Heteroptera: Miridae: Mirinae), with Description of a New Species from Japan Tomohide Yasunaga & Michael D

Tijdschrift voor Entomologie 159 (2016) 197–207

Review of the mirine plant bug genus Pachylygus (Heteroptera: Miridae: Mirinae), with description of a new species from Japan Tomohide Yasunaga & Michael D. Schwartz

The little-known plant bug genus Pachylygus Yasunaga, 1994, now considered nearly endemic to Japan proper and to include five monophagous species, is reviewed, diagnosed, and discussed. A new species, P. luteonotatus, is described on the basis of specimens recently collected from Nikko, central Honshu. All known congeners are diagnosed and an identification key to them is included. A checklist, with known distributions, is also provided. The Asian genus Peltidolygus Poppius, 1915 is proposed as the most closely related sister taxon of Pachylygus. Keywords Heteroptera; Miridae; Mirinae; Pachylygus; new species; Japan; hosts Tomohide Yasunaga*, Research Associate, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA. [email protected] Michael D. Schwartz, Visiting Scientist, Agriculture & Agri-Food Canada, 960 Carling Avenue, Ottawa, Ontario K1A 0C6, Canada. [email protected]

Introduction The present work reviews, diagnoses, and dis- The mirine plant bug genus Pachylygus (Mirinae: cusses Pachylygus, which is considered to be sister Mirini) was proposed by Yasunaga (1994), to accom- to Peltidolygus Poppius, 1915 on the basis of close modate three Japanese species described by Kerzhner examinations of the morphological characters and (1977) in Orthops Fieber, 1858. Subsequently, Yasu- available biological information. A new species, naga et al. (2002) added a new species from Shikoku, Pachylygus luteonotatus, is described, and all known southwestern Japan proper, and since then Pachylygus congeners are also diagnosed and keyed, with color has been represented by the four members basically digital images of live individuals. A checklist, with endemic to Japan, except for P. nigrescens (Kerzhner), known distributions, is provided. which was recorded from central Korea by Seong et al. (2009), based on a female. Material and methods Recently, through the courtesy of an enthusiastic colleague, Mr. Satoshi Maehara, we were able to ex- The depositories of material examined are abbrevi- amine a pair of specimens of an undescribed mirine ated in the text as follows: that can be accommodated within Pachylygus. This mirid was collected from Kalopanax septemlobus AMNH: American Museum of Natural History, (Thunb.) Koidz. (Araliaceae) [as K. pictus (Thunb.) New York, USA Nakai, S. Maehara, pers. comm.]; thus, reconfirm- NIAES: National Institute of Agro-Environmental ing that the species of the genus are monophagous Sciences, Tsukuba, Japan as stated below. SMCT: S. Maehara Collection, Tochigi, Japan

Tijdschrift voor Entomologie 159: 197–207, Figs 1–33. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 30 December 2016. DOI 10.1163/22119434-15903001 *Corresponding author Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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TYCN: T. Yasunaga Collection, Nagasaki, Japan characters: moderate size (total length 4.5–6.5 mm); ZMAS: Z oological Institute, Russian Academy of castaneous to dark basic coloration; silvery, reclining Sciences, St Petersburg. setae on dorsum; clearly punctate pronotum (except for P. japonicus, punctures shallow and sparse); Matrix code labels were attached to several type or tumid or arched scutellum; triangularly developed representative specimens used in this work; the la- sensory lobe of left paramere (Figs 13, 17, 20); en- bels uniquely identify each specimen, and are referred dosoma always with a frontal sclerotized plate (FSC) to as ‘unique specimen identifiers’ (USIs). The USI covering secondary gonopore, spinulate lateral scler- codes (e.g., AMNH_PBI 000123) comprise an insti- ite (LSC), and a single horn-like spiculum (SPL) (see tution and project code (AMNH_PBI) and a unique Figs 15, 18, 21, 27); narrow lateral lobe and moder- number (000123). These data were digitized on the ate interramal lobe on posterior wall (Figs 26, 31); Arthropod Easy Capture (formerly the Planetary and oval, thick-rimmed, laterally rectangular female Biodiversity Inventory) database maintained by the sclerotized ring (Figs 25, 28–30, 32, 33). AMNH (http://research.amnh.org/pbi/) and are also Distribution. Japan proper (two species also searchable on the ‘Heteroptera Species Pages’ (http:// known from Rishiri and Kunashiri Islands), Korea research.amnh.org/pbi/heteropteraspeciespage/). (P. nigrescens). All measurements are in millimeters (mm). Termi- Biology. Four of the five known members of Pachy- nology of the genitalia principally follows Schwartz lygus are associated with a deciduous broadleaf, Ka- & Foottit (1998) and Yasunaga & Schwartz (2007). lopanax septemlobus (Araliaceae); only P. japonicus In the synonymic lists for known taxa only selected is known to inhabit shrubs of a Japanese wild rose, references are cited, as comprehensive catalogs cover- Rosa multiflora Thunb. (Rosaceae). All species are as- ing the necessary literature are now available (Schuh sumed to have a univoltine life cycle and to overwin- 1995, 2002–2014, Kerzhner & Josifov 1999). Plant ter in the egg stage. names and authors are reported according to Tropi- Discussion. By virtue of its endemism to Japan, cos.org (Missouri Botanical Garden: http://www. Pachylygus may be regarded as a little known mir- tropicos.org). ine genus. Although previous work (Yasunaga 1994, 2001, Yasunaga et al. 2002) fragmentarily docu- mented this unique taxon, the systematic position Checklist of Pachylygus species, with of Pachylygus is insufficiently elucidated. During a known distributions closer reevaluation of characters, this genus is now presumed to be sister to Peltidolygus Poppius, a Pachylygus Yasunaga, 1994 small Asian genus including two Oriental elements P. anthrax Yasunaga, Schwartz et Chérot, 2002 — inhabiting warmer climates: P. puncticollis Poppius Japan (central highlands of Shikoku). from Sikkim, India and P. scutellatus (Yasunaga & P. festivus (Kerzhner, 1977) — Japan (Hokkaido, Lu) (Figs 7, 8) from SE China, Korean Jeju Island, Honshu, Shikoku [new record], Kunashiri & and SW Japan, including Tsushima and Ryukyu Is- Rishiri Islands). lands (Yasunaga et al. 2002). Pachylygus can be dis- P. japonicus (Kerzhner, 1977) — Japan (Honshu, tinguished from Peltidolygus externally by the almost Shikoku, Kyushu). impunctate or sometimes matte hemelytron with P. luteonotatus Yasunaga & Schwartz, n. sp.— uniformly or sparsely distributed, silvery vestiture; Japan (Central Honshu: Tochigi Pref.). in Peltidolygus, the hemelytron is shallowly but dis- P. nigrescens (Kerzhner, 1977) — Japan (Hok- tinctly punctate and with silvery vestiture arranged kaido, Honshu, Kunashiri & Rishiri Islands), in patches (Fig. 7). Korea (Chungcheongbuk-do). Both genera share the following characters: pronotum deeply and clearly punctate; visible silvery setae present on mesoscutum, base of scutellum, Taxonomy and hemelytron; and noticeably developed, shiny Genus Pachylygus Yasunaga hemelytron. In the male genitalia of Peltidolygus Pachylygus Yasunaga, 1994: 124 (n. gen.), type spe- (Figs 22−24), which are much smaller than those cies by original designation: Orthops japonicus Kerzh- of Pachylygus, the parameres are basically similar in ner, 1977; Schwartz 1994: 978 (discussion); Schuh shape (22, 23), and the endosoma (Fig. 24) also has 1995: 860 (cat.); Kerzhner & Josifov 1999 (cat.); a single spiculum and a conspicuously developed lat- Yasunaga 2001: 253 (diag.); Yasunaga et al. 2002: 9 eral sclerite but completely lacks the frontal sclero- (diag.); Seong et al. 2009: 2 (diag.). tized plate (FSC). The female genitalia of Peltidolygus are more simple and small sized. Nonetheless, based Diagnosis. Distinguished from other genera of on sharing several similar characters, such as in the the tribe Mirini by the following combination of pronotum, scutellum, hemelytron, parameres, and Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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Figs 1–8. Pachylygus spp. (1–6) and Peltidolygus scutellatus (7, 8), habitus images of live individuals. — 1, Pa. nigrescens, male (Rishiri Is., Hokkaido); 2, Pa. festivus, male (Rishiri Is.); 3, Pa. anthrax, male (Kochi Pref.); 4, Pa. japonicus, female (below) and male (Kumamoto Pref.); 5, ditto, male; 6, Pa. luteonotatus n. sp., holotype male (Tochigi Pref., photo courtesy of S. Maehara); 7, Pe. scutellatus, female (Nagasaki Pref.); 8, ditto, final (5th) instar nymph (Okinawa Is.).

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200 Tijdschrift voor Entomologie, volume 159, 2016 endosoma, Peltidolygus is considered most closely Pachylygus anthrax Yasunaga, Schwartz & Chérot, related to Pachylygus among other mirine genera 2002: 10 (n. sp.); Yasunaga & Yamada, 2014: 32 known from the eastern Palearctic and the Orien- (diag.). tal regions. The final-instar nymph of Peltidolygus scutellatus (Fig. 8) is similar in coloration, size and Diagnosis. Recognized by its comparatively small shape to Pachylygus nigrescens; however, the dorsum size (total body length 4.5−5.1 mm); almost wholly of the former is almost glabrous (cf. below, diagnosis fuscous body; and fuscous scutellum without pale for P. nigrescens). midline or apex. See Yasunaga et al. (2002) for There is a strong likelihood that Peltidolygus detailed description. scutellatus is polyphagous, as the adults (including a Biology. This species is associated with Kalopanax teneral individual) have been found on inflorescenc- pictus (Thunb.) Nakai (Araliaceae), but its distribu- es of the broadleaf evergreen trees, Meliosma arnotti- tion is currently restricted to central montane areas ana (Wight) Walp. (Sabiaceae), Schefflera heptaphylla of Shikoku. A univoltine life cycle is assumed for this (L.) Frodin (Araliaceae), and Fraxinus griffithii mirid. C.B.Clarke and Ligustrum japonicum Thunb. Material examined. Holotype ♂, Japan, Shikoku, (Oleaceae). Final-instar nymphs were discovered on Kochi Pref., Hongawa Vil., Teragawa, 1300–1400 m, Ligustrum japonicum and Schefflera heptaphylla flow- on Kalopanax pictus [= K. septemlobus], 24 Jul 1996, ers (Yasunaga & Takai, unpublished data), which T. Yasunaga & M. Takai (AMNH_PBI 00380415) may imply a close relationship with Pachylygus, as (TYCN). Paratypes: same data as for holotype, one of the breeding hosts belongs to the Araliaceae. 1♂ 1♀ (AMNH_PBI00380416−00380417) & On the other hand, all members of Pachylygus appear 3♂ 3♀ (TYCN); Ehime Pref.: Odamiyama, 11 Jul strictly monophagous. 1994, M. Takai, 1♂ (TYCN); Kochi Pref., Honga- Among the five currently known members of wa, Kamegamori, 10 Jul 1993, T. Befu, 1♂ (TYCN). Pachylygus, four species are associated with Ka- lopanax septemlobus, but only P. japonicus is a special- Pachylygus festivus (Kerzhner, 1977) ist of Rosa multiflora. Some structural modifications Figs 2, 29 are exhibited in this mirine, such as the weakly and Orthops festivus Kerzhner, 1977: 14 (n. sp.) sparsely punctate pronotum (Figs 4, 5) and the wide Pinalitus festivus: Kerzhner, 1988a: 70 (n. comb.); sclerotized ring with a sharp medial angle (Fig. 30). 1988b: 813, fig. 520 (key). Pachylygus festivus: Yasunaga 1994: 125 (n. comb.); Key to species of Pachylygus Yasunaga 2001: 253, fig. 286a, b (diag., habitus 1. Punctures on pronotum shallow and sparsely images). distributed (Figs 4, 5); inhabiting Rosa multiflora �� japonicus Diagnosis. Recognized by its small size (4.5–5.5 – Punctures on pronotum deep and densely dis- mm); castaneous basic coloration (Fig. 2); widely tributed (Figs 1–3, 9, 11); associated only with pale (or yellowish brown) head and mesal part of Kalopanax septemlobus �� 2 pronotum; a yellowish, mesal (usually continu- 2. Scutellum totally yellow; hemelytron fuscous, ous) stripe on fuscous scutellum; almost entirely contrastingly with two pairs of yellow marks on yellow legs; and rather ovoid female sclerotized anterior areas of corium and cuneus (Figs 9, 11) rings (Fig. 29). Often collected simultaneously with �� luteonotatus n. sp. P. nigrescens ; easily distinguished by the small size – Dorsum lacking conspicuous yellow marks �� 3 and paler general coloration. 3. All femora yellow; metafemur apically with two Biology. This mirid is associated only with Kalopanax reddish brown rings (Fig. 2) ��festivus septemlobus, and sometimes co-occurs with the as- – All femora dark brown to fuscous, usually with sumed sister taxon, P. nigrescens. two pale rings apically �� 4 Material examined. Holotype ♂: Japan, Kunashiri 4. Body smaller, less than 5.1 mm in total length; Island, Golovnina Volcano [= Mt. Tomari], 24 Jul dorsum almost entirely black (Fig. 3) �� anthrax 1973, I.M. Kerzhner (without USI, ZMAS); para- – Body larger, more than 5.5 mm in total type: same data as for holotype, 1♂ (AMNH_PBI length; dorsum largely fuscous but with apex 00380418) (TYCN). Additional material: Japan, of scutellum pale brown, and clavus and ante- Hokkaido, Kamikawa Subpref., Mts Taisetsu, Mt rior area of corium more or less brown (Fig. 1) Asahi-dake (500−800 m), 43.6458 N 142.7387 E, on ��nigrescens Kalopanax pictus [= K. septemlobus], 11 Jul 1994, T. Yasunaga, 1♂ (AMNH_PBI 00380419); Hokkaido, Pachylygus anthrax Yasunaga, Schwartz & Soya Subpref., Rishiri Island, Oniwaki, 45.1409 N Chérot 141.2913 E, on Kalopanax pictus, 28−29 Jul 1994, Figs 3, 28 T. Yasunaga, 2♂ 5♀ (TYCN); Hokkaido, Abashiri Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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Figs 9–12. Habitus images of Pachylygus luteonotatus n. sp. — 9, 10, holotype male (AMNH_PBI 00380413); 11, 12, paratype female (00380414).

Subpref., Shari Town, Mt. On’nebetsu, on Kalopanax 7♂ 7♀ (TYCN); Honshu, Aomori Pref., Towadako pictus [= K. septemlobus], 30 Aug–1 Sep 1995, T. Town, Yakeyama, 25 Jul 1987, T. Ichita, 1♂ (TYCN); Yasunaga, 2♂ 2♀ (TYCN); Hokkaido, Iburi Sub- Aomori Pref., Hiraka Town, G unmatai, 22 Aug pref., Tomakomai City, Takaoka, on Kalopanax pic- 1990, T. Ichita, 1♂ (TYCN); Shikoku, Ehime Pref., tus [= K. septemlobus], 29−31 Jul 1997, T. Yasunaga, Odamiyama, 11 Jun 1994, M. Takai, 1♀ (TYCN). Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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Figs 13–15. Male genitalia of Pachylygus luteonotatus n. sp., holotype, structures and terms in figures.

Pachylygus japonicus (Kerzhner) (Figs 4, 5); weakly and shallowly punctate prono- Figs 4, 5, 16–18, 30, 31 tum; yellow cuneus with a darkened base and apex; broadened base of endosomal spiculum (Fig. 18); Orthops japonicus Kerzhner, 1977 (n. sp.). and widened female sclerotized ring, with sharp in- Pinalitus festivus: Kerzhner 1988a: 70 (n. comb.); ner corner (Fig. 30). This species could be confused Yasunaga et al. 1993: 157, figs. 54a, b (diag., with some species of Apolygus China, because of habitus images). shiny dorsum and reduced pronotal punctures. It Pachylygus japonicus: Yasunaga 1994 (n. comb., is, however, distinguished by the noticeably tumid diag.); Yasunaga 2001: 254, fig. 287 (diag., habi- scutellum. tus image). Biology. This monophagous species is associated only with Rosa multiflora, instead of Kalopanax Diagnosis. Recognized by its moderate size (5.6–6.3 septemlobus. The adults appear from mid-May to mm); brown to purplish brown general coloration mid-June, and often aggregate on newly developed Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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Figs 16–24. Male genitalia of Pachylygus japonicus spp. (16–18), Pa. nigrescens spp. (19–21) and Peltidolygus scutellatus (22–24). — 16, 19, 22, right paramere; 17, 20, 23, left paramere; 18, 21, 24, endosoma. Scale bars 0.2 mm. Abbreviations: ASC, apical sclerite; FSC, frontal sclerite covering secondary gonopore; LSC, lateral sclerite; SMD, seminal duct; SPL, spiculum. leaves and stems of the host, which are sometimes Yamanashi Pref., Mt. Fuji, Shoji, 35.48 N 138.63 severely damaged by this mirid (Yasunaga 2001). E, 21 Jun 1984, M. Hayashi, 1♂ (AMNH_PBI Material examined. Holotype ♂, Japan, Tokyo, 00380422) (TYCN); Kyushu, Kumamoto Pref., Mt. Takao, 21 May 1930 (without USI, ZMAS). Izumi Village, Momiki, 32.502319 N 130.957720 Paratype: same data as for holotype, 1♀ (without E, on Rosa multiflora, 1 Jun 1989, T. Yasunaga, 1♂ USI, ZMAS). Additional material: Japan, Honshu, (AMNH_PBI 00380423) & 10♂ 10♀ (TYCN). Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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Figs 25–27. Female (25, 26) and male (27) genitalia of Pachylygus luteonotatus n. sp. — 25, bursa copulatrix; 26, posterior wall; 27, endosoma. Abbreviations: ASC, apical sclerite; FSC, frontal sclerite covering secondary gonopore; LSC, lateral sclerite; SMD, seminal duct; SPL, spiculum.

Pachylygus luteonotatus n. sp. and two pairs of yellow spots on hemelytron; dor- Figs 6, 9–15, 25–27 sal surface with sparsely distributed, brown, simple setae and silvery reclining setae. Head shiny dark Type material. Holotype ♂. Japan, Honshu, brown, vertical; vertex and frons narrowly yellow Tochigi Pref., Nikko City, Shobugahama, 36.75 N laterally. Antenna yellowish brown, slender, almost 139.45 E, N shore of Chuzenji Lake, 13 Jul 2009, linear; segment II with slightly darkened apex, lon- S. Maehara (AMNH_PBI 00380413) (AMNH). ger (♂)/ shorter (♀) than basal width of pronotum; Paratypes: Japan, 1♀, same data as for the holotype segments III and IV brown, filiform. Labium shiny (AMNH_PBI 00380414) (TYCN); 1♂, same data, reddish brown, short, reaching middle of meso- except for date, 6 Jul 2011 (SMCT) and 5♂ 1♀, 11 coxa; apical half of segment IV darkened. Pronotum Jul 2016 (SMTC, TYCN). dark brown, shining, clearly punctate, with yellow Diagnosis. Recognized readily by its conspicuous, mesal part and posterior margin, in ♀ additionally two-tone coloration — fuscous body clearly with with a pair of yellow spots posterior to calli; scutel- yellow marks as in Figs 6, 9, 11; no other described lum shiny yellow, narrowly darkened anteromesally; congener exhibits such a color pattern. collar yellow; pleura widely grayish brown, matte; Description. Body ovate, comparatively small, gen- propleuron shiny dark brown with yellow ventral erally fuscous, contrastingly with yellow scutellum 1/3; ostiolar peritreme yellow. Hemelytron fuscous,

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Figs 28–33. Female genitalia of Pachylygus spp., structures and terms in figures. — 28–30, 32, 33, bursa copulatrix; 31, posterior wall. weakly shining, with two pairs of noticeable yellow 0.59, 1.81, 0.95, 0.56/0.57, 1.45, 0.91, 0.76; length marks on anterior corium to exocorium and cuneus of labium 1.18/1.25; pronotal width 1.71/1.73; to apical margin of corium; apical part of clavus pale maximum width across hemelytron 2.28/2.47; and except for dark apex; apex of cuneus darkened; mem- lengths of metafemur, tibia and tarsus 1.67, 2.43, brane including veins smoky brown, with a pale, 0.53/1.62, 2.32, 0.57. semitransparent spot posterior to apex of cuneus. All Etymology. From Latin luteus (= yellow) combined coxae and legs uniformly yellow. Abdomen black- with notatus (= marked), referring to the noticeable ish brown. Male genitalia (Figs 13−15, 27): Basal yellow spots of this new species; an adjective. protuberance of left paramere narrow, rather sharp. Biology. This new species was collected from leaves Endosoma with developed FSC and an additional, of Kalopanax septemlobus in July. crest-like, apical sclerite, or ASC; lateral sclerite and spiculum small. Female genitalia (Figs 25, 26): Bursa Pachylygus nigrescens (Kerzhner) copulatrix similar in general shape and size to that Figs 1, 19–21, 32, 33 of P. festivus; sclerotized ring elongate-ovoid; posterior wall with rather wide lateral lobe and laterally Orthops nigrescens Kerzhner, 1977: 14 (n. sp.). rounded interramal lobe. Pinalitus nigrescens: Kerzhner 1988a: 70 (n. comb.); Measurements. ♂/♀: Total body length 4.8/5.0; 1988b: 813, fig. 520 (key). width of head across eyes 1.00/0.99; width of ver- Pachylygus nigrescens: Yasunaga 1994: 126 (n. tex 0.40/0.44; lengths of antennal segments I–IV: comb., diag.); Yasunaga 2001: 254, fig. 288a–c

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(diag., habitus images, incl. last-instar nymph); Laboratory, ARS-USDA, c/o National Museum of Seong et al. 2009: 2 (n. rec. from Korea). Natural History, Smithsonian Institution, Wash- ington, DC, USA) and Dr D. Polhemus (associate Diagnosis. Recognized by its large size (5.7−6.6 editor, TVE) for reviewing the manuscript. mm, usually more than 6 mm); generally dark coffee brown coloration; more or less pale mesial pronotum and scutellum, and anterior parts of corium References and clavus; narrowly pale anterior margin of cune- Kerzhner, I.M., 1977. Novye i maloizvestnye vidy polu- us; widely darkened femora; and broadened female zhestkokryrykh (Heteroptera) s Dal’nego Vostoka bursa copulatrix with thickened, laterally squared SSSR [New and little known species of Heteroptera sclerotized ring. A detailed redescription was pro- from the Far East of the USSR]. — Trudy Zoologitch- vided by Yasunaga (1994). The final-instar nymph seskogo Instituta Akademiya Nauk SSSR Leningrad, never has any dark pattern, and is recognized by the 62[1976]: 6–35. [In Russian] wholly light green, ovoid, tumid body, uniformly Kerzhner, I.M., 1988a. Novye i maloizvestnye poluzhest- distributed, dark, short vestiture on the dorsum, and kokrylye Nasekomye s Dal’nego Vostoka SSSR [New and little known heteropterous insects from the Soviet yellowish green antennae and legs (see Yasunaga, Far East]. — Akademiya Nauk SSSR [1987], Vladivo- 2001, fig. 288c). stok, USSR, 83 pp. [In Russian] Biology. This mirid is associated with Kalopanax pic- Kerzhner, I.M., 1988b. Sem. Miridae (Capsidae)— tus, and the late instars and adults are found from Slepnjaki. Opredelitel’ nasekomykh Dal’nego Vosto- mid-July to early August. In eastern Hokkaido, how- ka SSSR [Keys to the insects of the Soviet Far East], ever, the adults emerge from late August to early Sep- Vol. 2. — Nauka, Leningrad, USSR, pp. 778–857. tember, due to foggy weather and low temperatures [In Russian] between June and August, which may significantly Kerzhner, I.M. & M. Josifov, 1999. Miridae Hahn, 1833. delay the mirids’ development. — In: B. Aukema & C. Rieger (Eds), Catalogue of the Material examined. Holotype ♂: Japan, Ku- Heteroptera of the Palaearctic Region, Vol. 3, Cimico- nashiri Island, Golovnina Volcano [= Mt. Tomari], morpha II, pp. 1–576. The Netherlands Entomological 26 Jul 1973, I.M. Kerzhner (ZMAS). Paratypes: Society, Amsterdam, Netherlands. same data as for holotype except for date 24 Jul Schuh, R.T., 1995. Plant bugs of the world (Insecta: 1973, 1♂ (AMNH_PBI 00380420) (TYCN). Ad- Heteroptera: Miridae). Systematic catalog, distribu- ditional material: Japan, Hokkaido, Soya Subpref., tions, host list and bibliography. — The New York En- Rishiri Island, Oniwaki, 45.1409 N 141.2913 E, tomological Society, New York, NY, USA, xii + 1329 pp. on Kalopanax pictus [= K. septemlobus], 28−29 Jul Schuh, R.T., 2002–2014. On-line Systematic Catalog 1994, T. Yasunaga, 1♂ (AMNH_PBI 00380421); of Plant Bugs (Insecta: Heteroptera: Miridae). — Hokkaido, Abashiri Subpref., Shari Town, Mt. Available from: http://research.amnh.org/pbi/catalog/. On’nebetsu, on Kalopanax pictus [= K. septemlobus], Schwartz, M.D., 1994. Review of the genus Salignus Kelton and a character discussion of related genera 30 Aug–1 Sep 1995, T. Yasunaga, 1♂ 1♀ (TYCN); (Heteroptera: Miridae: Mirinae). — The Canadian Hokkaido, Ishikari Subpref., Atsuta Village, Hattari, Entomologist 126: 971–993. on flower ofKalopanax pictus [= K. septemlobus], 7 Schwartz, M.D. & R.G. Foottit, 1998. Revision of the Aug 1998 (last instar nymph; emerging on 9 Aug), Nearctic species of the genus Lygus Hahn, with review 1♀ (TYCN); Honshu, Nagano Pref., Saku City, of the Palearctic species (Heteroptera: Miridae). — Uchiyama, UV light trap, 22 Jul 1996, S. Sakurai, Memoirs on Entomology International, Vol. 10., vii + 2♂ 2♀ (TYCN); Honshu, Nara Pref., Tenkawa 428 pp. Associated Publishers, Gainesville, FL, USA. Village, Gyojagaeri, 2 Aug 1996, Y. Nakatani, 1♂ Seong, J., G.S. Lee & S.H. Lee, 2009. Two new records of 3♀(NIAES). the family Miridae (Hemiptera: Heteroptera) in Korea. — Korean Journal of Applied Entomology 48: 1–4. Yasunaga, T., 1994. Pinalitus Kelton (Heteroptera, Miri- Acknowledgements dae) and its allied genera of Japan, with descriptions of Special thanks are due to the late Dr I.M. Kerzhner new genera and species. — Japanese Journal of Ento- mology 62: 115–131. (ZMAS) for generously permitting TY to examine Yasunaga, T., 2001. Family Miridae Hahn, plant bugs. – In: related types in ZMAS (in 1998 and 2002) and of- T. Yasunaga, M. Takai & T. Kawasawa (Eds), A Field fering other specimens necessary to this study. We Guide to Japanese Bugs, Vol. II, pp. 1–96, 111–351. are much indebted to the following individuals for Zenkoku Noson Kyoiku Kyokai Publishing Co. Ltd., providing specimens for this study: Dr M. Hayashi Tokyo, Japan. [In Japanese] (Nango, Chigasaki City, Kanagawa); Dr Y. Nakatani Yasunaga, T. & M.D. Schwartz, 2007. Revision of the mir- (NIAES); Mr S. Sakurai (Niigata); Mr S. Maehara ine plant bug genus Philostephanus Distant and allies (Tochigi); and Mr T. Ichita (Aomori). Thanks are Heteroptera (Miridae: Mirinae: Mirini). — Tijdschrift extended to Dr T.J. Henry (Systematic Entomology voor Entomologie 150: 100−180. Downloaded from Brill.com10/04/2021 03:47:57PM via free access

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Yasunaga, T. & K. Yamada, 2014. Japanese or scientific Yasunaga, T., M.D. Schwartz & F. Chérot, 2002. New names proposed for mirid and anthocorid species, genera, species, synonymies, and combinations in the recently described or recorded from Japan (Heterop- “Lygus complex” from Japan, with discussion on Pelti- tera, Miridae and Anthocoridae). — Rostria 57: 25– dolygus Poppius and Warrisia Carvalho (Heteroptera: 38. [In Japanese] Miridae: Mirinae). — American Museum Novitates Yasunaga, T., M. Takai, I. Yamashita, M. Kawamura & 3378: 1–26, 68 figs. T. Kawasawa, 1993. A Field Guide to Japanese Bugs. Terrestrial Heteropterans. — Zenkoku Noson Kyoiku Received: May 2, 2016 Kyokai, Tokyo, Japan, 380 pp. [In Japanese] Accepted: October 3, 2016

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