Bull. Natl. Mus. Nat. Sci., Ser. A, 47(4), pp. 163–188, November 22, 2021
DOI: 10.50826/bnmnszool.47.4_163

Taxonomic Notes and New Distribution and Host Plant Records for
Sawﬂies and Woodwasps (Hymenoptera, Symphyta) of Japan VI
Hideho Hara1, Shinichi Ibuki2 and Akihiko Shinohara3
1

Nishi 4 Kita 3–4–29, Bibai, Hokkaido 072–0033, Japan
E-mail: harahideho@bell.ocn.ne.jp
2
Wami 1355–13, Nakagawa, Tochigi 324–0612, Japan
E-mail: banbi-fa@ktd.biglobe.ne.jp
3
Department of Zoology, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan
E-mail: shinohar@kahaku.go.jp(Received 9 September 2021; accepted 22 September 2021)

Abstract Two larch sawﬂies, Euura imperfecta (Zaddach, 1876) and E. itoi (Okutani, 1955)(Tenthredinidae), are redescribed. New distribution records are: Arge similis (Snellen van Vollenhoven, 1860) (Argidae) from Amami-Oshima Island and Phylloecus etorofensis (Takeuchi, 1955)(Cephidae), Apareophora nebuta Togashi, 1964, Ardis pallipes (Serville, 1823), Eriocampopsis
subtruncata Takeuchi, 1952, Eutomostethus pumicosus Seiyama, 1981, Stethomostus babai Togashi,
1984 and Tomostethus nigritus (Fabricius, 1804) (Tenthredinidae) from Hokkaido. Information on
the distribution of Phymatocera nipponica Togashi, 1958 (Tenthredinidae) is given. New hostrecords are Rosa multiﬂora Thunb. (Rosaceae) for Sterictiphora nipponica Takeuchi, 1939 (Argidae),Alnus japonica (Thunb.) Steud. (Betulaceae) and Juglans mandshurica Maxim. var. sachalinensis(Komatsu) Kitam. (Juglandaceae) for Tremex apicalis Matsumura, 1912 (Siricidae), Actinidia
polygama (Siebold et Zucc.) Planch. ex Maxim. (Actinidiaceae), Symplocarpus renifolius Schott ex
Tzvelev. (Araceae) and Impatiens textorii Miq. (Balsaminaceae) for Aglaostigma albicinctum(Takeuchi, 1953) (Tenthredinidae), Sanguisorba oﬃcinalis L. (Rosaceae) for Allantus meridionalis(Takeuchi, 1933) (Tenthredinidae), Quercus serrata Murray (Fagaceae) for Apethymus kunugi
Togashi, 2005 (Tenthredinidae), Sanguisorba oﬃcinalis L. (Rosaceae) for Corymbas nipponica
Takeuchi, 1936 (Tenthredinidae), Alnus hirsuta (Spach) Turcz. ex Rupr. var. hirsuta (Betulaceae) for
Masaakia longivaginata Takeuchi, 1950 (Tenthredinidae), Rubus hirsutus Thunb. (Rosaceae) for
Metallus albipes (Cameron, 1875) (Tenthredinidae), Rubus microphyllus L.f. (Rosaceae) for Metallus
pumilus (Klug, 1816) (Tenthredinidae), Rubus parvifolius L. (Rosaceae) for Monophadnoides
montanus Togashi, 1980 (Tenthredinidae) and Corylus sieboldiana Blume var. sieboldiana (Betulaceae) for Pristiphora fulviceps Takeuchi, 1933 (Tenthredinidae), Actinidia polygama (Siebold etZucc.) Planch. ex Maxim. (Actinidiaceae) and Glechoma hederacea L. subsp. grandis (A. Gray) H.
Hara (Lamiaceae) for Siobla ferox (Smith, 1874) (Tenthredinidae), Impatiens textorii Miq. (Balsaminaceae) and Circaea mollis Siebold et Zucc. (Onagraceae) for Siobla japonica Shinohara, Wei &Niu, 2013 (Tenthredinidae) and Agrimonia pilosa Ledeb. var. japonica (Miq.) Nakai (Rosaceae) for
Taxonus shinoharai Togashi, 2009 (Tenthredinidae). Information on the host plants of Strongylogastermoiwana Matsumura, 1912 (Tenthredinidae) is given. The larvae of Sterictiphora nipponica,
Aglaostigma albicinctum, Allantus meridionalis, Apethymus kunugi, Corymbas nipponica, Masaakia
longivaginata, Monophadnoides montanus, Siobla ferox, Siobla japonica, Strongylogaster moiwana
and Taxonus shinoharai are brieﬂy described and their color photographs are given for the ﬁrst time.
Key words: Argidae, Cephidae, Siricidae, Tenthredinidae, redescriptions, new distribution
records, new host plant records.
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Introduction

Results and Discussion

In the sixth paper of the series on Japanese
sawﬂies and woodwasps, we treat 26 species ofthe families Argidae, Cephidae, Siricidae andTenthredinidae. The redescriptions of Euura
imperfecta (Zaddach, 1876) and E. itoi (Okutani,1955), a well-known larch pest, are presented,
because their previous descriptions are partly
wrong and therefore it has been diﬃcult to distinguish them correctly. These two species havebeen included in Larinematus Zhelochovtsev,
1988, the E. imperfecta group, etc. (e.g., Zhelochovtsev and Zinovjev, 1988; Zinovjev, 1993;
Lacourt, 2020). We make some comments on thisgroup. Information on distribution, host plants
and larvae is also given for two species of Argidae, one species each of Cephidae and Siricidae
and 20 species of Tenthredinidae.

ArgidaeArge similis (Snellen van Vollenhoven, 1860)

Materials and Methods
The material used in this study is kept in the
National Museum of Nature and Science, Tsukuba. Morphological examination was undertaken with a Leica MS5 and an Olympus SZ60
stereo binocular microscopes and Olympus BH-2
light microscope. Photographs were taken with
Canon PowerShot S95, Casio EX-ZR1000,
Olympus TG-4, Olympus TG-5, Panasonic
DMC-FZ28 and Ricoh GR DIGITAL 2 digital
cameras and a Sony DSC-RX100 digital camera
with a Leica MS5 microscope or an Olympus
BH-2 light microscope. The digital images were
processed and arranged with GIMP 2.10 and
Adobe Photoshop Elements 9 and 12 software.
Rearing was done in rooms in Bibai, Hokkaido,
and Nakagawa, Tochigi Prefecture, where thetemperature and day length were not controlled.
For the morphological terminology, we generally
follow Viitasaari (2002).

Japanese name: Ruri-churenji

Material examined. KYUSHU: Kagoshima
Pref.: 1 ♂, Amami-Oshima Island, Amami,
Nesebu, 7. IV. 2019, M. Kimura.
Distribution. Japan: Southern Kuril Islands,
Hokkaido, Honshu, Izu Islands, Oki Islands,
Awaji Island, Shikoku, Kyushu, Tsushima Island,
Tanegashima Island, Yakushima Island, Tokara
Islands, Amami-Oshima Island (new record),
Okinawa Island. East Palaearctic and Oriental
regions.
Remarks. This species is widely distributed in
the East Palaearctic and Oriental regions including various parts of Japan (e.g., Hara, 2019), but
it has not been recorded from Amami-OshimaIsland until now.
Sterictiphora nipponica Takeuchi, 1939
Japanese name: Kuwagata-habachi(Fig. 1A)

Material examined. HONSHU: Tochigi Pref.:1 ♀, Nakagawa, Hanatate-toge, 36°47′N
140°12′E, coll. larva, on Rosa multiﬂora, 25. V.
2015, mat. 30. V., em. 28. III. 2016, S. Ibuki.
Larva. Late instar larva pale green (Fig. 1A);
setae inconspicuous; spiracles dark green; thoracic legs apically brownish.
Host plants. Rosaceae: Rosa luciae Rochebr.
et Franch. ex Crèp. (Okutani, 1967a), R. multiﬂora Thunb. (new record).
Life history. This species is univoltine. The
larva is solitary and makes a cocoon in the soil.
Hibernation is done in the cocoon.
Remarks. The previously unknown larva of
Sterictiphora nipponica is very similar to the larvae of other congeners in appearance (cf. Macek
et al., 2020). The known host plants are onlyRosa.

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Fig. 1. A, Late instar larva of Sterictiphora nipponica, 25. V. 2015, photographed by Ibuki. B, Phylloecus etorofensis, female, photographed by Hara. C, Dead female of Tremex apicalis on trunk of Alnus japonica, 23. V.
2021, photographed by Shinohara. D, Apethymus kunugi, female, photographed by Hara.

CephidaePhylloecus etorofensis (Takeuchi, 1955)
Japanese name: Chishima-kiberi-kukibachi(Fig. 1B)

Material examined. HOKKAIDO: 1 ♀,Fukagawa, Takadomari, 43°55′N 142°03′E, 25.
VI. 2019, H. Hara.
Distribution. Japan: Hokkaido (new record),
Etorofu Island (Takeuchi, 1955), Kunashiri
Island (Sundukov, 2015). Russia: European part
(Zhelochovtsev and Prochorova, 1976) to Siberia
and southern Primorye (Verzhutskij, 1981). Fin-land (Taeger et al., 2006).
Host plants. ?Rosaceae: Rosa spp. (Sundukov,
2015, 2017; see comments below).
Remarks. Takeuchi (1955) described this species from Etorofu Island, southern Kuriles, and it
was later recorded from Moskva, Jakutsk andsouthern Primorye, Finland and Kunashiri Island,
southern Kuriles (see references above). The
specimen listed above represents the ﬁrst distribution record from the main island of Hokkaido.
This is a mainly black species, with characteristic
yellow marks on the lower inner orbit, mandible,
and each side of the abdominal terga 3 and 4(Fig. 1B).

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Sundukov (2015, 2017) just noted “Личинки впобегах розовых [Larvae in pink shoots] (Rosaceae)” (2015) and “Host plant: Rosa spp. (Rosaceae)” (2017) without giving source of information. This host record may need conﬁrmation.
Siricidae
Tremex apicalis Matsumura, 1912
Japanese name: Kuro-hiraashi-kibachi(Fig. 1C)

Host plants. Betulaceae: Alnus japonica(Thunb.) Steud. (new record), Betula (Shinohara
and Hara, 2020). Cornaceae: Swida (Kuramitsu
et al., 2019). Eupteleaceae: Euptelea (Kuramitsu
et al., 2019). Fagaceae: Quercus (Okutani,
1967a), Lithocarpus (Shinohara and Hara, 2020).
Juglandaceae: Juglans mandshurica Maxim. var.
sachalinensis (Komatsu) Kitam. (new record).
Magnoliaceae: Magnolia (Kuramitsu et al.,2019). Oleaceae: Fraxinus (Kuramitsu et al.,2019). Rosaceae: Cerasus (Okutani, 1967a).
Salicaceae: Populus (Okutani, 1967a). Sapindaceae: Acer (Okutani, 1967a).
Remarks. On May 23, 2021, Shinohara foundseveral dead females of this species on the trunksat the height of 1.5–2.5 m of dead trees of theabove two newly recorded species standing on the
broad riverbed of the Tone River in Koya (about
7 m), Moriya City, Ibaraki Prefecture. Apparently,
those females (one of them in Fig. 1C) died after
oviposition as often observed for siricid females.
This is the ﬁrst record of Alnus japonica (Betulaceae) and Juglans mandshurica var. sachalinensis (Juglandaceae) as hosts of Tremex apicalis.
This woodwasp is now known to attack various
broadleaved trees of eleven families.
Tenthredinidae
Aglaostigma albicinctum (Takeuchi, 1953)
Japanese name: Urashimasou-habachi

1 ♂, Wami, 36°45′N 140°09′E, Nakagawa, coll.
larva, on Arisaema serratum, 10. VI. 2014, mat. 13.
VI., em. 6. IV. 2015, S. Ibuki; 1 ♀, same locality,
coll. larva (Fig. 2C), on Actinidia polygama, 22. VI.
2016, mat. 26. VI., em. 26. IV. 2017, S. Ibuki; 1 ♀,same locality, coll. larva (Fig. 2B, D, F), on Impatiens textorii, 28. V. 2016, mat. 10. VI., em. 24. IV.
2017, S. Ibuki; 1 ♂, same locality, coll. larva (Fig.
2A), on Symplocarpus renifolius, 31. V. 2018, mat.
13. VI., em. 21. IV. 2019, S. Ibuki.
Larva. Early or middle instar (Fig. 2A–B): headpale grayish brown (Fig. 2A, probably earlier
instar) or gray (Fig. 2B, probably later instar);trunk and legs creamy white. Late instar (Fig.
2C–E): head blackish; trunk and legs creamy
white, most of dorsal region brown with paired
whitish lines dorsally, and supraspiracular, epipleural and hypopleural regions of each segment
and two or three terminal segments blackish.
Mature larva (Fig. 2F): entirely vivid pale green.
Host plants. Actinidiaceae: Actinidia polygama (Siebold et Zucc.) Planch. ex Maxim. (newrecord). Araceae: Arisaema spp. (Okutani,
1967b), Symplocarpus renifolius Schott ex
Tzvelev. (new record). Balsaminaceae: Impatiens
textorii Miq. (new record).
Life history. This species has one generation ayear. In lower mountains in central Honshu, theadults emerge in late April to May and the larval
feeding stages are in May to June. The larva issolitary and has an extra molt on maturity before
entering the soil.
Remarks. Okutani (1967b) recorded Arisaema
spp. (Araceae) as hosts of Aglaostigma albicinctumand here we newly recognize three species of three
diﬀerent plant families as hosts of this sawﬂy basedon rearing. This is a polyphagous species. See Okutani and Takeuchi (1953) and Okutani (1959) for a
detailed description of the larva and life history.
Allantus meridionalis (Takeuchi, 1933)
Japanese name: O-shiroobi-kuro-habachi(Fig. 2A–F)(Fig. 2G–J)

Material examined. HONSHU: Tochigi Pref.:Material examined. HONSHU: Tochigi Pref.:Taxonomic Notes and New Records in Japanese Symphyta VI

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Fig. 2. A–F, Aglaostigma albicinctum: A, early or middle instar larva on Symplocarpus renifolius, 31. V. 2018; B,
early or middle instar larva on Impatiens textorii, 29. V. 2016; C, late instar larva on Actinidia polygama, 25.
VI. 2016; D, late instar larva on I. textorii, 30. V. 2016; E, late instar larva on A. polygama, 1. VII. 2016; F,mature larva, 10. VI. 2016. G–J, Allantus meridionalis: G, early instar, 20. IX. 2015; H, J, middle or late instar
larva, 12. X. 2015; I, mature larva, 13. X. 2015. K–M, Apethymus kunugi: K, probably late instar larva, 10. V.
2019; L, same larva, 12. V. 2019; M, same larva, matured, and cast skin above, 17. V. 2019. N–P, Corymbas
nipponica: N, middle instar larva, 17. VI. 2016; O, same larva, late instar, 26. VI. 2016; P, same larva,
matured, and cast skin above, 29. VI. 2016. All photographed by Ibuki.

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1 ♀, Nakagawa, Wami, 33°46′N 140°10′E, coll.
larva (Fig. 2G), on Sanguisorba oﬃcinalis, 20.
IX. 2015, mat. 21. X., em. 16. XII. 2015, S.
Ibuki; 1 ♀, same data but coll. larva, 25. IX.
2015, mat. 14. X., em. 16. XI. 2015.
Larva. Early or middle instar (Fig. 2G): headpale gray; trunk and legs creamy white. Late instar(Fig. 2H, J): head pale pink; trunk and legs palecreamy white, with dorsal region pale greenish
gray; thoracic legs tinted with pink. Mature larva(Fig. 2I): similar to late instar; head more yellowish.
Host plants. Rosaceae: Rosa spp. (Okutani,
1967b), Fragaria × ananassa (Weston) Duchesne
ex Rozier (Okutani, 1967b), Sanguisorba oﬃcinalis L. (new record).
Life history. This is a multivoltine species. The
larva is solitary, though a few larvae may occur
on the same leaf (Fig. 2G). The larva has an extra
molt on maturation and enters a rotten wood, notsoil, to make a cell for diapause (Kondo, 1969).
Remarks. Okutani (1959) described the larvaof this species as Emphytus albicinctus Matsumura
and Kondo (1969) published his observations on
the life history of this species.
Apareophora nebuta Togashi, 1964
New Japanese name: Hime-shimotsuke-maru-habachiMaterial examined. HOKKAIDO: 1 ♂,Fukagawa, Takadomari, 26. V.–6. VI. 2007, by
Malaise trap, H. Hara; 1 ♀, Fukagawa, Takadomari, 43°55′N 142°03′E, 2. VI. 2018, H. Hara;
1 ♀, same data but 29. V. 2019.
Distribution. Japan: Hokkaido (new record),
Honshu (Togashi, 1964).
Remarks. This species has been only known
from Honshu, Japan (Hara and Shinohara, 2017). It
is here recorded from Hokkaido for the ﬁrst time.
Apethymus kunugi Togashi, 2005
New Japanese name: Kunugi-habachi(Figs. 1D, 2K–M)

Material examined. HONSHU: Tochigi Pref.:1 ♀ (Fig. 1D), Koisago, 36°46′N 140°08′E,
Nakagawa, coll. larva (Fig. 2K–L), on Quercus
serrata, 10. V. 2019, mat. 17. V. (Fig. 2M), em.
18. X. 2019, S. Ibuki.
Larva. Late instar (Fig. 2K–L): head dark
opaque gray; trunk and legs creamy white, dorsal
region dark gray with creamy white line along
middorsal line and each thoracic and abdominal
segment with blackish spot in laterodorsal
region. Mature larva (Fig. 2M): head paleorange; trunk and legs yellowish white, dorsal
region dark gray, each segment with blackish
spot in laterodorsal region.
Host plants. Fagaceae: Quercus acutissima
Carruth. (Togashi, 2005), Q. serrata Murray(new record).
Life history. This is a univoltine species withthe adult emergence in autumn, hibernation in
the egg stage, and the larval feeding stages inspring. The larva is a solitary external feeder ofthe host leaves (Fig. 2 L). On maturity, the reared
larva had an extra molt (Fig. 2M) and went into
the soil.
Remarks. Togashi (2005) described this species from the holotype which was reared from alarva feeding on Q. acutissima (Togashi, 2005),
but he did not give any further information about
the larva and life history. Here we newly record
Q. serrata as a host of A. kunugi. Other speciesof Quercus may also serve as the host of thissawﬂy.
Ardis pallipes (Serville, 1823)
Japanese name: Bara-kuki-habachiMaterial examined. HOKKAIDO: 1 ♂,Fukagawa, Takadomari, 26. V.–6. VI. 2007, by
Malaise trap, H. Hara; 1 ♀, same locality andcollector, 9. VI. 2010; 1 ♀, same data but,
43°55′N 142°03′E, 31. V. 2020.
Distribution. Japan: Hokkaido (new record),
Honshu (Okutani, 1959, as A. brunniventris)(Takeuchi, 1952 recorded this species from Japanfor the ﬁrst time, but he did not detail the locality). Palaearctic and Nearctic regions (Smith,Taxonomic Notes and New Records in Japanese Symphyta VI

1969, as A. brunniventris; Sundukov, 2017).
Remarks. This species is here recorded from
Hokkaido for the ﬁrst time.

Remarks. This species is here recorded from
Hokkaido for the ﬁrst time.
Euura imperfecta (Zaddach, 1876)

Corymbas nipponica Takeuchi, 1936

Japanese name: Karamatsu-madara-habachi

Japanese name: Huto-koshijiro-habachi(Figs. 3A–G, L–O, 4A–N, 5A–P, 6A–C, 7A–D)(Fig. 2N–P)

Material examined. HONSHU: Tochigi Pref.:1 ♀, Nakagawa, Wami, 33°46′N 140°10′E, coll.
larva (Fig. 2N), on Sanguisorba oﬃcinalis, 12.
VI. 2016, mat. 29. VI. (Fig. 2P), em. 26. IV.
2017, S. Ibuki.
Larva. Middle instar (Fig. 2N): head creamy
white; trunk pale, covered with snow-whitewoolly wax. Late instar (Fig. 2O): head dark yellow; trunk pale greenish, covered with snowwhite woolly wax; thoracic legs dark yellow.
Mature larva (Fig. 2P): head dark yellow; trunk
pale green, without wax covering.
Host plants. Rosaceae: Rubus spp. (Okutani,
1967b), Geum japonicum Thunb. (Okutani,
1967b), Sanguisorba oﬃcinalis L. (new record).
Life history. This is a univoltine species withthe adult emergence in April to May and the larval feeding stages in May to June in lowlands ofcentral Honshu. The larva is a solitary external
feeder of the host leaves (Fig. 2N–O). On maturity, the reared larva had an extra molt (Fig. 2P)
and went into the soil.
Remarks. This is the ﬁrst record of Sanguisorba oﬃcinalis as the host plant of C. nipponica. For a detailed description of the larva, see
Okutani (1959).
Eriocampopsis subtruncata Takeuchi, 1952
Japanese name: Sarashinashoma-habachiMaterial examined. HOKKAIDO: 1 ♀, Bibai,
Tomei, 10. V. 2017, H. Hara.
Distribution. Japan: Hokkaido (new record),
Honshu (Takeuchi, 1952). Korean Peninsula (Leeet al., 2019). Russia: Primorye (Zhelochovtsev,
1968).

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Nematus imperfectus Zaddach, 1876: 80.
Pachynematus imperfectus: Jörgensen, 1906: 350;
Benson, 1958: 233; Ito, 1959; Pschorn-Walcher andEichhorn, 1963: 62; Verzhutskii, 1966: 138;
Verzhutskii, 1981: 210.
Pachynematus ? imperfectus: Takizawa, 1957: 27.
[Unidentiﬁed species 2]: Hara and Kitagawa, 1986: 21.
Nematus (Larinematus) imperfectus: Zhelochovtsev andZinovjev, 1988: 169.
Pikonema imperfectum: Zinovjev, 1993: 21.
Larinematus imperfectus: Lacourt, 1999: 151; Lacourt,
2020: 428.
Pachynematus (Larinematus) imperfectus: Taeger et al.,
2010: 438; Macek et al., 2020: 532.
Euura imperfecta: Prous et al., 2014: 46; Shinohara andHara, 2015: 174; Hara, 2019: 73; Hara, 2020: 89, 349.
For more references, see Lacourt (1999).

Redescription (female and male). Length 5.5–
6.5 mm in female, 5.5 mm in male. Female yellow, red yellow or brown and black (Fig. 3A–B,
E–G); head capsule with peculiar black markings(Fig. 4A–D, L–N); antenna black; thorax partly
black; mesepisternum black on epicnemium andoften ventrally or ventrolaterally (Fig. 4B, F–G);
legs with coxae basally black and tarsi and apexof hind tibia sometimes slightly darkened; abdomen dorsally or mostly black; valvula 3 always
black. Male mostly black (Fig. 3C–D); legs yellow from apices of femora to tarsi but tibiae andtarsi slightly darkened apically; subgenital plateyellow. In both sexes, wings very slightly yellowish (Fig. 3A, F, C); stigma almost uniformly
pale brown to brown, sometimes basally pale;veins mostly brown to black; vein C mostly orbasally yellow.
Head in dorsal view with length behind eye
0.6–0.8 × eye length in female, 0.4–0.5 × inmale (Fig. 4A, H, L); length behind lateral ocellus 2.0–2.8 × length of lateral ocellus in female,
1.7–1.8 × in male. Frontal area with lateral and170

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Fig. 3. A–G, L–O, Euura imperfecta: A–B, female, Hokkaido, dorsal and ventral views; C–D, male, Hokkaido,
dorsal and ventrolateral views; E–F, female, Honshu, dorsal and ventrolateral views; G, female, Honshu, ventrolateral view; L, egg, Hokkaido; M, semiﬁnal instar larva, lateral view; N–O, ﬁnal instar larva, lateral anddorsal views; M–O, Hokkaido, Bibai, 1986. H–K, Euura itoi: H–I, female, dorsal and ventrolateral views;
J–K, male, dorsal and ventrolateral views. All photographed by Hara.

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Fig. 4. A–N, Euura imperfecta: A–G, female, Hokkaido; H–K, male, Hokkaido; L–N, female, Honshu. O–X,
Euura itoi: O–U, female; V–X, male. A, H, L, O, V, Head, dorsal view; B, I, M, P, W, dorsomedial part ofhead anterodorsal view; C, J, N, Q, X, head, anterior view; D, K, R, Y, head, lateral view; E, S, torulus and its
surroundings, anterolateral view; F, T, left mandible, outer view; G, U, right mandible, outer view. All photographed by Hara.

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Fig. 5. A–P, Euura imperfecta: A–H, female, Hokkaido; I–K, male, Hokkaido; L–P, female, Honshu. Q–AA,
Euura itoi: Q–X, female; Y–AA, male. A, I, Q, Y, Posteromedial part of thorax, dorsal view; B, J, L, R, Z, tarsal claw, anterior or posterior view; C, S, anteromedial part of fore wing; D, T, apical part of abdomen, ventrolateral view; E, M, U, abdomen apex, lateral view; F, N, V, ditto, posterolateral view; G, O, W, ditto, dorsal
view; H, P, X, ditto, posterior view; K, AA, apical part of abdomen, dorsal view. M–N, P, Ovipositor removed;K, genitalia removed. All photographed by Hara.

Taxonomic Notes and New Records in Japanese Symphyta VI

anterior ridges usually distinct (Fig. 4B, I, M);anterior ridge not grooved medially. Distance
between eyes at torulus 1.4–1.5 × eye height infemale, 1.2 × in male (Fig. 4C, J, N). Area justabove torulus almost ﬂat and triangular, laterallywith long straight furrow (Fig. 4E). Dorsal tentorial pit obscure. Narrow ridge around torulus dorsally widely vague or disappearing. Paraantennalﬁeld normal, covered with setae except for narrow medial margin. Clypeus distinctly concave
roundly on ventral margin (Fig. 4C, J, N); depthof ventral emargination 0.3–0.7 × median heightof clypeus; clypeus with width 3.3–3.9 × maximum height; maximum height 0.7–0.9 × torulus
height. Malar space length 1.2–1.4 × medianocellus width in female, 0.8–0.9 × in male.
Antenna length 2.1–2.5 × head width in female,
3.0 × in male (Fig. 3A, C, E); ﬂagellomere 1
0.8–0.9 × as long as major axis of eye, simple infemale, slightly convex basally on ventral margin
in lateral view in male (Fig. 4K); ﬂagellomere 2
1.1–1.3 × as long as ﬂagellomere 1. Left mandible 1.1–1.2 × as long as right one, and in outerview sharply tapering basally, thin from middle
to apex, with middle part same thickness or
slightly thickening towards apex (Fig. 4F). Right
mandible rather sharply tapering on basal halfand gradually tapering on apical half (Fig. 4G).
Basal outer surfaces of both mandibles notwidely ﬂattened or concave. Maxillary palpus
rather short, with palpomere 2 0.7–0.8 × as longas palpomere 3 and palpomere 6 0.7–0.8 × aslong as torulus height.
Mesoscutellar appendage with length 0.9–
1.6 × minor axis of cenchrus (Fig. 5A, I). Metascutellum length 0.9–1.3 × minor axis of cenchrus. Mesepisternum with groove along anterioredge extending into epicnemium. Epicnemiumwith ventral edge distinctly grooved. Katepimeron entirely glabrous. Anterior fore tibial spurwith velum. Hind tibia normal, 0.8 × as broad ashind femur in anterior view; posterior hind tibial
spur 0.9–1.2 × as long as apical breadth of hindtibia, 0.4–0.5 × as long as hind tarsomere 1. Hind
tarsus 0.8–1.0 × as long as hind tibia. Tarsal
claws narrow, with small inner tooth (Fig. 5B, J,

173L); depth of concavity between apical and inner
teeth 0.3–0.5 × distance between these teeth.
Fore wing with cell Sc 0.2–0.6 × as wide as vein
C at level of base of vein Rs＋M (Fig. 5C). Hind
wing with section of vein 1A between cell 1Aand crossvein cu-a 1.9–2.1 × as long as crossvein
cu-a.
In female abdomen, sternum 7 with posterior
margin deeply concave beside median projection(Fig. 5D). Tergum 9 normal in size, in lateralview 0.8–1.6 × as long as tergum 8 at level ofspiracle 8 (Fig. 5E, M). Cercus 3.0–5.0 × as longas wide, posteriorly extending to or slightly
beyond apex of valvula 3 (Fig. 5E, G, M, O).
Ovipositor sheath 0.3 × as long as abdomen (Fig.
3B, G), 0.6–0.7 × as long as hind tibia; valvula 3
in lateral view with apical edge nearly truncate or
widely rounded (Fig. 4E, M); both valvulae 3
combined apically slightly or distinctly concave,with apicolateral ridge sharp (Fig. 5F–G, M–P).
Lance in lateral view with dorsal margin slightly
concave (Fig. 6A); radix with subdorsal carina;
annuli distinctly oblique. Lancet with radix 1.2–
1.3 × as long as lamnium (Fig. 6B, C); lamniumwith 14–15 annuli; annuli arched; ctenidial setae
absent on ventrobasal and dorsoapical parts oflamnium; annulus 1 usually without ctenidial setae;
ctenidial setae about as long as length of annulus;
ventral margin of lamnium not serrate, without
distinct denticles.
In male abdomen, procidentia with sharply
deﬁned median area (Fig. 5K); tergal hollow
with anterior edge distinctly ridged laterally.
Subgenital plate 0.4–0.5 × as long as abdomen,
0.7–0.8 × as long as hind tibia (Fig. 3D), gradually tapering and narrowly rounded apically (Fig.
5K). Genitalia with harpe in ventral view about
as long as wide (Fig. 7A); parapennis apicallypointed; penis valve with paravalva apically protruding below valvispina (Fig. 7B–D); valvispina
narrow and acute.
Head capsule mostly distinctly or slightly
microsculptured and inconspicuously punctured;clypeus not microsculptured. Thorax predominantly microsculptured; medial part of mesopostnotum, whole metapostnotum, etc. not microscu-

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Fig. 6. Female ovipositor. A–C, Euura imperfecta: A–B, Hokkaido; C, Honshu. D–E, Euura itoi. A, D, Ovipositor; B–C, E, lancet. All photographed by Hara.

Taxonomic Notes and New Records in Japanese Symphyta VI

175

Fig. 7. Male genitalia. A–D, Euura imperfecta, Hokkaido; E–I, E. itoi. A, E–F, Genital capsule: A, ventral view
(penis valve removed); E, F, dorsal and ventral views. B–D, G–I, Penis valve, lateral view (left dorsal). All
photographed by Hara.

lptured; punctures generally inconspicuous;
mesoscutum, propleuron and ventrolateral part ofmesepisternum with small punctures. Abdomen
microsculptured; sterna slightly or inconspicuously microsculptured.
Immature stages. Egg long oval (Fig. 3 L).
Semiﬁnal instar larva (Fig. 3M): head and legs
pale yellow brown; legs basally narrowly black;trunk pale green, with slightly blackish subdorsal

stripe. Final instar larva (Fig. 3N–O): 15 mm
long, as in semiﬁnal larva, but trunk with distinct
blackish green subdorsal stripe and narrow darkgreen pleural stripe. Cocoon: 8 mm long, brown,
double walled; outer wall thin. For more information, see Jörgensen (1906), Verzhutskii (1966)
and Macek et al. (2020).
Material examined. HOKKAIDO: 1 ♀ (Fig.
6A), Shintoku, Shintoku, 8. V. 2012, H. Hara

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(cited by Hara, 2020); 1 ♀ (Figs. 5B, 6B), Bibai,
9. V. 1961, K. Kamijo; 1 ♀, same locality, 7. V.
1987, H. Hara; 1 ♀ (Figs. 5I, 7A, C–D), samedata but 3. V. 1988; 1 ♂, same data but 6. V.
1988; 1 ♀, same data but 8. V. 1988; 1 ♂ (Figs.
3C–D, 4H–K, 5J–K, 7B), Hayakita, 13. V. 1986,
H. Hara; 1 ♀, same data but 11. V. 1987; 1 ♀
(Fig. 5D), same locality, coll. cocoon in litter
under Larix kaempferi, 30. IV. 1987, em. 25. V.
1987, H. Hara; 1 ♀, same locality but 19. V.
1987, H. Hara; 1 ♀ (Figs. 4D, F–G, 5A, C),same data but 6. V. 1988; 3 ♀ (Figs. 3A–B, 4A–C,
E, 5E–H), same locality, coll. litter under Larix
kaempferi, em. 28, 30. IV. 1992, H. Hara, and theirprogeny (Fig. 3 L). — HONSHU: Nagano Pref.:3 ♀, Ueda, Sanada-machi, Soehi, 30. IV. 1956,
Takeuchi (probably reared); 1 ♀, Aoki-toge, coll.
larva on Larix kaempferi 1956, em. 19. III. 1957,
Ito (cited by Ito, 1959).
Distribution. Japan: Hokkaido (Shinohara andHara, 2015), Honshu (Takizawa, 1957; Ito,
1959). Russia: European part, Siberia (Verzhutskii, 1981; Zhelochovtsev and Zinovjev,
1988). Europe (Zaddach, 1876; Benson, 1958;
Taeger et al., 2018).
Host plants. Pinaceae: Larix decidua Mill.
(Jörgensen, 1906), L. kaempferi (Lamb.) Carrière(Takizawa, 1957; Ito, 1959), L. sibirica Ledeb.
(Verzhutskii, 1966, as “лиственницы сибирской”).
Life history. In our observation in Hokkaido,
Japan, this sawﬂy has one generation a year; theadults were collected in May; under rearing condition, the female laid her eggs singly on needles(Fig. 3 L); the cocoons were found within the litter in larch forests. For more information, see
Jörgensen (1906), Pschorn-Walcher and Eichhorn (1963), Verzhutskii (1966) and Macek et al.
(2020).
Remarks. Euura imperfecta is mainly characterized by the following features: area just above
torulus almost ﬂat and triangular and laterallywith long straight furrow (Fig. 4E); female sternum 7 with posterior margin deeply concave
beside median projection (Fig. 5D); male procidentia with median area sharply deﬁned (Fig.
5K). The following character states are also useful

to distinguish this species from other congenersof Euura and similar genera: tarsal claws withsmall inner tooth (Fig. 5B, J, L) and depth
between apical and inner teeth about half or lessof distance between apices of these teeth; narrowridge around torulus widely vague or disappearing dorsally (Fig. 4E); mandibles asymmetrical,
in outer view left mandible with middle partsame thickness or slightly thickening towards
apex (Fig. 3F), right mandible continuously
tapering from base to apex (Fig. 4G); katepimeron entirely glabrous; in female, valvulae 3 nottapering and apically concave (Fig. 5F–H, N–P);
in lancet, lamnium without serrulae and denticles
on ventral margin; in male, tergal hollow of tergum 8 with anterior edge distinctly ridged laterally (Fig. 5K).
Zhelochovtsev in Zhelochovtsev and Zinovjev(1988) established Larinematus Zhelochovtsev,
1988 as a subgenus of Nematus Panzer, 1801 forthree species, E. imperfecta (Fig. 3A–D), E. itoi
(Fig. 3H–K) and one unstated species. Zinovjev(1993) considered this subgenus a species groupof Pikonema Ross, 1937. Lacourt (1999, 2020)
raised Larinematus to the rank of genus. Roller
and Haris (2008) described a new species under“Pachynematus (Larinematus)”. Taeger et al.
(2010) and Macek et al. (2020) treated Larinematus as a subgenus of Pachynematus Konow,
1890. On the other hand, Prous et al. (2014) synonymized Larinematus, Pachynematus andPikonema with Euura Newman, 1837. Actually,
the features of Larinematus stated by previous
authors are not stable or diﬃcult to recognize,
judging from Japanese specimens of these two
species. Zhelochovtsev and Zinovjev (1988),
Zinovjev (1993), Macek et al. (2020) and
Lacourt (2020) distinguished Larinematus or theE. imperfecta group from other species of former
Pachynematus mainly by the relative lengths ofhind tibia and tarsus and the shape of the ovipositor sheath. Zhelochovtsev and Zinovjev (1988),
Macek et al. (2020) and Lacourt (2020) statedthat Larinematus has the hind tarsus as long as or
longer than the hind tibia. Benson (1958) alsocharacterized E. imperfecta by a long hind tarsus.

Taxonomic Notes and New Records in Japanese Symphyta VI

However, in the Japanese specimens of the two
species, the hind tarsus is often distinctly shorter
than the hind tibia in E. imperfecta (Fig. 3B) andalways so in E. itoi (Fig. 3I) (the ratio is 0.83–
0.97 : 1.0 in E. imperfecta and 0.78–0.86:1.0 in E.
itoi). Zhelochovtsev and Zinovjev (1988) statedthat the ovipositor sheath of Larinematus waswith “опорной площадкой” (probably refers to awide concavity at the apex; Fig. 5F, N) with aﬁgure of the ovipositor sheath of E. imperfecta.
Macek et al. (2020) also adopted this feature forPachynematus (Larinematus). However, inEuura itoi, the apex of the ovipositor sheath ismostly ﬂattened, dorsally narrowly elevated andwith a median carina (Fig. 5V, X). Lacourt(2020) wrote that the ovipositor sheath was
incised when viewed from above as a genericfeature of Larinematus. The sheaths combined
are apically incised in dorsal view in E. imperfecta (Fig. 5G, O) but truncate in E. itoi (Fig.
5W). It is impossible to separate Larinematus or
the E. imperfecta group from other species offormer Pachynematus except for the host plant.
Furthermore, there are some signiﬁcant diﬀerences between these two species apart from theirmorphological diﬀerences. The antenna of E.
imperfecta shows distinct sexual diﬀerence. The
male antenna is longer than that of the female(Fig. 3A, C), and the male ﬂagellomere 1 isbasally convex on the ventral margin (Fig. 4K)
while the female ﬂagellomere 1 is simple. On theother hand, there is no distinct structural diﬀerence between the female and male antennae of E.
itoi. The life cycle of E. imperfecta is univoltine
while that of E. itoi is multivoltine. These two
species do not appear to be closely related.
Apart from the relative lengths of hind tibia
and tarsus, Japanese specimens of E. imperfecta
well agree with the concept of this species by
Benson (1958). He wrote “Underthorax with atleast mesosternum black”. In the Japanese specimens, the mesepisternum is often not darkened
ventrally. The Japanese specimens also wellagree with the description by Muche (1974)
except for the color of the stigma. He wrote“Stigma braun mit breiter weißer Basis”, but the177

Japanese specimens have the uniformly brownstigma (Fig. 3A, C, E).
The Hokkaido larva agrees with the description of the European larva by Jörgensen (1906).
The trunks of these mature larvae are green withtwo dark longitudinal stripes (subdorsal and pleural stripes) (Fig. 3N–O; p. 533-ﬁg. 3 in Macek etal., 2020). However, according to Verzhutskii(1966), the Siberian larva has the trunk with only
one dark longitudinal stripe (subdorsal stripe)
and the ventral side is distinctly paler than thedorsal side (Рис. 34-8 and Рис. 43 in Verzhutskii,1966). Furthermore, the Honshu larva quite differs from the Hokkaido, European and Siberian
larvae. According to Ito (1959), the Honshu larva
is wholly purplish black in the semiﬁnal instar
and younger larvae and has the head blackish
brown and the trunk dark brown with a pale
brown middorsal line in the ﬁnal instar larva.
Therefore, Ito (1959) questioned identifying the
Honshu specimens with E. imperfecta and statedthe identiﬁcation was tentative. We have notfound any diﬀerence between the Hokkaido andHonshu adults and so consider them conspeciﬁc.
More material and genetical study will be needed
to interpret the large larval color variation.
Euura itoi (Okutani, 1955)
Japanese name: Karamatsu-aka-habachi(Figs. 3H–K, 4O–Y, 5Q–AA, 6D–E, 7E–I)
Pachynematus itoi Okutani, 1955: 98; Takizawa, 1957:
17; Pschorn-Walcher and Eichhorn, 1963: 59; PschornWalcher and Zinnert, 1971: 348; Muche, 1974: 86;Verzhutskii, 1981: 211; Xiao et al., 1992: 131;
Takizawa, 1994: 341; Park et al., 2007: 2.
“Species ignorata”: Verzhutskii, 1966: 143 (see PschornWalcher and Zinnert, 1971).
Nematus (Pikonema) itoi: Zhelochovtsev, 1976: 82.
Nematus (Larinematus) itoi: Zhelochovtsev and Zinovjev,
1988: 169.
Pikonema itoi: Zinovjev, 1993: 21.
Larinematus itoi: Lacourt, 1999: 152; Lacourt, 2020: 429.
Pachynematus (Larinematus) itoi: Taeger et al., 2010:
438; Macek et al., 2020: 532.
Euura itoi: Shinohara and Hara, 2015: 174; Hara, 2019:73; Hara, 2020: 89, 350.

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Redescription (female and male). Length 6.5–
8.0 mm in female, 5.0–6.5 mm in male. Red yellow (Fig. 3H–K). Female: head capsule black
only on ocellar area (Fig. 4O–R); antenna black,with ﬂagellomere 1 red yellow ventrally or
entirely; thorax dorsally with three large blackspots, ventrally black except for narrow or wide
dorsal part of mesepisternum (Fig. 3H–I); legs
black from coxae to femora of fore and middle
legs and on most of hind leg; abdomen only
darkened on narrow anteromedial part of dorsum
and valvula 3. Male: head capsule darkened on
medial or most part of frons, medial part of vertex and most of occiput (Fig. 4V–Y); antenna red
yellow; thorax black except for most of pronotum and tegula (Fig. 3J–K); legs darkened on
bases of coxae, basal part of fore femur and apical part of hind tarsus, often also on all trochanters, basal parts of middle and hind femora andwhole hind tarsus; abdomen dorsally darkened
except for lateral and apical parts. Wings
strongly tinged with brown on basal two thirds;
veins brown to black; stigma yellow, darkened
on wide base and narrow margins.
Head in dorsal view with length behind eye
0.8–0.9 × eye length in female, 0.6–0.7 × in male(Fig. 4O, V); length behind lateral ocellus
3.7–4.3 × length of lateral ocellus in female, 2.8–
3.0 × in male. Frontal area with distinct lateraland anterior ridges (Fig. 4P, W); anterior ridge
not grooved medially. Distance between eyes attorulus 1.6–1.7 × eye height in female, 1.4–
1.5 × in male (Fig. 4Q, X). Area above torulus
concave, often with small tubercle (Fig. 4S). Narrow ridge around torulus wholly distinct. Paraantennal ﬁeld normal, covered with setae except for
narrow medial margin (Fig. 3S). Clypeus very
deeply concave angularly or roundly on ventral
margin (Fig. 4Q, X); depth of ventral emargination 1.4–1.7 × median height of clypeus; clypeus
with width 2.7–3.2 × maximum height; maximum height 1.0–1.3 × torulus height. Malar
space length 0.8–1.4 × median ocellus width.
Antenna length 2.7–3.1 × head width; ﬂagellomere 1 1.0–1.2 × as long as major axis of eye,simple in both sexes (Figs. 3H–K, 4Y); ﬂagello-mere 2 1.0–1.2 × as long as ﬂagellomere 1. Leftmandible 1.1–1.2 × as long as right one, in outerview sharply tapering basally, thin from middle toapex, with middle part slightly thickening
towards apex (Fig. 4T). Right mandible rather
sharply tapering on basal half and gradually narrowing on apical half (Fig. 4U). Basal outer surfaces of both mandibles not widely ﬂattened orconcave. Maxillary palpus with palpomere 2 0.6–
0.8 × as long as palpomere 3 and palpomere 6
0.6–0.9 × as long as torulus height.
Mesoscutellar
appendage
length0.5–
0.9 × minor axis of cenchrus (Fig. 5Q, Y).
Metascutellum length 1.2–1.6 × minor axis ofcenchrus. Mesepisternum with groove along
anterior edge extending into epicnemium. Epicnemium with ventral edge distinctly grooved.
Katepimeron glabrous, usually narrowly covered
with setae posteriorly. Anterior fore tibial spurwith velum. Hind tibia usual, 0.7–0.9 × as broadas hind femur in anterior view; posterior hind tibial spur 1.0–1.3 × as long as apical breadth ofhind tibia, 0.5–0.7 × as long as hind tarsomere 1.
Hind tarsus 0.8–0.9 × as long as hind tibia (Fig.
3I). Tarsal claws narrow, with small inner tooth(Fig. 5R, Z); depth of concavity between apicaland inner teeth about 0.5 × or less distance
between these teeth. Fore wing with cell Sc 0.7–
1.2 × as wide as vein C at level of base of vein
Rs＋M (Fig. 5S). Hind wing with section of vein
1A between cell 1A and crossvein cu-a 1.1–
1.4 × as long as crossvein cu-a.
In female abdomen, sternum 7 with posterior
margin moderately concave beside median projection (Fig. 5T). Tergum 9 normal in size, in lateral view 1.0–1.7 × as long as tergum 8 at levelof spiracle 8 (Fig. 5U). Cercus 4.0–7.0 × as longas wide, posteriorly extending far beyond valvula
3 (Fig. 5U, W). Ovipositor sheath 0.3–0.4 × aslong as abdomen, 0.5 × as long as hind tibia (Fig.
3I); valvula 3 in lateral view short and wide,
almost truncate apically (Fig. 5U), in dorsal viewroundly convex laterally, truncate apically (Fig.
5W). Lance in lateral view with dorsal margin
slightly rounded (Fig. 6D); radix with subdorsal
carina; lamnium with annuli distinctly oblique.

Taxonomic Notes and New Records in Japanese Symphyta VI

Lancet with radix 0.9 × as long as lamnium (Fig.
6E); lamnium with 12–13 annuli; basal annuli
straight; middle and apical annuli slightly arched;ctenidia absent; ventral margin of lamnium serrate; serrulae with minute denticles.
In male abdomen, tergum 8 with procidentia
much protruding posteriorly, nearly rectangle(Fig. 5AA) and tergal hollow indistinctly deﬁned.
Subgenital plate 0.5 × as long as abdomen, 0.8–
0.9 × as long as hind tibia, gradually narrowing
towards apex, with apex narrowly rounded or
narrowly truncate. Genitalia with harpe in ventral
view longer than wide (Fig. 7F); valviceps withmany spinules centrally (Fig. 7G–H); parapennis
acute; valvispina tapering with pointed apex;
paravalva not protruding apically.
Head, thorax and abdomen generally smoothand shiny, with punctures minute or inconspicuous. Frons, vertex, pronotum, median mesoscutallobe and most of abdomen very slightly microsculptured. Mesoscutellar appendage laterally
microsculptured. Mesopostnotum microsculptured, with narrow medial part not or slightly
microsculptured. Metapostnotum not microsculptured.
Material examined. Holotype of Pachynematus itoi Okutani, 1955 (Figs. 3H–I, 4O–R, 5Q–V,
X): ♀ with a label written as “Soehi-mura,
Nagano-ken, 13-VIII-1955” and a piece of red
paper with nothing written on it. Paratypes of
P. itoi: 1 ♀ (Fig. 6E) and 1 ♂ (Fig. 7E–F, H–I)
with the same label as the holotype; 1 ♀ (Fig.
6D) and 1 ♂, “Soehi-mura, Nagano-ken,
13-VIII-1955, [Karamatsu (in Japanese; ＝Larix
kaempferi)]”; 2 ♀ (Fig. 5W), “Ueda, Naganoken, 15-VIII-1955”; 1 ♂ (Figs. 3J–K, 4V–Y, 5Y–AA) with a label written as “Ueda, Nagano-ken,
15-VIII-1955” and a piece of red paper withnothing written on it.
The type series of P. itoi mentioned by Okutani (1955) is “Holotype: 1 female, Soehi-mura,
Nagano Prefecture, Aug. 13, 1955, T. Ito leg.
Allotype: 1 male, Ueda, Nagano Prefecture, Aug.
15, 1955, T. Ito leg. Paratypes: 2 males and 3
females, same data as holotype; 2 females, samedata as allotype”. Although there are no specimens

179

with the type label of Pachynematus itoi in Okutaniʼs collection now kept in the National
Museum of Nature and Science, Tsukuba, wehave found the above ﬁve females and threemales agreeing with the original description in
the museum. The female and male with a pieceof red paper are safely considered the holotype
and the allotype stated by Okutani (1955),
respectively, judging from the ﬁgures in the original description (compare Fig. 3H, J with ﬁgs. 6,
7 in Okutani, 1955, respectively).
Other material examined. JAPAN: HONSHU:
Gunma Pref.: 4 ♀, Mt. Asama-yama, VIII. 1956,
Takeuchi. —Nagano Pref.: 1 ♀, Ueda, em. 21.
VI. 1956, K. Ozawa; 4 ♀, Ueda, Sanada-machi,
Soehi, mid-VII–late VIII. 1955, host: Larix
kaempferi; 5 ♀5 ♂ (Fig. 6G), same locality, V.
1956, Takeuchi (probably reared); 1 ♀, Chikuhoku, Aoki-toge, 29. VI. 1956, T. Ito; 4 ♀
(Fig. 3S–U), Utsukushigahara, 16–17. VIII.
1961, T. Naito; 2 ♀, Matsumoto, Shimashima, 7.
VI. 1929, Takeuchi; 1 ♂, Kiso-machi, Shinkai, 7.
VIII. 1956, T. Ito; 5 ♀, Kiso-machi, Fukushima,em. 28–30. VI. 1956, T. Ito.
Distribution. Japan: Honshu (Okutani, 1955).
Korea (Park et al., 2007). China (Xiao et al.,1992). Russia: Magadan Oblast, Siberia, Ural
(Zhelochovtsev, 1976; Verzhutskii, 1981). East
Europe (Pschorn-Walcher and Eichhorn, 1963;
Pschorn-Walcher and Zinnert, 1971).
Host plants. Pinaceae: Larix decidua Mill.
(Pschorn-Walcher and Eichhorn, 1963), L. gmelinii (Rupr.) Rupr. (Xiao et al., 1992), L. kaempferi(Lamb.) Carrière (Okutani, 1955), L. olgensis A.
Henry (Xiao et al., 1992).
Life history and immature stages. See
Takizawa (1957, 1994), Pschorn-Walcher andEichhorn (1963), Verzhutskii (1966, as “Species
ignorata”), Pschorn-Walcher and Zinnert (1971)
and Park et al. (2007).
Remarks. Euura itoi is distinguished fromother species of Euura and similar genera by thefollowing features: head capsule and abdomen
almost entirely red yellow in female, predominantly so in male (Fig. 3H, J); wings strongly
tinged with brown on basal two thirds; stigma

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yellow, widely darkened basally, narrowly
darkened marginally; malar space length
0.8–1.4 × median ocellus width; left mandible
longer than right mandible, in outer view withmiddle part slightly thickening towards apex
(Fig. 4T); right mandible continuously taperingfrom base to apex (Fig. 4U); maxillary palpus
not shortened, with palpomere 2 0.6–0.8 × aslong as palpomere 3; mesoscutellar appendage
0.5–0.9 × as long as minor axis of cenchrus (Fig.
5Q, Y); in hind leg, tibia narrower than femur inanterior view and tarsus 0.8–0.9 × as long as
tibia (Fig. 3I); tarsal claws with small inner tooth(Fig. 5R, Z); in female, cercus posteriorly
extending far beyond valvula 3 and valvula 3
short and apically truncate in dorsal view (Fig.
5W); lancet without ctenidia (Fig. 6E); in male,procidentia nearly rectangular (Fig. 5AA); penis
valve with many spinules on center of valviceps(Fig. 7G–I).
In the key to species of Pachynematus by
Muche (1974), our specimens of E. itoi including
the holotype do not go to this species, because allof them have the femora not entirely pale (atleast the base of the fore femur is darkened) (Fig.
3I, K), the stigma yellow with the wide dark base(Fig. 3H, I) and the mesoscutellum widely ﬂat(Fig. 5Q, Y). The keys by Zhelochovtsev andZinovjev (1988), Macek et al. (2020) and
Lacourt (2020) do not work either for the reasonsstated under the remarks of E. imperfecta above.
Eutomostethus pumicosus Seiyama, 1981
Japanese name: Suge-ko-maru-habachiMaterial examined. HOKKAIDO: 1 ♂,Fukagawa, Takadomari, 43°55′N 142°03′E, 9.
VI. 2020, H. Hara.
Distribution. Japan: Hokkaido (new record);
Honshu, Shikoku, Kyushu (Seiyama, 1981).
Remarks. This species is only known fromJapan (Naito, 2019) but has not been recorded
from Hokkaido up to now.

Masaakia longivaginata Takeuchi, 1950
Japanese name: Hime-yashabushi-habachi(Fig. 8A)

Material examined. HONSHU: Tochigi Pref.:1 ♂, Nakagawa, Yamata, 36°42′N 140°12′E, coll.
larva (Fig. 8A), on Alnus hirsuta var. hirsuta, 16.
V. 2019, em. 9. IV. 2020, S. Ibuki.
Larva. Late instar larva (Fig. 8A) pale green;
trunk with semitransparent branched spines; thoracic spines apically brownish. Mature larva
12 mm long; head pale greenish yellow; trunk
pale green, without spines; legs pale yellow
green, with claws brownish.
Host plants. Betulaceae: Alnus hirsuta (Spach)Turcz. ex Rupr. var. hirsuta (new record), A. pendula Matsum. (Togashi, 1979).
Life history. The sawﬂy has one generation ayear. The larva is solitary and has an extra larval
molt. The ﬁnal instar larva makes a cocoon in thesoil. Hibernation is done in the cocoon.
Remarks. The larva has not been described sofar. The known host plants are only Alnus.
Metallus albipes (Cameron, 1875)
Japanese name: Kiichigo-hamuguri-habachiMaterial examined. HONSHU: Wakayama
Pref.: 1 ♀, Wakayama, Okawa, coll. larva mining leaf of Rubus hirsutus, 14. XI. 2020, em. 19.
III. 2021, M. Murase.
Host plant. Rosaceae: Rubus crataegifolius
Bunge (Okutani, 1967b), R. hirsutus Thunb.
(new record) in Japan; R. fruticosus L., R. idaeus
L. in Europe (Taeger et al., 1998).
Remarks. This sawﬂy feeds on various speciesof Rubus.
Metallus pumilus (Klug, 1816)
Japanese name: Kumaichigo-hamuguri-habachiMaterial examined. HONSHU: Wakayama
Pref.: 1 ♀, Wakayama, Okawa, coll. larva mining leaf of Rubus microphyllus, 14. XI. 2020, em.

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181

Fig. 8. A–C, Late instar larvae: A, Masaakia longivaginata, 21. V. 2019; B, Monophadnoides montanus, 18. VI.
2019; C, Strongylogaster moiwana, 30. V. 2017. All photographed by Ibuki.

25. XII. 2020, M. Murase.
Host plants. Rosaceae: Rubus crataegifolius
Bunge (Okutani, 1970), R. microphyllus L.f.
(new record) in Japan; R. caesius L., R. fruticosus L., R. idaeus L., R. saxatilis L. in Europe(Taeger et al., 1998).
Remarks. This species feeds on various speciesof Rubus.
Monophadnoides montanus Togashi, 1980
New Japanese name: Nawashiro-ichigo-maru-habachi(Fig. 8B)

Material examined. HONSHU: Tochigi Pref.:1 ♂, Nakagawa, Wami, 36°45′N 140°9′E, coll.
larva, on Rubus parvifolius, 16. VI. 2019, em. 9.
IV. 2020, S. Ibuki; 1 ♂, same data but 36°46′N
140°10′E, coll. larva (Fig. 8B) on Rubus parvifolius, 16. VI. 2019, mat. 24. VI., em. 16. IV. 2020.
Larva. Late instar larva (Fig. 8B): head pale

brownish green; trunk entirely pale green except
for green middorsal stripe, with many branchedspines; legs, spines and spiracles whitish. Maturelarva: 12 mm long, pale green, without spines.
Host plant. Rosaceae: Rubus parvifolius L.
(new record).
Life history. This species has one generation ayear. The larva is solitary and eats the inside ofleaves. The larva performs the extra larval molt
and then makes a cocoon in the soil. Hibernation
is done in the cocoon.
Remarks. The larva and host plant arerecorded for the ﬁrst time.
Phymatocera nipponica Togashi, 1958
Japanese name: Higenaga-kuro-habachiMaterial examined. HOKKAIDO: 1 ♂, Bibai,
Koshunai, 43°17′N 141°51′E, 24. V. 2015, H.
Hara.

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Distribution. Japan: Hokkaido (Matsumura,
1912, as “Phymatocerus aterrimus Klug”,
according to Togashi, 1958; Togashi, 1997; present study), Honshu (Yano, 1932, as Phymatocera
aterrima, according to Togashi, 1958), Shikoku(Togashi, 1974), Kyushu (Togashi, 1972). Korea(Doi, 1938, as “Phymatocera aterrima KLUG
[Higenaga-kuro-habachi (in Japanese)]”); Sakhalin (Haris, 2006); Kuril Islands (Sundukov,
2017).
Remarks. Although Naito (2019, 2020a) did
not include Hokkaido in the distribution andNaito (2020a) questioned its occurrence in Hokkaido, Hara collected the above male in central
Hokkaido. It agrees with the original descriptionof Togashi (1958) and goes to this species in the
key to the Japanese species of Phymatocera by
Togashi (2004).
Pristiphora fulviceps Takeuchi, 1933
Japanese name: Kigao-higenaga-habachiMaterial examined. HONSHU: Tochigi Pref.:2 ♀, Nakagawa, Bato, 36°44′N 140°9′E, coll.
larva, on Corylus sieboldiana var. sieboldiana,
17. V. 2019, mat. 18. V., em. 19. IV. 2020, S.
Ibuki.
Host plants. Betulaceae: Carpinus cordata
Blume (Hara et al., 2018), Corylus sieboldiana
Blume var. sieboldiana (new record).
Remarks. The host plants belong to the subfamily Coryloideae.
Siobla ferox (Smith, 1874)
Japanese name: O-koshiaka-habachi(Fig. 9A–G)

Material examined. HONSHU: Tochigi Pref.:1 ♂, Nakagawa, Oyamada, coll. larva, on Actinidea polygama, 14. VI. 2016, mat. 18. VII., em. 2.
V. 2017, S. Ibuki; 1 ♂, Nakagawa, Bicchuzawa,coll. larva (Fig. 9A), on Actinidea polygama, 22.
VI. 2016, diet changed to Glechoma hederacea
subsp. grandis 27. VI., mat. 9. VIII. (Fig. 9D),

em. 6. V. 2017, S. Ibuki; 1 ♀, Nasukarasuyama,coll. larva, on Actinidea polygama, 15. VI. 2016,
mat. ?, em. 6. V. 2017, S. Ibuki.
Larva. Middle or late instar (Fig. 9A–C, E–F):
head dark purplish gray to pale yellowish gray;trunk and legs creamy white, dorsal region pale
gray or pale yellowish gray, sparsely covered
with minute black spots, each segment withpaired yellowish dorsolateral thorns. Maturelarva (Fig. 9D, G): blackish, shiny; thoracic legs
dark yellowish.
Host plants. Actinidiaceae: Actinidia polygama (Siebold et Zucc.) Planch. ex Maxim. (newrecord). Asteraceae: Cirsium pendulum Fisch. ex
DC. (Okutani, 1967b). Balsaminaceae: Impatiens
spp. (Okutani, 1967b). Lamiaceae: Glechomahederacea L. subsp. grandis (A. Gray) H. Hara.
(new record). Polygonaceae: Polygonum cuspidatum Siebold et Zucc. (Okutani, 1967b), Rumex
sp. (Okutani, 1967b).
Life history. This is a univoltine species. The
larva is a solitary external feeder on various
plants. The larval period is very long, about 40
days (Okutani, 1959) or more than 48 days (seebelow). On maturity, the reared larva had an
extra molt and went into the soil.
Remarks. This is the ﬁrst record of Actinidia
polygama (Actinidiaceae) and Glechoma hederacea subsp. grandis (Lamiaceae) as the hosts of S.
ferox. The known hosts of this species now
include more than six species of ﬁve families ofplants. For a detailed description of the larva, see
Okutani (1959). Okutani (1959) noted that thelarval period of this species was about 40 days.
However, one of our cases of rearing indicatedthat the period may be much longer. The larva
collected on June 22, 2016 (Fig. 9A, probably
middle instar) and reared indoors matured on
August 9 (Fig. D). The rearing period from middle instar to maturity in this case lasted 48 days.
The full larval period of this individual is estimated at 60 days at least.

Taxonomic Notes and New Records in Japanese Symphyta VI

183

Fig. 9. A–G, Siobla ferox: A, probably middle instar larva on Glechoma hederacea subsp. grandis, 27. VI. 2016;
B, same larva, probably late instar, 12. VII. 2016; C, same larva, 16. VII. 2016; D, same larva, matured, 9.
VIII. 2016; E. probably middle instar larva on Actinidia polygama, 14. VI. 2016; F. probably middle instar larvae on A. polygama, 19. VI. 2016; G. same larva, 15. VII. 2016. H–I, Siobla japonica: H, last instar larva, 5.
VI. 2019; I, mature larva and cast skin above, 9. VI. 2019; J–M, Taxonus shinoharai: J, middle and late instar
larvae on Agrimonia pilosa var. japonica, 29. IX. 2019; K, late instar, 29. IX. 2019; L, same larva, matured,
30. IX. 2019; M, male adult, just emerged, 26. X. 2019. All photographed by Ibuki.

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Siobla japonica Shinohara, Wei and Niu, 2013
Japanese name: Nihon-koshiaka-habachi(Fig. 9H–I)

Material examined. HONSHU: Tochigi Pref.:1 ♀, Nakagawa, Wami, 33°45′N 140°09′E, coll.
larva, on Impatiens textorii or Circaea mollis, 5. VI.
2019, mat. 8. VI., em. 8. IV. 2020, S. Ibuki; 1 ♂,Nakagawa, Wami, 33°45′N 140°09′E, coll. larva,
on Impatiens textorii or Circaea mollis, 5. VI. 2019,
mat. 9. VI. (Fig. 9I), em. 11. IV. 2020, S. Ibuki.
Larva. Late instar (Fig. 9H): head purplish
black; trunk and legs creamy white, dorsal region
dark gray, sparsely covered with irregular blackspots, each segment with paired yellowish dorsolateral thorns, and black line along ventral margin of pleural region. Mature larva (Fig. 9I):
blackish, shiny; thoracic legs creamy white.
Host plants. Balsaminaceae: Impatiens textorii
Miq. (new record). Onagraceae: Circaea mollis
Siebold et Zucc. (new record).
Life history. This is a univoltine species. The
larva is a solitary external feeder and, whenmatured, it has an extra molt before entering the soil.
Remarks. This species is fairly common incentral Honshu (Shinohara et al., 2013), but its
immature stages and host plants were totally
unknown. Like the preceding species, S. ferox, S.
japonica seems to be a polyphagous species withlong larval feeding period. The larva of this species may be distinguished from that of S. ferox
by the dark color pattern of the head and dorsal
region of the trunk in the feeding stages (compare A–C, E–F with H in Fig. 9) and the creamy
white thoracic legs of the mature larva (compare
D, G with I in Fig. 9).
Stethomostus babai Togashi, 1984
Japanese name: Baba-maru-habachiMaterial examined. HOKKAIDO: 1 ♀, Bibai,
Tomei, 43°20′N 141°54′E, 10. V. 2017, H. Hara;
1 ♀, same data but 11. V. 2019, H. Hara; 1 ♂,same data but 15. V. 2019; 1 ♂, same data but 8.
V. 2020.

Distribution. Japan: Hokkaido (new record),
Honshu (Togashi, 1984), Shikoku (Naito, 2019).
China (Nie and Wei, 1999).
Remarks. This species is here recorded from
Hokkaido for the ﬁrst time.
Strongylogaster moiwana Matsumura, 1912
Japanese name: Iwagane-zenmai-habachi(Fig. 8C)

Material examined. HONSHU: Tochigi Pref.:1 ♂, Nakagawa, Oyamada, coll. larva, on Coniogramme japonica, 30. V. 2017, mat. 7. VI., em.
30. III. 2018, S. Ibuki.
Larva. Late instar larva (Fig. 8C): head pale
brown, black on vertex and lateral part; trunk andlegs pale yellowish green.
Host plants. Onocleaceae: Onoclea sensibilis
L. var. interrupta Maxim. (Naito, 2020a). Pteridaceae: Coniogramme intermedia Hieron.
(Naito, 2002, 2004, 2020a), C. japonica (Thunb.)
Diels (Okutani, 1967b; Naito, 1980, 1996; present study).
Remarks. As the host, Okutani (1967b) andNaito (1980, 1996) recorded Coniogrammejaponica, but Naito (2002, 2020a) and Naito etal. (2004) mentioned C. intermedia instead of C.
japonica without any comment. We conﬁrmed C.
japonica as the host by rearing the larva.
Taxonus shinoharai Togashi, 2009
New Japanese name: Kinmizuhiki-habachi(Fig. 9J–M)

Material examined. HONSHU: Tochigi Pref.:1 ♂,Nasukarasuyama,
Ogane,
36°39′N
140°06′E, coll. larva, on Agrimonia pilosa var.
japonica, 24. IX. 2019, mat. 6. X., em. 13. I.
2020, S. Ibuki; 1 ♂, Nakagawa, Wami, 36°45′N
140°09′E, coll. larva, on Agrimonia pilosa var.
japonica, 26. IX. 2019 (Fig. 9J[larger larva], K),
mat. 30. IX. (Fig. 9 L), em. 26. X. 2019 (Fig.
9M), S. Ibuki; 1 ♀, same locality, coll. larva, on
Agrimonia pilosa var. japonica, 26. IX. 2019

Taxonomic Notes and New Records in Japanese Symphyta VI(Fig. 9J[smaller larva]), mat. 11. X., em. 23. I.
2020, S. Ibuki.
Larva. Middle instar (Fig. 9J[smaller larva]):head brownish white, with large median spot andsmaller lateral spot on vertex black; trunk andlegs creamy white, dorsal region pale greenish,
each segment with small black spot laterally.
Late instar (Fig. 9J[larger larva], K): similar to
middle instar. Mature larva (Fig. 9 L): yellowish
white, shiny, with small blackish lateral spots oneach segment of trunk.
Host plants. Rosaceae: Agrimonia pilosa
Ledeb. var. japonica (Miq.) Nakai (new record).
Life history. This is a multivoltine species. In
our rearing indoors, one larva that matured on
September 30, 2019, emerged as a male adult on
October 26, 2019, another larva that matured on
October 6, 2019, emerged as a male adult on January 13, 2020, and another larva that matured on
October 11, 2019, emerged as a female adult on
January 23, 2020. The larva is a solitary external
feeder and, when matured, it has an extra molt
before entering the soil.
Remarks. This species was ﬁrst described as
Taxonus fulvipes Togashi, 2000, but a new namewas given to it in Blank et al. (2009) because
Taxonus fulvipes Togashi, 2000, is a junior primary homonym of Taxonus fulvipes Cameron,
1899.
The female specimen listed above agrees withthe holotype of Taxonus fulvipes Togashi, 2000,
but does not run to this species in the keys by
Togashi (2000) and Naito (2020) beause of theerroneous statement in the keys. Taxonus shinoharai runs to the couplet 7 in Togashiʼs (2000)
key and the couplet 11 in Naitoʼs (2020) key,both of which read “Mesoscutellum black with asmall milky white macula; last abdominal tergite
milky white” (leading to couplet 8 in Togashiʼs
key and couplet 12 in Naitoʼs key) in the ﬁrst lineand “Mesoscutellum and last abdominal tergite
black” (leading to couplet 9 in Togashiʼs key andcouplet 12 in Naitoʼs key) in the second line. In
T. shinoharai, the mesoscutellum is entirely
black and the last abdominal tergite is milky
white, thus agreeing neither of the two lines of185

the couplet in both keys. Taxonus shinoharai isin the second line in the couplet 9 in Togashiʼs
key and in the couplet 13 in Naitoʼs key, where
the last abdominal tergite is correctly describedas milky white.
This species was described from a pair of
specimens collected in Tokyo in 1990 andTogashi (2002) recorded it from Tochigi Prefecture.
Tomostethus nigritus (Fabricius, 1804)
Japanese name: Yachidamo-habachiMaterial examined. HOKKAIDO: 1 ♀, Abira,Oiwake, 42°52′N 141°48′E, 12. V. 2020, H. Hara.
Distribution. Japan: Hokkaido (new record),
Honshu (Abe and Togashi, 1989). Palaearctic
and Oriental regions.
Remarks. The ﬁrst record of this species fromJapan is Takeuchi (1952), but he did not state thedetails of the distribution. This speccies is here
recorded from Hokkaido for the ﬁrst time.
Acknowledgements
We thank Masaaki Kimura (Naha) andMasumi Murase (Wakayama) for the gift ofmaterial and David R. Smith (United States
Department of Agriculture, Washington, D. C.)for his helpul comments on the manuscript.
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