Bull. Natl. Mus. Nat. Sci., Ser. A, 38(3), pp. 125–143, August 22, 2012

Arge obesa n. sp. (Hymenoptera, Argidae) Feeding on Carpinus andOstrya in Japan, with Taxonomic Notes on Hylotoma japonica
Hideho Hara1, Akihiko Shinohara2 and Shin-ichi Ibuki3
1

Forestry Research Institute Doto Station, Hokkaido Research Organization,
Shintoku, Hokkaido, 081–0038 Japan
E-mail: hara-hideho@hro.or.jp
2
Department of Zoology, National Museum of Nature and Science,
4–1–1, Amakubo, Tsukuba, Ibaraki, 305–0005 Japan
E-mail: shinohar@kahaku.go.jp
31355–13, Wami, Nakagawa, Tochigi, 324–0612 Japan(Received 21 May 2012; accepted 27 June 2012)

Abstract Arge obesa Hara and Shinohara, n. sp. is described from Hokkaido, Honshu, Shikoku
and Kyushu, Japan. The larva feeds on Carpinus cordata Blume, C. laxiﬂora (Siebold et Zucc.)
Blume, C. tschonoskii Maxim., and Ostrya japonica Sarg. (Betulaceae). The immature stages andlife history are also described. The sawﬂy is a multivoltine species with polymodal adult emergence. The lectotype of Hylotoma japonica Marlatt, 1898, is designated in order to ﬁx its synonymy with A. jonasi (Kirby, 1882).
Key words : Argidae, Arge obesa, new species, Carpinus, Ostrya, Hylotoma japonica, lectotype
designation, Japan.

Introduction
Larvae of the sawﬂy genus Arge are externalleaf feeders of eudicot trees and herbs such as the
Berberidaceae, Betulaceae, Caplifoliaceae, Ericaceae, Rosaceae, Sabiaceae, Salicaceae, andUlmaceae (Lorenz and Kraus, 1957; Okutani,
1967; Smith, 1989; Liston, 1995; Taeger et al.,
1998; Shinohara et al., 2011). The Betulaceae areutilized by several species in Japan. Larvae ofA. aenea Hara and Shinohara, 2008, A. pullata(Zaddach, 1859), A. solowiyofka (Matsumura,1911) and some unidentiﬁed or undescribed species feed on Betula (Hara and Shinohara, 2008a,
2008b; Hara et al., 2007; unpublished data). Arge
hakusana Togashi, 1997, may be associated with
Alnus (Togashi, 1997). We have found a newArge whose larva feeds on Carpinus and Ostrya.
This species is described here as A. obesa n. sp.
Hylotoma japonica Marlatt, 1898, describedfrom ﬁve syntype specimens from Honshu, Japan(Marlatt, 1898), was regarded as a junior synonym of A. jonasi (Kirby, 1882) by Konow(1906) and subsequent authors (Takeuchi, 1919,
1932, 1939; Gussakovskij, 1935; Taeger et al.,2010). Our examination of the syntypes has
shown that three syntypes are A. jonasi, but theother two are A. obesa and a species of the Spinarge fulvicornis group (Hara and Shinohara,2006), respectively. For the sake of stability, we
designate a syntype identiﬁed with A. jonasi as
the lectotype of Hylotoma japonica.
Materials and Methods
The material used in this study is kept in the
National Museum of Nature and Science, Tsukuba, unless otherwise indicated. Abbreviations
for the depositories are: HSC–H. Suda Collection, Sakura; HU–Hokkaido University, Sapporo;
KU–Kobe University, Kobe; MNHAH–Museumof Nature and Human Activities, Hyogo, Sanda;

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MYC–M. Yamada Collection, Kuroishi; NAC–
N. Aoki Collection, Abiko; OPU–Osaka Prefecture University, Sakai; USNM–National Museumof Natural History, Washington, D. C.
Observations of morphology were made with a
Leica MS5 stereo binocular microscope andmeasurements of each structure were taken withan ocular micrometer. Photographs were takenwith a ﬁlm camera, OLYMPUS OM-4, digital
cameras, Canon EOS Kiss Digital X, Konica
Minolta DiMAGE A200, Nikon E990, and Ricoh
Caplio GX100, Keyence Digital Microscope
VHX-900, and AnMo Electronics Dinolite digital
microscope. The digital images were processed
and arranged with Adobe Photoshop Elements
7.0 software.
Rearings were done in a room without air-conditioning in Bibai and Shintoku by Hara, in aroom without air-conditioning in Nakagawa by
Ibuki, and in an air-conditioned room in Tokyo
by Shinohara. In the rearing rooms in Bibai and
Shintoku, the temperature and light were almost
the same as in the open-air condition, but thehibernating individuals were moved in March or
April into an air-conditioned room, where thetemperature was about 10–25°C. In the rearing
room in Tokyo, the temperature was kept at
20–25°C. Both in Nakagawa and Tokyo, the light
was usually on for about 16 hours a day during
the feeding period; the larvae were kept indoors
but without air-conditioning during winter.
Results
Arge obesa Hara and Shinohara, n. sp.
[Japanese name: Maru-unmon-churenji](Figs. 1–9, Table 1)
Hylotoma japonica Marlatt, 1898, p. 504 (part).
Arge abelivora Okutani, 1956, p. 97 (part).
Arge suzukii: Shinohara and Hara, 2008, p. 30 (part, not
Matsumura, 1912).

Female (Fig. 1A–F). Length 6.5–9.5 mm.
Black, with faint metallic bluish reﬂection, often
faint brassy reﬂection on clypeus; subanal area
narrowly or widely blackish centrally. Flagellum

brown to blackish brown, with colorless reﬂection. Mandible apically reddish. Palpi darkbrown, apically sometimes somewhat pale. Legs
black, with colorless reﬂection; fore leg brownish on apex of femur; fore tibia and tarsus white
to brown, each not or slightly darkened apically;
middle and hind tibiae and tarsi white, each darkened apically, rarely hind tarsus darkened
throughout; hind femur rarely narrowly whitish
at base; hind tibia black on apical third to fourth;tibial spurs dark brown to black; claws apicallybrownish. Wings hyaline, slightly yellowish
basally and slightly brownish apically on fore
wing, and slightly brownish on hind wing, withdark transverse band below stigma, very rarely
entirely brownish (Fig. 1G); dark band below
stigma, if present, extending to posterior wing
margin, covering all of cells Rs and 2M; stigma
dark brown, basally narrowly pale, or yellow andcentrally darkened; veins dark brown to black,yellow on C, Sc, and most of section of R1 basal
to stigma; vein C darkened basally on anterior
margin. Setae whitish, brownish dorsally and on
sawsheath; wings with setae blackish, yellowish
on basal membranous areas of forewing.
Surface generally smooth and polished; punctures on anterior part of head relatively dense anddistinct, generally separated from each other.
Head in dorsal view (Fig. 3A) slightly concave
at outer orbits, with width across eyes slightly
narrower than or about as wide as width at genae.
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, with anterior furrow shallow andblunt, and lateral furrow distinct anteriorly,
sometimes indistinct throughout (Fig. 3A). Ocellar area weakly concave between ocelli (Fig. 3A,
D). Frontal area nearly ﬂat, not or slightly raisedanteriorly (Fig. 3B, D), rarely medially or posteromedially shallowly concave as in Fig. 3E;
lateral ridges converging anteriorly. Distance
between median fovea and front ocellus 1.2–
2.0×width of front ocellus. Median fovea almost
touching frontal area, very small or indistinct,
sometimes with weak minute tubercle below

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127

Fig. 1. Arge obesa n. sp. — A, B, Female, holotype (HH040810B, host Carpinus cordata), dorsolateral and ventral views; C, female, paratype (SI091005, host C. cordata), Nakagawa, dorsolateral view; D, do., (SI100707-2,
host C. tschonoskii); E, F, female, paratype (HH100829A, host Ostrya japonica), Shintoku, dorsolateral andventral views; G, female wings, paratype, Omogo; H, male, paratype (HH040810B), Iwamizawa, dorsolateralview; I, do., (HH100829A), Shintoku; J, do., Iwamizawa; K, L, ﬁnal instar larva (HH040810B), Iwamizawa,
dorsal and lateral views, photographed indoors on 11 August 2004; M, do., semiﬁnal instar, lateral view. K–M,
Photographs taken by H. Hara.

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Arge obesa n. sp. Feeding on Carpinus and Ostrya in Japan(Fig. 3D); weak depression sometimes present
below this tubercle. Interantennal carinae bluntor sharp, dorsally roundly curved medially andbecoming weak or inconspicuous, fused or separated from each other, ventrally weakened andseparated from each other (Fig. 3F), or fused into
carinate median supraclypeal ridge (Fig. 3H).
Median supraclypeal ridge rounded (Fig. 3F) or
bluntly carinate (Fig. 3H); side slope frommedian ridge rounded or ﬂattened medially androunded laterally, slightly rugulose. Malar space
0.7–1.1×width of front ocellus. Clypeus distinctly sunk below supraclypeal carina, ﬂattened
ventrally (Fig. 3B, F); ventral margin widelyroundly concave medially. Antennal length
1.0–1.2×maximum head width; ﬂagellum (Fig.
3I, J) scarcely compressed, distinctly curved
basally and narrowly rounded at apex in innerlateral (dorsal) view. Right mandible not incisedon inner margin (Fig. 3H).
In forewing, cell 1Rs2 with anterior length
0.9–1.1×posterior length and crossvein 3r-m
roundly curved (Fig. 1A, C–E); in both wings,
wing margin between veins Rs and Cu not ciliate, with width of marginal glabrous area about
2.0×width of vein M and length of marginal
setae about 1.5×width of vein M (Fig. 3M).
Abdomen with ﬁrst to fourth terga glabrous
above; ﬁfth and sixth, sometimes also seventh
terga medially almost glabrous; more posterior
terga setose. Seventh sternum with posterior margin medially roundly produced and sunk. Sawsheath in posterodorsal view (Fig. 4A–C) about
as long as or wider than long, basally widely
sunk, dorsally rounded, with apical margin narrowly or moderately rounded, lateral margin
weakly or moderately rounded, medial margin

129

edged at narrow base, and basal median lobe
small, in lateral view (Fig. 4D–F) with ventral
margin, except for basal convexity, roundly convex, dorsal margin nearly straight or slightly
roundly convex, and apex rounded; inner surface
spinose.
Lance without membranous areas (Fig. 4G, H)
and groups of minute setae along ventral margin(Fig. 4L); apical crest somewhat roughly or
ﬁnely serrate on dorsal margin (Fig. 4I–K). Lancet with dorsal margin slightly roundly convex(Fig. 5A–D), ventral margin rounded, distinctly
serrate, dorsally with inconspicuous membranousarea (Fig. 5I), apically with wide non-annulatearea, and with about 18–20 serrulae; marginal
sensillae long; longitudinal rows of minute setae
present before ﬁrst annulus, narrowing dorsally;basal two rows dorsally far apart from dorsal
margin of lancet; sclerotized part on ﬁrst annulus
small, far apart from dorsal margin of lancet;
sclerotized part on second annulus narrowing or
disappearing dorsally; middle annuli each withthree to six sensory pores except for ventral ones;
basal annuli ventrally curved basally; middle andapical annuli nearly straight, oblique, with ventral end located apical to dorsal end; serrulae
ﬁnely dentate (Fig. 5E–H), subtriangular withlong posterior slope.
Male. Length 5.5–7.0 mm. Coloration as infemale, but wings always entirely brownish, with
faint dark mark below stigma (Fig. 1H–J); veins
C and R1 basal to stigma sometimes mostly
black (Fig. 1J); middle tibia rarely entirely pale;
genital capsule brown to dark brown, with harpe
black (Fig. 6C, D). Head in dorsal view withwidth across eyes about as long as or slightly
longer than width at genae. Interantennal carinae

Fig. 2. Arge obesa n. sp. — A, Late instar larvae (AS100920A) and infested leaves of Carpinus laxiﬂora, Mt.
Haguro-san, photographed on 20 September 2010; B, C, part of larvae in A; D, ﬁnal instar larva (Ph910717B)
on Ostrya japonica, Shintoku, photographed on 17 July 1991; E, remains of egg shells and ﬁrst instar larvae(AS090927A) on Carpinus cordata, Nakagawa, photographed on 27 September 2009; F, G, remains of egg
shells and ﬁrst instar larvae, enlargement of E; H–J, ﬁnal instar larva (AS090926) on C. laxiﬂora, Nasukarasuyama, photographed indoors on 7 October 2009; I., do., apex of abdomen (arrow indicate hook-shaped
spines); K, L, ﬁnal instar larva (AS060929) on C. laxiﬂora, Kobe, photographed on 29 September 2006; M,
hook-shaped spines on subanal lobe in exuvia of ﬁnal instar larva (AS100920A, male, black-headed larva).
Photographs taken by A. Shinohara, except for D and M photographed by H. Hara.

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Fig. 3. A, Head, dorsal view; B, C, do., lateral view; D, E, ocellar and frontal areas, anterodorsal view; F, G,
supraclypeal area and clypeus, anterior view; H, supraclypeal are to mandibles, anterior view; I, J, female
antenna, inner lateral and ventral views; K, L, male antenna, inner lateral and ventral views; M, apical marginof forewing (vM: vein M). — A, B, D, F, I, J, M, Holotype; C, E, G, K, L, male paratype (brother of holotype); H, female, paratype, Iwamizawa.

sharp, dorsally blunt. Distance between eyes
1.1–1.3×vertical diameter of eye; eye with vertical diameter 1.4–1.6×horizontal diameter. Malar
space 0.7–0.9×width of front ocellus. Antennal

length 1.3–1.7×maximum head width; ﬂagellum
distinctly and slightly compressed (Fig. 3K, L),basally curved or gently curved throughout. In
forewing, crossvein 3r-m sometimes somewhat

Arge obesa n. sp. Feeding on Carpinus and Ostrya in Japan

131

Fig. 4. A–C, Sawsheath, posterodorsal view; D–F, do., lateral view; G, ovipositor, lateral view; H, lance, lateralview; I–K, apical part of lance, lateral view; L, ventral part of middle of lance (arrowed part in G enlarged).
— A, D, I, Holotype (HH040810B, host Carpinus cordata); B, E, paratype (SI091005, host C. cordata),
Nakagawa; C, F, H, K, paratype (HH100829A, host Ostrya japonica), Shintoku; G, L, paratype (SI100707-2,
host C. tschonoskii), Nakagawa; J, paratype (host C. tschonoskii), Sasayama. E, G, L, Reversed images.

angularly curved. Fifth to eighth abdominal terga
sparsely setose on wide medial areas. Subgenital
plate in ventral view with posterior margin apically widely truncate, slightly concave (Fig. 6D),or bilobate (Fig. 6N).
Genitalia as in Fig. 6; gonostipes in ventral
view narrowing apically, with medial margin
almost straight and apical width about as wide asbasal width of harpe (Fig. 6B, D, F, H). Harpe in
dorsal view tapering apically, about 1.5×as longas wide, with apex pointed. Valviceps in dorsal
view (Fig. 6A, C, E, G) with round lateral lobe atapex, in lateral view (Fig. 6I–M) nearly triangular, with apex nearly truncate and its dorsal andventral ends narrowly rounded, and dorsal
apodemal projection (dap) very small and far

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Hideho Hara, Akihiko Shinohara and Shin-ichi Ibuki

Fig. 5. A–D, Lancet; E–H, middle serrulae; I, dorsal part of lancet (arrowed part in B enlarged). — A, E, Holotype (HH040810B, host Carpinus cordata); B, F, I, paratype (SI091005, host C. cordata), Nakagawa; C, G,
paratype (SI100628, host C. laxiﬂora), Nakagawa; D, H, paratype (HH100829A, host Ostrya japonica), Shintoku. Each numeral indicates serrula number. A–C, E–I, Reversed images.

apart from ergot (erg).
Larva. Early to middle instar (Figs. 1M, 2G):
Head brownish yellow to pale brown, sometimes
with dark dorsomedial stripe, rarely with dark
stripe on gena; legs pale yellow, sometimes dark-

ened on coxae; trunk pale greenish yellow to palegreen, rather densely covered with pale setae,
sometimes dorsally with scattered black spots
and with apically blackish subspiracular lobe.
Final instar (Figs. 1K, L, 2B–D, H–J, K, L): In

Arge obesa n. sp. Feeding on Carpinus and Ostrya in Japan

133

Fig. 6. A, E, G, Male genitalia, dorsal view; B, F, H, do., ventral view; C, D, male genitalia and apical abdominal
segment; I–L, penis valve, lateral view (left dorsal); M, valviceps, lateral view (left dorsal) (dap: dorsal
apodeme, erg: ergot); N, apex of subgenital plate, ventral view; A–N, paratypes. — A, B, I, M, Brother ofholotype, HH040810B, host Carpinus cordata; C, D, AS100920A (emerged from black-headed larva), host C.
laxiﬂora, Mt. Haguro-san; E, F, K, SI100707-2, host C. tschonoskii, Nakagawa; G, H, L, N, Ph910717B, host
Ostrya japonica, Shintoku; J, SI100628, host C. laxiﬂora, Nakagawa.

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pale specimens, head pale greenish yellow tobrownish yellow, with or without dark dorsomedial stripe, legs pale yellow to yellowish green,sometimes slightly darkened basally, and trunk
pale whitish green to yellowish green, with orwithout pair of dark spots along dorsomedianline of thorax (Figs. 1K, L, 2B, D); head sometimes with dark dorsomedian stripe anteriorly
branching and extending towards eyes and withdark genal stripe (Fig. 2H); trunk sometimes withpair of dark spots along dorsomedian line fromprothorax to eighth abdominal segment (Fig. 2H,K); in dark specimens, head black, legs palegreen, basally black, trunk pale green, covered
with many small black spots dorsally and ventrally, with subspiracular lobe apically blackish(Fig. 2C); setae pale brownish, often partly
blackish.
Structure (late instar): Length 15.0 mm;
antenna cylindrical, rather short (Fig. 7C); clypeus with two pairs of setae; labrum with twopairs of setae; mandible with three setae on outersurface; maxillary palp with four palpomeres;
palpifer with two setae; labial palp with three
palpomeres; ﬁrst to ninth abdominal segments
each with three annulets (Fig. 7A); prolegs on
second to sixth and tenth segments, those on second to sixth elongate (Figs. 1L, 2J, 7A); trunk
rather densely setose; subspiracular lobe distinctly roundly produced (Fig. 7E, F); tenth tergum in dorsal view rounded apically, basallywith bulbous lobe (Fig. 7G, H) (this lobe usuallyinconspicuous as in Fig. 2H, K, probably deﬂating); subanal lobe slightly extending posteriorly
beyond suranal lobe, and with pair of hookshaped spines (Figs. 2I, M, 7G). In early to middle instars, spine on subanal lobe conical (Fig.
7I).
Cocoon. Creamy to pale brown. Length 8.0–
11.0 mm in female, 6.0–8.0 mm in male. Elongate oval, double walled; outer wall netted; innerwall parchment like.
Distribution Japan (Hokkaido, Honshu, Shikoku, Kyushu).
Material examined. Holotype (Fig. 1A, B): ♀,labelled “JAPAN, Hokkaido, Kurisawa, Manji,

em. 2004/8/28–9/2, H. HARA” and “Part.
HH040810B, 11 larvae on branch of Carpinus
cordata, col. 2004/8/10, cocoon. 8/12–13”.
Deposited in the National Museum of Nature andScience, Tsukuba. Paratypes: HOKKAIDO–2♂
(Ph910629C), Tokachi, Shintoku, larvae coll. 29.
VI. 1991, coc. VII., em. 27. VII. 1991, Host:
Ostrya japonica, H. Hara; 1♂ (Ph910629C), do.,but em. 15. V. 1992; 1♂ (Ph910717B), same
locality and host, larva coll. 17. VII. 1991, coc.
18. VII., em. 24. IV. 1992, H. Hara; 2♂, Mikasa,
Tomatsu, 22–29. VII. 2002, H. Hara; 1♂, Mt.
Yubari-dake, 10. VIII. 1966, T. Kumata (HU);
1♂, Iwamizawa, Manji, 7–23. VII. 2003, H.
Hara & N. Ishihama; 1♀7♂, do., but 23. VII.–7.
VIII. 2003; 1♀ (HH030830A), same locality,from 7 gregarious larvae coll. 30. VIII. 2003,
coc. 1–5. IX., em. 20. IX. 2003, Host: Carpinus
cordata, H. Hara; 1♀3♂(HH040810B), samedata as holotype; 1♀, same locality, 2. VII.
2006, on Carpinus cordata, 24 eggs on leaf margins of two adjascent leaves nearby, H. Hara; 1♂
(HH050806A), Mikasa, Ikushunbetsu, from 8
gregarious larvae coll. 6. VIII. 2005, coc. 11.
VIII., em. 2–4. IX. 2005, Host: Carpinus
cordata, H. Hara; 1♀, Sapporo, 6. VII. 1954,
M. Konishi (HU); 1♀, Sapporo, Moiwa, 3.
VIII. 1911, Matsumura (HU); 1♂, Sapporo,
Maruyama, 24. VI. 1930, K. Igarashi (HU); 1♂,
Sapporo, Ashiribetsu, 5. VIII. 1962, S. Takagi(HU); 1♀, Tomakomai, 20. VIII. 1957, K.
Kamijo (HU). HONSHU–Aomori Pref.: 1♀,
Fukaura, “Azumagawa”, 2. VII. 1989, M.
Yamada (MYC). Yamagata Pref.: 1♂
(AS100920A), Tsuruoka, Mt. Haguro-san, 320m,
larva coll. 20. IX. 2010, mat. 22–24. IX., em. 18.
V. 2011, Host: Carpinus laxiﬂora, A. Shinohara;
1♂ (AS100920A), do., but em. 19. V. 2011; 1♂
(AS100920A), do., but em. 27. V. 2011; 1♂
(AS100920B), do., but mat. 2–3. X., em. 20. X.
2010. Tochigi Pref.: 1♀, Nasushiobara, “Uwanohara”, 28. V. 2008, E. Katayama; 1♀, Utsunomiya, Mt. Hagurosan, 3. V. 1976, T. Saito; 1♀
(SI091005), Nakagawa, Bato, Sukusukunomori,
160 m, larva coll. 5. X. 2009, coc. 13. X., em. 7.
VI. 2010, Host: Carpinus cordata, S. Ibuki; 1♀

Arge obesa n. sp. Feeding on Carpinus and Ostrya in Japan

135

Fig. 7. A–H, Final instar larva (HH040810B), Iwamizawa; I, middle instar larva, Nakagawa. — A, Entire larva,
lateral view; B, head, anterior view; C, antenna, anterior view; D–F, ﬁrst to third abdominal segments, lateral,
dorsal and ventral views; G, I, ninth and tenth abdominal segment, lateral view; H, do., dorsal view.

(SI091005), do., but em. 10. VI. 2010, deposited
21 eggs (hatched 26. VI. but all died); 1♂
(SI091005), do., but em. 21. VI. 2010; 1♂
(SI091005), do., but em. 23. VI. 2010; 1♀
(SI091005), do., but em. 26. VI. 2010; 6♂
(SI100911), do., but larvae coll. 11. IX. 2010,
coc. 20–21. IX., em. 12. X. 2010; 1♀
(SI100628), Nakagawa, Wami, 230m, larva coll.

28. VI. 2010, coc. 9. VII., em. 31. VII. 2010,
Host: Carpinus laxiﬂora, S. Ibuki; 1♀1♂
(SI100628), do., but em. 2. IX. 2010; 1♀, Nakagawa, Bato, Bicchuzawa, 150 m, 8. V. 2008, A.
Shinohara; 1♀ (SI100707-2), Nakagawa, Bato,
Bicchuzawa, 125m, larva coll. 7. VII. 2010 on
Carpinus tschonoskii, reared with Carpinus laxiﬂora, coc. 16. VII., em. 12. IX. 2010, S. Ibuki;

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Hideho Hara, Akihiko Shinohara and Shin-ichi Ibuki

1♂ (SI100707-2), do., but em. 16. IX. 2010; 1♀
(SI100707-3), same locality, larva coll. 7. VII.
2010 on Carpinus tschonoskii, reared with Carpinus cordata, coc. 23. VII., em. 13. IX. 2010, S.
Ibuki; 1♂ (SI100912), same locality, larva coll.
12. IX. 2010, coc. 20. IX., em. 12. X. 2010,
Host: Carpinus cordata, S. Ibuki; 1♀
(SI110914), same locality, larva coll. 14. IX.
2011, coc. 28. IX., em. 26. X. 2011, Host: Carpinus cordata, S. Ibuki; 1♂ (SI110914), do., coc.
27. IX., em. 15. XI. 2011; 1♂ (SI110914), do.,but coc. 28. IX., em. 19. XI. 2011; 1♀
(SI110914), do., but coc. 28–29. IX., em. 25. XI.
2011; 1♂ (SI110914), do., but coc. 27–28. IX.,
em. 1. XII. 2011. Gunma pref.: 3♂ (AS100909),
Katashina, Marunuma, 1400 m, larvae coll. 9. IX.
2010, mat. 19–20. IX., em. 5. X. 2010, Host:
Carpinus cordata, A. & N. Shinohara; 1♂
(AS100909), do., but em. 8. X. 2010. Chiba
Pref.: 2♀19♂, Sakura, Iida, 31. V. 1974, H.
Suda (HSC); 1♀, Sakura, Miroku-machi–
Kaburagi-machi, 5. VI. 1974, H. Suda (HSC);
1♀, Sakura, Mogami, 5. VI. 1974, H. Suda
(KU); 1♂, Sakura, Yamanozaki, 31. V. 1974, H.
Suda (KU); 1♀, Kashiwa, Neto, 6. IX. 2008, N.
Aoki (NAC); 1♀1♂ in copulation, Matsudo,“15. 5. 30”. J. Yoshikawa; 1♀, do., but “2600Fig. 8.

5-30”; 1♂, Ichikawa, “2597 6-5”, J. Yoshikawa;
1♀1♂, do., but “2600 5-29”; 12♂, do., but “155 29”; 1♀, Funabashi, Narashinohara, 29. VIII.
1958, H. Suda (HSC). Tokyo Met.: 1♀,
Musashino, Gotenyama, 12. VII. 2000, H. Takahashi; 1♀, Hachioji, Mt. Takaosan, 1917. V. 27(KU). Kanagawa Pref.: 1♀, Sagamihara,
Kamimizo, 27. V. 1990, M. Tao; 1♀, Oiso, 28.
VII. 1974, K. Kimura. Yamanashi Pref.: 1♂,“Yutsubo”, 29. V. 1932, Takeuchi (OPU). Nagano
Pref.: 1♀, Kamikochi, 5. VII. 1927, K. Sato;1♂, Matsumoto, Shimashima-dani, 13. VII.
1928, K. Sato. 1♀, same locality, 30. VI. 1976,
A. Shinohara. Ishikawa Pref.: 1♀, Nakanoto,
Kashima, Ida, 17. VII. 1996, I. Togashi. Gifu
Pref.: 1♀ (paralectotype of Hylotoma japonica;
see below), “25, 6, 24 [Gifuyama]” “Japan Mitsukuri” (USNM). Mie Pref.: 1♀, “Hirakura, 36. 7.
8”, M. Matsuura (KU). Kyoto Pref.: 1♀,“Kibune”, 13. VI. 1930, Takeuchi (OPU). Hyogo
Pref.: 1♀, Sayo, Onemi, 11. VI. 1995, T. Morita(MNHAH); 4♀, Sasayama, “Kogane”, em. 3.
VIII. 1965, Host Carpinus tschonoskii, T. Okutani (KU); 1♀ (paratype of Arge abelivora Okutani, 1956), Sasayama, Fujioka, 12. VI. 1954, T.
Okutani (KU). Hiroshima Pref.: 1♀, “Sandankyo, Furinkan–Koeda”, 11. VI. 1939, T. Nakani-

Number of adults collected aﬁeld and those emerged by rearings in lowlands of Honshu, Japan.

Arge obesa n. sp. Feeding on Carpinus and Ostrya in Japan

Fig. 9.

137

Cocoon durations of non-hibernating individuals under rearing conditions.

shi (OPU). SHIKOKU–Ehime Pref.: 1♀,
Omogo-kei, 27. V. 1986, A. Shinohara; 1♀,
Omogo, Ishizuchi Mts., Tsuchigoya, 23. V. 1999,
M. Shiraishi. KYUSHU–Nagasaki Pref.: 2♀,
Sasebo, Mt. Kunimi-yama, 16. VI. 1950, T. Shirozu (OPU). Other material: Larvae in ethanol(see Table 1 for collection data): 3 late instar larvae (part of Ph910629C), ﬁxed on 18. VII. 1991;
2 mature larva (part of HH030830A); 3 lateinstar larvae (part of HH040810B), ﬁxed on 12.
VIII. 2004; 3 late instar larvae (part of
HH060820A); 1 middle instar larva (part of
HH060820B); 2 late instar larvae (part of
HH060820C); 3 late instar larvae (part of
HH060820D); 3 late instar larvae (part of
HH060820E); 2 mature larvae (part of
HH070814A), ﬁxed on 27. VIII. 2004; 2 lateinstar larvae (part of AS080614), ﬁxed on 30. VI.
2008; 10 middle instar larvae, Nakagawa, Bato,coll. 7. VII. 2010, S. Ibuki; 10 late instar larvae(part of AS101007), ﬁxed on 7. X. 2010.
Etymology. The speciﬁc epithet is from theLatin obesus, meaning plump, referring to therelatively plump body.
Host plants. Betulaceae: Carpinus cordata
Blume, C. laxiﬂora (Siebold et Zucc.) Blume, C.
tschonoskii Maxim. and Ostrya japonica Sarg.
Field observations and rearing records. Adults
were collected from late June to late August in
Hokkaido, and from early May to early September in Honshu, most frequently in May and Junein the lowlands of Honshu (Fig. 8). Two groupsof eggs or their remains were observed in theﬁeld. The number of eggs per group was 22–24.

Eggs were laid along leaf margins of one or two
adjacent leaves in rows (Fig. 2E). Larvae were
found from late June to late August in Hokkaido,
and from middle June to early July and early
September to middle October in Honshu (Table1). They were usually gregarious (n＝1–20), but
separately fed on leaves in all instars (Fig. 2A,
E). Early instar larvae fed on leaves leaving thelateral and main veins, while late instar larvae
left only main veins. Larvae spun cocoons withinsoil in the laboratory.
Rearings are summarized in Table 1. In Hokkaido, larvae became adults in the same year or
the following year. This variation was observed
even in the same group of larvae (Ph910629C).
Cocoon durations were about 17–23 days in nonhibernating individuals and about 281–306 daysin hibernating individuals. In Honshu, larvae collected in June and July became adults in late Julyand early to middle September in the same year.
Larvae collected in September and Octoberbecame adults in October to December of thesame year or May and June in the following year.
Cocoon durations in non-hibernating individuals
greatly varied, ranging from 15 to 65 days. The
adult emergence was clearly bimodal (Fig. 9right), the ﬁrst cluster with almost the same
cocoon duration as non-hibernating individualsof Hokkaido (Fig. 9 left). Cocoon durations inhibernating individuals were about 237–256
days. The durations were variable among the larvae of the groups SI100628 (22–55 days) and
SI110914 (29–65 days), but were rather stablewithin each of the groups AS100920A (all hiber-

138

Hideho Hara, Akihiko Shinohara and Shin-ichi Ibuki
Table 1.
Locality

Prefecture Details

Shintoku, 200 m

Mikasa, Ikushunbetsu, 100 m

Hokkaido

Yamagata

Iwamizawa,
Manji, 210 m

Ostryajaponica

Carpinus
cordata

Carpinus
cordata

Larval group
code*

Found

Number oflarvaeCocooned

Emerged (number
and sex**)27. VII. 1991 (2♂)
15. V. 1992 (1♂)

Ph910629C

29. VI. 1991

6

VII.

Ph910717B

17. VII. 1991

unrecorded

18. VII.

HH100829A

29. VIII. 2010

unrecorded

1–5. IX.

HH050806A

6. VIII. 2005

8

11. VIII.

2–4. IX. 2005 (1♂)

HH030830A

30. VIII. 2003

7

1–5. IX.

20. IX. 2003 (1♀)

HH040810B

10. VIII. 200411

12–13. VIII.

28. VIII.–2. IX.
2004 (2♀3♂)

HH060820A

20. VIII. 20069

22–25. VIII. dead in cocoons

HH060820B

20. VIII. 2006

6

dead before
maturity

HH060820C

20. VIII. 20064

dead before
maturity

HH060820D

20. VIII. 20068

22–25. VIII. dead in cocoons

HH060820B

20. VIII. 20066

22–25. VIII. dead in cocoons

24. IV. 1992 (1♂)
6–7. VI. 2011(1♀1♂)

HH070814A

14. VIII. 2007

7

24. VIII.

dead in cocoons

Mukawa,Hobetsu, 100 m

Carpinus
cordata

HH070814B

14. VIII. 2007

4

27. VIII.

dead in cocoons

HH070814C

14. VIII. 2007

2

24. VIII.

dead in cocoons

Urakawa,
Nishicha, 30 m

Carpinus
cordata

HH070813A

13. VIII. 2007

930. VIII.

dead in cocoons

Mt. Haguro-san,
320 m

Carpinus
laxiﬂora

AS100920A

20. IX. 2010

unrecorded

22–24. IX.

18. V. 2011 (1♂)
19. V. 2011 (1♂)
27. V. 2011 (1♂)

AS100920B

20. IX. 2010

unrecorded

2–3. X.

20. X. 2010 (1♂)

Nakagawa,
Sukusukunomori,160 mTochigi

Host plant

Summary of rearings.

Nakagawa, Wami,
230 m

Nakagawa,
Bicchuzawa,
125 m

SI091005

5. X. 20098

13. X.

7. VI. 2010 (1♀)
10. VI. 2010 (1♀)
21. VI. 2010 (1♂)
23. VI. 2010 (1♂)
26. VI. 2010 (1♀)

SI100911

11. IX. 20109

20–21. IX.

12. X. 2010 (6♂)

Carpinus
laxiﬂora

SI100628

28. VI. 2010

ca. 10

9. VII.

31. VII. 2010 (1♀)
2. IX. 2010(1♀1♂)

Carpinustschonoskii

SI100707-1

7. VII. 201012from 14.
VII.

dead in cocoons

Carpinustschonoskii/
C. laxiﬂora

SI100707-2***

7. VII. 20103

16. VII.

12. IX. 2010 (1♀)
16. IX. 2010 (1♂)

Carpinustschonoskii/
C. cordata

SI100707-3***

7. VII. 20103

23. VII.

13. IX. 2010 (1♀)

Carpinus
cordataArge obesa n. sp. Feeding on Carpinus and Ostrya in Japan
Table 1.
Locality

Host plant

Prefecture Details

Tochigi

Nakagawa,
Bicchuzawa,
125 m

Carpinus
cordata

Larval group
code*

139

(Continued)
Found

Number oflarvaeCocooned

Emerged (number
and sex**)

AS090927A

27. IX. 200910

9–13. X.

dead in cocoons

AS091018

18. X. 200914

20–22. X.

dead in cocoons

SI100715

15. VII. 20101

16. VII.

dipterous parasite
emerged 12. IX.

SI100912

12. IX. 20105

20. IX.

12. X. 2010 (1♂)

AS101007

7. X. 2010

unrecorded

7–12. X.

dead in cocoons

SI110914

14. IX. 2011

unrecorded

27–29. IX.

26. X. 2011 (1♀)
15. XI. 2011 (1♂)
19. XI. 2011 (1♂)
25. XI. 2011 (1♀)
1. XII. 2011 (1♂)

Nasukarasuyama,
Ryumon-no-taki,
90 m

Carpinus
laxiﬂora

AS090926

26. IX. 20098

10–11. X.

dead in cocoons

Gunma

Katashina,
Marunuma,1400 m

Carpinus
cordata

AS100909

9. IX. 20106

19–20. IX.

5. X. 2010 (3♂)
8. X. 2010 (1♂)

SaitamaNamegawa, 50 m

Carpinustschonoskii/
C. laxiﬂora

AS080614

14. VI. 200820

25–30. VI.

dead in cocoons

Mie

Matsuzaka,
Iitaka-cho, Kikaji,
400m

Carpinus
laxiﬂora

AS091011

11. X. 20094

dead before
maturity

Hyogo

Kobe, Yamadamachi, 450m

Carpinus
laxiﬂora

AS060929A

29. IX. 2006

1

1. X.

dead in cocoons* Reared by Hara in Bibai or Shintoku (Ph, HH), by Shinohara in Tokyo (AS) and by Ibuki in Nakagawa (SI).
** Other specimens died or were ﬁxed in larval stage.
*** Larvae collected on C. tschonoskii, but reared on C. laxiﬂora (SI100707-2) and C. cordata (SI100707-3),
respectively.

nated), SI091005 (all hibernated), and AS100909(15–19 days).
Discussion
Comparison with Related Species
Arge obesa is one of the blue-black species
having hyaline wings with a black transverseband on the forewing, a basally pale hind tibia
and pale setae on the mesopleuron, though themale has entirely dark wings without a conspicuous transverse band. This species is characterized
by the lancet whose annuli are oblique and theirventral ends are located apical to the dorsal ends
in the female (Fig. 5A–D), and the almost trian-

gular and apically truncate valviceps (in lateralview) in the male (Fig. 6I–M). A combination of
the following characters is also useful to distinguish this species: Abdomen without pale areas(Fig. 1); frontal area nearly ﬂat, sometimes withshallow medial or posteromedial concavity,
sometimes slightly raised anteriorly (Fig. 3D, E);
median fovea very small, often inconspicuous;
median ridge of supraclypeal area rounded or
bluntly carinate (Fig. 3F–H); clypeus distinctly
sunk below supraclypeal area, and ﬂattened ventrally (Fig. 3B, C, F–H); right mandible without
notch on inner margin (Fig. 3H); in both wings,
wing margins between veins Rs and Cu not ciliate, with wide marginal glabrous area (Fig. 3M);

140

Hideho Hara, Akihiko Shinohara and Shin-ichi Ibuki

in female, seventh sternum normal; sawsheath inposterodorsal view robust and basally widely
sunk (Fig. 4A–C); lance without linear membranous areas (Fig. 4G, H); in male genitalia,
gonostipes narrowing apically in ventral view(Fig. 6B, D, F, H).
In Gussakovskijʼs (1935) key to the Palearctic
species, the female of A. obesa may run tothe couplet 151/152, containing A. potanini
Jakowlew, 1891, and “A. fulvicornis Mocs.”(=Spinarge fulvicornis (Mocsáry, 1909)), and themale to the couplet 160/161, containing A. ciliaris (Linné, 1767) and “A. rufocincta Gussakovskij, 1935” (=A. longicornis KuznetzovUgamskij, 1927; see Shinohara et al., 2012). In
Takeuchiʼs (1939) key to East Asian species, A.
obexa usually runs to the couplet 25, which contains A. kobayashii Takeuchi, 1931, and A. jonasi(Kirby, 1882), and a part of the male to “A. rufocincta” (=A. longicornis) or A. mali (Uchiyama,1906). The female of this species is distinguishedfrom A. potanini by the black, not bluish legs
(green blue in the latter; see Jakowlew, 1891),
from Spinarge fulvicornis by the ﬁfth abdominal
tergum without a median suture (with a mediansuture in the latter; see Hara and Shinohara,2006), from A. kobayashii and A. jonasi by thesmaller size, 6.5–9.5 mm (usually more than
10 mm in the latter two) and the apically roundedsawsheath in posterodorsal view (nearly pointed
in the latter two; see Hara et al., 2007). The maleis distinguished from A. ciliaris, A. longicornis,
A. kobayashii, A. jonasi and A. mali by the combination of the following characters: Reﬂection
faintly metallic bluish; middle and hind tibiae
basally pale; wings distinctly infuscated all over;frontal area not or slightly raised anteriorly; apical wing margins not ciliate; harpe narrow in
dorsal view (Benson, 1951; Hara et al., 2007;
Shinohara et al., 2012). This species is also similar to A. sinensis Wei, 2003 in Wei and Nie(2003) from China, but differs from the latter inhaving the widely white middle tibia, the robust
sawsheath, and the characteristic valviceps statedabove. In A. sinensis, the middle tibia has nowhite areas, the sawsheath is narrower (ﬁg.

28-174A in Wei and Nie, 2003), and the valviceps is rounded apically and with a large ventral
lobe basally in lateral view (ﬁg. 28-174C in Wei
and Nie, 2003).
The type series of A. obesa includes a paralectotype of Hylotoma japonica Marlatt, 1898 (=A.
jonasi (Kirby, 1882); see discussion below) and aparatype of Arge abelivora Okutani, 1956 (=A.
suzukii Matsumura, 1912). Arge obesa is separated from A. suzukii by the entirely black abdomen, the rounded or bluntly carinate supraclypeal ridge, the robust sawsheath, and theglabrous outer margin of the wings (Shinohara
and Hara, 2008).
The larva of A. obesa is easily distinguishedfrom those of Japanese congeners by the hostplant and the pair of conical or hook-shaped
spines on the subanal lobe (Figs. 2I, M, 7G, I). A
pair of subanal spines are also found in the North
American A. coccinea (Fabricius, 1804) associated with Rhus (Smith, 1989) within the genus
and in the Sterictophorinae of the Argidae(Lorenz and Kraus, 1957).
The extremely dark color variant of the last
instar larva as noted in the description (Fig. 2C)
has been found so far on Mt. Hagurosan,
Yamagata Prefecture, northern Honshu, only.
Further search for the larvae in various localities
covering the whole range of this species may
reveal the existence of geographical variations oflarval characters as in the case of A. enkianthus
Hara and Shinohara, 2011 (Hara and Shinohara,2011).
Life History
Arge obesa probably has a multivoltine life
cycle with polymodal adult emergence. This general life history pattern is rather commonly foundin the genus (Shinohara and Hara, 2008, 2009;
Shinohara et al., 2009; Hara and Shinohara,
2011; Kawasaki et al., 2012). This species
showed different polymodal adult emergence
patterns between Hokkaido and Honshu populations under rearing conditions (Fig. 9). In Hokkaido, the larvae collected in the ﬁeld partly
became adults within the same year without dia-Arge obesa n. sp. Feeding on Carpinus and Ostrya in Japanpause (ﬁrst cluster) and partly became adults inthe following year after winter diapause or hibernation (second cluster) even in the same larvalgroup. This bimodal pattern is therefore formedby the presence or absence of winter diapause
in each individual. In Honshu, however, weobserved two clusters of adult emergence before
hibernation (Fig. 9) and a cluster of adult emergence after hibernation (Fig. 8). The individualsof the ﬁrst cluster of emergence in Honshu had
no diapause, as in those of Hokkaido population,and the cocoon durations in the ﬁrst clusters were
almost identical in both areas (Fig. 9). The individuals of the second cluster of emergence probably had short summer diapause. The adult emergence after hibernation (third cluster) may havepossibly occurred as a result of prolonged diapause caused by low temperature.
Kawasaki et al. (2012) detailed a trimodal
adult emergence of the summer generation of A.
nigronodosa (Motschulsky, 1860), and considered that the ﬁrst and second clusters of emergence consisted of individuals that had genetically different cocoon durations without
diapause, while the third emergence occurred
after summer diapause without genetically determined timing. The proportion of the third emergence within the progeny of a female was genetically determined; namely, every motherproduced both non-diapausing and diapausing
individuals. Our rearings of A. obesa showed thatthe cocoon durations within each larval groupwere stable or variable. All individuals in thegroups HH100829A, AS100920A, and SI091005
went into diapause and those in the groups
HH040810B and AS100909 did not, whereas
both non-diapausing and diapausing individuals
were contained in the groups Ph910629C,
SI100628 and SI110914 (Table 1). The polymodal adult emergence in A. obesa may be caused
by genetic polymorphism. However, our rearing
material of A. obesa is very small and rearing
conditions are not constant. More extensive anddetailed study will be needed to conﬁrm our supposition.
In lowlands of Honshu, adult occurrences in141

the ﬁeld agree with adult emergence in rearings
at least during May through September (Fig. 8).
The delayed emergence in October to December
in rearings apparently occurs only under laboratory conditions.
This is the ﬁrst record of Arge associated with
Carpinus and Ostrya in the Old World (cf.
Lorenz and Kraus, 1957, Okutani, 1967, Liston,
1995, Taeger et al., 1998). In North America,“Arge cerulea (Norton, 1864)” (= A. smithi
Blank, Liston and Taeger, 2009) and A. sp. areassociated with Carpinus americana Michx., andA. sp. with Ostrya sp. (Smith, 1989). All those
two or three species belong to the A. clavicornis
group, whose larvae are characterized by thepresence of prolegs on the second to seventh oreighth and tenth abdominal segments. Arge
obesa does not belong to this species group, as
the larva has prolegs on the second to sixth andtenth abdominal segments.
Lectotype Designation of Hylotoma japonica
Marlatt, 1898
In the original description of Hylotoma japonica, Marlatt (1898) wrote “Type. ― No. 3838,
U.S.N.M.” “Described from ﬁve specimens, twomales and three females” and “(Gifu zuzushi),(No. 12).” He did not designate a holotype. We
examined ﬁve specimens preserved under thename of H. japonica in USNM. Actually, they
consist of four females and one male of three different species, but we regard them as the syntypes of Hylotoma japonica. Only two females
have type labels, but one of them is erroneous.
One female is labeled “♀ Type No. 3838” andagrees with the original description. Anotherfemale bears the label, “♂ Type No. 3839”,
which is the number given to Lophyrus japonicus
Marlatt, 1898, described in the same paper, andthe sex is incorrect. The former female is the
only specimen labeled Hylotoma japonica. We
designate the female as the lectotype here,
although the female is directly pinned, and,
therefore, does not ﬁt Marlattʼs (1898) statementin the preface, “All of the specimens are mounted

142

Hideho Hara, Akihiko Shinohara and Shin-ichi Ibuki

on large ﬂat cards”. Another female having thetype label and one male are also directly pinned.
The pinnings were not subsequently done, judging from their conditions. In mounted specimens(two females), the locality data [Gifu] or[Gifuyama] (in Japanese letters) are written on
the underside of the mount cardboard, while eachof the three pinned specimens has a thinner paperlabel noted [Gifu], and the handwritings on themount cardboards and the thinner paper labels
are similar to each other.
Lectotype (here designated): ♀ directly
pinned, with four thin paper labels, “22, 5, 20[Gifu] (in Japanese letters) ♀” “Japan Mitsukuri”“Hylotoma japonica Marlatt ♀” and “♀ TypeNo. 3838 U.S.N.M.”. Paralectotypes: 1♀
directly pinned, with three thin paper labels, “22,5, 9 [Gifu] ♀” “Japan Mitsukuri” and “♂ TypeNo. 3839 U.S.N.M.”; 1♀ mounted on a card
written “26, 5, 25 [Gifu] ♀ #25”; 1♀ mounted
on a card written “25, 6, 24 [Gifuyama]” and athin paper label “Japan Mitsukuri” (the specimen
was removed from a mount card and directly
pinned in the present study); 1♂ directly pinned,
with two thin paper labels, “20, 6, 19 [Gifu] ♂”and “Japan Mitsukuri”. All in USNM.
The lectotype and the former two paralectotypes are A. jonasi. The third paralectotype(female) is A. obesa described in this paper. This
paralectotype disagrees with the original description of Hylotoma japonica by its smaller size,
length 8 mm (12 mm for the female and 8 mm forthe male in the original description). Marlatt(1898) might have regarded it as a male. The lastparalectotype (male) belongs to the group of Spinarge fulvicornis (Mocsáry, 1909), whose species are not distinguishable in males (Hara andShinohara, 2006).
Acknowledgements
We thank D. R. Smith (USNM) for his carefulreading of the manuscript and the loan of material, and N. Aoki (Abiko), Y. Hashimoto(MNHAH), T. Hirowatari (OPU), E. Katayama(Otawara), K. Kimura (Tokyo), K. Maeto (KU),

M. Ohara (HU), T. Saito (Nikko), N. Shinohara(Tokyo), H. Suda (Sakura), H. Takahashi(Hachioji), M. Tao (Sagamihara), I. Togashi(Hakusan) and M. Yamada (Kuroishi) for making
material available.
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