido University, Sapporo, on September 12, 2006. One female emerged on April
25, 2007.
E, AS090702 (Fig. 11B, D). A group of 19
young larvae were found on Ulmus davidiana
var. japonica in Nopporo Forest Park, 50 m alt.,
Ebetsu, Hokkaido, on July 2, 2009. They all diedwithin a week.
F, HH080717H (Fig. 11C). Several eggs werecollected on Ulmus davidiana var. japonica in
Kikyo, Hakodate, on July 17, 2008. They hatchedon July 24 and the larvae made cocoons on August 8–11. One male and three females emergedon August 31–September 10 of the same year.
G, AS061008A (Fig. 11F). A group of 71
young and middle-sized larvae were found feeding on Ulmus parvifolia in Kamigo-cho, Yokohama-shi, Kanagawa Prefecture, Honshu, on October 8, 2006. They were apparently a mixture ofdifferent instars derived from a few clutches. A
total of 28 larvae reached maturity, 20 larvae
were fixed in ethanol, and all others died. The 28
larvae spun cocoons on October 15 to November
5. Two males and 13 females emerged on November 14–29. The remaining 13 larvae died in
the cocoons.
H, AS080804 (Fig. 11A). A group of about 30
young larvae were found feeding on Ulmus davidiana var. japonica near the coast of Shirakaba-ko Lake, 1420 m alt., Nagano Prefecture,
Honshu, on August 4, 2008. Mainly because theshort of fresh leaves, some larvae died and the remaining 25 larvae were fixed in ethanol on August 12.
I, AS080925B. A group of about 20 larvae
were collected from the leaves of Ulmus davidiana var. japonica in Nikko, 590 m alt., Tochigi
Prefecture, Honshu, on September 25, 2008.
They reached maturity on October 1–5 and six
males emerged on October 18–25 of the same
year. Others did not survive.
J, AS071103A. Four almost mature larvae
feeding on Ulmus parvifolia were found in
Godai-san,Kochi-shi,
Kochi
Prefecture,
Shikoku, on November 3, 2007. They were foundon nearby leaves but not forming a group. They

273

matured on November 4–5 and three females
emerged on April 23–25 and one female on May
21, 2008.
Life history. The general life cycle of thisspecies is as follows. The female deposits eggs
along the margin of the leaf in a row. The
hatched larvae cling to the margin of the leaf andfeed on it in a group (Fig. 11A–B). When the larvae become large, the group may become veryloose. The mature larva goes down to the groundand spins a cocoon, in which it pupates often
after a diapause of various lengths. After a shortperiod of pupa, the adult emerges.
Takeuchi (1932) first noted simply that the larvae of A. captiva fed on Ulmus sp. The life history of this species was briefly described by Okutani (1959), Okuno et al. (1977) and Hara andShinohara (2005) in Japan, by Saito (1941) and
Lee and Chung (1997) in Korea, and by Dai(1979), Yang and Bianweihui (1983), and Xiao etal. (1992) in China. There are two generations ayear in Japan (Okuno et al., 1977), Korea (Saito,
1941), and in China (Dai, 1979; Yang and Bianweihui, 1983), though Okutani (1959) noted “twoor three generations a year” for Japanese populations.
Our rearing records as well as the adult collection records may suggest a complex and variable
life history of the species, not as simple and regular as the one described by previous authors.
The rearing data obtained, though rather poor,
may show that one life cycle of this species lasts
about 50–60 days without prolonged diapause(see rearing cases A, B, F and G above). All thelarvae, except for part of those found in late autumn (rearing case J), became adults in the laboratory within the same year. In northern areas,
this species probably has two generations a year,
whereas there are potentially three or four generations a year in southern and warmer areas in its
distributional range. The adult emergence of thisspecies may show a polymodal pattern (Knerer,1993) (see rearing case J above), which is rather
common in Arge species (e.g., Shinohara andHara, 2009).
The adult collection records (see specimens

274

Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

examined) may indicate that the spring emergence of the adults (e.g., in early June to earlyJuly in Sapporo, Hokkaido, Japan, in late April tolate May in lowlands of central Honshu, Japan, inlate May to early June in lowlands of Gyeonggido, Korea) is largest in scale but the emergence
period is long and may not be unimodal. In Sapporo, Hokkaido, the larvae in various stages andadults were found together in late June to earlyJuly. These larvae were the offspring of the adults
that had occurred a few weeks before (earlyJune). The adults found in late June to early July
probably belonged to the same generation as theadults that oviposited in early June, the individuals that emerged from the hibernated larvae. In
summer through autumn, the occurrence of theadults and larvae in the field is rather irregular,
and it is difficult to determine to which generation the individual adults or larvae actually belong. As a result of polymodal adult emergence,
the adults found in early summer may be the offspring of the spring adult (first generation of theyear) or possibly a late occurrence of the hibernated individual (last generation of the previousyear). In warmer areas, the adults and larvae
found in autumn may be the individuals of thethird or even fourth generations probably mixed
with late-occurring individuals of earlier generations. The actual dynamics of populations anddetails of the life history of A. captiva are still little known.
Remarks. The characters given in the foregoing key will distinguish the adults of A. captiva
from those of the other two species of the captiva
group. The larvae are gregarious leaf-feeders on
elm and A. captiva is the only known species of
the genus associated with elm in eastern Asia.
The larvae of A. macrops and A. siamana are unknown. In general appearance, the larvae of A.
captiva resemble those of A. pullata, which are
gregarious leaf-feeders on birch. Besides the definite differences in host plants, the late instar larvae of A. captiva differ from those of A. pullata
in the non-metallic black areas (with slight bluishmetallic luster in A. pullata), the presence of
black spot on the subanal lobe (black spot absent

on the subanal lobe in A. pullata), and the presence of prolegs on the second to sixth and tenthabdominal segments (on second to eighth andtenth abdominal segments in A. pullata).
Arge captiva is a widespread species showing
large variations in the color of thorax. The continental specimens, including the type material of
Hylotoma flavicollis Cameron, 1876, from Hong
Kong (Fig. 1H–I), A. sanguinolenta Mocsáry,
1909, from the Russian Far East (Fig. 1J–K), A.
kolthoffi Forsius, 1927, from Jiangsu, China (Fig.
1L) and A. captiva rufoscutellata Takeuchi,
1927, from Korea (Fig. 1M), are paler in color
pattern; the females usually have the pronotum,
mesonotum, and dorsal part of mesopleuron entirely reddish. The Japanese females, including
the type specimen of Hylotoma captiva Smith,
1874 (Fig. 1G), are more variable, having various
amount of reddish areas on the thorax, but evenin the palest specimens the posterior part of themesoscutellum is marked with black. The Japanese specimens often have no reddish areas on thethorax, while such entirely dark females are rareon the continent. After examining a long series of
specimens from various localities (see material
examined), we have concluded that these taxa are
all conspecific and are difficult to distinguisheven at the subspecies level.
The systematic position of A. flavicollis wasparticularly in confusion. This taxon was first described under the name of Hylotoma flavicollis
by Cameron (1876) on the basis of the female
from Hong Kong. The description was very
short, only five lines long. Kirby (1882) redescribed it, again in a few lines, based on one
female (“Type of species”) and noted “closely allied to H. captiva”. Konow (1908) gave a moredetailed description of the female apparently
based on his new material from “Tientsin”. Forsius (1927) followed Konow’s interpretation of
the species. Gussakovskij (1935) described boththe male and female and recorded the species
from North China, Ussuri, Khabarovsk andTransbaikalia. Takeuchi (1939) recorded A. flavicollis from Taiwan and synonymized A. vulnerata Mocsáry, 1909, with it. More recently, SainiArge captiva Species-groupand Thind (1995) redescribed A. flavicollis using
the Indian and Chinese material. Based on Sainiand Thind’s redescription, Wei and Wen (2000)
synonymized A. flavicollis with A. captiva(Smith, 1874), whereas Wei and Nie (2003) treated them as separate subspecies of A. captiva andSaini et al. (2006) again treated them as separate
species. All these works, except for the originaldescription by Cameron (1876) and the redescription by Kirby (1882), were not based on
the examination of the type material.
The main source of confusion may probably
date back to the misinterpretation of A. flavicollis
by Takeuchi (1939) who grouped it with A. rejecta, a species having a sharp median carina in the
supraclypeal area, in his key. He also synonymized A. vulnerata Mocsáry, 1909, a very
close relative of A. rejecta, with A. flavicollis.
Our examination of the type material has revealed that A. flavicollis is a junior synonym of
A. captiva, whose supraclypeal area has no sharp
median carina.
Cameron (1876) did not designate a holotype
of Hylotoma flavicollis and did not mention thenumber of specimens he had before him. Kirby(1882) listed only one specimen of this species as
the “Type of species”. We consider this to be thelectotype designation.
We have also examined the type material of
A. sanguinolenta Mocsáry, 1909, A. kolthoffi
Forsius, 1927, and A. captiva rufoscutellata
Takeuchi, 1927, and found that they all show no
significant differences from the lectotype of A.
captiva. As discussed by Blank et al. (2009), A.
captiva var. watanabei Takeuchi, 1932, is an infrasubspecific unavailable name. The “types”(holotype, allotopotype and four paratopotypes)
are very dark specimens of A. captiva collectedwith the other “normal” specimens in the sameplace on the same day.
Arge praesternalis Malaise, 1934
Arge praesternalis Malaise, 1934: 474; Saini and Thind,
1995: 106; Saini et al., 2006: 598.

Material examined.

INDIA: 1?, “India, U.

275

P., Bartot, 900 m, 13–6–83, M. Saini” “Argepraesternalis Mal., det. Saini” (USNM); 1?,“India: Himachal Pradesh: Kasol 1600 m, June30, 1984, M. Saini” “Arge praesternalis, H. P.,
Kasol – 1600 m, 30. 6. 1984” (USNM).
Distribution. Northern India.
Remarks. We were not able to study theholotype of A. praesternalis Malaise, 1934, described from Assam, India. Also without seeing
Malaise’s holotype, Saini and Thind (1995) treated A. praesternalis and A. flavicollis as two distinct species. The three characters used to separate the species are the malar space length/median ocellus diameter ratio, the 1st/2–4th metatarsomere lengths ratio, and the shape of the mesoscutellum. In our view, the differences in thesecharacters are not stable enough to distinguishspecies. We have examined two northern Indian
male specimens identified with A. praesternalis
by Saini (see above) and found no significant differences from the specimens of A. captiva in extragenitalic characters. Unfortunately the two Indian specimens have no genitalia. According to
Saini and Thind’s (1995) illustrations (figs. 52,
66), the lancet and the penis valve of their “A.
praesternalis” show some differences from thoseof A. captiva. We tentatively treat A. praesternalis as a valid species, though, judging from theoriginal description, this taxon may possibly be asynonym of A. captiva. We need more material
from India and particularly a study of Malaise’s
holotype to clarify the true identity of this taxon.
Acknowledgments
We wish to thank G. Broad and C. Taylor(BMNH), Y. Hashimoto (MNHAH), T. Hirowatari (OPU), H.-S. Lee (Suwon), S.-H. Lee(SNU), R. Matsumoto (OMNH), T. Naito and K.
Maeto (KU), M. Ohara (HU), P. Perkins (MCZ),
W. Pulawski (CAS), S. Schmidt (ZSSM), D. R.
Smith (USNM), H. Suda (Sakura), A. Taeger and
S. Blank (SDEI), H. Takahashi (Hachioji), H.
Vårdal and G. Lindberg (NRMS), L. Vilhelmsen(SNM), M.-C. Wei (CSFU), H. Yoshida (Kobe)
and L. Zombori (HNHMB) for making the mate-

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Akihiko Shinohara, Hideho Hara and Jung-Wook Kim

rial available for the present study. We also thank
B.-K. Byun (Pocheon), S.-W. Park (Suwon), N.
Shinohara (Tokyo) and X.-H. Wang (CSFU) for
their help in various ways. This study is partly
supported by the Grant-in-aid for Scientific Research No. 20580165 from the Japan Society for
the Promotion of Science.
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