Zootaxa 3779 (1): 020–032 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press
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ISSN 1175-5326 (print edition)
ZOOTAXA
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http://dx.doi.org/10.11646/zootaxa.3779.1.5 http://zoobank.org/urn:lsid:zoobank.org:pub:79B81076-9D53-4D21-912E-721703237DB2
Review of the Diplazontinae (Hymenoptera, Ichneumonidae) of the Kuril islands, with descriptions of two new species SERAINA KLOPFSTEIN Swedish Museum of Natural History, Box 50007, SE–104 05 Stockholm, Sweden and School of Earth and Environmental Sciences, The University of Adelaide, North Terrace, Darling Building, Adelaide, SA 5005. E-mail: [email protected]
Abstract A sample of 346 specimens of Diplazontinae parasitoid wasps from the Kuril islands was studied. Twenty-six species are reported, Tymmophorus gelidus Dasch for the first time for the Eastern Palaearctic. Two new species are described, Diplazon kurilensis sp. n. and Homotropus formosus sp. n. Diplazon urupensis Uchida is removed from synonymy (stat. rev.), and Promethes persulcatus Nakanishi is suggested as a synonym of Promethes bridgmani Fitton. Reasons are discussed for the large proportion of species with a Holarctic or even multi-regional distribution in the sample, which amounts to17 of the species or 65%. Key words: parasitoid wasps, faunistics, Russian Far East, distributional patterns
Introduction The Diplazontinae are a moderately-sized subfamily of parasitoids of the family Ichneumonidae; 347 species are currently recognized worldwide (Yu et al., 2012). Where reliable host records are available, they point to a rather specialized host range of mostly aphidophagous hoverflies (e.g., Schneider, 1951; Fitton & Rotheray, 1982; Rotheray, 1984; Ngamo Tinkeu & Hance, 1997). Most diplazontine species have been described from temperate regions (Yu et al., 2012). Although the tropics are severely understudied, two locally extensive studies suggest that the subfamily does not exhibit an increase in species richness with decreasing latitude as it can be observed in other insect groups (Dasch, 1964b; Gauld et al., 1997), which is in line with the relative scarcity of aphids and aphidophagous syrphids in the tropics. In the Holarctic region, both the Nearctic (Dasch, 1964a) and the Western Palaearctic faunas (Klopfstein, in press) have been revised, while the Eastern Palaearctic is only partially covered by taxonomic and faunistic studies (Uchida, 1957; Nakanishi, 1967, 1978, 1979, 1985, 1986; Kasparyan & Manukyan, 1987; Manukyan, 1987, 1988; Kasparyan & Manukyan, 1989; Manukyan, 2007; Klopfstein, 2011). Summarizing distributional patterns of Palaearctic diplazontines, Manukyan (1995) pointed out that a high proportion of diplazontine species occurs in several faunistic regions; of the 130 Palaearctic species known at the time; 38% were either cosmopolitan, multiregional or at least Holarctic. His assessment was however hindered by a lack of taxonomic revisions which compared the Nearctic to the Palaearctic faunas and a gap of knowledge in part of the Eastern Palaearctic region. Even though only a part of the Nearctic species were studied in the revision of the Western Palaearctic fauna, Klopfstein (in press) proposed eight new synonyms between Nearctic and Western Palaearctic taxa, and more can be expected in the future (Dasch, 1964a). Manukyan's estimate is thus probably an underestimate. I recently received 346 specimens from the Kuril islands, most of which had been collected by the International Kuril Island Project (www.burkemuseum.org/static/okhotskia/ikip/, Sauter, 1996). This material includes 26 species, two of which are here newly described. Given the geographic location of the Kuril islands, between Russian Kamchatka and the Japanese island Hokkaido (Fig. 1), this material is especially interesting with respect to an assessment of the distributional ranges in Diplazontinae.
20 Accepted by J. T. Jennings: 21 Jan. 2014; published: 13 Mar. 2014
Material and methods I obtained 346 specimens from the Kuril islands from the Texas A&M University Insect Collection (College Station, Texas). Additional material was studied for comparison at the Swedish Museum of Natural History (Stockholm). Junior synonyms are only given if they are proposed in the present paper, as accurate lists of synonyms are given in the catalogue by Yu et al. (2012). For terminology of morphology, I follow Townes (1969). All measurements were taken with a Leica Wild M10 stereo-microscope with a 10x ocular including an eye-piece micrometer. Specimens were photographed with a Canon EOS 7D with a Canon Macro lens EF 100mm 1:2.8 USM, and stacked using Helicon Focus pro x64. All specimens, including the type material, are stored at the Texas A&M University Insect Collection (TAMU).
FIGURE 1. Map of the Kuril islands and adjacent Hokkaido and Kamchatka. Modified from Wikimedia Commons.
Descriptions of new species Diplazon kurilensis sp. n. (Fig. 2) This species belongs to the pectoratorius group of species in the genus Diplazon, which unites the species with rather weak notauli, reduced carinae on the propodeum, epicnemical carina reduced ventrally, weak transverse impressions on the tergites, and a white hind tibia with only the apex but not the base darkened. According to recent molecular and morphological phylogenetic studies (Klopfstein et al., 2010; Klopfstein et al., 2011), this species group is sister to all the other species in the genus. The other three species currently known in the group are D. urupensis Uchida from the Kuril islands (see below), D. pectoratorius (Thunberg) with a Holarctic, Oriental and Neotropic distribution, and the Nearctic and Eastern Palaearctic D. albotibialis Dasch. From these species, D. kurilensis can be distinguished by its small size, reduced sculpture on the mesoscutum and metasoma, and brown coloration of the metasoma. DIPLAZONTINAE OF THE KURIL ISLANDS
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Etymology. This species is named “kurilensis” after the type locality. Description. Fore wing length 3.2–3.6 mm (3.6 in holotype) in females, 4.1 mm in the male. Antenna with 15–16 (16) flagellomeres in females, all flagellomeres with multiporous plate sensillae sparse but also present ventrally, apical flagellomere long and enlarged, thus appearing clubbed; 18 flagellomeres in the male, without tyloids. Face strongly coriaceous and matt, impunctate, without vertical impressions. Clypeus with apical margin thin, slightly impressed on the sides after a basal elevation, resulting in the lateral areas being slightly concave. Pronotum and propleuron smooth with some sparse punctures. Mesoscutum with weak to distinct notauli; smooth and shining with very sparse punctures, mostly in the front and on the sides. Mesopleuron with epicnemical carina reduced ventrally; with weak and sparse punctures and some very irregular, weak sculpture. Scutellum only carinate at base, smooth and shining. Metapleuron mostly smooth, with some punctures in the front. Propodeum only with metapleural carina and hind part of the lateral longitudinal carinae present, rugulously sculptured near the base, more rugose and partly smooth on petiolar area; propodeal spiracle not enlarged. Fore wing without areolet; hind wing with 2 basal hamuli. Legs including coxae smooth and shining, hind tibia without thickened setae. Female metasoma dorsoventrally depressed, tapered towards the apex, apical margins of the tergites straight to even slightly concave, with transverse impressions weak to almost indistinct on tergites 1–3. First tergite 1.3–1.45 (1.45) times as long as wide, with median dorsal carinae only present on about basally, finely coriaceous and matt; second tergite 0.7 times as long as wide, finely coriaceous and matt; spiracle of second and third tergites on dorsal part, above lateral fold. Ovipositor sheaths about 0.3 times as long as hind tibia, pointed and closed at the tip, coriaceous and matt and with numerous setae on whole surface.
FIGURE 2. Habitus of the female holotype (A) and male paratype (B) of Diplazon kurilensis sp. n.
Coloration of females. Antenna brown. Head and mesosoma dark brown, with yellow inner eye margins, yellow on clypeus, mouthparts, hind corner of pronotum, tegula, subtegular ridge, shoulder mark, and mesepimeron; scutellum yellow on anterior corners, orange centrally. Legs yellow, all coxae yellow, hind coxa with some orange basally, hind femur orange, hind tibia white with apex light brown, hind tarsus brown. Metasoma brown, progressively lighter towards apex which is orange, end margins of tergites whitish. Coloration of male. As in females but additionally with yellow over entire face, scape, pedicel and flagellomeres ventrally, propleuron, mesosternum, mesopleuron in front of epicnemical carina and with a line on lower mesopleuron; all coxae entirely yellow. Types material. Holotype ♀: Russia, Kuril islands: Kharimkotan, Northwest corner of island. N49°08.60'/ E154°28.10', 28. VII. 2000. leg. D.J. Bennett. Paratypes, one ♀ and one #: Russia, Kuril islands: Matua Island, inland from Dvoinaya Bay. N48°03.95'/E153°15.63', 31. VII. 2000. leg. D.J. Bennett.
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Homotropus formosus sp. n. (Fig. 3) This species belongs to the pictus species group (according to Dasch, 1964a). Its black coxae and large size in combination with a strongly coriaceous and matt mesosoma distinguish it from most species there-in. In terms of size, it could be confused with Homotropus megaspis Thomson and H. momoi (Uchida); from the former, it differs by the dark coxae, and from both by the coriaceous sculpture of the mesosoma. The otherwise similar Homotropus areolaris (Uchida) differs by the coloration of the stigma in the fore wing, the coloration of the fore and mid coxae, and the more extensive coriaceous sculpture.
FIGURE 3. Habitus of the female holotype (A) and male paratype (B) of Homotropus formosus sp. n.
Etymology. The name “formosus” is intended to reflect the overall appearance of the species. Description. Fore wing length 5.1–5.5 mm (5.1 in holotype) in females, 5.8 mm in the male. Antenna with 21– 22 (21) flagellomeres in females, apical flagellomeres with multiporous plate sensilla sparse but also present ventrally; 23 flagellomeres in males, with narrow, long tyloids on flagellomeres 7 to 15. Face strongly coriaceous and matt, distinctly punctate, especially centrally, without vertical impressions. Clypeus with apical margin thin, strongly impressed, resulting in the central area being convex. Pronotum and propleuron with strong and dense punctures, smooth or finely coriaceous in-between. Mesoscutum without notauli; finely coriaceous and matt in the females, less coriaceous and mostly shining in the male, in both sexes with dense punctures on whole surface. Mesopleuron with epicnemical carina strong and complete ventrally; strongly coriaceous and matt with dense punctures except for small smooth area around the speculum in females, in the male matt only on lower half, more shining on upper half between the punctures. Scutellum only carinate at base, rather strongly convex, smooth between the strong punctures. Metapleuron with dense punctures, but also some smooth areas. Propodeum only with metapleural carina and hind part of the lateral longitudinal carinae present, densely and rugulously punctate near the base, more rugose and partly smooth on petiolar area; propodeal spiracle not enlarged. Fore wing areolet sometimes large, almost rhombic, vein 3rs-m mostly well pigmented; hind wing with 2–3 basal hamuli. Legs including coxae coriaceous and matt, hind tibia with thickened, scale-like hairs on outer surface. Female metasoma dorsoventrally depressed, tapered towards the apex, apical margins of the tergites straight or convex, without transverse impressions. First tergite 1.1–1.3 (1.1) times as long as wide, with median dorsal carinae only present on about basal fourth, finely coriaceous and matt; second tergite 0.6–0.7 times as long as wide, basally with some strong longitudinal wrinkles, finely coriaceous and matt and with a few, very sparse and weak punctures; second tergite 0.8–0.9 times length of first tergite; spiracle of second and third tergites on dorsal part, above lateral fold. Ovipositor sheaths about 0.3 times as long as hind tibia, more or less parallel-sided and truncate close to the tip, smooth and shining and with some setae close to the tip. Coloration of females. Antenna black. Head and mesosoma black, with small yellow or orange central face DIPLAZONTINAE OF THE KURIL ISLANDS
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patch, yellow on clypeus, mouthparts, hind corner of pronotum, tegula, subtegular ridge, large shoulder mark, and upper mesepimeron; scutellum entirely black. Fore wing with stigma lighter at base. Legs orange, all coxae black, fore coxa apically with some yellow, trochanters basally black, apically yellow, trochantelli yellow, femora orange, hind tibia white with apex, base and a small subbasal spot dark, hind tarsus dark. Metasoma entirely black. Coloration of males. As in females but additionally with yellow over entire face, scape, pedicel and flagellomeres ventrally, propleuron, mesosternum, mesopleuron in front of epicnemical carina and with a spot on apical lower corner; fore and mid coxae entirely and hind coxa apically yellow, hind femur with dark marks at base and yellow at tip. Type material. Holotype ♀: Russia, Kuril islands: Urup Island, Inland from Aleutka Bay. N45°56.18'/ E150°09.39', 07. VIII. 2000. leg. D.J. Bennett. Paratypes: one ♀, same data as holotype; one ♂: Japan, Hokkaido, Mt. Yuubari, Yuubari city. 01. VII. 1995. leg. Ito Gen.
Annotated species list Diplazon pectoratorius (Thunberg) Russia, Kuril islands: Iturup Island, Inland Konservnaya Bay; N45°19.79'/E147°59.75', 19. VIII. 1996, leg. P. Oberg. 2♂. Paramushir Island, inland from Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. B.K. Urbain. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°36.62'/E154°49.21', 27.VII.2000, leg. D.J. Bennett. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°36.08'/E154°48.96', 27.VII.2000, leg. D.J. Bennett. 1♀. Shiashkotan, Inland SW Zakatnaya Bay; N48°46.83'/E154°01.85', 29. VII. 2000, leg. T.R. Anderson. 1♀. Urup Island, inland of Ukromnaya Bay; N45°35.43'/E149°31.91', 20. VIII. 1996, leg. B.K. Urbain. 6♂. Holarctic, Oriental and Neotropic; known from Russia and Japan. Diplazon urupensis Uchida (stat. rev.) Russia, Kuril islands: Paramushir Island, Inland from Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. B.K. Urbain. 1♀. Shiashkotan, inland SW Zakatnaya Bay; N48°46.83'/E154°01.85', 29. VII. 2000, leg. T.R. Anderson. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 5♀. This species was described from the Kuril islands by Uchida (1935). In a later publication (Uchida, 1957), he then synonymized it with D. pectoratorius. However, after studying the present specimens and consulting Uchida's holotype and allotype of the species (deposited at the Hokkaido University, Sapporo; label data of the holotype: "Uruppu; K.Doi" / "Holotype; Diplazon; urupensis; (Uch.)"), I decided to reinstate D. urupensis as a valid species (stat. rev.). The specimens are clearly smaller and more slender than D. pectoratorius, lack the orange coloration of the mesopleuron and mesosternum (except for a few males), and have weaker and sparser punctures on the mesoscutum and mesopleuron. It remains to be shown whether the Nearctic D. albotibialis Dasch (1964a), which has also been reported from Eastern and Western Russia (Manukyan, 1987, 2007), is a distinct species or a junior synonym of D. urupensis. Known only from the Kuril islands. Diplazon varicoxa (Thomson) Russia, Kuril islands: Urup Island, inland from Aleutka Bay; N45°56.18'/E150°09.39', 07. VIII. 2000, leg. D.J. Bennett. 1♀. Urup Island, inland from Aleutka Bay; N45°56.10'/E150°09.70', 07. VIII. 2000, leg. T.R. Anderson. 1♀. Urup Island, Inland of Chernoburka Bay; N45°36.26'/E149°34.55', 09. VIII. 2000, leg. D.J. Bennett. 1♂. Iturup Island, inland of Medvezh'ya Bay; N45°27.27'/E148°49.76', 05. VIII. 1998, leg. B.K. Urbain. 1♂. Iturup Island, Inland of Medvezh'ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. B.K. Urbain. 1♀; same data, but leg. B.K. Urbain, 1♂. Iturup Island, inland of Medvezh'ya Bay; N45°25.75'/E148°49.75', 05. VIII. 1998, leg. B.K. Urbain. 1♀. Iturup Island, Inland of Konservnaja Bay; N45°20.04'/E147°59.84', 12. VIII. 1999, leg. D.J. Bennett. 1♂; same data, but 30. VII. 1997, leg. B.K. Urbain:. 1♀. Iturup Island, Nr. Konservnaya Bay; N45°20.02'/ E148°00.03', 30. VII. 1997, leg. B.K. Urbain. 2♀. Iturup Island, inland of Konservnaja Bay; N45°20.01'/ E147°59.91', 31. VIII. 1995, leg. N. Minakawa. 1♂; same data, but leg. N. Minakawa: 2♂. Iturup Island, 4km east Kitovyi Village; N45°15.90'/E147°55.90', 29. VII. 1997, leg. J. Schweikert. 3♂. Iturup Island, North of Cape
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Kanonerka; N45°11.31'/E148°15.43', 03. VIII. 1998, leg. K. Kavun. 1♂. Iturup Island, inland of Kuybyshevskiy Bay; N45°04.49'/E147°39.07', 13. VIII. 1999, leg. B.K. Urbain. 5♀. Iturup Island, Near Sernozavoskaya Bay; N44°58.29'/E147°53.69', 02. VIII. 1998, leg. B.K. Urbain. 1♂. Iturup Island, Nr. Sernozavoskaya Bay; N44°57.88'/E147°53.89', 02. VIII. 1998, leg. D.J. Bennett. 1♂. Iturup Island, Nr. Sernozavoskaya Bay; N44°57.84'/E147°53.83', 02. VIII. 1998, leg. B.K. Urbain. 1♂. Iturup Island, Atsonupuri Peninsula base; N44°47.60'/E147°11.42', 14. VIII. 1999, leg. B.K. Urbain. 1♂. Iturup Island, Atsonupuri Peninsula base; N44°47.34'/E147°11.34', 14. VIII. 1999, leg. B.K. Urbain. 3♂. Iturup Island, Atsonupuri Peninsula base; N44°46.41'/E147°11.20', 14. VIII. 1999, leg. B.K. Urbain. 23♂. Iturup Island, Atsonupuri Peninsula base; N44°46.31'/E147°11.32', 15. VIII. 1999, leg. B.K. Urbain. 7♂. Iturup Island, Atsonupuri Peninsula base; N44°46.01'/E147°11.58', 14. VIII. 1999, leg. D.J. Bennett. 2♂. Iturup Island, Env. Tikhaya River mouth; N44°43.38'/E147°12.77', 15. VIII. 1999, leg. D.J. Bennett. 2♀. Kunashir Island, 1km E of Tyatina R. Mouth; N44°16.32'/E146°10.34', 28. VIII. 1997, leg. Y. Marusik. 2♂. Kunashir Island, Goby Hot Springs environment; N44°00.39'/E145°41.07', 17. VIII. 1999, leg. D.J. Bennett. 1♂. Palaearctic and Oriental; known from Russia and Japan. Fossatyloides gracilentus (Holmgren) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 1♂. Holarctic; also known from the Russian Far East. Homotropus elegans (Gravenhorst) Russia, Kuril islands: Raikoke Island, inland from Eastern side; N48°17.87'/E153°15.67', 30. VII. 2000, leg. D.J. Bennett. 1♀. Raikoke Island, inland from Eastern side; N48°17.81'/E153°15.68', 30. VII. 2000, leg. T.R. Anderson. 13♂. Raikoke Island, inland from Eastern side; N48°17.73'/E153°15.59', 30. VII. 2000, leg. T.R. Anderson. 1♂. Raikoke Island, inland from Eastern side; N48°17.71'/E153°15.64', 30. VII. 2000, leg. D.J. Bennett. 1♂. Holarctic; reported from Russia, including its easternmost provinces. Homotropus nigritarsus (Gravenhorst) Russia, Kuril islands: Paramushir Island, Nr. Putyatino settlement; N50°44.19'/E156°08.82', 04. VIII. 1997, leg. B.K. Urbain. 1♀, 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.54'/E156°05.71', 30. VII. 1999, leg. B.K. Urbain. 1♂. Alaid Island, inland of Alaidskaya Bay; N50°29'/E155°39', 25. VII. 1999, leg. A.S. Lelej. 1♂. Antsiferova Island, inland of near Vydar Rock; N50°12.36'/E154°57.73', 15. VIII. 1997, leg. B.K. Urbain. 1♀. Paramushir Island, Nr. Vasil'yeva Peninsula; N50°02.87'/E155°23.46', 03. VIII. 1996, leg. B.K. Urbain. 3♀. Kharimkotan, Northwest corner of island; N49°08.75'/E154°27.64', 28. VII. 2000, leg. D.J. Bennett. 1♂. Holarctic and Neotropical; reported from Russia, including its easternmost provinces. Homotropus pallipes (Gravenhorst) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 1♂. Iturup Island, Near Sernozavoskaya Bay; N44°58.29'/E147°53.69', 02. VIII. 1998, leg. B.K. Urbain. 1♀. Shikotan Island, SW shore of Delfin Bay; N43°50'/E146°44', 11. IX. 1997, leg. Y. Marusik. 1♂. Holarctic and Neotropical; reported from Russia, including its easternmost provinces. Homotropus pictus (Gravenhorst) Russia, Kuril islands: Paramushir Island, Nr. Putyatino settlement; N50°44.19'/E156°08.82', 04. VIII. 1997, leg. B.K. Urbain. 1♂. Paramushir Island, Env. Utyesnaya River; N50°37.73'/E156°08.21', 01. VIII. 1996, leg. M. Ohara. 1♂. Makanrushi Island, inland from Zakat Bay; N49°44.37'/E154°24.84', 18. VIII. 1997, leg. J. Schweikert. 1♀. Kharimkotan, Northwest corner of island; N49°08.95'/E154°28.54', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corner of island; N49°08.93'/E154°28.09', 28. VII. 2000, leg. T.R. Anderson. 2♀. Kharimkotan, Northwest corner of island; N49°08.67'/E154°28.29', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corner of island; N49°08.60'/E154°28.16', 28. VII. 2000, leg. D.J. Bennett. 1♀, 1♂. Matua Island, inland from Dvoinaya Bay; N48°03.95'/E153°15.63', 31. VII. 2000, leg. D.J. Bennett. 2♂. Matua
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Island, Inland of Dvoinaya Bay; N48°03.12'/E153°14.80', 03. VIII. 1999, leg. B.K. Urbain. 2♂. Rasshua Island, inland near Malenkaya Bay; N47°43.26'/E152°58.38', 04. VIII. 1999, leg. B.K. Urbain. 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.86'/E152°16.20', 02. VIII. 2000, leg. D.J. Bennett. 3♀. Simushir Island, inland of NE Broutona Bay; N47°07.82'/E152°16.40', 02. VIII. 2000, leg. T.R. Anderson. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.77'/E152°16.28', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.27'/E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 10♀. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 3♀. Simushir Island, inland of NE Broutona Bay; N47°07.01'/E152°16.17', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of NE Broutona Bay; N47°06.93'/E152°15.92', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of Malaya Bay; N47°05.38'/E152°08.10', 18. VIII. 1995, leg. B.K. Urbain. 1♀. Simushir Island, inland of Malaya Bay; N47°05.18'/E152°07.81', 18. VIII. 1995, leg. M. Ohara. 2♀. Palaearctic; reported from Russia, including its easternmost provinces. Homotropus signatus (Gravenhorst) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 2♂. Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 3♂. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♂. Matua Island, inland of Dvoinaya Bay; N48°04'/E153°15', 03. VIII. 1999, leg. A.S. Lelej. 1♀, 1♂. Rasshua Island, W. Coast of Beloye Lake; N47°43.30'/E152°59.52', 12. VIII. 1995, leg. Y. Marusik. 1♂. Rasshua Island, SSW part; near "Arches"; N47°43.19'/E152°58.80', 12. VIII. 1995, leg. M. Ohara. 1♀. Ketoi Island, East of Cape Storozheva; N47°22.64'/E152°27.42', 15. VIII. 1995, leg. B.K. Urbain. 1♀. Ketoi Island, East of Cape Storozheva; N47°22.60'/E152°27.76', 15. VIII. 1995, leg. M. Ohara. 1♂. Ketoi Island, East of Cape Storozheva; N47°22.54'/ E152°27.47', 15. VIII. 1995, leg. N. Minakawa. 4♀, 2♂. Ketoi Island, inland of Diany Bay; N47°18.72'/ E152°26.07', 06. VIII. 1999, leg. D.J. Bennett. 2♂. Simushir Island, inland of NE Broutona Bay; N47°07.86'/ E152°16.20', 02. VIII. 2000, leg. T.R. Anderson. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.77'/ E152°16.28', 02. VIII. 2000, leg. D.J. Bennett. 2♀, 7♂. Simushir Island, inland of NE Broutona Bay; N47°07.27'/ E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 1♀, 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.20'/ E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 1♀, 2♂. Simushir Island, Inland of Dushnaya Bay; N47°05'/E152°12', 09. VIII. 1999, leg. A.S. Lelej. 1♂. Simushir Island, inland of Dushnaya Bay; N47°04.76'/E152°12.19', 09. VIII. 1999, leg. B.K. Urbain. 1♀, 1♂. Chirpoi Island, inland SW Peschanaya Bay; N46°32.05'/E150°53.51', 03. VIII. 2000, leg. D.J. Bennett. 1♀, 1♂. Urup Island, inland from Aleutka Bay; N45°56.08'/E150°09.98', 07. VIII. 2000, leg. T.R. Anderson. 2♀. Iturup Island, inland of Slavnaja Bay; N45°29.47'/E148°37.13', 06. VIII. 1998, leg. D.J. Bennett. 1♀, 1♂. Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 1♀. Kunashir Island, Near Lake Aliger; N44°02.88'/E145°44.48', 11. VIII. 1998, leg. D.J. Bennett. 1♂. Kunashir Island, Environs of Lake Aliger; N44°02.83'/E145°44.47', 09. IX. 1995, leg. P. Oberg. 4♀. Tanfilyeva, Inland of Tanfil'Yevka Bay; N43°26.62'/ E145°55.89', 19. VIII. 1998, leg. B.K. Urbain. 1♂. Holarctic; reported from Russia and Japan. Phthorima compressa (Desvignes) Russia, Kuril islands: Iturup Island, inland of Slavnaja Bay; N45°29.47'/E148°37.13', 05. VIII. 1998, leg. D.J. Bennett. 1♂. Shikotan Island, inland of Otradnaya Bay; N43°52.14'/E146°47.67', 12. VIII. 1998, leg. B.K. Urbain. 1♀. Tanfilyeva, inland Tanfil'Yevka Bay; N43°26.62'/E145°55.89', 19. VIII. 1998, leg. D.J. Bennett. 1♂. Anuchina Island, inland of Bolshoye Bay; N43°22.26'/E146°00.36', 19. VIII. 1998, leg. B.K. Urbain. 1♂. Holarctic; reported from Russia. Promethes bridgmani Fitton Promethes persulcatus Nakanishi, 1986 is a junior synonym, syn. nov. Russia, Kuril islands: Paramushir Island, inland of Rifovaya Bay; N50°29.84'/E156°05.80', 30. VII. 1999, leg. B.K. Urbain. 1♂. The type of Promethes persulcator Nakanishi could not be obtained, but based on the detailed description (Nakanishi, 1986) and on this male from the Kuril islands, these two species are clearly synonymous. While P.
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bridgmani has only been reported from Europe, P. persulcator has been recorded in Japan and the Russian Far East. Palaearctic, including Japan and the Russian Far East. Promethes sulcator (Gravenhorst) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. J. Schweikert. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♀, 4♂. Paramushir Island, inland of Rifovaya Bay; N50°29.43'/E156°05.52', 30. VII. 1999, leg. D.J. Bennett. 1♀. Paramushir Island, Base Vasil'Yeva Peninsula; N50°03.31'/E155°23.60', 16. VIII. 1997, leg. B.K. Urbain. 1♂. Rasshua Island, inland near Malenkaya Bay; N47°42.77'/E152°58.16', 04. VIII. 1999, leg. B.K. Urbain. 2♀. Simushir Island, inland of NE Broutona Bay; N47°07.27'/E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 2♂. Simushir Island, inland of NE Broutona Bay; N47°07.01'/E152°16.17', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Urup Island, inland of Ukromnaya Bay; N45°35.43'/E149°31.91', 20. VIII. 1996, leg. B.K. Urbain. 2♀. Iturup Island, inland of Konservnaja Bay; N45°20.04'/E147°59.84', 12. VIII. 1999, leg. D.J. Bennett. 1♂. Iturup Island, 4km east Kitovyi Village; N45°16.07'/E147°56.38', 29. VII. 1997, leg. B.K. Urbain. 1♂. Iturup Island, North of Cape Kanonerka; N45°11.12'/E148°14.57', 03. VIII. 1998, leg. D.J. Bennett. 1♂. Iturup Island, Nr. Sernozavoskaya Bay; N44°57.84'/E147°53.83', 02. VIII. 1998, leg. B.K. Urbain. 1♀. Kunashir Island, Env. Of Cape Sukacheva; N44°04.50'/E145°51.95', 20. VIII. 1997, leg. Y. Marusik. 1♂. Shikotan Island, inland from Gorobets Bay; N43°92.64'/E146°42.48', 18. VIII. 1998, leg. B.K. Urbain. 1♀. Shikotan Island, SW shore of Delfin Bay; N43°50'/E146°44', 11. IX. 1997, leg. Y. Marusik. 1♀. Holarctic and Oriental; recorded from Russia and Japan. Sussaba aciculata (Ruthe) Russia, Kuril islands: Paramushir Island, Env. Utyesnaya River; N50°37.73'/E156°08.21', 01. VIII. 1996, leg. M. Ohara. 1♀. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♀. Paramushir Island, inland of Rifovaya Bay; N50°29'/E156°06', 30. VII. 1999, leg. A.S. Lelej. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°22.26'/E155°36.17', 13. VIII. 1997, leg. B.K. Urbain. 1♂. This species differs from the Japanese species Sussaba nigra Nakanishi by the strongly coriaceous median lobe of the mesonotum and from Sussaba dajsetsuzana Nakanishi by the lack of a black area in the tyloids of males (Nakanishi, 1979). Holarctic; reported from the Russian Far East. Sussaba cognata (Holmgren) Russia, Kuril islands: Alaid Island, inland of Alaidskaya Bay; N50°49.56'/E155°39.88', 25. VII. 1999, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.84'/E156°05.80', 30. VII. 1999, leg. B.K. Urbain. 1♀. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°36.62'/E154°49.21', 27.VII.2000, leg. D.J. Bennett. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°35.96'/E154°49.11', 27.VII.2000, leg. D.J. Bennett. 1♂. Kharimkotan, Northwest corder of island; N49°08.95'/E154°28.54', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corder of island; N49°08.67'/E154°28.29', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corder of island; N49°08.60'/E154°28.10', 28. VII. 2000, leg. D.J. Bennett. 1♀. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.87'/E154°02.08', 29. VII. 2000, leg. T.R. Anderson. 1♀. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.83'/E154°01.85', 29. VII. 2000, leg. D.J. Bennett. 1♀. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.77'/E154°02.01', 29. VII. 2000, leg. D.J. Bennett. 2♀. Matua Island, inland from Dvoinaya Bay; N48°04.17'/E153°15.28', 31. VII. 2000, leg. D.J. Bennett. 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.86'/E152°16.20', 02. VIII. 2000, leg. T.R. Anderson. 3♀. Simushir Island, inland of NE Broutona Bay; N47°07.85'/E152°16.34', 02. VIII. 2000, leg. T.R. Anderson. 4♀. Simushir Island, inland of NE Broutona Bay; N47°07.82'/E152°16.40', 02. VIII. 2000, leg. T.R. Anderson. 3♀, 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.77'/E152°16.28', 02. VIII. 2000, leg. D.J. Bennett. 2♀. Simushir Island, inland of NE Broutona Bay; N47°07.27'/E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 3♀, 3♂. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Simushir Island, inland of NE
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Broutona Bay; N47°07.01'/E152°16.17', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Brat Chirpoev, inland from Uglovaya Bay; N46°27.24'/E150°48.50', 03. VIII. 2000, leg. D.J. Bennett. 4♀. Shikotan Island, Krabozavodskoe Village; N43°50.10'/E146°45.23', 11.–18. IX. 1997, leg. Y. Marusik. 1♀. Shiashkotan, SW shore of Delfin Bay; N43°44.90'/E146°36.41', 13. IX. 1997, leg. Y. Marusik. 1♂. This common and very widespread species exhibits a large range of variation both in color and ultrasculpture which has led to the proposition of several subspecies (Dasch, 1964a). Most of the specimens from the Kuril islands have yellow coxae, rather extensive yellow coloration on the face and epicnemium, and a smooth mesoscutum; they would thus be associated with the subspecies S. cognata albicoxa (Thomson) or S. cognata faceata Dasch. However, some specimens have the base of the hind coxae dark. Nakanishi (1979) discussed the color variation of this species in Japan and concluded that no clear separations can currently be drawn between the subspecies. Furthermore, proposing subspecies that occur in sympatry, as is the case in most of the range of S. cognata, is in conflict with a biological species concept (Mayr, 1942); the color variation in S. cognata is thus likely to represent phenotypic variation only and is not indicative of separate evolutionary lineages. Holarctic and Oriental; recorded from Russia and Japan. Sussaba dorsalis (Holmgren) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. T.R. Anderson. 1♀. Urup Island, inland of Smuglyi Bay; N46°01.46'/E149°59.02', 24. VIII. 1995, leg. N. Minakawa. 1♂. Urup Island, inland from Aleutka Bay; N45°56.10'/E150°09.70', 07. VIII. 2000, leg. T.R. Anderson. 1♀, 2♂. Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 1♂. Holarctic; recorded from Russia, including the Eastern provinces. Sussaba mongolica Klopfstein Figure 4A: Habitus. Russia, Kuril islands: Simushir Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.84'/E156°05.80', 30. VII. 1999, leg. B.K. Urbain. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.67'/E156°06.84', 30. VII. 1999, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.54'/E156°05.71', 30. VII. 1999, leg. B.K. Urbain. 1♀, 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.43'/E156°05.52', 30. VII. 1999, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29'/E156°06', 30. VII. 1999, leg. A.S. Lelej. 2♀, 1♂. Matua Island, inland of Dvoinaya Bay; N48°04.37'/E153°15.98', 31. VII. 2000, leg. T.R. Anderson. 1♂. Matua Island, inland of Dvoinaya Bay; N48°04.15'/E153°15.70', 03. VIII. 1999, leg. D.J. Bennett. 1♂. Simushir Island, inland of NE Broutona Bay; N47°06.93'/E152°15.92', 02. VIII. 2000, leg. D.J. Bennett. 1♂. This species can be distinguished from the Japanese Sussaba nigra Nakanishi by the bi-colored tyloids in the male and from Sussaba dajsetsuzana Nakanishi by the smooth and punctured mesoscutum and the shorter and more stout petiole. Described from Mongolia (Klopfstein, 2011). Sussaba pulchella (Holmgren) Russia, Kuril islands: Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Kunashir Island, 1km E of Tyatina R. Mouth; N44°16.32'/E146°10.34', 28. VIII. 1997, leg. Y. Marusik. 1♂. Kunashir Island, environs of Lake Aliger; N44°02.83'/E145°44.47', 09. IX. 1995, leg. P. Oberg. 1♀. Kunashir Island, inland of Aalekhina Bay; N43°55.23'/E145°32.04', 19. VIII. 1999, leg. D.J. Bennett. 1♂. Shikotan Island, inland from Del'Fin Bay; N43°45.11'/E146°37.41', 15. VIII. 1998, leg. K. Kavun. 1♂. Kunashir Island, between 1st and 2nd km of Golovino highway; N/E, 01. IX. 1997, leg. Y. Marusik. 1♂. Holarctic and Oriental; recorded from Russia and Japan. Sussaba sugiharai (Uchida) (Fig. 4B) Russia, Kuril islands: Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 3♂. Shikotan Island, inland of Zvezdnaya Bay; N43°46.34'/E146°36.36', 16. VIII. 1998, leg. D.J. Bennett. 1♂. Tanfilyeva, inland Tanfil'Yevka Bay; N43°26.79'/E145°55.32', 19. VIII. 1998, leg. B.K. Urbain. 1♂.
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Tanfilyeva, inland Tanfil'Yevka Bay; N43°26.62'/E145°55.89', 19. VIII. 1998, leg. D.J. Bennett. 1♀, 3♂. Kunashir Island, between 1st and 2nd km of Golovino highway; N/E, 01. IX. 1997, leg. Y. Marusik. 2♀. Eastern Palaearctic and Oriental; recorded from the Russian Far East and Japan.
FIGURE 4. Habitus of a female Sussaba mongolica (A) and a female Sussaba sugiharai (B).
Syrphoctonus tarsatorius (Panzer) Russia, Kuril islands: Paramushir Island, inland of Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. J. Schweikert. 1♀. Iturup Island, inland of Medvezh'ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. B.K. Urbain. 1♀. Iturup Island, Atsonupuri Peninsula base; N44°46.31'/E147°11.32', 15. VIII. 1999, leg. B.K. Urbain. 1♂. Iturup Island, Atsonupuri Peninsula base; N44°46.01'/E147°11.58', 14. VIII. 1999, leg. D.J. Bennett. 1♂. Kunashir Island, Environs "Goby" Hot Springs; N44°00.34'/E147°40.99', 27. VII. 1997, leg. N. Minakawa. 1♀. Holarctic and Oriental; recorded from Russia and Japan. Syrphophilus bizonarius (Gravenhorst) Russia, Kuril islands: Paramushir Island, inland E. Vasilyeva Bay; N50°01.84'/E155°23.88', 25. VII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of Dushnaya Bay; N47°04.49'/E152°11.52', 09. VIII. 1999, leg. D.J. Bennett. 1♂. Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 1♀. Iturup Island, inland Medvezh'Ya Bay; N45°25.75'/E148°49.75', 05. VIII. 1998, leg. B.K. Urbain. 1♀. Iturup Island, inland Konservnaya Bay; N45°19.79'/E147°59.75', 19. VIII. 1996, leg. P. Oberg. 1♀. Iturup Island, inland of Kuybyshevskiy Bay; N45°05.12'/E147°40.26', 13. VIII. 1999, leg. D.J. Bennett. 1♀. Iturup Island, Atsonupuri Peninsula base; N44°46.31'/E147°11.32', 15. VIII. 1999, leg. B.K. Urbain. 1♀. Kunashir Island, BtwLks Aliger Lagunnoye; N44°03.25'/E145°44.91', 11. VIII. 1998, leg. B.K. Urbain. 1♂. Kunashir Island, Near Lake Serebryanoye; N44°03.07'/E145°49.25', 22. VIII. 1998, leg. B.K. Urbain. 1♂. Holarctic and Oriental; recorded from Russia and Japan. Syrphophilus scabriculus (Holmgren) Russia, Kuril islands: Onekotan Island, inland of Nemo Bay; N49°36.24'/E154°49.81', 24. VII. 1999, leg. D.J. Bennett. 1♂. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.77'/E154°02.01', 29. VII. 2000, leg. D.J. Bennett. 1♀. Holarctic; recorded from Russia.
Tymmophorus gelidus Dasch(Fig. 5A) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 1♀. Although very similar in coloration to the widespread Tymmophorus obscuripes (Holmgren), this Northern species is distinguished by the short antennae, two strongly coriaceous bands starting at the notauli and continuing towards the scutellum, and some additional characters (see Dasch, 1964a). This species has been described from
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Canada and Greenland and has recently been reported from Northern Europe (Klopfstein, in press). Its presence on the northern Kuril island of Paramushir suggests a circumpolar distribution. Holarctic; new for Russia and the Eastern Palaearctic.
FIGURE 5. Habitus of a female Tymmophorus gelidus (A) and a female Tymmophorus obscuripes (B).
Tymmophorus obscuripes (Holmgren) (Fig. 5B) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. T.R. Anderson. 1♀. Holarctic; known from Russia. Tymmophorus suspiciosus (Brischke) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 2♂. Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. T.R. Anderson. 2♂; same data, but leg. D.J. Bennett: 1♂. Urup Island, inland from Aleutka Bay; N45°56.10'/ E150°09.70', 07. VIII. 2000, leg. T.R. Anderson. 3♂. Kunashir Island, Near Lake Allger; N44°02.88'/E145°44.48', 11. VIII. 1998, leg. D.J. Bennett. 2♀, 2♂. Palaearctic; known from Russia. Because this species has only recently been removed from synonymy with Tymmophorus erythrozonus (Förster) (which was often referred to under the name Tymmophorus rufiventris (Gravenhorst), which is a primary homonym (Klopfstein, in press)), it is not possible currently to associate the distributional records to either of them. The current distribution is thus only based on the records made under the name T. suspiciosus.
Discussion The 26 species of diplazontine wasps are probably a good sample of the diversity present on the Kuril islands, even though additional species might turn up when more sampling is done especially on the larger islands. For comparison, 33 species have been recorded from the Russian province Sakhalin Oblast to which the Kuril islands belong (Manukyan, 2007), 37 from Russian Kamchatka Oblast and 45 species from Japan (Yu et al., 2012). Except for Diplazon urupensis and D. varicoxa that were reported from the Kuril islands in a treatment of a limited sample of ichneumonids (Uchida, 1935), these are the first records for the Kuril islands. For one species, Tymmophorus gelidus Dasch, the present record fills a gap in the hitherto known distribution, i.e., Canada and Northern Europe. Given its occurrence on the Kuril islands, it is likely that it will be found in the Northern parts of the Eastern Palaearctic as well.
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In terms of distributional patterns, the proportion of very widespread species in this sample is striking. Only four of the species have been recorded exclusively from the Eastern Palaearctic, of which D. urupensis might still turn out to be a synonym of a Holarctic species (see note under this species). Sussaba sugiharai has an Eastern Palaearctic and Oriental distribution, and four species are Palaearctic. The remaining 17 species (65%) are Holarctic or even multiregional. This proportion is much higher than the 38% recorded by Manukyan (1995), a discrepancy that is probably caused by a combination of several factors. First, progress in diplazontine systematics has exposed several new synonyms between Nearctic and Palaearctic species since Manukyan's analysis (e.g., Klopfstein, in press). Second, if there is a positive relationship between population density and range, then the limited sample analyzed here might have overlooked some rare species with a restricted distributional range. And third, islands are areas where species with more effective dispersal abilities and thus wider distributions are overrepresented. Future accumulation of taxonomic and distributional data for Diplazontinae will help disentangling the reasons for the high proportion of widely distributed taxa on the Kuril islands, and host record data will help investigating the proximate reasons for the comparatively large distribution ranges.
Acknowledgements I would like to thank Prof. Robert Wharton for inspiring this study and making the material accessible. John Jennings helped improve an earlier version of this manuscript. This work was supported by the Swiss National Science Foundation grants PBBEP3_130173 and PA00P3_145377 to S. Klopfstein.
References Dasch, C.E. (1964a) Ichneumon-flies of America north of Mexico 5: Subfamily Diplazontinae. Memoirs of the American Entomological Institute, 3, 1–304. Dasch, C.E. (1964b) The neotropic Diplazontinae. Contributions to the American Entomological Institute, 1, 1–77. Fitton, M.G. & Rotheray, G.E. (1982) A key to the European genera of diplazontine ichneumon-flies, with notes on the British fauna. Systematic Entomology, 7, 311–320. http://dx.doi.org/10.1111/j.1365-3113.1982.tb00448.x Gauld, I.D., Wahl, D., Bradshaw, K. & Hanson, W.S. (1997) The Ichneumonidae of Costa Rica, 2. Introduction and keys to species of the smaller subfamilies, Anomaloninae, Ctenopelmatinae, Diplazontinae, Lycorininae, Phrudinae, Tryphoninae (excluding Netelia) and Xoridinae, with an appendices on the Rhyssinae. Memoirs of the American Entomological Institute, 57, 1–485. Kasparyan, D.R. & Manukyan, A.R. (1987) A new genus of the ichneumonid wasps of the subfamily Diplazontinae (Hymenoptera, Ichneumonidae) from the eastern Palaearctic region (in Russian). Entomologicheskoye obozreniye, 66, 841–844. Kasparyan, D.R. & Manukyan, A.R. (1989) A new genus of parasitic wasps (Hymenoptera: Ichneumonidae Diplazontinae) from the eastern Palaearctic region. Entomological Review, 67, 14–17. Klopfstein, S. (2011) A review of the Diplazontinae of Mongolia (Hymenoptera: Ichneumonidae). Zootaxa, 2790, 35–53. Klopfstein, S. (2014) Revision of the Western Palaearctic Diplazontinae (Hymenoptera, Ichneumonidae), Zootaxa, in press. Klopfstein, S., Kropf, C. & Quicke, D.L.J. (2010) An evaluation of phylogenetic informativeness profiles and the molecular phylogeny of Diplazontinae (Hymenoptera, Ichneumonidae). Systematic Biology, 59, 226–241. Klopfstein, S., Quicke, D.L.J., Kropf, C. & Frick, H. (2011) Molecular and morphological phylogeny of Diplazontinae (Hymenoptera, Ichneumonidae). Zoologica Scripta, 40, 379–402. http://dx.doi.org/10.1111/j.1463-6409.2011.00481.x Manukyan, A.R. (1987) On the systematics of Ichneumonids of the genus Diplazon Nees (Hymenoptera, Ichneumonidae) of the fauna of the USSR. In: Ler, P.A. & Storozheva, N.A. (Eds.), New data on the systematics of insects of the Far East. Acad. Sci., USSR., Vladivostok, pp. 66–72. [in Russian] Manukyan, A.R. (1988) Review of the genera Sussaba Cameron and Xestopelta Dasch (Hymenoptera, Ichneumonidae) of the USSR fauna. Proceedings of the Zoological Institute, 175, 44–54. [in Russian] Manukyan, A.R. (1995) The geographic distribution of the Diplazontinae (Hymenoptera, Ichneumonidae) in the Palaearctic region, with description of two new species. Acta Zoologica Fennica, 199, 55–60. Manukyan, A.R. (2007) Diplazontinae. In: Lelej, A.S. (Ed.), Neuropteroidea, Mecoptera, Hymenoptera. Pt 5. Dalnauka, Vladivostok, pp. 718–732. [in Russian] Mayr, E. (1942) Systematics and the Origin of Species, from the Viewpoint of a Zoologist. Harvard University Press, Cambridge, 334 pp.
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Nakanishi, A. (1967) Description of a new species of the genus Diplazon Nees from the Ryukyus (Hymenoptera, Ichneumonidae). Kontyu, 35, 34–35. Nakanishi, A. (1978) A new genus and two new species of the Diplazontinae (Hymenoptera: Ichneumonidae) from Japan. Kontyu, 46, 429–432. Nakanishi, A. (1979) Studies on the genus Sussaba Cameron of Japan (Hymenoptera: Ichneumonidae). Sieboldia, 4, 297–315. Nakanishi, A. (1985) Discovery of the Diplazontine genera Bioblapsis, Campocraspedon and Tymmophorus from Japan (Hymenoptera, Ichneumonidae). Kontyu, 53, 28–33. Nakanishi, A. (1986) A study of the genus Promethes Foerster from Japan (Hymenoptera, Ichneumonidae). Sieboldia, 5, 119– 129. Ngamo Tinkeu, L.S. & Hance, T. (1997) Reduction of predatory efficiency of hoverflies (Diptera, Syrphidae) in cereal fields due to parasitism. Mededelingen Faculteit Landbouwkundige En Toegepaste Biologische Wetenschappen Universiteit Gent, 62, 469–472. Rotheray, G.E. (1984) Host relations, life cycles and multiparasitism in some parasitoids of aphidophagous Syrphidae (Diptera). Ecological Entomology, 9, 303–310. http://dx.doi.org/10.1111/j.1365-2311.1984.tb00853.x Sauter, K.F. (1996) Treasure islands: Modern-day Charles Darwins put ashore on the remote islands of the Kuril Archipelago. Popular Science, 49, 58–61. Schneider, F. (1951) Einige physiologische Beziehungen zwischen Syrphidenlarven und ihren Parasiten. Zeitschrift für Angewandte Entomologie, 33, 150–162. http://dx.doi.org/10.1111/j.1439-0418.1952.tb00661.x Uchida, T. (1935) Beitraege zur Kenntnis der Ichneumonidenfauna der Kurilen. Insecta Matsumurana, 9, 108–122. Uchida, T. (1957) Beiträge zur Kenntnis der Diplazoninen-Fauna Japans und seiner Umgegenden (Hymenoptera, Ichneumonidae). Journal of the Faculty of Agriculture, 50, 225–265. Yu, D.S., Van Achterberg, C. & Horstmann, K. (2012) Taxapad 2012, Ichneumonoidea 2011, Ottawa, Ontario, Canada. Database on flash-drive. Available from: http://www.taxapad.com (accessed 14 February 2014)
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Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3779.1.5 http://zoobank.org/urn:lsid:zoobank.org:pub:79B81076-9D53-4D21-912E-721703237DB2
Review of the Diplazontinae (Hymenoptera, Ichneumonidae) of the Kuril islands, with descriptions of two new species SERAINA KLOPFSTEIN Swedish Museum of Natural History, Box 50007, SE–104 05 Stockholm, Sweden and School of Earth and Environmental Sciences, The University of Adelaide, North Terrace, Darling Building, Adelaide, SA 5005. E-mail: [email protected]
Abstract A sample of 346 specimens of Diplazontinae parasitoid wasps from the Kuril islands was studied. Twenty-six species are reported, Tymmophorus gelidus Dasch for the first time for the Eastern Palaearctic. Two new species are described, Diplazon kurilensis sp. n. and Homotropus formosus sp. n. Diplazon urupensis Uchida is removed from synonymy (stat. rev.), and Promethes persulcatus Nakanishi is suggested as a synonym of Promethes bridgmani Fitton. Reasons are discussed for the large proportion of species with a Holarctic or even multi-regional distribution in the sample, which amounts to17 of the species or 65%. Key words: parasitoid wasps, faunistics, Russian Far East, distributional patterns
Introduction The Diplazontinae are a moderately-sized subfamily of parasitoids of the family Ichneumonidae; 347 species are currently recognized worldwide (Yu et al., 2012). Where reliable host records are available, they point to a rather specialized host range of mostly aphidophagous hoverflies (e.g., Schneider, 1951; Fitton & Rotheray, 1982; Rotheray, 1984; Ngamo Tinkeu & Hance, 1997). Most diplazontine species have been described from temperate regions (Yu et al., 2012). Although the tropics are severely understudied, two locally extensive studies suggest that the subfamily does not exhibit an increase in species richness with decreasing latitude as it can be observed in other insect groups (Dasch, 1964b; Gauld et al., 1997), which is in line with the relative scarcity of aphids and aphidophagous syrphids in the tropics. In the Holarctic region, both the Nearctic (Dasch, 1964a) and the Western Palaearctic faunas (Klopfstein, in press) have been revised, while the Eastern Palaearctic is only partially covered by taxonomic and faunistic studies (Uchida, 1957; Nakanishi, 1967, 1978, 1979, 1985, 1986; Kasparyan & Manukyan, 1987; Manukyan, 1987, 1988; Kasparyan & Manukyan, 1989; Manukyan, 2007; Klopfstein, 2011). Summarizing distributional patterns of Palaearctic diplazontines, Manukyan (1995) pointed out that a high proportion of diplazontine species occurs in several faunistic regions; of the 130 Palaearctic species known at the time; 38% were either cosmopolitan, multiregional or at least Holarctic. His assessment was however hindered by a lack of taxonomic revisions which compared the Nearctic to the Palaearctic faunas and a gap of knowledge in part of the Eastern Palaearctic region. Even though only a part of the Nearctic species were studied in the revision of the Western Palaearctic fauna, Klopfstein (in press) proposed eight new synonyms between Nearctic and Western Palaearctic taxa, and more can be expected in the future (Dasch, 1964a). Manukyan's estimate is thus probably an underestimate. I recently received 346 specimens from the Kuril islands, most of which had been collected by the International Kuril Island Project (www.burkemuseum.org/static/okhotskia/ikip/, Sauter, 1996). This material includes 26 species, two of which are here newly described. Given the geographic location of the Kuril islands, between Russian Kamchatka and the Japanese island Hokkaido (Fig. 1), this material is especially interesting with respect to an assessment of the distributional ranges in Diplazontinae.
20 Accepted by J. T. Jennings: 21 Jan. 2014; published: 13 Mar. 2014
Material and methods I obtained 346 specimens from the Kuril islands from the Texas A&M University Insect Collection (College Station, Texas). Additional material was studied for comparison at the Swedish Museum of Natural History (Stockholm). Junior synonyms are only given if they are proposed in the present paper, as accurate lists of synonyms are given in the catalogue by Yu et al. (2012). For terminology of morphology, I follow Townes (1969). All measurements were taken with a Leica Wild M10 stereo-microscope with a 10x ocular including an eye-piece micrometer. Specimens were photographed with a Canon EOS 7D with a Canon Macro lens EF 100mm 1:2.8 USM, and stacked using Helicon Focus pro x64. All specimens, including the type material, are stored at the Texas A&M University Insect Collection (TAMU).
FIGURE 1. Map of the Kuril islands and adjacent Hokkaido and Kamchatka. Modified from Wikimedia Commons.
Descriptions of new species Diplazon kurilensis sp. n. (Fig. 2) This species belongs to the pectoratorius group of species in the genus Diplazon, which unites the species with rather weak notauli, reduced carinae on the propodeum, epicnemical carina reduced ventrally, weak transverse impressions on the tergites, and a white hind tibia with only the apex but not the base darkened. According to recent molecular and morphological phylogenetic studies (Klopfstein et al., 2010; Klopfstein et al., 2011), this species group is sister to all the other species in the genus. The other three species currently known in the group are D. urupensis Uchida from the Kuril islands (see below), D. pectoratorius (Thunberg) with a Holarctic, Oriental and Neotropic distribution, and the Nearctic and Eastern Palaearctic D. albotibialis Dasch. From these species, D. kurilensis can be distinguished by its small size, reduced sculpture on the mesoscutum and metasoma, and brown coloration of the metasoma. DIPLAZONTINAE OF THE KURIL ISLANDS
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Etymology. This species is named “kurilensis” after the type locality. Description. Fore wing length 3.2–3.6 mm (3.6 in holotype) in females, 4.1 mm in the male. Antenna with 15–16 (16) flagellomeres in females, all flagellomeres with multiporous plate sensillae sparse but also present ventrally, apical flagellomere long and enlarged, thus appearing clubbed; 18 flagellomeres in the male, without tyloids. Face strongly coriaceous and matt, impunctate, without vertical impressions. Clypeus with apical margin thin, slightly impressed on the sides after a basal elevation, resulting in the lateral areas being slightly concave. Pronotum and propleuron smooth with some sparse punctures. Mesoscutum with weak to distinct notauli; smooth and shining with very sparse punctures, mostly in the front and on the sides. Mesopleuron with epicnemical carina reduced ventrally; with weak and sparse punctures and some very irregular, weak sculpture. Scutellum only carinate at base, smooth and shining. Metapleuron mostly smooth, with some punctures in the front. Propodeum only with metapleural carina and hind part of the lateral longitudinal carinae present, rugulously sculptured near the base, more rugose and partly smooth on petiolar area; propodeal spiracle not enlarged. Fore wing without areolet; hind wing with 2 basal hamuli. Legs including coxae smooth and shining, hind tibia without thickened setae. Female metasoma dorsoventrally depressed, tapered towards the apex, apical margins of the tergites straight to even slightly concave, with transverse impressions weak to almost indistinct on tergites 1–3. First tergite 1.3–1.45 (1.45) times as long as wide, with median dorsal carinae only present on about basally, finely coriaceous and matt; second tergite 0.7 times as long as wide, finely coriaceous and matt; spiracle of second and third tergites on dorsal part, above lateral fold. Ovipositor sheaths about 0.3 times as long as hind tibia, pointed and closed at the tip, coriaceous and matt and with numerous setae on whole surface.
FIGURE 2. Habitus of the female holotype (A) and male paratype (B) of Diplazon kurilensis sp. n.
Coloration of females. Antenna brown. Head and mesosoma dark brown, with yellow inner eye margins, yellow on clypeus, mouthparts, hind corner of pronotum, tegula, subtegular ridge, shoulder mark, and mesepimeron; scutellum yellow on anterior corners, orange centrally. Legs yellow, all coxae yellow, hind coxa with some orange basally, hind femur orange, hind tibia white with apex light brown, hind tarsus brown. Metasoma brown, progressively lighter towards apex which is orange, end margins of tergites whitish. Coloration of male. As in females but additionally with yellow over entire face, scape, pedicel and flagellomeres ventrally, propleuron, mesosternum, mesopleuron in front of epicnemical carina and with a line on lower mesopleuron; all coxae entirely yellow. Types material. Holotype ♀: Russia, Kuril islands: Kharimkotan, Northwest corner of island. N49°08.60'/ E154°28.10', 28. VII. 2000. leg. D.J. Bennett. Paratypes, one ♀ and one #: Russia, Kuril islands: Matua Island, inland from Dvoinaya Bay. N48°03.95'/E153°15.63', 31. VII. 2000. leg. D.J. Bennett.
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Homotropus formosus sp. n. (Fig. 3) This species belongs to the pictus species group (according to Dasch, 1964a). Its black coxae and large size in combination with a strongly coriaceous and matt mesosoma distinguish it from most species there-in. In terms of size, it could be confused with Homotropus megaspis Thomson and H. momoi (Uchida); from the former, it differs by the dark coxae, and from both by the coriaceous sculpture of the mesosoma. The otherwise similar Homotropus areolaris (Uchida) differs by the coloration of the stigma in the fore wing, the coloration of the fore and mid coxae, and the more extensive coriaceous sculpture.
FIGURE 3. Habitus of the female holotype (A) and male paratype (B) of Homotropus formosus sp. n.
Etymology. The name “formosus” is intended to reflect the overall appearance of the species. Description. Fore wing length 5.1–5.5 mm (5.1 in holotype) in females, 5.8 mm in the male. Antenna with 21– 22 (21) flagellomeres in females, apical flagellomeres with multiporous plate sensilla sparse but also present ventrally; 23 flagellomeres in males, with narrow, long tyloids on flagellomeres 7 to 15. Face strongly coriaceous and matt, distinctly punctate, especially centrally, without vertical impressions. Clypeus with apical margin thin, strongly impressed, resulting in the central area being convex. Pronotum and propleuron with strong and dense punctures, smooth or finely coriaceous in-between. Mesoscutum without notauli; finely coriaceous and matt in the females, less coriaceous and mostly shining in the male, in both sexes with dense punctures on whole surface. Mesopleuron with epicnemical carina strong and complete ventrally; strongly coriaceous and matt with dense punctures except for small smooth area around the speculum in females, in the male matt only on lower half, more shining on upper half between the punctures. Scutellum only carinate at base, rather strongly convex, smooth between the strong punctures. Metapleuron with dense punctures, but also some smooth areas. Propodeum only with metapleural carina and hind part of the lateral longitudinal carinae present, densely and rugulously punctate near the base, more rugose and partly smooth on petiolar area; propodeal spiracle not enlarged. Fore wing areolet sometimes large, almost rhombic, vein 3rs-m mostly well pigmented; hind wing with 2–3 basal hamuli. Legs including coxae coriaceous and matt, hind tibia with thickened, scale-like hairs on outer surface. Female metasoma dorsoventrally depressed, tapered towards the apex, apical margins of the tergites straight or convex, without transverse impressions. First tergite 1.1–1.3 (1.1) times as long as wide, with median dorsal carinae only present on about basal fourth, finely coriaceous and matt; second tergite 0.6–0.7 times as long as wide, basally with some strong longitudinal wrinkles, finely coriaceous and matt and with a few, very sparse and weak punctures; second tergite 0.8–0.9 times length of first tergite; spiracle of second and third tergites on dorsal part, above lateral fold. Ovipositor sheaths about 0.3 times as long as hind tibia, more or less parallel-sided and truncate close to the tip, smooth and shining and with some setae close to the tip. Coloration of females. Antenna black. Head and mesosoma black, with small yellow or orange central face DIPLAZONTINAE OF THE KURIL ISLANDS
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patch, yellow on clypeus, mouthparts, hind corner of pronotum, tegula, subtegular ridge, large shoulder mark, and upper mesepimeron; scutellum entirely black. Fore wing with stigma lighter at base. Legs orange, all coxae black, fore coxa apically with some yellow, trochanters basally black, apically yellow, trochantelli yellow, femora orange, hind tibia white with apex, base and a small subbasal spot dark, hind tarsus dark. Metasoma entirely black. Coloration of males. As in females but additionally with yellow over entire face, scape, pedicel and flagellomeres ventrally, propleuron, mesosternum, mesopleuron in front of epicnemical carina and with a spot on apical lower corner; fore and mid coxae entirely and hind coxa apically yellow, hind femur with dark marks at base and yellow at tip. Type material. Holotype ♀: Russia, Kuril islands: Urup Island, Inland from Aleutka Bay. N45°56.18'/ E150°09.39', 07. VIII. 2000. leg. D.J. Bennett. Paratypes: one ♀, same data as holotype; one ♂: Japan, Hokkaido, Mt. Yuubari, Yuubari city. 01. VII. 1995. leg. Ito Gen.
Annotated species list Diplazon pectoratorius (Thunberg) Russia, Kuril islands: Iturup Island, Inland Konservnaya Bay; N45°19.79'/E147°59.75', 19. VIII. 1996, leg. P. Oberg. 2♂. Paramushir Island, inland from Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. B.K. Urbain. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°36.62'/E154°49.21', 27.VII.2000, leg. D.J. Bennett. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°36.08'/E154°48.96', 27.VII.2000, leg. D.J. Bennett. 1♀. Shiashkotan, Inland SW Zakatnaya Bay; N48°46.83'/E154°01.85', 29. VII. 2000, leg. T.R. Anderson. 1♀. Urup Island, inland of Ukromnaya Bay; N45°35.43'/E149°31.91', 20. VIII. 1996, leg. B.K. Urbain. 6♂. Holarctic, Oriental and Neotropic; known from Russia and Japan. Diplazon urupensis Uchida (stat. rev.) Russia, Kuril islands: Paramushir Island, Inland from Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. B.K. Urbain. 1♀. Shiashkotan, inland SW Zakatnaya Bay; N48°46.83'/E154°01.85', 29. VII. 2000, leg. T.R. Anderson. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 5♀. This species was described from the Kuril islands by Uchida (1935). In a later publication (Uchida, 1957), he then synonymized it with D. pectoratorius. However, after studying the present specimens and consulting Uchida's holotype and allotype of the species (deposited at the Hokkaido University, Sapporo; label data of the holotype: "Uruppu; K.Doi" / "Holotype; Diplazon; urupensis; (Uch.)"), I decided to reinstate D. urupensis as a valid species (stat. rev.). The specimens are clearly smaller and more slender than D. pectoratorius, lack the orange coloration of the mesopleuron and mesosternum (except for a few males), and have weaker and sparser punctures on the mesoscutum and mesopleuron. It remains to be shown whether the Nearctic D. albotibialis Dasch (1964a), which has also been reported from Eastern and Western Russia (Manukyan, 1987, 2007), is a distinct species or a junior synonym of D. urupensis. Known only from the Kuril islands. Diplazon varicoxa (Thomson) Russia, Kuril islands: Urup Island, inland from Aleutka Bay; N45°56.18'/E150°09.39', 07. VIII. 2000, leg. D.J. Bennett. 1♀. Urup Island, inland from Aleutka Bay; N45°56.10'/E150°09.70', 07. VIII. 2000, leg. T.R. Anderson. 1♀. Urup Island, Inland of Chernoburka Bay; N45°36.26'/E149°34.55', 09. VIII. 2000, leg. D.J. Bennett. 1♂. Iturup Island, inland of Medvezh'ya Bay; N45°27.27'/E148°49.76', 05. VIII. 1998, leg. B.K. Urbain. 1♂. Iturup Island, Inland of Medvezh'ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. B.K. Urbain. 1♀; same data, but leg. B.K. Urbain, 1♂. Iturup Island, inland of Medvezh'ya Bay; N45°25.75'/E148°49.75', 05. VIII. 1998, leg. B.K. Urbain. 1♀. Iturup Island, Inland of Konservnaja Bay; N45°20.04'/E147°59.84', 12. VIII. 1999, leg. D.J. Bennett. 1♂; same data, but 30. VII. 1997, leg. B.K. Urbain:. 1♀. Iturup Island, Nr. Konservnaya Bay; N45°20.02'/ E148°00.03', 30. VII. 1997, leg. B.K. Urbain. 2♀. Iturup Island, inland of Konservnaja Bay; N45°20.01'/ E147°59.91', 31. VIII. 1995, leg. N. Minakawa. 1♂; same data, but leg. N. Minakawa: 2♂. Iturup Island, 4km east Kitovyi Village; N45°15.90'/E147°55.90', 29. VII. 1997, leg. J. Schweikert. 3♂. Iturup Island, North of Cape
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Kanonerka; N45°11.31'/E148°15.43', 03. VIII. 1998, leg. K. Kavun. 1♂. Iturup Island, inland of Kuybyshevskiy Bay; N45°04.49'/E147°39.07', 13. VIII. 1999, leg. B.K. Urbain. 5♀. Iturup Island, Near Sernozavoskaya Bay; N44°58.29'/E147°53.69', 02. VIII. 1998, leg. B.K. Urbain. 1♂. Iturup Island, Nr. Sernozavoskaya Bay; N44°57.88'/E147°53.89', 02. VIII. 1998, leg. D.J. Bennett. 1♂. Iturup Island, Nr. Sernozavoskaya Bay; N44°57.84'/E147°53.83', 02. VIII. 1998, leg. B.K. Urbain. 1♂. Iturup Island, Atsonupuri Peninsula base; N44°47.60'/E147°11.42', 14. VIII. 1999, leg. B.K. Urbain. 1♂. Iturup Island, Atsonupuri Peninsula base; N44°47.34'/E147°11.34', 14. VIII. 1999, leg. B.K. Urbain. 3♂. Iturup Island, Atsonupuri Peninsula base; N44°46.41'/E147°11.20', 14. VIII. 1999, leg. B.K. Urbain. 23♂. Iturup Island, Atsonupuri Peninsula base; N44°46.31'/E147°11.32', 15. VIII. 1999, leg. B.K. Urbain. 7♂. Iturup Island, Atsonupuri Peninsula base; N44°46.01'/E147°11.58', 14. VIII. 1999, leg. D.J. Bennett. 2♂. Iturup Island, Env. Tikhaya River mouth; N44°43.38'/E147°12.77', 15. VIII. 1999, leg. D.J. Bennett. 2♀. Kunashir Island, 1km E of Tyatina R. Mouth; N44°16.32'/E146°10.34', 28. VIII. 1997, leg. Y. Marusik. 2♂. Kunashir Island, Goby Hot Springs environment; N44°00.39'/E145°41.07', 17. VIII. 1999, leg. D.J. Bennett. 1♂. Palaearctic and Oriental; known from Russia and Japan. Fossatyloides gracilentus (Holmgren) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 1♂. Holarctic; also known from the Russian Far East. Homotropus elegans (Gravenhorst) Russia, Kuril islands: Raikoke Island, inland from Eastern side; N48°17.87'/E153°15.67', 30. VII. 2000, leg. D.J. Bennett. 1♀. Raikoke Island, inland from Eastern side; N48°17.81'/E153°15.68', 30. VII. 2000, leg. T.R. Anderson. 13♂. Raikoke Island, inland from Eastern side; N48°17.73'/E153°15.59', 30. VII. 2000, leg. T.R. Anderson. 1♂. Raikoke Island, inland from Eastern side; N48°17.71'/E153°15.64', 30. VII. 2000, leg. D.J. Bennett. 1♂. Holarctic; reported from Russia, including its easternmost provinces. Homotropus nigritarsus (Gravenhorst) Russia, Kuril islands: Paramushir Island, Nr. Putyatino settlement; N50°44.19'/E156°08.82', 04. VIII. 1997, leg. B.K. Urbain. 1♀, 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.54'/E156°05.71', 30. VII. 1999, leg. B.K. Urbain. 1♂. Alaid Island, inland of Alaidskaya Bay; N50°29'/E155°39', 25. VII. 1999, leg. A.S. Lelej. 1♂. Antsiferova Island, inland of near Vydar Rock; N50°12.36'/E154°57.73', 15. VIII. 1997, leg. B.K. Urbain. 1♀. Paramushir Island, Nr. Vasil'yeva Peninsula; N50°02.87'/E155°23.46', 03. VIII. 1996, leg. B.K. Urbain. 3♀. Kharimkotan, Northwest corner of island; N49°08.75'/E154°27.64', 28. VII. 2000, leg. D.J. Bennett. 1♂. Holarctic and Neotropical; reported from Russia, including its easternmost provinces. Homotropus pallipes (Gravenhorst) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 1♂. Iturup Island, Near Sernozavoskaya Bay; N44°58.29'/E147°53.69', 02. VIII. 1998, leg. B.K. Urbain. 1♀. Shikotan Island, SW shore of Delfin Bay; N43°50'/E146°44', 11. IX. 1997, leg. Y. Marusik. 1♂. Holarctic and Neotropical; reported from Russia, including its easternmost provinces. Homotropus pictus (Gravenhorst) Russia, Kuril islands: Paramushir Island, Nr. Putyatino settlement; N50°44.19'/E156°08.82', 04. VIII. 1997, leg. B.K. Urbain. 1♂. Paramushir Island, Env. Utyesnaya River; N50°37.73'/E156°08.21', 01. VIII. 1996, leg. M. Ohara. 1♂. Makanrushi Island, inland from Zakat Bay; N49°44.37'/E154°24.84', 18. VIII. 1997, leg. J. Schweikert. 1♀. Kharimkotan, Northwest corner of island; N49°08.95'/E154°28.54', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corner of island; N49°08.93'/E154°28.09', 28. VII. 2000, leg. T.R. Anderson. 2♀. Kharimkotan, Northwest corner of island; N49°08.67'/E154°28.29', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corner of island; N49°08.60'/E154°28.16', 28. VII. 2000, leg. D.J. Bennett. 1♀, 1♂. Matua Island, inland from Dvoinaya Bay; N48°03.95'/E153°15.63', 31. VII. 2000, leg. D.J. Bennett. 2♂. Matua
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Island, Inland of Dvoinaya Bay; N48°03.12'/E153°14.80', 03. VIII. 1999, leg. B.K. Urbain. 2♂. Rasshua Island, inland near Malenkaya Bay; N47°43.26'/E152°58.38', 04. VIII. 1999, leg. B.K. Urbain. 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.86'/E152°16.20', 02. VIII. 2000, leg. D.J. Bennett. 3♀. Simushir Island, inland of NE Broutona Bay; N47°07.82'/E152°16.40', 02. VIII. 2000, leg. T.R. Anderson. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.77'/E152°16.28', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.27'/E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 10♀. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 3♀. Simushir Island, inland of NE Broutona Bay; N47°07.01'/E152°16.17', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of NE Broutona Bay; N47°06.93'/E152°15.92', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of Malaya Bay; N47°05.38'/E152°08.10', 18. VIII. 1995, leg. B.K. Urbain. 1♀. Simushir Island, inland of Malaya Bay; N47°05.18'/E152°07.81', 18. VIII. 1995, leg. M. Ohara. 2♀. Palaearctic; reported from Russia, including its easternmost provinces. Homotropus signatus (Gravenhorst) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 2♂. Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 3♂. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♂. Matua Island, inland of Dvoinaya Bay; N48°04'/E153°15', 03. VIII. 1999, leg. A.S. Lelej. 1♀, 1♂. Rasshua Island, W. Coast of Beloye Lake; N47°43.30'/E152°59.52', 12. VIII. 1995, leg. Y. Marusik. 1♂. Rasshua Island, SSW part; near "Arches"; N47°43.19'/E152°58.80', 12. VIII. 1995, leg. M. Ohara. 1♀. Ketoi Island, East of Cape Storozheva; N47°22.64'/E152°27.42', 15. VIII. 1995, leg. B.K. Urbain. 1♀. Ketoi Island, East of Cape Storozheva; N47°22.60'/E152°27.76', 15. VIII. 1995, leg. M. Ohara. 1♂. Ketoi Island, East of Cape Storozheva; N47°22.54'/ E152°27.47', 15. VIII. 1995, leg. N. Minakawa. 4♀, 2♂. Ketoi Island, inland of Diany Bay; N47°18.72'/ E152°26.07', 06. VIII. 1999, leg. D.J. Bennett. 2♂. Simushir Island, inland of NE Broutona Bay; N47°07.86'/ E152°16.20', 02. VIII. 2000, leg. T.R. Anderson. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.77'/ E152°16.28', 02. VIII. 2000, leg. D.J. Bennett. 2♀, 7♂. Simushir Island, inland of NE Broutona Bay; N47°07.27'/ E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 1♀, 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.20'/ E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 1♀, 2♂. Simushir Island, Inland of Dushnaya Bay; N47°05'/E152°12', 09. VIII. 1999, leg. A.S. Lelej. 1♂. Simushir Island, inland of Dushnaya Bay; N47°04.76'/E152°12.19', 09. VIII. 1999, leg. B.K. Urbain. 1♀, 1♂. Chirpoi Island, inland SW Peschanaya Bay; N46°32.05'/E150°53.51', 03. VIII. 2000, leg. D.J. Bennett. 1♀, 1♂. Urup Island, inland from Aleutka Bay; N45°56.08'/E150°09.98', 07. VIII. 2000, leg. T.R. Anderson. 2♀. Iturup Island, inland of Slavnaja Bay; N45°29.47'/E148°37.13', 06. VIII. 1998, leg. D.J. Bennett. 1♀, 1♂. Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 1♀. Kunashir Island, Near Lake Aliger; N44°02.88'/E145°44.48', 11. VIII. 1998, leg. D.J. Bennett. 1♂. Kunashir Island, Environs of Lake Aliger; N44°02.83'/E145°44.47', 09. IX. 1995, leg. P. Oberg. 4♀. Tanfilyeva, Inland of Tanfil'Yevka Bay; N43°26.62'/ E145°55.89', 19. VIII. 1998, leg. B.K. Urbain. 1♂. Holarctic; reported from Russia and Japan. Phthorima compressa (Desvignes) Russia, Kuril islands: Iturup Island, inland of Slavnaja Bay; N45°29.47'/E148°37.13', 05. VIII. 1998, leg. D.J. Bennett. 1♂. Shikotan Island, inland of Otradnaya Bay; N43°52.14'/E146°47.67', 12. VIII. 1998, leg. B.K. Urbain. 1♀. Tanfilyeva, inland Tanfil'Yevka Bay; N43°26.62'/E145°55.89', 19. VIII. 1998, leg. D.J. Bennett. 1♂. Anuchina Island, inland of Bolshoye Bay; N43°22.26'/E146°00.36', 19. VIII. 1998, leg. B.K. Urbain. 1♂. Holarctic; reported from Russia. Promethes bridgmani Fitton Promethes persulcatus Nakanishi, 1986 is a junior synonym, syn. nov. Russia, Kuril islands: Paramushir Island, inland of Rifovaya Bay; N50°29.84'/E156°05.80', 30. VII. 1999, leg. B.K. Urbain. 1♂. The type of Promethes persulcator Nakanishi could not be obtained, but based on the detailed description (Nakanishi, 1986) and on this male from the Kuril islands, these two species are clearly synonymous. While P.
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bridgmani has only been reported from Europe, P. persulcator has been recorded in Japan and the Russian Far East. Palaearctic, including Japan and the Russian Far East. Promethes sulcator (Gravenhorst) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. J. Schweikert. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♀, 4♂. Paramushir Island, inland of Rifovaya Bay; N50°29.43'/E156°05.52', 30. VII. 1999, leg. D.J. Bennett. 1♀. Paramushir Island, Base Vasil'Yeva Peninsula; N50°03.31'/E155°23.60', 16. VIII. 1997, leg. B.K. Urbain. 1♂. Rasshua Island, inland near Malenkaya Bay; N47°42.77'/E152°58.16', 04. VIII. 1999, leg. B.K. Urbain. 2♀. Simushir Island, inland of NE Broutona Bay; N47°07.27'/E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 2♂. Simushir Island, inland of NE Broutona Bay; N47°07.01'/E152°16.17', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Urup Island, inland of Ukromnaya Bay; N45°35.43'/E149°31.91', 20. VIII. 1996, leg. B.K. Urbain. 2♀. Iturup Island, inland of Konservnaja Bay; N45°20.04'/E147°59.84', 12. VIII. 1999, leg. D.J. Bennett. 1♂. Iturup Island, 4km east Kitovyi Village; N45°16.07'/E147°56.38', 29. VII. 1997, leg. B.K. Urbain. 1♂. Iturup Island, North of Cape Kanonerka; N45°11.12'/E148°14.57', 03. VIII. 1998, leg. D.J. Bennett. 1♂. Iturup Island, Nr. Sernozavoskaya Bay; N44°57.84'/E147°53.83', 02. VIII. 1998, leg. B.K. Urbain. 1♀. Kunashir Island, Env. Of Cape Sukacheva; N44°04.50'/E145°51.95', 20. VIII. 1997, leg. Y. Marusik. 1♂. Shikotan Island, inland from Gorobets Bay; N43°92.64'/E146°42.48', 18. VIII. 1998, leg. B.K. Urbain. 1♀. Shikotan Island, SW shore of Delfin Bay; N43°50'/E146°44', 11. IX. 1997, leg. Y. Marusik. 1♀. Holarctic and Oriental; recorded from Russia and Japan. Sussaba aciculata (Ruthe) Russia, Kuril islands: Paramushir Island, Env. Utyesnaya River; N50°37.73'/E156°08.21', 01. VIII. 1996, leg. M. Ohara. 1♀. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♀. Paramushir Island, inland of Rifovaya Bay; N50°29'/E156°06', 30. VII. 1999, leg. A.S. Lelej. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°22.26'/E155°36.17', 13. VIII. 1997, leg. B.K. Urbain. 1♂. This species differs from the Japanese species Sussaba nigra Nakanishi by the strongly coriaceous median lobe of the mesonotum and from Sussaba dajsetsuzana Nakanishi by the lack of a black area in the tyloids of males (Nakanishi, 1979). Holarctic; reported from the Russian Far East. Sussaba cognata (Holmgren) Russia, Kuril islands: Alaid Island, inland of Alaidskaya Bay; N50°49.56'/E155°39.88', 25. VII. 1999, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.84'/E156°05.80', 30. VII. 1999, leg. B.K. Urbain. 1♀. Paramushir Island, inland of Rifovaya Bay; N50°29.70'/E156°05.68', 30. VII. 1999, leg. B.K. Urbain. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°36.62'/E154°49.21', 27.VII.2000, leg. D.J. Bennett. 1♀. Onekotan Island, inland and South of Nemo Bay; N49°35.96'/E154°49.11', 27.VII.2000, leg. D.J. Bennett. 1♂. Kharimkotan, Northwest corder of island; N49°08.95'/E154°28.54', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corder of island; N49°08.67'/E154°28.29', 28. VII. 2000, leg. T.R. Anderson. 1♀. Kharimkotan, Northwest corder of island; N49°08.60'/E154°28.10', 28. VII. 2000, leg. D.J. Bennett. 1♀. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.87'/E154°02.08', 29. VII. 2000, leg. T.R. Anderson. 1♀. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.83'/E154°01.85', 29. VII. 2000, leg. D.J. Bennett. 1♀. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.77'/E154°02.01', 29. VII. 2000, leg. D.J. Bennett. 2♀. Matua Island, inland from Dvoinaya Bay; N48°04.17'/E153°15.28', 31. VII. 2000, leg. D.J. Bennett. 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.86'/E152°16.20', 02. VIII. 2000, leg. T.R. Anderson. 3♀. Simushir Island, inland of NE Broutona Bay; N47°07.85'/E152°16.34', 02. VIII. 2000, leg. T.R. Anderson. 4♀. Simushir Island, inland of NE Broutona Bay; N47°07.82'/E152°16.40', 02. VIII. 2000, leg. T.R. Anderson. 3♀, 1♂. Simushir Island, inland of NE Broutona Bay; N47°07.77'/E152°16.28', 02. VIII. 2000, leg. D.J. Bennett. 2♀. Simushir Island, inland of NE Broutona Bay; N47°07.27'/E152°16.33', 02. VIII. 2000, leg. D.J. Bennett. 3♀, 3♂. Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Simushir Island, inland of NE
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Broutona Bay; N47°07.01'/E152°16.17', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Brat Chirpoev, inland from Uglovaya Bay; N46°27.24'/E150°48.50', 03. VIII. 2000, leg. D.J. Bennett. 4♀. Shikotan Island, Krabozavodskoe Village; N43°50.10'/E146°45.23', 11.–18. IX. 1997, leg. Y. Marusik. 1♀. Shiashkotan, SW shore of Delfin Bay; N43°44.90'/E146°36.41', 13. IX. 1997, leg. Y. Marusik. 1♂. This common and very widespread species exhibits a large range of variation both in color and ultrasculpture which has led to the proposition of several subspecies (Dasch, 1964a). Most of the specimens from the Kuril islands have yellow coxae, rather extensive yellow coloration on the face and epicnemium, and a smooth mesoscutum; they would thus be associated with the subspecies S. cognata albicoxa (Thomson) or S. cognata faceata Dasch. However, some specimens have the base of the hind coxae dark. Nakanishi (1979) discussed the color variation of this species in Japan and concluded that no clear separations can currently be drawn between the subspecies. Furthermore, proposing subspecies that occur in sympatry, as is the case in most of the range of S. cognata, is in conflict with a biological species concept (Mayr, 1942); the color variation in S. cognata is thus likely to represent phenotypic variation only and is not indicative of separate evolutionary lineages. Holarctic and Oriental; recorded from Russia and Japan. Sussaba dorsalis (Holmgren) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. T.R. Anderson. 1♀. Urup Island, inland of Smuglyi Bay; N46°01.46'/E149°59.02', 24. VIII. 1995, leg. N. Minakawa. 1♂. Urup Island, inland from Aleutka Bay; N45°56.10'/E150°09.70', 07. VIII. 2000, leg. T.R. Anderson. 1♀, 2♂. Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 1♂. Holarctic; recorded from Russia, including the Eastern provinces. Sussaba mongolica Klopfstein Figure 4A: Habitus. Russia, Kuril islands: Simushir Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.84'/E156°05.80', 30. VII. 1999, leg. B.K. Urbain. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.67'/E156°06.84', 30. VII. 1999, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.54'/E156°05.71', 30. VII. 1999, leg. B.K. Urbain. 1♀, 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29.43'/E156°05.52', 30. VII. 1999, leg. D.J. Bennett. 1♂. Paramushir Island, inland of Rifovaya Bay; N50°29'/E156°06', 30. VII. 1999, leg. A.S. Lelej. 2♀, 1♂. Matua Island, inland of Dvoinaya Bay; N48°04.37'/E153°15.98', 31. VII. 2000, leg. T.R. Anderson. 1♂. Matua Island, inland of Dvoinaya Bay; N48°04.15'/E153°15.70', 03. VIII. 1999, leg. D.J. Bennett. 1♂. Simushir Island, inland of NE Broutona Bay; N47°06.93'/E152°15.92', 02. VIII. 2000, leg. D.J. Bennett. 1♂. This species can be distinguished from the Japanese Sussaba nigra Nakanishi by the bi-colored tyloids in the male and from Sussaba dajsetsuzana Nakanishi by the smooth and punctured mesoscutum and the shorter and more stout petiole. Described from Mongolia (Klopfstein, 2011). Sussaba pulchella (Holmgren) Russia, Kuril islands: Simushir Island, inland of NE Broutona Bay; N47°07.20'/E152°16.02', 02. VIII. 2000, leg. D.J. Bennett. 1♂. Kunashir Island, 1km E of Tyatina R. Mouth; N44°16.32'/E146°10.34', 28. VIII. 1997, leg. Y. Marusik. 1♂. Kunashir Island, environs of Lake Aliger; N44°02.83'/E145°44.47', 09. IX. 1995, leg. P. Oberg. 1♀. Kunashir Island, inland of Aalekhina Bay; N43°55.23'/E145°32.04', 19. VIII. 1999, leg. D.J. Bennett. 1♂. Shikotan Island, inland from Del'Fin Bay; N43°45.11'/E146°37.41', 15. VIII. 1998, leg. K. Kavun. 1♂. Kunashir Island, between 1st and 2nd km of Golovino highway; N/E, 01. IX. 1997, leg. Y. Marusik. 1♂. Holarctic and Oriental; recorded from Russia and Japan. Sussaba sugiharai (Uchida) (Fig. 4B) Russia, Kuril islands: Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 3♂. Shikotan Island, inland of Zvezdnaya Bay; N43°46.34'/E146°36.36', 16. VIII. 1998, leg. D.J. Bennett. 1♂. Tanfilyeva, inland Tanfil'Yevka Bay; N43°26.79'/E145°55.32', 19. VIII. 1998, leg. B.K. Urbain. 1♂.
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Tanfilyeva, inland Tanfil'Yevka Bay; N43°26.62'/E145°55.89', 19. VIII. 1998, leg. D.J. Bennett. 1♀, 3♂. Kunashir Island, between 1st and 2nd km of Golovino highway; N/E, 01. IX. 1997, leg. Y. Marusik. 2♀. Eastern Palaearctic and Oriental; recorded from the Russian Far East and Japan.
FIGURE 4. Habitus of a female Sussaba mongolica (A) and a female Sussaba sugiharai (B).
Syrphoctonus tarsatorius (Panzer) Russia, Kuril islands: Paramushir Island, inland of Severo-Kurilsk; N50°40.71'/E156°05.31', 05. VIII. 1997, leg. J. Schweikert. 1♀. Iturup Island, inland of Medvezh'ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. B.K. Urbain. 1♀. Iturup Island, Atsonupuri Peninsula base; N44°46.31'/E147°11.32', 15. VIII. 1999, leg. B.K. Urbain. 1♂. Iturup Island, Atsonupuri Peninsula base; N44°46.01'/E147°11.58', 14. VIII. 1999, leg. D.J. Bennett. 1♂. Kunashir Island, Environs "Goby" Hot Springs; N44°00.34'/E147°40.99', 27. VII. 1997, leg. N. Minakawa. 1♀. Holarctic and Oriental; recorded from Russia and Japan. Syrphophilus bizonarius (Gravenhorst) Russia, Kuril islands: Paramushir Island, inland E. Vasilyeva Bay; N50°01.84'/E155°23.88', 25. VII. 2000, leg. D.J. Bennett. 1♀. Simushir Island, inland of Dushnaya Bay; N47°04.49'/E152°11.52', 09. VIII. 1999, leg. D.J. Bennett. 1♂. Iturup Island, inland Medvezh'Ya Bay; N45°26.70'/E148°49.68', 05. VIII. 1998, leg. D.J. Bennett. 1♀. Iturup Island, inland Medvezh'Ya Bay; N45°25.75'/E148°49.75', 05. VIII. 1998, leg. B.K. Urbain. 1♀. Iturup Island, inland Konservnaya Bay; N45°19.79'/E147°59.75', 19. VIII. 1996, leg. P. Oberg. 1♀. Iturup Island, inland of Kuybyshevskiy Bay; N45°05.12'/E147°40.26', 13. VIII. 1999, leg. D.J. Bennett. 1♀. Iturup Island, Atsonupuri Peninsula base; N44°46.31'/E147°11.32', 15. VIII. 1999, leg. B.K. Urbain. 1♀. Kunashir Island, BtwLks Aliger Lagunnoye; N44°03.25'/E145°44.91', 11. VIII. 1998, leg. B.K. Urbain. 1♂. Kunashir Island, Near Lake Serebryanoye; N44°03.07'/E145°49.25', 22. VIII. 1998, leg. B.K. Urbain. 1♂. Holarctic and Oriental; recorded from Russia and Japan. Syrphophilus scabriculus (Holmgren) Russia, Kuril islands: Onekotan Island, inland of Nemo Bay; N49°36.24'/E154°49.81', 24. VII. 1999, leg. D.J. Bennett. 1♂. Shiashkotan, inland of SW Zakatnaya Bay; N48°46.77'/E154°02.01', 29. VII. 2000, leg. D.J. Bennett. 1♀. Holarctic; recorded from Russia.
Tymmophorus gelidus Dasch(Fig. 5A) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 1♀. Although very similar in coloration to the widespread Tymmophorus obscuripes (Holmgren), this Northern species is distinguished by the short antennae, two strongly coriaceous bands starting at the notauli and continuing towards the scutellum, and some additional characters (see Dasch, 1964a). This species has been described from
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Canada and Greenland and has recently been reported from Northern Europe (Klopfstein, in press). Its presence on the northern Kuril island of Paramushir suggests a circumpolar distribution. Holarctic; new for Russia and the Eastern Palaearctic.
FIGURE 5. Habitus of a female Tymmophorus gelidus (A) and a female Tymmophorus obscuripes (B).
Tymmophorus obscuripes (Holmgren) (Fig. 5B) Russia, Kuril islands: Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. T.R. Anderson. 1♀. Holarctic; known from Russia. Tymmophorus suspiciosus (Brischke) Russia, Kuril islands: Paramushir Island, Northeast corner of island; N50°43.92'/E156°08.24', 04. VIII. 1997, leg. T. I. Ritchie. 2♂. Shumshu Island, inland of Baykovo Village; N50°42.97'/E156°12.29', 24. VII. 2000, leg. T.R. Anderson. 2♂; same data, but leg. D.J. Bennett: 1♂. Urup Island, inland from Aleutka Bay; N45°56.10'/ E150°09.70', 07. VIII. 2000, leg. T.R. Anderson. 3♂. Kunashir Island, Near Lake Allger; N44°02.88'/E145°44.48', 11. VIII. 1998, leg. D.J. Bennett. 2♀, 2♂. Palaearctic; known from Russia. Because this species has only recently been removed from synonymy with Tymmophorus erythrozonus (Förster) (which was often referred to under the name Tymmophorus rufiventris (Gravenhorst), which is a primary homonym (Klopfstein, in press)), it is not possible currently to associate the distributional records to either of them. The current distribution is thus only based on the records made under the name T. suspiciosus.
Discussion The 26 species of diplazontine wasps are probably a good sample of the diversity present on the Kuril islands, even though additional species might turn up when more sampling is done especially on the larger islands. For comparison, 33 species have been recorded from the Russian province Sakhalin Oblast to which the Kuril islands belong (Manukyan, 2007), 37 from Russian Kamchatka Oblast and 45 species from Japan (Yu et al., 2012). Except for Diplazon urupensis and D. varicoxa that were reported from the Kuril islands in a treatment of a limited sample of ichneumonids (Uchida, 1935), these are the first records for the Kuril islands. For one species, Tymmophorus gelidus Dasch, the present record fills a gap in the hitherto known distribution, i.e., Canada and Northern Europe. Given its occurrence on the Kuril islands, it is likely that it will be found in the Northern parts of the Eastern Palaearctic as well.
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In terms of distributional patterns, the proportion of very widespread species in this sample is striking. Only four of the species have been recorded exclusively from the Eastern Palaearctic, of which D. urupensis might still turn out to be a synonym of a Holarctic species (see note under this species). Sussaba sugiharai has an Eastern Palaearctic and Oriental distribution, and four species are Palaearctic. The remaining 17 species (65%) are Holarctic or even multiregional. This proportion is much higher than the 38% recorded by Manukyan (1995), a discrepancy that is probably caused by a combination of several factors. First, progress in diplazontine systematics has exposed several new synonyms between Nearctic and Palaearctic species since Manukyan's analysis (e.g., Klopfstein, in press). Second, if there is a positive relationship between population density and range, then the limited sample analyzed here might have overlooked some rare species with a restricted distributional range. And third, islands are areas where species with more effective dispersal abilities and thus wider distributions are overrepresented. Future accumulation of taxonomic and distributional data for Diplazontinae will help disentangling the reasons for the high proportion of widely distributed taxa on the Kuril islands, and host record data will help investigating the proximate reasons for the comparatively large distribution ranges.
Acknowledgements I would like to thank Prof. Robert Wharton for inspiring this study and making the material accessible. John Jennings helped improve an earlier version of this manuscript. This work was supported by the Swiss National Science Foundation grants PBBEP3_130173 and PA00P3_145377 to S. Klopfstein.
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