Research Article |
Corresponding author: Kyohei Watanabe ( himebati-love@hotmail.co.jp ) Academic editor: Jose Fernandez-Triana
© 2021 Kyohei Watanabe, Rikio Matsumoto.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Watanabe K, Matsumoto R (2021) Revision of the genus Xanthopimpla Saussure, 1892 (Hymenoptera, Ichneumonidae, Pimplinae) from Japan. Deutsche Entomologische Zeitschrift 68(2): 269-297. https://doi.org/10.3897/dez.68.69768
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Japanese species of the genus Xanthopimpla Saussure, 1892 are revised. A total of 15 species are recorded from Japan, including three new species, X. nipponensis sp. nov. X. sylvicola sp. nov. and X. yoshimurai sp. nov. and three species, X. honorata (Cameron, 1899), X. minuta Cameron, 1905 and X. trias Townes & Chiu, 1970, newly recorded from Japan. Host, habitat, overwintering and distribution patterns of Japanese Xanthopimpla species are discussed.
Asia, biogeography, bionomics, new species, parasitoid wasp, Ryukyu Islands, taxonomy
Xanthopimpla Saussure, 1892 is a large genus belonging to subfamily Pimplinae and containing 265 species from all zoogeographical regions (
In this study, the dried specimens deposited in the following collections were examined:
NIAES Institute for Agro-Environmental Sciences, NARO, Tsukuba, Ibaragi, Japan;
NSMT National Museum of Nature and Science, Tsukuba, Ibaragi, Japan;
A stereomicroscope (SMZ800: Nikon, Tokyo) was used for observation. Photographs (Figs
Morphological terminology follows
There was some variation in the black maculation on the body amongst specimens identified as X. clavata Krieger, 1914. To clarify the variability amongst species, molecular phylogenetic analysis was conducted for X. clavata and some closely-related species, based on the partial sequences of the mitochondrial gene, cytochrome c oxidase subunit I (COI) and the nuclear gene, 28S rDNA (28S). All specimens used for DNA analysis were collected adults and preserved in 99.5% ethanol. Total genomic DNA was extracted from the right middle legs using a DNeasy Blood and Tissue Kit (Qiagen, The Netherlands) according to the manufacturer’s protocol for animal tissues. Voucher specimens and their extracted genomic DNA were deposited in the Osaka Museum of Natural History. The DNA samples were stored in a freezer (at -40 °C) and the voucher specimens were dried and mounted. The partial sequences of one mitochondrial gene, cytochrome c oxidase subunit I (COI) and one nuclear gene, 28S rDNA (28S), were amplified using the primers designed by
Information on the individual ID, accession numbers, collecting date, site and depository of each DNA-sequenced specimen of Japanese Xanthopimpla (X. clavata, X. yoshimurai sp. nov., X. trias, X. naenia and X. niponensis sp. nov.) shown in Fig.
Voucher ID | Species | Accession number | Sex | Date | Information on collecting sites, person and depository | |
---|---|---|---|---|---|---|
COI | 28S | |||||
Pol083 | Xanthopimpla yoshimurai | LC632431 | LC633946 | ♀ | 30.XII.2011 | Koyodai, Matsuyama, Ehime, R.M. ( |
Pol145 | Xanthopimpla clavata | LC632432 | LC633947 | ♀ | 8.V.2013 | Yona, Okinawajima, Okinawa, R.M. ( |
Pol192 | Apecthis rufata, outgroup | LC632433 | LC633948 | ♂ | 1.VIII.2013 | Bekanbeushi, Akkeshi, Hokkaido, R.M. ( |
Pol311 | Xanthopimpla clavata | LC632434 | LC633949 | ♀ | 31.XII.2014 | Koyodai, Matsuyama, Ehime, R.M. ( |
Pol372 | Xanthopimpla clavata | LC632435 | LC633950 | ♀ | 15.V.2015 | Yatacho, Yamatokouriyama, Nara, R.M. ( |
Pol378 | Xanthopimpla clavata | LC632436 | LC633951 | ♀ | 1.VI.2015 | Byakugouji, Nara, Nara, R.M. ( |
Pol379 | Xanthopimpla clavata | LC632437 | LC633952 | ♀ | 1.VI.2015 | Byakugouji, Nara, Nara, R.M. ( |
Pol386 | Xanthopimpla clavata | LC632438 | LC633953 | ♂ | 21.VI.2015 | Urabudake, Yonagunjimai, Okinawa, R.M. ( |
Pol388 | Xanthopimpla clavata | LC632439 | LC633954 | ♂ | 24.VI.2015 | Shirahama, Iriomotejima, Okinawa, R.M. ( |
Pol401 | Xanthopimpla clavata | LC632440 | LC633955 | ♀ | 25.XII.2015 | Kasugajinja, Sanda, Hyogo, R.M. ( |
Pol557 | Xanthopimpla trias | LC632441 | LC633956 | ♀ | 12.I.2017 | Amakashinooka, Asuka, Nara, R.M. ( |
Pol657 | Xanthopimpla naenia | LC632442 | LC633957 | ♂ | 7.IX.2017 | Heijo Palace site, Nara, Nara, R.M. ( |
Pol664 | Xanthopimpla clavata | LC632443 | LC633958 | ♀ | 4.XII.2017 | Arimafuji, Sanda, Hyogo, R.M. ( |
Pol665 | Xanthopimpla yoshimurai | LC632444 | LC633959 | ♀ | 4.XII.2017 | Arimafuji, Sanda, Hyogo, R.M. ( |
Pol711 | Xanthopimpla nipponensis | LC632445 | LC633960 | ♀ | 19.I.2019 | Yatacho, Yamatokoriyama, Nara, R.M. ( |
Pol712 | Xanthopimpla clavata | LC632446 | – | ♀ | 19.I.2019 | Yatacho, Yamatokoriyama, Nara, R.M. ( |
Pol713 | Xanthopimpla yoshimurai | LC632447 | LC633961 | ♀ | 13.I.2019 | Taishoike, Ide, Kyoto, R.M. ( |
Pol718 | Xanthopimpla nipponensis | LC632448 | LC633962 | ♀ | 21.I.2019 | Arimafuji, Sanda, Hyogo, R.M. ( |
Pol719 | Xanthopimpla clavata | LC632449 | LC633963 | ♀ | 21.I.2019 | Arimafuji, Sanda, Hyogo, R.M. ( |
Pol720 | Xanthopimpla yoshimurai | LC632450 | LC633964 | ♀ | 21.I.2019 | Arimafuji, Sanda, Hyogo, R.M. ( |
Pol721 | Xanthopimpla clavata | LC632451 | LC633965 | ♀ | 28.I.2019 | Hirae, Ishigakijima, Okinawa, R.M. ( |
Pol724 | Xanthopimpla clavata | LC632452 | LC633966 | ♂ | 29.I.2019 | Hirae, Ishigakijima, Okinawa, R.M. ( |
Pol726 | Xanthopimpla clavata | LC632453 | LC633967 | ♀ | 30.I.2019 | Hirae, Ishigakijima, Okinawa, R.M. ( |
Pol732 | Xanthopimpla clavata | LC632454 | LC633968 | ♂ | 27.IV.2019 | Arakawa, Ishigakijima, Okinawa, R.M. ( |
Pol738 | Xanthopimpla naenia | LC632455 | LC633969 | ♂ | 12.V.2019 | Aonogahara, Ono, Hyogo, R.M. ( |
Pol826 | Xanthopimpla trias | LC632456 | LC633970 | ♀ | 14.I.2020 | Amakashinooka, Asuka, Nara, R.M. ( |
Pol827 | Xanthopimpla trias | LC632457 | LC633971 | ♀ | 5.II.2020 | Hattori-ryokuchi, Osaka,Osaka, R.M. ( |
Pol851 | Xanthopimpla trias | LC632458 | LC633972 | ♀ | 20.V.2020 | Kasugayama, Nara, Nara, R.M. ( |
Pol923 | Xanthopimpla naenia | LC632459 | LC633973 | ♂ | 3.XI.2020 | Heijo Palace site, Nara, R.M. ( |
Polymerase chain reactions (PCRs) were conducted in 30 μl final volumes using ExTaq Hot Start (Takara Bio Inc., Japan). The PCR programme for COI comprised initial denaturation for 5 min at 94–95 °C followed by 35–40 cycles of denaturation at 94 °C for 15 s, annealing at 46 °C for 15 s, extension at 72 °C for 15 s and a final extension at 72 °C for 10 min. The reaction conditions for the 28S rRNA fragment were the same, except the annealing temperature was modified to 50 °C and 57 °C and the extension time was modified to 30 s and 60 s, respectively. The PCR products were purified using Illustra Exo-ProStar (GE Healthcare, USA). The purified PCR products were mixed with primers and sent to the CDM Center (Takara Bio Inc.) and run on an ABI 3730xl DNA Analyzer (Applied Biosystems). All sequences generated in this study were submitted to INSD under accession numbers LC632431–LC632459 and LC633946–LC633973, as summarised in Table
The forward and reverse sequences were checked, assembled and edited using Seaview (
In total, 15 species of Xanthopimpla were identified in Japan (Table
List of Japanese Xanthopimpla. P_CHI: Palaearctic part of China; HON: Honshu; IZU: Izu Islands; SHI: Shikoku; KYU; Kyushu (including Tsushima Is.); YAK; Yakushima Is. (including Kuchinoerabu Is.); TOK: Tokara Isls.; AMA: Amami Isls. (including Amamioshima Is., Kakeroma Is. and Tokunoshima Is.); OKI: Okiwana Isls. (including Okinoerabu Is., Yoron Is., Okinawa Is. and Miyako Is.); YAE: Sakishima Isls. (including Ishigaki Is., Taketomi Is., Iriomote Is. and Yonaguni Is.); FOR: Taiwan; O_CHI: Oriental part of China; VIE: Vietnam. “?” is doubtful record.
Palaearctic region | Oriental region | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
P_CHI | HON | IZU | SHI | KYU | YAK | TOK | AMA | OKI | YAE | FOR | O_CHI | VIE | |
Xanthopimpla Saussure, 1892 | |||||||||||||
The brachycentra species group | |||||||||||||
clavata Krieger, 1914 | | | | | | | | | | | | | |
yoshimurai sp. nov. | | | |||||||||||
The citrina species group | |||||||||||||
flavolineata Cameron, 1907 | | | | | | ||||||||
The incompleta species group | |||||||||||||
naenia Morley, 1913 | | | | | | | | | | | | ||
The occidentalis species group | |||||||||||||
honorata honorata (Cameron, 1899) | | | | | | | |||||||
nipponensis sp. nov. | | | | ||||||||||
The punctata species group | |||||||||||||
punctata (Fabricius, 1781) | | | | | | | | | | ||||
The regina species group | |||||||||||||
brullei Krieger, 1899 | ? | ||||||||||||
konowi Krieger, 1899 | | | | | | ||||||||
pedator (Fabricius, 1775) | | | | | | | | | | ||||
The stemmator species group | |||||||||||||
modesta modesta (Smith, 1860) | | | | ||||||||||
stemmator (Thunberg, 1822) | | | | | |||||||||
The terebatrix species group | |||||||||||||
sylvicola sp. nov. | | | | ||||||||||
The trunca species group | |||||||||||||
minuta Cameron, 1905 | | | | | | | |||||||
trias Townes & Chiu, 1970 | | | | | | | |
Subfamily Pimplinae Wesmael, 1845
Tribe Pimplini Wesmael, 1845
Xanthopimpla
Saussure, 1892 in Grandidier: pl. 13, figs. 1, 2, 3. Type: Xanthopimpla hova Saussure, 1982. Designated by
Chloropimpla Saussure, 1892 in Grandidier: pl. 13, fig. 5. Type: Chloropimpla dorsigera Saussure, 1892. Monotypic.
Notopimpla Krieger, 1899: 106. Type: Pimpla terminalis Brullé, 1846. Monotypic.
Neopimploides Viereck, 1912: 151. Type: Neopimploides syleptae Viereck, 1912 (= Ichneumon punctatus Fabricius, 1781). Original designation.
Austrapophua
Girault, 1926: 135. Type: Austrapophua xanthopimploides Girault, 1925 (= Xanthopimpla rhopaloceros Krieger, 1915). Designated by
This genus can be distinguished from other pimpline genera by the following character states: clypeus divided into dorsal and ventral parts by a transverse suture (Fig.
Japanese Xanthopimpla A, B. X. nipponensis sp. nov.; C. X. pedator (Fabricius, 1775); D. X. punctata (Fabricius, 1781); E. X. stemmator (Thunberg, 1822); F. X. sylvicola sp. nov.; G. X. trias Townes & Chiu, 1970; H, I. X. yoshimurai sp. nov. A, C–H. dorsal habitus; B. mesoscutum, dorsal view; I. propodeum and T I, dorsal view.
Japanese Xanthopimpla A, H. X. clavata Krieger, 1914; B, J. X. konowi Krieger, 1899; C. X. modesta modesta (Smith, 1860); D, K, N, P. X. nipponensis sp. nov.; E. X. stemmator (Thunberg, 1822); F, L. X. sylvicola sp. nov.; G, M. X. yoshimurai sp. nov.; I. X. minuta Cameron, 1905; O. X. naenia Morley, 1913 (D, F, G, K–M, P. holotype; N. paratype) ― A, B. head, frontal view; C–G. head, dorsal view; H, I. mesonotum, dorsal view; J–M. scutellum, lateral view; N. mesosternum, ventral view; O, P. areolet of right wing.
1 | Mesosoma and metasoma entirely yellow (Fig. |
2 |
– | Mesosoma and metasoma with some black markings (e.g. Fig. |
3 |
2 | Area between oceller area and eye without black markings (Fig. |
X. flavolineata Cameron, 1907 |
– | Area between oceller area and eye with large black markings (Fig. |
X. modesta modesta (Smith, 1860) (in part) |
3 | Areolet absent (vein 3rs-m completely absent) (Fig. |
X. naenia Morley, 1913 |
– | Areolet present (vein 3rs-m at least partly present) (Fig. |
4 |
4 | Scutellum subconically elevated in lateral view (Fig. |
5 |
– | Scutellum not subconically elevated in lateral view (Fig. |
7 |
5 | Punctuation on T III to T V relatively dense, i.e. T IV having the punctures on its sublateral black spots separated by 0.3–1.3 × their diameter and T III having numerous punctures between its black spots (Fig. |
X. pedator (Fabricius, 1775) |
– | Punctuation on T III to T V sparser i.e. T IV having the punctures on its sublateral black spots separated by 1.3–6.0 × their diameter and T III having few or no punctures between its black spots (Fig. |
6 |
6 | Black spots on T IV each with about 20 punctures (Fig. |
X. konowi Krieger, 1899 |
– | Black spots on T IV each with about 10 punctures. Hind tibia with 1 to 4 pre-apical bristles, scattered from apex towards middle of tibia | X. brullei Krieger, 1899 |
7 | Hind tibia entirely yellow, without a basal black area (Fig. |
8 |
– | Hind tibia with a narrow, but conspicuous basal black area (Fig. |
9 |
8 | Area between oceller area and eye black (Fig. |
X. modesta modesta (Smith, 1860) (in part) |
– | Area between oceller area and eye yellow (Fig. |
X. stemmator (Thunberg, 1822) |
9 | Propodeum without carinae and black spots (Fig. |
X. trias Townes & Chiu, 1970 |
– | Propodeum at least partly with carinae and sometimes with black spots (Fig. |
10 |
10 | Propodeum with lateral section of anterior transverse carina joined to area superomedia at or near posterior angle of the area (Fig. |
X. punctata (Fabricius, 1775) |
– | Propodeum with lateral section of anterior transverse carinae joined to area superomedia anterior to posterior angle of the area (Fig. |
11 |
11 | Notauli long, extending posteriorly beyond centre of mesoscutum, their posterior ends joined with each other (Fig. |
X. minuta Cameron, 1905 |
– | Notauli short, not extending posteriorly beyond centre of mesoscutum, their posterior ends not joined with each other (Fig. |
12 |
12 | Posterior transverse carina of propodeum incomplete medially (Fig. |
X. honorata honorata (Cameron, 1899) |
– | Posterior transverse carina of propodeum complete (weak in X. sylvicola, but this species has black spots on propodeum) (Fig. |
13 |
13 | Longest bristle on hind tarsal claw distinctly widened next to apex, with a mucronate apex (Fig. |
X. sylvicola sp. nov. |
– | Longest bristle on hind tarsal claw slender, not widened next to apex (Fig. |
14 |
14 | Mesoscutum smooth in front of scuto-scutellor groove (Fig. |
X. nipponensis sp. nov. |
– | Mesoscutum with fine punctures in front of scuto-scutellor groove. Propodeum with a pair of black markings (Fig. |
15 |
15 | Black spots of propodeum triangular (Fig. |
X. clavata Krieger, 1914 |
– | Black spots of propodeum semicircular (Fig. |
X. yoshimurai sp. nov. |
Xanthopimpla brullei
Krieger, 1899: 88;
This species belongs to the regina species group sensu
Japan (Okinawa Is.?). Outside Japan, this species has been recorded from Indonesia, Malaysia and Philippines (
Unknown in Japan.
Although
Xanthopimpla clavata
Krieger, 1914: 40, 91;
Xanthopimpla minomensis
Uchida, 1932: 157;
This species belongs to the brachycentra species group sensu
Propodeum of Japanese Xanthopimpla, dorsal view A. X. clavata Krieger, 1914; B. X. flavolineata Cameron, 1907; C. X. honorata honorata (Cameron, 1899); D. X. konowi Krieger, 1899; E. X. minuta Cameron, 1905; F. X. modesta modesta (Smith, 1860); G. X. naenia Morley, 1913; H. X. nipponensis sp. nov.; I. X. pedator (Fabricius, 1775); J. X. punctata (Fabricius, 1781); K. X. stemmator (Thunberg, 1822); L. X. sylvicola sp. nov.; M. X. trias Townes & Chiu, 1970; N. X. yoshimurai sp. nov. (H, L, N. holotype).
Japanese Xanthopimpla ― A. X. clavata Krieger, 1914; B. X. flavolineata Cameron, 1907; C. X. honorata honorata (Cameron, 1899); D. X. konowi Krieger, 1899; E. X. minuta Cameron, 1905; F. X. modesta modesta (Smith, 1860); G, H. X. naenia Morley, 1913; I, P. X. nipponensis sp. nov.; J. X. pedator (Fabricius, 1775); K. X. punctata (Fabricius, 1781); L. X. stemmator (Thunberg, 1822); M, Q. X. sylvicola sp. nov.; N. X. trias Townes & Chiu, 1970; O, R. X. yoshimurai sp. nov. (I, M, O–R. holotype) A–O. left hind leg, lateral view; P–R. left hind tarsal claw, lateral view.
Type series: [Honshu] 1 F (holotype of X. minomensis), Osaka Pref., Mt. Minoo, 1 Jul 1920, C. Teranishi leg. (
Japan (Honshu, Izuoshima Is., Shikoku, Kyushu, Tsushima Is., Yakushima Is., Amamioshima Is., Kakeroma Is., Tokunoshima Is., Okinawa Is., Miyako Is., Ishigaki Is., Taketomi Is., Iriomote Is. and Yonaguni Is.). Outside Japan, this species has been recorded from China, Malaysia and Taiwan (
In Japan, adults were collected in all months, except for November. In Honshu, winter is passed in the stage of adult (Fig.
This is the first record of this species from Nakanoshima Is., Kakeroma Is., Tokunoshima Is., Taketomi Is. and Yonaguni Is. Both
Xanthopimpla flavolineata
Cameron, 1907a: 48;
Xanthopimpla emaculata
Szépligeti, 1908: 256;
Xanthopimpla immaculata
Morley, 1913: 115. Synonymised by
Xanthopimpla hyaloptila
Krieger, 1915: 35;
Xanthopimpla xanthostigma
Giraut, 1925: 38;
Xanthopimpla xara
Cheesman, 1936: 179;
Metopius sesamiae
Rao, 1953: 184. Synonymised by
Xanthopimpla sesamiae:
This species belongs to the citrina species group sensu
Japan: [Ryukyu Isls.] 1 F, Okinawa Pref., Ishigaki Is., Yonehara, 15 Oct 1981, K. Konishi leg. (NIAES); 1 F, Okinawa Pref., Ishigaki Is., Mt. Omotodake, 19 Jan 1998, K. Takahashi leg. (NIAES); 1 M, Okinawa Pref., Iriomote Is., Otomi-rindo, 20 Nov 1980, E. Nishida leg. (
Japan (Ishigaki Is., Iriomote Is. and Yonaguni Is.). Outside Japan, this species has been recorded from Australia, Bangladesh, China, India, Indonesia, Laos, Malaysia, Nepal, New Caledonia, Pakistan, Palau, Papua New Guinea, Philippines, Solomon Islands, Sri Lanka, Taiwan, Truk Islands, Vanuatu and Vietnam (
In Japan, adults were collected in January, March, May and from October to December.
In Japan, distribution of this species is restricted in Yaeyama Islands of Ryukyus.
This species belongs to the occidentalis species group sensu
This is the first record of this species from Japan. This species is divided into three subspecies, X. honorata honorata (Cameron, 1899), X. atriculinata Chao, 1997 and X. honorata munda Krieger, 1915. All Japanese materials are identified as X. honorata honorata.
Pimpla honorata Cameron, 1899: 170.
Xanthopimpla cera
Cameron, 1908: 38;
Xanthopimpla kriegeriana
Cameron, 1908: 38. Synonymised by
Xanthopimpla binghami
Cameron, 1908: 39. Synonymised by
Xanthopimpla erythroceros
Krieger, 1915: 32;
Xanthopimpla erythroceros var. assamensis
Krieger, 1915: 99. Synonymised by
Xanthopimpla honorata honorata:
Xanthopimpla
sp. A:
This subspecies can be distinguished from other subspecies by the propodeum and T I with black markings and the hind slope of vertex entirely yellow.
Japan: [Ryukyu Isls.] 2 F, Kagoshima Pref., Amamioshima Is., Mt. Yuwan, 3 May 1953, T. Shiraki leg. (NIAES); 2 F, Kagoshima Pref., Amamioshima Is., Yuwan, 7 May 1953, T. Shiraki leg. (NIAES); 1 F, Kagoshima Pref., Amamioshima Is., Shinokawa, 15 May 1953, T. Shiraki leg. (NIAES); 1 F, Okinawa Pref., Okinawa Is., Kunigami Vil., Mt. Yonahadake, 6 Apr 1979, K. Ohara leg. (
Japan (Amamioshima Is., Okinawa Is. and Iriomote Is.). Outside Japan, this subspecies has been recorded from China, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Philippines, Singapore, Taiwan, Thailand and Vietnam (
In Japan, adults were collected in April, May, June, October and November. In Iriomote Is., the first author collected this species in the forest path with sunlight. We record a host, Pithecops corvus ryukyuensis Shirozu, 1964 (Lepidoptera, Lycaenidae), here.
This is the first record of this subspecies from Japan. No characteristics unique to the Japanese population were detected.
Xanthopimpla konowi
Krieger, 1899: 87;
Xanthopimpla iaponica
Krieger, 1899: 81;
Xanthopimpla anthereae
Cameron, 1911: 46;
Xanthopimpla watsoni
Cameron, 1911: 46;
Xanthopimpla princeps
Krieger, 1914: 43;
Xanthopimpla dux
Krieger, 1914: 43;
Xanthopimpla formosensis
Krieger, 1914: 43. Synonymised by
Xanthopimpla macrodactyla
Krieger, 1914: 42. Synonymised by
Xanthopimpla grandis
Cushman, 1925: 43. Synonymised by
Xanthopimpla theophilae
Rao, 1953: 159;
Xanthopimpla japonica
(!): Matsumura, 1912: 138;
This species belongs to the regina species group sensu
Japanese Xanthopimpla A, F. X. nipponensis sp. nov.; B, G. X. sylvicola sp. nov.; C, H. X. yoshimurai sp. nov.; D. X. konowi Krieger, 1899; E. X. pedator (Fabricius, 1775) (A–C, F. holotype; G, H. paratype) A–C. T I, dorsal view; D, E. T III, dorsal view; F–H. apex of ovipositor, lateral view.
Japan: [Ryukyu Isls.] 1 F, Okinawa, 7 Jul 329, Matsumura leg. (
Japan (Okinawa Is., Ishigaki Is. & Iriomote Is.). Outside Japan, this species has been recorded from China, India, Indonesia, Malaysia, Myanmar, Taiwan, Thailand, Togo and Vietnam (
In Japan, adults were collected in March, May, June, July and October. In Iriomote Is., the first author collected this species in the forest edge. Dendrolimus spectabilis (Butler, 1877) (Lepidoptera, Lasiocampidae), is recorded as a host of this species in Japan (
At least part of the records of “X. iaponica” and “X. japonica” (misspelling of iaponica) may be due to misidentification of this species and some of the rest may be of X. pedator. We could not examine the voucher specimens of
Xanthopimpla minuta Cameron, 1905a: 137.
Xanthopimpla ischnoceros
Krieger, 1915: 23. Synonymised by
Xanthopimpla ischnoceros var. assamensis Krieger, 1915: 135.
Xanthopimpla ischnoceros assamensis:
Xanthopimpla ischnoceros ischnoceros:
Xanthopimpla
sp. D:
This species belongs to the trunca species group sensu
Japan: [Ryukyu Isls.] 1 F, Kagoshima Pref., Yakushima Is., Shiratani, 10 Jul–8 Aug 2000, T. Murata leg. (MT) (
Japan (Yakushima Is., Amamioshima Is. and Okinawa Is.). Outside Japan, this species has been recorded from China, India, Malaysia, Nepal, Philippines, Sri Lanka, Taiwan, Thailand and Vietnam (
In Japan, adults were collected from June to November. Host is unknown.
This is the first record of this species from Japan. Although this species is divided into five subspecies, X. minuta minuta Cameron, 1905, X. minuta aurangabadensis Patil & Nikam, 1995, X. minuta lita Townes & Chiu, 1970, X. minuta lotipes Townes & Chiu, 1970 and X. minuta quadrula Chao, 1997,
This species belongs to the stemmator species group sensu
This species is divided into two subspecies, X. modesta modesta (Smith, 1860) and X. modesta microcephala Krieger, 1914. All Japanese materials are identified as the former.
Pimpla modesta Smith, 1860: 64.
Xanthopimpla latebalteata
Cameron, 1903: 137;
Xanthopimpla kuchingensis
Cameron, 1905b: 119. Synonymised by
Xanthopimpla dohrni
Krieger, 1915: 34. Synonymised by
Xanthopimpla dohrni var. sukabumensis
Krieger, 1915: 34. Synonymised by
Xanthopimpla dohrni var. novarae
Krieger, 1915: 34. Synonymised by
Xanthopimpla modesta:
Xanthopimpla modesta modesta:
This subspecies can be distinguished from other subspecies by the central part of hind slope of vertex black, the mesoscutum and metasomal tergites usually with black or dark brown markings.
[Ryukyu Isls.] Japan: 1 F, Okinawa Pref., Ishigaki Is., Omoto-Path, 6 Jul 1998 T. Matsumura leg. (NIAES); 1 F, Okinawa Pref., Iriomote Is., Nakamagawa-rindo, 8 Aug 1995, R. Matsumoto leg. (
Japan (Ishigaki Is. and Iriomote Is.). Outside Japan, this subspecies has been recorded from Indonesia, Malaysia, Philippines, Singapore, Thailand, Taiwan and Vietnam (
In Japan, adults were collected in March and July. Host is unknown in Japan, whereas
In Japan, distribution of this species is restricted to Yaeyama Islands of Ryukyus. A black spot of mesoscutum is sometimes absent in Japanese specimens.
Xanthopimpla naenia
Morley, 1913: 115;
Xanthopimpla imperfecta
Krieger, 1915: 23;
This species belongs to the incompleta species group sensu
Japan: [Honshu] 1 M (DNA-Pol-657), Nara Pref., Nara City, Saki-cho, Heijo Palace Site, 7. Sep 2017, R. Matsumoto leg. (
Japan (Honshu, Shikoku, Kyushu, Yakushima Is., Amamioshima Is., Tokunoshima Is., Okinawa Is., Ishigaki Is. and Iriomote Is.). Outside Japan, this species has been recorded from China, India, Malaysia, Philippines, Taiwan and Vietnam (
In Japan, adults were collected from April to November. In Tokunoshima Is., the first author collected this species along the edge of forest.
This is the first record of this species from Ishigaki Is. and Iriomote Is.
Xanthopimpla brachyparea:
Xanthopimpla
sp. nov.:
Xanthopimpla
sp. C:
Holotype: F, Japan, Honshu, Kanagawa Pref., Odawara City, Iriuda, Mt. Ishigaki-yama, 4 Sep 2014, K. Watanabe leg. (
This species belongs to the terebatorix species group sensu
Female (n = 18). Body covered with silver setae, polished, largely smooth, length 7.3–11.2 (HT: 10.5) mm,
Head 0.53 × length of width in dorsal view. Clypeus almost flat in lateral view, sparsely punctate, except for ventral margin, 0.63–0.67 (HT: 0.65) × length of maximum width. Face 1.0 × length of maximum width, sparsely punctate laterally (separated by ca. 1.0–2.0 × their diameter) and densely punctate medially (separated by ca. 0.2 × their diameter). Frons with a conspicuous convexity medially. Length of malar space 0.3 × length of basal mandibular width. OD: POL: OOL = 1.0: 0.7–0.75 (HT: 0.7): 0.75–0.85 (HT: 0.85). Antenna longer than fore wing. Flagellum with 33–35 (HT: 34) flagellomeres. Length of FL I 4.0 × length of maximum depth in lateral view, 1.48–1.54 (HT: 1.48) × length of F II.
Mesosoma. Epomia short. Front end of notaulus with a sharp-edged transverse crest. Notauli not reaching past centre of mesoscutum, their posterior ends not joined with each other. Mesoscutum sparsely and finely punctate anteriorly, smooth posteriorly (Fig.
Metasoma. T I 1.0–1.1 (HT: 1.1) × length of maximum width, largely smooth, with a weak transverse depression subapically (Fig.
Colouration (Figs
Male (n = 1). Similar to female. Length of malar space 0.2 × length of basal mandibular width. Pre-apical bristles of mid-tibia 3. Apical bristle of mid-tibia 4. T II with a pair of conspicuous black spots.
Japan (Honshu, Shikoku and Kyushu).
In Japan, adults were collected in February, May to October and December. In Honshu, winter is passed in the stage of adult (Fig.
The specific name is from Nippon (= Japan).
Ichneumon punctator
Linnaeus, 1767: 935. Name preoccupied by Müller 1766. Synonymised by
Ichneumon pedator Fabricius, 1775: 828.
Ichneumon multipunctor
Thunberg, 1822: 262. Synonymised by
Xanthopimpla scutata
Krieger, 1899: 85;
Xanthopimpla punctatrix
Schulz, 1906: 114. Synonymised by
Xanthopimpla braueri
Krieger, 1914: 43;
Xanthopimpla manilensis
Krieger, 1914: 43;
Xanthopimpla multipunctor:
Xanthopimpla iaponica:
Xanthopimpla japonica
(!):
Xanthopimpla punctator:
Xanthopimpla pedator:
This species belongs to the regina species group sensu
Japan: [Kyushu] 1 F, Kagoshima Pref., Kagoshima City, Uearatacho, 21 Aug 1964, K. Hashimoto leg., 24 Aug. em. from Melacosoma neustria testacea Motsc. (
Japan (Kyushu, Nakanoshima Is., Amamioshima Is., Tokunoshima Is., Okinawa Is., Miyako Is. and Ishigaki Is.). Outside Japan, this species has been recorded from Bangladesh, China, France, India, Indonesia, Malaysia, Myanmar, Pakistan, Philippines, Singapore, Taiwan and Vietnam (
In Japan, adults were collected from April to August and October.
This is the first record of this species from Kyushu, Kuchinoerabujima Is, Takarajima Is., Tokunoshima Is. and Miyako Is. We examined the voucher specimens of “X. japonica” used in
Ichneumon punctatus Fabricius, 1781: 437.
Ichneumon punctator
Thunberg, 1822: 262. Synonymised by
Pimpla transversalis
Vollenhoven, 1879: 146. Synonymised by
Pimpla ceylonica
Cameron, 1899: 165. Synonymised by
Xanthopimpla ruficornis
Krieger, 1899: 103. Synonymised by
Zanthopimpla
(!) appendiculata Cameron, 1902: 51. Synonymised by
Xanthopimpla brunneciornis
Cameron, 1903: 139. Synonymised by
Xanthopimpla kandyensis
Cameron, 1905b: 136. Synonymised by
Xanthopimpla maculiceps
Cameron, 1905c: 37. Synonymised by
Xanthopimpla lissonota
Cameron, 1906: 115. Synonymised by
Xanthopimpla kriegeri
Szépligeti, 1908: 255. Synonymised by
Neopimploides syleptae
Viereck, 1912: 151. Synonymised by
Xanthopimpla tibialis
Morley, 1913: 124. Synonymised by
Xanthopimpla szepligetii
Krieger, 1914: 131. Synonymised by
Phygadenon
(!) punctator Ishida, 1915: 106. Synonymised by
Xanthopimpla pyraustae
Rao, 1953: 163;
Xanthopimpla trimaculata:
Xanthopimpla punctata:
This species belongs to the punctata species group sensu
Japan: [Ryukyu Isls.] 1 M, Kagoshima Pref., Tokara Isls., Nakanoshima Is., Satomura, 5–8 Jun 2005, T. Mita leg. (YPT); 1 F, Kagoshima Pref. Amamioshima Is., Apr 1954, T. Kumata leg. (
Japan (Shikoku, Nakanoshima Is., Amamioshima Is., Tokunoshima Is., Okinoerabu Is., Yoron Is., Okinawa Is., Miyako Is., Ishigaki Is., Iriomote Is. and Yonaguni Is.). Outside Japan, this species has been recorded from Afghanistan, Bangladesh, China, Guam, India, Indonesia, Laos, Malaysia, Mariana Is., Mauritius, Myanmar, Nepal, Nigeria, Pakistan, Papua New Guinea, Philippines, Poland, Russia, Singapore, Sri Lanka, Taiwan, Thailand, Togo and Vietnam (
In Japan, adults were collected from March to October. In Iriomote Is., the first author collected this species in the open forests around crop fields of sugar cane, the forest edge and the forest path in sunlight.
This is the first record of this species from Nakanoshima Is., Yoron Is. and Yonaguni Is.
Ichneumon stemmator Thunberg, 1822: 262.
Pimpla integrata
Smith, 1860: 140. Synonymised by
Xanthopimpla thoracalis
Krieger, 1899: 95. Synonymised by
Xanthopimpla maculifrons
Cameron, 1903: 138;
Xanthopimpla bimaculata
Cameron, 1906: 116. Synonymised by
Xanthopimpla nursei
Cameron, 1907b: 592;
Xanthopimpla maculifrons
Cameron, 1907b: 591. Name preoccupied. Synonymised by
Xanthopimpla facialis
Szépligeti, 1908: 256. Synonymised by
Xanthopimpla stemmator var. confluens
Krieger, 1914: 27. Synonymised by
Xanthopimpla stemmator var. doleschali
Krieger, 1915: 34. Synonymised by
Xanthopimpla transfuga
Krieger, 1915: 38;
Habropimpla sesamiae
Rao, 1953: 166. Synonymised by
Xanthopimpla sesamiae:
Xanthopimpla stemmator:
This species belongs to the stemmator species group sensu
Japan: [Ryukyu Isls.] 1 M, Okinawa Pref., Ishigaki Is., Yoshihara, 17 Sep 1967, K. Mizusawa leg. (
Japan (Ishigaki Is., Iriomote Is. and Hateruma Is.). Outside Japan, this species has been recorded from China, India, Indonesia, Laos, Malaysia, Mauritius, Pakistan, Philippines, Reunion, Singapore, South Africa, Sri Lanka, Taiwan, Thailand and Vietnam (
In Japan, adults were collected in March, May to November. In Iriomote Is., the first author collected this species in the open forests around crop fields of sugar cane.
This is the first record of this species from Hateruma Is. In Japan, distribution of this species is restricted to Yaeyama Islands of Ryukyus.
Xanthopimpla sp. E: Watanabe, 2011: 18.
Holotype: F, Japan, Kagoshima Pref., Amamioshima Is., Yamato Vil., Oodana, 29 Jun 2011, K. Watanabe leg. (
This species belongs to the terebatrix species group sensu
Female (n = 7). Body covered with silver setae, except for some dark brown setae on mesoscutum, polished, largely smooth, length 7.8–9.4 (HT: 9.2) mm.
Head 0.54–0.56 (HT: 0.56) × length of width. Clypeus almost flat in lateral view, sparsely punctate, except for ventral margin, 0.56–0.59 (HT: 0.56) × length of maximum width. Face 0.91–0.95 (HT: 0.91) × length of maximum width, densely punctate. Frons with a slight convexity medially. Length of malar space 0.2–0.25 (HT: 0.2) × length of basal mandibular width. OD: POL: OOL = 1.0: 0.7–0.8 (HT: 0.8): 0.55–0.6 (HT: 0.6). Antenna longer than fore wing. Flagellum with 33–35 (HT: 35) flagellomeres. Length of FL I 3.3–3.6 (HT: 0.3) × length of maximum depth in lateral view, 1.43 × length of F II.
Mesosoma. Epomia very short. Front end of notaulus with a sharp-edged transverse crest. Notauli not reaching past centre of mesoscutum, their posterior ends not joined with each other. Mesoscutum sparsely and finely punctate, its anterior end protruded anteriorly. Scutellum sparsely and finely punctate, roundly convex, with a lateral carina that reaches apex (Fig.
Metasoma. T I 1.0–1.1 (HT: 1.1) × length of maximum width, largely smooth, with a weak transverse depression subapically (Fig.
Colouration (Figs
Male. Unknown.
Japan (Honshu, Kyushu and Amamioshima Is.).
In Japan, adults were collected from May to July, October and December. The authors collected this species along a path inside the broad-leaved forest in Kyushu and Amamioshima Is. Host is unknown.
The species name is from Latin “silva” + “colo”, which is based on the habitat of this species.
Xanthopimpla trias Townes & Chiu, 1970: 242.
Xanthopimpla sp. B: Watanabe, 2011: 17.
This species belongs to the trunca species group sensu
Type series: Taiwan: 1 F (paratype), Koshun, 25 Apr–25 May 1918, J. Sonan, K. Miyake & M. Yoshino leg. (
Japan (Honshu, Kyushu, Yakushima Is., Amamioshima Is. and Okinawa Is.). Outside Japan, this species has been recorded from India, Nepal, Taiwan, Thailand and Vietnam (
In Japan, adults were collected in January, February and from May to October. Although a small number of individuals have been observed, winter is passed in the stage of adult in Honshu (Fig.
This is the first record of this species from Japan. No characteristics unique to the Japanese population were detected.
Holotype: F, Japan, Honshu, Kyoto Pref., Maizuru City, Nyo, 10–30 Aug 2010, T. Murao leg. (MT) (
This species belongs to the brachycentra species group sensu
Female (n = 29). Body covered with silver setae, polished, largely smooth, length 7.9–10.4 (HT: 10.4) mm.
Head 0.51–0.53 (HT: 0.51) × length of width. Clypeus slightly convex in lateral view, sparsely punctate, except for ventral margin, 0.59–0.61 (HT: 0.61) × length of maximum width. Face 0.95–1.05 (HT: 0.98) × length of maximum width, punctate. Frons without a conspicuous convexity medially. Length of malar space 0.15–0.2 (HT: 0.2) × length of basal mandibular width. OD: POL: OOL = 1.0: 0.5–0.7 (HT: 0.7): 0.5–0.75 (HT: 0.7). Antenna longer than fore wing. Flagellum with 31–34 (HT: 34) flagellomeres. Length of FL I 5.0 × length of maximum depth in lateral view, 1.54 × length of F II.
Mesosoma. Epomia very short. Front end of notaulus with a sharp-edged transverse crest. Notauli not reaching past centre of mesoscutum, their posterior ends not joined with each other. Mesoscutum sparsely and finely punctate, its anterior end weakly protruded anteriorly. Scutellum sparsely and finely punctate, roundly convex, with a lateral carina that reaches apex (Fig.
Metasoma. T I 1.0–1.1 (HT: 1.0) × length of maximum width, largely smooth, with a weak transverse depression subapically (Fig.
Colouration (Figs
Male. Unknown.
Japan (Honshu, Shikoku and Kyushu).
In Japan, adults were collected in January, February, June to September and December. Winter is passed in the stage of adult (Fig.
The specific name is after Hiroyuki Yoshimura (Sanda City), who collected part of the paratypes and first noticed the differences in body maculation from X. clavata.
Although this species is morphologically very similar to X. clavata and these two species can be distinguished from each other by mainly body colouration, the difference in colouration is quite stable. Furthermore, the DNA sequences of COI and 28S rRNA are considerably different from each other and both species formed distinct clades with high supporting values in phylogenetic analysis, respectively (Fig.
Phylogenetic trees showing the relationships of five species of Japanese Xanthopimpla (X. clavata, X. yoshimurai sp. nov., X. trias, X. naenia and X. niponensis sp. nov.) See Table
The host records of Japanese Xanthopimpla remain poorly documented. According to
1) Parasitoids of stem borers
Xanthopimpla flavolineata, X. modesta modesta, X. punctata and X. stemmator belong to this category. All these species have a long ovipositor for the genus to attack the lepidopterous borers in plant stems. They are, thus, important or potentially important natural enemies of crop pests. In Japan, X. flavolineata, X. punctata and X. stemmator are collected in somewhat open areas (e.g. meadows, open forest and crop fields).
2) Parasitoids of large moths with exposed habitats
Species of the regina group belong to this category, which parasitises the larvae and pupae of larger moths, such as Saturniidae and Lasiocampidae, in cocoons in somewhat exposed habitats. These species are rather large and have a robust body and long ovipositor. In Japan, X. konowi and X. pedator are collected in forest areas (e.g. Mt. Omoto-dake and Shiramizu of Ishigakijima Island) where their hosts are abundant.
3) Parasitoids of bagworm moths
Xanthopimpla naenia seems to be a specialist parasitoid of bagworms (Psychidae). Nipponopsyche fuscescens and Manatha sp. were recorded as the hosts of this species in Japan. Bagworms carry unique portable cases that function as defensive shelters. Possibly because of the difficulty in being utilized as a host, they are attacked by specialised parasitoids (e.g. Sericopimpla Kriechbaumer, 1895 and Paraphylax Förster, 1869). However, how a few species of Xanthopimpla became parasitic on psychids is unclear.
The hosts of other species of Japanese Xanthopimpla are almost unknown. Given that most species are found in forest habitats, they possibly use hosts that are abundant in the forest, such as microlepidopterans, in leaf rolls and cocoons. Xanthopimpla clavata was observed to lay eggs in cocoons of Galleria mellonella (Linnaeus, 1758), which were experimentally exposed to the wasp in a cage. The offspring emerged successfully. Another male was reared from an unidentified small lepidopterous pupa collected in the field. These observations suggest that this species may utilie a wide range of lepidopterans as hosts.
The genus Xanthopimpla generally thrives in the tropics and subtropics. Data from collected specimens in the Ryukyus indicate that the adults are active almost all year round and that many species are possibly multivoltine originally. However, their activity is restricted by the low temperature of winter. To the best of our knowledge, in Honshu, Shikoku and northern Kyushu, which have a winter season, most of the species, including X. clavata, X. nipponensis, X. yoshimurai and X. trias, pass the winter in the adult stage. The wintering adults rest under the leaves of broad-leaved evergreen trees (Fig.
All wintering individuals were female. This is similar to other ichneumonids overwintering as adults under evergreen leaves, such as Zatypota maculata (Matsumoto & Takasuka, 2010) (
Most Japanese Xanthopimpla species are also distributed in Southeast Asian countries, whereas a few species are unknown beyond Japan and provisionally considered as Japanese endemics. The biogeographical border of the Palaearctic and Oriental Regions is called the Tokara Gap (Watase line), which lies between Yakushima Island and the Tokara Islands (the north-eastern islands of Amamioshima Island). Thus, the faunas of Yakushima Island and Amamioshima Island are usually very different from each other. However, the range of distribution of any Japanese Xanthopimpla is not limited by this line, except for that of X. honorata honorata. A total of seven species, X. clavata, X. minuta, X. naenia, X. pedator, X. punctata, X. sylvicola and X. trias, are distributed both in the Palearctic and Oriental parts of Japan beyond the Gap (Table
Xanthopimpla nipponensis and X. yoshimurai are endemic to Japan at present, but detailed distribution data for many more species are needed for a comprehensive faunal study of this group in Asia.
We cordially thank David Wahl (