Tijdschrift voor Entomologie 158 (2015) 21–47

Review of the mirine plant bug genus Phytocoris Fallén in Japan (Hemiptera: Heteroptera: Miridae: Mirinae), with descriptions of eight new species Tomohide Yasunaga & Michael D. Schwartz

The mirine plant bug genus Phytocoris Fallén of Japan is reviewed; twelve species are recognized. Eight new species are proposed, diagnosed, and described: P. amateras, P. hasegawai, P. izanagii, P. izanamiae, P. kerzhnerianus, P. minakatai, P. miyamotoi, and P. protobothrops. Four known species are diagnosed, and two poorly recognized species, P. ohataensis Linnavuori and P. pallidicollis Kerzhner, are also redescribed. Phytocoris scotinus Kerzhner is proposed as a junior synonym of P. ohataensis. The male and female genitalic structures are figured and the biological information is documented for most treated species. A checklist and a key are provided to aid in the identifications of all Japanese species. Keywords: Heteroptera; Miridae; Mirinae; Phytocoris; new species; Japan; taxonomy; biology Tomohide Yasunaga*, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA; Plant Protection Division, Myanmar Ministry of Agriculture & Irrigation, c/o Japan International Cooperation Agency (JICA), Myanmar Office, 701 Sakura Tower, No. 339, Bogyoke Aung San Road, Kyauktada, Yangon. [email protected] Michael D. Schwartz, Agriculture & Agri-Food Canada Environmental Health, K.W. Neatby Building 20, Room 3121, 960 Carling Avenue, Central Experimental Farm, Ottawa, ON. Canada K1A 0C6. [email protected]

Introduction found in conifer/deciduous mixed forests of the Phytocoris Fallén, of the plant bug subfamily Mirinae, Nepalese Himalayas at more than 3,500 m altitude; is the largest genus in the Heteroptera, with more also in Taiwan, a few were collected only in decidu- than 500 described species worldwide (Schuh 1995, ous, montane forests. Therefore, Phytocoris species 2002–2014, Stonedahl 1988). About 80% of the are assumed not to inhabit tropical or subtropical species are known from the Holarctic Region, but climate zones where Adelphocorisella and Creontiades quite a few species placed in Phytocoris from the trop- are commonly present. ics or subtropics appear not to be true members (pos- Approximately 300 species are known to occur in sibly, representatives of Adelphocorisella Miyamoto & the Nearctic Region (Stonedahl 1988), and new Yasunaga, Creontiades Distant, or other related gen- species are still being described in the western era, which require revision). Palearctic Region, particularly from Mediterranean Previous Japanese academic projects in SE Asia areas. On the other hand, around 40 species have have collected numerous relevant mirine specimens been reported from eastern Asia, where Phytocoris which do not include any true member of Phytocoris. does not show significant species diversity (Kerzhner In subtropical Asia, two Phytocoris species were 1988, Yasunaga 2001, Zheng et al. 2004). In Japan,

Tijdschrift voor Entomologie 158: 21–47, Figs 1–102. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 20 October 2015. DOI 10.1163/22119434-15812048 *Corresponding author Downloaded from Brill.com10/06/2021 05:01:49PM via free access

22 Tijdschrift voor Entomologie, volume 158, 2015 only five species are currently known, but the pres- or http://www.discoverlife.org for additional infor- ence of more than a few undescribed taxa was pre- mation on specimens examined. dicted previously (Yasunaga 2001). All measurements are in millimeters. The syn- In our continuing efforts to classify Japanese spec- onymic lists for known taxa only provide selected imens of Phytocoris, twelve species were recognized references, as comprehensive catalogs are now avail- and eight of them are confirmed to be undescribed. able (Schuh 1995, 2002–2014 online catalog, In addition, P. scotinus Kerzhner is found to be con- Kerzhner & Josifov 1999, Aukema et al. 2013). We specific withP. ohataensis Linnavuori, although these refrain from using subgenera or species-groups for two species were considered distinct and distin- Japanese Phytocoris species, because the majority of guished from each other by obvious different color the subgenera were proposed mainly by Wagner patterns that are herein regarded as intraspecific (1954, 1968, 1974, etc.) for classification of the variations. western Palearctic faunas, and the Nearctic species- The present paper is organized to aid in the cor- groups (Stonedahl 1988) also are not satisfactorily rect identification of JapanesePhytocoris species, applicable to the eastern Palearctic congeners. clarifying undetermined or undescribed ones. Terminology of the genitalia mainly follows Cassis Eight species are described as new, and four known (2008), Stonedahl (1988) (male genitalia), and species are diagnosed and/or redescribed. Phytocoris Yasunaga & Schwartz (2007) (female genitalia). is diagnosed and discussed, and P. scotinus is syn- Some terms used in this paper for the male genitalia onymized with P. ohataensis, which is now con- are partly modified from Kerzhner (1988) and firmed to be distributed in the circumferential Stonedahl (1988). The following abbreviations are areas of the Sea of Japan. The male and female used in the text and figures to indicate the male and genitalic structures are figured and biological female genitalic structures properly: BSP: basal scler- information is documented for most treated otized process; Comb: comb-shaped lobal sclerite; species. A checklist of Japanese species with DLP: dorsal labiate plate; DV: dorsal view; LBS: known distributions and a key to these species are lobal sclerite; PML: primary lobe; RBL: right basal provided. lobe; SGP: secondary gonopore; SPGC: sclerotized perimeter of genital chamber; SR: sclerotized ring; VV: ventral view. Materials and methods About 1500 specimens (including those of exotic spe- cies deposited in the Natural History Museum, Taxonomy London, US National Museum of Natural History, Genus Phytocoris Fallén Washington, DC, and Zoological Museum, University Phytocoris Fallén, 1814: 10 (n. gen.). Type species: of Helsinki, Finland) were examined; most of them Cimex populi L., subsequent designation by were collected by the senior author and/or his col- Westwood, 1840; Schuh, 1995: 863 (cat.); leagues acknowledged below. The depositories of mate- Kerzhner & Josifov, 1999: 136 (cat.); rial examined are abbreviated in the text as follows: Yasunaga, 2001: 255 (diag.); Zheng et al., AMNH American Museum of Natural 2004: 462 (diag., key to Chinese spp.). History, New York, NY, USA NIAES National Institute of Agro- Diagnosis. In eastern Asia, Phytocoris is recognized by Environmental Sciences, Tsukuba, the following combination of characters: Body elon- Japan gate, subparallel-sided (except for rather ovoid bra- NSMT National Science Museum, Tokyo, chypterous female as in Fig. 14); dorsal coloration Japan more or less mottled (every individual usually exhibit- TKPM Tokushima Prefectural Museum, ing a unique pattern, cf. Figs 44–47, 50); dorsum Tokushima, Japan matte or weakly shining, with two or three types of TYCN T. Yasunaga Collection, Nagasaki, Japan vestiture; eye as a rule large; vertex without particular ZMAS Zoological Institute, Russian Academy structure (not sulcate or concaved); frons sometimes of Sciences, St Petersburg, Russia striolate; antenna long, more than body length, with Matrix code labels are attached to the holotype slender and linear segments; labium long, reaching ­specimens, which uniquely identify each specimen, or exceeding much beyond apex of metacoxa; leg and are referred to as ‘unique specimen identifiers’ more or less mottled and/or partly annulated; meta- (USIs). The USI codes (e.g., AMNH_PBI 012345) femur long, slightly to moderately flattened; parameres comprise an institution and project code (AMNH_ with indentation, notch or spines; endosoma with a PBI) and a unique number (012345). Please visit comb-shaped sclerite broadly fused with a membranous the ­website of the Planetary Biodiversity Inven­ sac, or primary lobe (PML) and sometimes with an tory (PBI) Project (http://research.amnh.org/pbi/), additional lobal sclerite (LBS); secondary gonopore Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 23

Figs 1–8. Japanese Phytocoris species, live individuals. – 1, P. amateras, male, from Shikoku; 2, ditto, female, from N. Honshu; 3, P. miyamotoi, male, from Hokkaido; 4, 5, ditto, male, from Shikoku; 6, P. protobothrops, male, from Amami-Oshima Island; 7, 8, ditto, female, from Okinawa Island.

rounded to crescent, thick-rimmed; sclerotized ring Distribution. Known from all zoogeographical elongate ovoid; and bursa copulatrix with usually con- regions, but records from tropical and subtropical tinuous, rounded sclerotized perimeter (SPGC). The climate zones need verification; most species occur- most detailed generic diagnosis and redescription are ring in temperate zones of the Holarctic Region. provided by Stonedahl (1988). Distinguished from Biology. Stonedahl (1988) indicated that many spe- its related genera, Adelphocoris, Adelphocorisella or cies of Phytocoris have nocturnal habits, as evidenced Creontiades, primarily by some characters above indi- by the large number of species that are attracted to cated by italicized texts. light at night. Specimens of all Japanese species Downloaded from Brill.com10/06/2021 05:01:49PM via free access

24 Tijdschrift voor Entomologie, volume 158, 2015

Figs 9–17. Japanese Phytocoris species, live individuals. – 9, P. izanagii, male from Honshu; 10, ditto, female, from Honshu; 11, P. izanamiae, male, from N. Honshu; 12, ditto, female, from N. Honshu; 13, P. nowickyi, male, from Hokkaido; 14, ditto, brachypterous female, from Hokkaido; 15, ditto, final instar nymph; 16,P. pallidicollis, final instar nymph, from Hokkaido; 17, P. longipennis, just emerged adult, from Hokkaido.

also were collected using UV light traps, and occa- Stonedahl, such nocturnal members may retreat to sionally hundreds of individuals (often including the base of the host plant or to nearby ground litter. multiple species) visit a light trap screen per night. Some species were observed to sit firmly on the Sweeping and beating methods in the daytime nev- bark or branches during the day in Hokkaido, ertheless have produced only a few specimens of Japan (e.g., P. ohataensis on conifer branches; P. lon- limited species (e.g., P. nowickyi on bush clover, gipennis on bark or branches of deciduous alder, wormwood or willow leaves and P. pallidicollis elm or oak). The mottled, brownish color pattern on Japanese lime, Tilia japonica). As assumed by of such bark inhabitants is cryptic, well matching Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 25

Figs 18–28. Japanese Phytocoris species, live individuals. – 18, 21, P. ohataensis, female, from Shikoku; 19, ditto, male, from Hokkaido; 20, ditto, female, from Hokkaido; 22, P. minakatai, male, from Kumano, Honshu; 23, P. kerzhnerianus, male, from Shikoku; 24, 25, P. longipennis, female, from Hokkaido; 26, P. pallidicollis, male, from Hokkaido; 27, 28, ditto, male, from Shikoku.

the surface of the bark or branch, and making the individuals are liable to fade rapidly to brownish bug nearly invisible (by human vision). In some after death. Kerzhner (1988) and Yasunaga (2001) species (e.g., P. ohataensis), every specimen has a briefly commented on the biology for several more or less different, unique mottled pattern (cf. known species occurring in Japan and the Russian Figs 18–21, 44–47, 50); vivid olive or light green Far East. Downloaded from Brill.com10/06/2021 05:01:49PM via free access

26 Tijdschrift voor Entomologie, volume 158, 2015

Figs 29–43. Dorsal (33, 35) and ventral habitus images of Japanese Phytocoris species. – 29–31, P. izanagii; 32, P. izanamiae, holotype; 33, 34, P. kerzhnerianus, holotype; 35–37, P. minakatai; 38, 39, P. hasegawai; 40, P. amateras; 41, 42, P. miyamotoi; 43, P. protobothrops.

One generation per year is assumed for most Discussion Japanese Phytocoris species that appear to hibernate Although this genus is easily distinguished from any in the egg stage, but P. protobothrops occurring on related genera by superficial characters alone, the subtropical islands obviously has a bivoltine life identifications of species are sometimes severely cycle. In Japan, the immature forms were confirmed perplexing, due to the broad intraspecific variations only for P. nowickyi (Fig. 15) and P. pallidicollis in coloration and size. As pointed out by Stonedahl (Fig. 16). (1988), the phylogenetic relationship of Phytocoris to Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 27

Figs 44–53. Dorsal and ventral (48, 49, 51) habitus images of Phytocoris species. – 44–50, P. ohataensis, exhibiting various color pattern; 51, P. pallidicollis; 52, 53, P. shabliovskii.

other mirine genera is yet to be clearly demonstrated. characters diagnosed above. Regarding the Asian In Japan and adjacent regions, however, Phytocoris faunas, Phytocoris, Adelphocoris and Adelphocorisella appears to be closely related to Adelphocoris Reuter, are likely to have derived from the same lineage, Adelphocorisella Miyamoto & Yasunaga, and judging from the similar shape of the male genitalia, Creontiades Distant (see Yasunaga 1990a, b, 1997, particularly the endosoma often with a comb-shaped 1998, Miyamoto & Yasunaga 1993 for these genera); sclerotized process, which can possibly be regarded as members of Phytocoris can be distinguished from a synapomorphy. In Adelphocorisella, a lobal sclerite those of such allied genera by the combination of replaces the comb-shaped sclerite, which seems to Downloaded from Brill.com10/06/2021 05:01:49PM via free access

28 Tijdschrift voor Entomologie, volume 158, 2015 show the primitive character status within these – Total body length less than 4.3 (mostly around genera. 4) times as long as basal width of pronotum; As mentioned above, the subgenera and species- dorsal coloration not as above ���������������������� 2 groups are not satisfactorily applicable to the classifi- 2. Smaller species; body shorter than 5.3 (♂)/5.6 cation of Japanese species. To demonstrate the (♀) mm; antennal segment I nearly equal in phylogenetic position and to create more reliable length to width of head including eyes ����� 3 subdivisions of this tremendous, cosmopolitan – Moderate to larger species; body longer than genus, further comprehensive work on a global basis 5.5 (♂)/6.3 (♀) mm; length of antennal seg- would be required, far beyond the scope of this study ment I significantly greater than width of head and previous regional revisions. including eyes ����������������������5 3. Basic coloration of dorsum shiny stramine- ous or yellowish brown, often widely Checklist of Japanese species of tinged with red, not distinctly mottled Phytocoris, with known distributions (Figs 1, 2) �����������������P. amateras n. sp. – Dorsum matte or weakly shining, gener- 1. P. amateras Yasunaga & Schwartz, n. sp. — ally brown or dark brown, significantly Japan (Honshu, Shikoku, Kyushu). mottled (generally whitish in teneral speci- 2. P. hasegawai Yasunaga & Schwartz, n. sp. — mens ������������������������������ 4 Japan (Ryukyus: Ishigaki Island). 4. Hemelytron widely dark brown, less mot- 3. P. izanagii Yasunaga & Schwartz, n. sp. — tled; anterior corium and base of cuneus Japan (Honshu, Kyushu). sometimes tinged with red; occurring in 4. P. izanamiae Yasunaga & Schwartz, n. sp. — temperate and cold temperate zones (Figs 3–5) Japan (Honshu, Shikoku). ������������������������� P. miyamotoi n. sp. 5. P. kerzhnerianus Yasunaga & Schwartz, – Hemelytron mottled; apical part of corium n. sp. — Japan (Honshu, Shikoku). with a L- or slash-shaped dark mark; inhabit- 6. P. longipennis Flor, 1861 — Japan (Hokkaido, ing subtropical Amami and Okinawa islands S. Kuril Islands), Trans-Palearctic; the Nearctic (Figs 6–8) �������������P. protobothrops n. sp. populations (Canada) are due to recent intro- 5. Body mostly or entirely sanguineous, reddish duction (Aukema et al. 2013). brown or orange red (pink or whitish orange 7. P. minakatai Yasunaga & Schwartz, n. sp. — in teneral specimens) (Figs 11–14) ��������� 6 Japan (Honshu, Shikoku, Kyushu), Yakushima – Body stramineous, brownish or fuscous, Island. only with limited reddish pattern �����������7 8. P. miyamotoi Yasunaga & Schwartz, n. sp. — 6. Male vertex much wider than an eye in dorsal Japan (Hokkaido, Honshu, Shikoku). view; antennal segment III more than 1.5 9. P. nowickyi Fieber, 1870 — Japan (Hokkaido, times longer than basal width of pronotum; Honshu, S. Kuril Islands); Trans- hemelytron usually mottled with dark pat- Palearctic. tern; female brachypterous (Figs. 4, 13) ����� 10. P. ohataensis Linnavuori, 1963 (= P. scotinus ���������������������������������P. nowickyi Kerzhner, 1977, n. syn.) — Japan (Hokkaido, – Male vertex almost equal in width to eye in Honshu, Shikoku, Kyushu, Rishiri, Tsushima dorsal view; antennal segment III shorter than & S. Kuril Islands); Korea; Russian Far basal width of pronotum; hemelytron some- East. times mottled with white or pale spots, with- 11. P. pallidicollis Kerzhner, 1977 — Japan out darkened pattern; female macropterous (Hokkaido, Honshu, Shikoku, Kyushu, (Figs 11, 12) �������������� P. izanamiae n. sp. S. Kuril Islands); Korea; Russian Far East. 7. Pronotum (at least its posterior part) 12. P. protobothrops Yasunaga & Schwartz, clearly fuscous or blackish brown (Figs 9, 10, n. sp. — Japan (Ryukyus: Amami & Okinawa 18–22) �����������������������������������8 Islands). – Pronotum concolorously castaneous, grayish, whitish or pale brown, not clearly infuscate; posterior part sometimes obscurely mottled, Key to the Japanese species of Phytocoris usually with two dark spots formed by dark 1. Body very slender like a stenodemine, with setae (Figs 23–28) ������������������������� 10 total length more than 4.6 times as long as 8. Antennal segment III as long as basal width of basal width of pronotum; dorsum shiny strami- pronotum; labium reaching abdominal ster- neous, with longitudinal, orange-red stripes num VII (Figs 48, 49); hemelytron often from head to hemelytron (Figs 38, 39) tinged with olive or green (Figs 18, 45, 46, 50) ��������������������� P. hasegawai n. sp. �������������������������������� P. ohataensis Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 29

– Antennal segment III shorter than basal Tokai Region: Aichi Pref., Mennoki, 15 Aug 1985, width of pronotum; labium reaching base of T. Ieki, 1 ♂ (TYCN). Kinki Region: Kumano area, abdominal sternum VI or shorter, not much Wakayama Pref., Hatenashi Mts., Mt. Ando, 1,100 exceeding apex of metacoxa; hemelytron basi- m alt., light trap, 11 Aug 2002, T. Yasunaga & S. cally dark brown �����������������������������9 Gotoh, 1 ♂, 2 ♀ (TYCN); Tanabe City, Komori, 9. Antennal segment I longer than 1.6 (♂)/2.0 light trap, 20 Aug 1997, S. Gotoh, 1 ♂ (TYCN); (♀) mm, greater than 1.43 times as long Tanabe City, Ohtoh-Hinotani, 30 Aug 1988, S. as head width including eyes; base of Gotoh, 1 ♀ (TYCN); Nara Pref., Kami-kitayama, cuneus usually creamy or whitish brown Mt. Wasamata, N34.2186 E135.9844, light trap, 17 (Figs 9, 10) ����������������P. izanagii n. sp. Aug 1997, Y. Nakatani, 2 ♂ (NIAES); same locality, – Antennal segment I shorter than 1.5 (♂)/1.6 16 Aug 1993, N. Hirai, 2 ♂ (TYCN); Nara Pref., (♀) mm, less than 1.33 times as long as Kami-kitayama, Odaigahara, light trap, 16 Aug head width including eyes; base of 2004, K. Yamada, 1 ♂, 1 ♀ (TKPM); same locality, cuneus obscured or darkened (Figs 22, 35) 14 Sep 2004, K. Yamada (TKPM); Osaka Pref., Mt. �����������������������������������������P. minakatai n. sp. Katsuragi, light trap, 31 Aug 1997, 1 ♂ (TYCN). 10. Antennal segment III longer than basal width Chugoku Region: same data as for holotype, 4 ♂, 16 of pronotum; apical inner part of corium ♀ (AMNH, TYCN). Shikoku, Kochi Pref., Monobe, with a dark, slash-like mark (Figs 24, 25) Befu, light trap, 6 Aug 1994, M. Takai, 7 ♂, 2 ♀ ������������������������������������������������� P. longipennis (TYCN); same data, except for date 30 Jul 1994, 1 ♂ – Antennal segment III shorter than basal width (TYCN); same locality, light trap, 5 Sep 1997, T. of pronotum; apical inner part of corium circu- Yasunaga & M. Takai, 1 ♂ (TYCN); same locality, larly or widely darkened (Figs 23, 26–28, 35) light trap, 14 Aug & 19 Sep 2010, B. Shishido, 2 ♂ ������������������������������������������������������������������ 11 (TYCN); Kochi Pref., Monobe, Nishikuma, 11 Sep 11. General coloration brown or darker; hemely- 1999, M. Takai 2 ♂ (TYCN); Kochi Pref., tron widely brown or fuscous, mottled scatter- Tosashimizu City, Imanoyama, 15 Sep 2001, M. ing with pale portions (Figs 23, 35) �������������� Takai, 1 ♂, 1 ♀ (TYCN). Kyushu, Fukuoka Pref., ����������������������������������� P. kerzhnerianus n. sp. Yame-gun, Hirozo, 31 Aug 1959, 2 ♂ (NIAES); Mt. – General coloration whitish brown; hemely- Hikosan (Hikosan Biol. Lab., Kyushu Univ.), tron widely whitish or creamy brown, with N33.482046, E130.908898, light trap, 3–4 Aug dark pattern limited to inner margin of corium 1988, T. Yasunaga, 1 ♂, 2 ♀ (TYCN); Oita Pref.: and/or apical inner part of cuneus (Figs 26–28) Mts. Kuju, Mt. Kurodake, 15 Sep 1985, S. Nomura, ������������������������������������������������� P. pallidicollis 2 ♂, 5 ♀ (TYCN); Shonai Town, Shiramizu, light trap, 27–28 Jul 1995, Y. Nakatani, 6 ♂, 12 ♀ Phytocoris amateras Yasunaga & Schwartz, n. sp. (NIAES, TYCN). Figs 1–2, 40, 54, 64, 69 Diagnosis. Recognized by its small-sized, slender body; shiny dorsum; pale stramineous brown basic Type material. Holotype, ♂: Japan, Honshu, coloration; reddish brown lateral margins of prono- Chugoku Region: Hiroshima/Shimane Pref., Mt. tum; creamy yellow scutellum; widely reddish mesal Ungetsu (Uzutsuki), 900 m alt., N34.80 E132.23, hemelytron; and a dark, small spot on paracuneus on Larix leptolepis, 5 Sep 1997, T. Yasunaga (Figs 1–2); a pointed process on left paramere sen- (AMNH_PBI 00380366) (AMHN). Paratypes: sory lobe (Fig. 54); and simple endosoma, with small Japan, Honshu, Tohoku Region: Aomori Pref., comb-shaped sclerite and lacking lobal sclerite Kuroishi City, Sakaimatsu, 30 Aug 1986, T. Ichita, 2 (Fig. 54). Currently, any reliable sister species of P. ♂ (TYCN); Aomori Pref., Nishimeya Village, 15 amateras is yet to be confirmed. Aug 1998, T. Ichita, 3 ♂, 1 ♀ (TYCN); Yamagata Description. Body generally stramineous or pale Pref., Yamagata City, Minorigaoka, at light, 13−30 brown, partly tinged with red, slender, small-sized; Aug 1989, K. Watanabe, 3 ♂, 1 ♀ (TYCN); dorsal surface weakly shining, less mottled, with uni- Fukushima Pref., Aizuwakamatsu City, Makoshi, 30 formly distributed, simple, dark, semierect setae and Jul 1994, K. Nakano, 1 ♂ (TYCN). Koushin’etsu rather sparsely distributed, sericeous, reclining setae. Region: Niigata Pref., Gosen City, Ronze-Shinden, Head shiny yellowish brown; extreme apex of clyp- 16 Jul 1994, K. Nakano, 2 ♂ (TYCN); Niigata Pref., eus brown. Antenna dark brown; segment I reddish Niitsu City, Nanoka, 24 Aug 1976, S. Sakurai, 1 ♂ brown, speckled with small, yellow spots. Labium (TYCN). Kanto Region: Gunma Pref., Riku Village, shiny brown, reaching abdominal sternum VI; apical 13 Aug 1997, K. Takahashi, 2 ♂ (TYCN); Tochigi 1/2−2/3 of segment IV dark reddish brown. Lateral Pref., Mt. Yamizosan, light trap, 23 Aug 1993, and posterior margins of pronotum more or less K. Konishi, 3 ♂ (TYCN); Tokyo, Okutama, Nippara, darker or reddish; collar a little broader than anten- 21 Sep 1976, M. Tomokuni, 4 ♂, 4 ♀ (NSMT). nal segment II; scutellum shiny creamy yellow, Downloaded from Brill.com10/06/2021 05:01:49PM via free access

30 Tijdschrift voor Entomologie, volume 158, 2015

Figs 54–58. Male genitalia of Japanese Phytocoris species. – 54, P. amateras; 55, P. miyamotoi; 56, P. nowickyi; 57, P. protobothrops; 58, P. hasegawai.

almost immaculate, smooth; pleura and all coxae endosoma with small comb, lacking LBS. Female creamy yellow; propleuron and episternum some- genitalia as in Fig 69. Sclerotized ring small, what tinged with gray. Hemelytron widely pale red- narrowed. dish brown, less shiny than pronotum and scutellum, Measurements. ♂/♀: Total length of body 4.4– weakly mottled with obscure, grayish pattern; basal 5.1/4.5–5.2; head width including eyes 0.85– inner corner and sometimes median inner margin of 0.91/0.79–0.89; vertex width 0.24–0.25/0.30–0.43; cuneus each with a dark spot; apical 1/3 of cuneus lengths of antennal segments I–IV 0.78–0.91, 2.35– usually tinged with red; membrane including veins 2.63, 1.10–1.21, 0.82–0.89/0.83–0.98, 2.10–2.36, pale smoky brown. Leg reddish brown; all femora 1.07–1.13, 0.90–0.98; labial length 2.20– and basal parts of all tibiae with scattered pale, small 2.36/2.27–2.65; mesal length of pronotum includ- spots; tibial spines prominent, each longer than ing collar 0.68–0.76/0.71–0.86; basal width of diameter of respective tibia; all tarsi brown. Abdomen pronotum 1.25–1.30/1.22–1.40; maximum width varying from yellowish brown to grayish brown, across hemelytron 1.45–1.70/1.47–1.67; and sometimes tinged with red. Male genitalia: as in lengths of metafemur, tibia and tarsus 2.49–2.55, Figs 54, 64. Left paramere with a pointed process on 3.55–4.02, 0.58–0.62/2.57–2.82, 3.57–4.02, sensory lobe dorsally; right paramere simple, straight; 0.56–0.62. Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 31

Figs 59–63. Male genitalia of Japanese Phytocoris species. – 59, P. izanamiae; 60, P. kerzhnerianus; 61, P. izanagii; 62, P. longipennis; 63, P. pallidicollis.

Etymology. Named for Amateras(u)-Oomikami, a additional (possibly coniferous) plants since no larch major mythological goddess, symbolic of the sun, in is present at several localities cited above. Collecting the traditional Japanese Shinto religion; her father is records suggest P. amateras has one generation per Izanagi (see izanagii below); noun in apposition. year. Adults appear from August to October. Biology. Most specimens were collected using light traps, but more than 20 adults including some ten- Phytocoris hasegawai Yasunaga & Schwartz, n. sp. eral individuals were captured from a Japanese larch, Figs 38–39, 58, 65 Larix leptolepis (Sieb. et Zucc.) Gordon (Pinaceae) Type material. Holotype, ♂: Japan, Ryukyus, Ishigaki which is assumed to be one of its breeding hosts. Island, Hirai (probably misreading or misspelling of However, this new species must be associated with ‘Hirae’), black-light trap, 6 Mar 1973, H. Hasegawa Downloaded from Brill.com10/06/2021 05:01:49PM via free access

32 Tijdschrift voor Entomologie, volume 158, 2015

Figs 64–72. Male endosoma (64–68) and female bursa copulatrix (69–72) of Japanese Phytocoris species. – 64, 69, P. amateras; 65, P. hasegawai; 66, 70, P. miyamotoi; 67, 71, P. nowickyi; 68, 72, P. protobothrops.

(AMNH_PBI 00380367) (NIAES). Paratypes: head, pronotum, scutellum to similarly striped Same data as for holotype, 10 ♂, 2 ♀ (NIAES, hemelytron; a deep red spot at paracuneus (Fig. 38); TYCN). simple form of parameres (Fig. 58); and Diagnosis. Distinguished from all Japanese conge- Adelphocoris-like comb-shaped endosomal sclerite ners by its very slender, pale-colored, stenodemine- (Fig. 65). like body; shiny dorsum with several noticeable, Description. Body with stenodemine-like form, sanguineous or orange-red stripes running from generally pale brown, small-sized, elongate, slender; Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 33 dorsal surface shining, with several noticeable, Phytocoris izanagii Yasunaga & Schwartz, n. sp. sanguineous or orange-red stripes running from Figs 9–10, 29–31, 73 pronotum to scutellum, with sparsely distributed, sericeous, reclining setae (Fig. 38) and partly with Type material. Holotype ♂: Japan, Honshu, pale erect or semierect setae on head, pronotum Chugoku Region: Hiroshima Pref., Geihoku (=cur- and hemelytron. Head shiny yellowish brown, rent Kitahiroshima) Town, Yawata, Chojabaru, light rounded in front, with two sanguineous stripes; eye trap, 10 & 11 July 1994, K. Yoshizawa (AMNH_ small; vertex wide; frons with reddish striae (ante- PBI 00380368) (AMNH). Paratypes: Japan, rior end of sanguineous stripe from pronotum). Honshu, Aomori Pref., Mt. Iwaki-san, 1,200−1,625 Antenna yellow-stramineous; segment I with two m, 28 Jun 1986, T. Ozaki, 1 ♀ (TYCN); Gunma red stripes that are sometimes interrupted or Pref., Riku Village, 11 Jul 1999, K. Takahashi, 1 ♀ reduced; segment II partly tinged with red; seg- (TYCN); Tochigi Pref., Nikko, Yumoto, 6 Jun 2004, ments III and IV brown. Labium pale reddish S. Maehara, 1 ♀ (TYCN); Nagano Pref., Nagawa brown, reaching abdominal sternum V or VI; apical Village, Kisojihara, light trap, 9 Jul 1995, half of segment IV darkened (Fig. 39). Pronotum Y. Nakatani, 1 ♀ (NIAES); Yamanashi Pref., with four sanguineous stripes and lateral margins; Daibosatsu-toge, 14 Jul 1961, S. Katsuya, 1 ♀ calli weakly swollen; collar about as thick as anten- (NIAES); Okayama Pref., Kawakami, Kamiyubune, nal segment I; pleura with a sanguineous stripe 11 Jul 2003, B. Shihido, 1 ♂ (TYCN); Hiroshima from propleuron to episternum; scutellum rather Pref., same data as for holotype, 3 ♂, 10 ♀ (AMNH, flat. Hemelytron with reddish stripes not confluent TYCN). Kyushu: Kumamoto Pref., Izumi-mura, to those from pronotum to scutellum; inner part of Mt. Yamaingiri, 1,100 m, light trap, 26–27 Jun corium and medial clavus somewhat matte or 1992, T. Yasunaga, 1 ♀ (TYCN). roughened, pale reddish brown, weakly speckled Diagnosis. Recognized by its largest size among the with yellow spots; apical inner corner of corium known east Asian congeners; fuscous and brown- with a deep red spot; membrane pale smoky brown mottled basic coloration (Figs 9–10); relatively short with apically reddish veins. All coxae and legs yel- labium not exceeding apex of metacoxa (Figs 29–31); lowish brown; metafemur with reddish, reticular almost continuously whitish base of cuneus; and dis- stripe apically; tibial spines pale brown, short. tinctive male and female genitalic structures. Similar Abdomen shiny stramineous; sterna laterally with a to darkened individuals of P. minakatai, from which pair of sanguineous stripes along rows of spiracles. this new species can be distinguished by the charac- Male genitalia as in Figs 58, 65. Parameres simple ters in the key (couplet 9). in form; left paramere semi-circularly curved; endo- Description. Body elongate oval, large; basic color- soma with typical, Adelphocoris-like comb that is ation chocolate brown or darker; dorsal surface fused basally with PML, lacking LBS. Female geni- matte, usually speckled with pale portions but almost talia not examined as two available females appear wholly fuscous in some male individuals, with uni- teneral. formly distributed, sericeous, reclining setae and Measurements. ♂/♀: Total length of body 5.7– sparsely distributed, dark, short, semierect setae. 6.6/5.9–6.0; head width including eyes 0.87– Head dark brown; vertex slightly pruinose; maxillary 0.90/0.80–0.87; vertex width 0.36–0.39/0.43–0.50; plate pale brown. Antenna chocolate brown; seg- lengths of antennal segments I–IV 1.23–1.33, 2.55– ment I usually with a few yellowish spots; basal and 2.68, 1.78–1.85, 1.26–1.30/1.44–1.51, 2.67–2.77, apical 1/3 parts of segment II, and extreme base of 1.66–1.76, 1.16–1.33; labial length 2.82– segment III yellowish brown. Labium shiny choco- 2.90/2.85–2.87; mesal length of pronotum includ- late brown, reaching or slightly exceeding apex of ing collar 0.64–0.65/0.61–0.65; basal width of metacoxa. Pronotum weakly shining, with narrowly pronotum 1.23–1.25/1.26–1.27; maximum width pale posterior margin; collar about as thick as anten- across hemelytron 1.56–1.70/1.63–1.70; and nal segment II; scutellum weakly shining, usually lengths of metafemur, tibia and tarsus 3.38–3.54, with pale brown corners; pleura widely dark brown; 4.92–5.17, 0.64–0.74/3.38–3.39, 5.07–5.08, ostiolar peritreme yellowish brown. Hemelytron 0.67–0.68. more or less speckled with pale brown portions; basal Etymology. Named for the late Mr. Hitoshi Hasegawa 1/4−1/3 of cuneus whitish or pale brown; membrane (NIAES ex-staff, heteropterist), who collected all including veins smoky brown, speckled with pale, specimens designated as type series; a noun in geni- semitransparent portions. All coxae shiny yellowish tive case. brown, with dark brown spots basally, but sometimes Biology. All available specimens were collected using widely darkened. Leg chocolate brown; femora and/ a UV light trap four decades ago. Neither biological or tibiae more or less speckled with pale spots; meta- information nor further material is currently femur usually with a pale brown ring subapically; available. each tibia usually with two yellowish annulations; Downloaded from Brill.com10/06/2021 05:01:49PM via free access

34 Tijdschrift voor Entomologie, volume 158, 2015

Figs 73–76. Male endosoma (73, 75) and female bursa copulatrix (74, 76) of Japanese Phytocoris species. – 73, 74, P. izanagii; 75, 76, P. izanamiae.

tibial spines rather short, less than diameter of laterally projected anterior rim and reduced posterior respective tibia; all tarsi brown. Abdomen widely or half. almost entirely shiny chocolate brown, or ventral Measurements. ♂/♀: Total length of body 6.9– surface irregularly speckled with pale brown por- 7.1/8.3–9.1; head width including eyes 1.13– tions. Male genitalia as in Figs 61, 73. Generally 1.17/1.17–1.27; vertex width 0.40–0.44/0.49–0.53; large in size; right paramere very long, compared lengths of antennal segments I–IV 1.63–1.73, 3.29– with left one, somewhat curved medially with inden- 3.39, 1.53–1.60, 1.23–1.27/2.09–2.13, 3.56–3.76, tation (Fig. 61); PML of endosoma large, with 1.56–1.82, 0.98–1.30; labial length 3.04–3.07/ comb-shaped sclerite and broad, apically rounded 3.75–4.09; mesal length of pronotum including col- LBS; SGP thick-rimmed, heart-shaped (Fig. 73). lar 1.16–1.20/1.32–1.45; basal width of pronotum Female genitalia as in Fig. 74. Bursa copulatrix 1.84–1.94/2.15–2.50; maximum width across widened; sclerotized ring with toughened and ­hemelytron 2.27–2.43/2.79–3.23; and lengths of

Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 35 metafemur, tibia and tarsus 3.38–3.69, 5.38–5.54, Nara Pref., Mt. Wasamata, light trap, 11 Aug 1994, 0.67–0.80/3.75–4.09, 5.84–6.03, 0.80–0.87. N. Hirai, 1 ♂ (TYCN); same locality, 9 Aug 1995, Y. Etymology. Named for Izanagi, the famous, Japanese Nakatani, 1 ♂ (NIAES); Nara Pref., Kamikitayama, mythological god, the father of Amaterasu, and the Odaigahara, light trap, 16 Aug 2004, K. Yamada, 3 Creator of the Japan Archipelago; a noun in genitive ♀ (TKPM). Chugoku Region: Tottori Pref., Mt. case. Hoki-Daisen, 12 Jul 1994, K. Yoshizawa, 2 ♂ Biology. Unknown; collection records suggest this (TYCN). Shikoku, Kochi Pref., Hongawa, Yosakoi- species has one generation per year. toge, light trap, 1 Aug 1998, T. Befu, 2 ♂ (TYCN); Kochi Pref., Hongawa, Teragawa, 17 Jul 1999, M. Takai, 1 ♂ (TYCN); Kochi Pref., Monobe, Befu, Phytocoris izanamiae Yasunaga & Schwartz, n. sp. light trap, 31 Aug 1998, T. Befu, 3 ♂ (TYCN); Figs 11–12, 32, 59, 75–76 Tokushima Pref., Kitoh-son, 15 Aug 1998, M. Takai, Type material. Holotype ♂: Japan, Honshu, Kinki 1 ♂ (TYCN). Region: Kumano area, Wakayama Pref., Hatenashi Diagnosis. Recognized by its medium size; bright Mts., Mt. Ando, 1100 m alt., light trap, 11 Aug orange-red or sanguineous general coloration 2002, T. Yasunaga & S. Gotoh (AMNH_PBI (Figs 11, 12); and less mottled dorsum (teneral 00380370) (AMNH). Paratypes: Japan, Honshu, individuals sometimes speckled with white spots Tohoku Region: Aomori Pref.: Gunmatai, Hiraka as in Fig. 12). In eastern Asia, this new species Town, 22 Aug 1990, T. Ichita, 2 ♀ (TYCN); Mt. may be confused with P. nowickyi, from which it is Osorezan, 1 Aug 1953, A. Habu, 1 ♂ (NIAES); easily distinguished by the characters in the key Nishimeya-mura, Imoritai, 22 Jul 1989, T. Ichita 1 (couplet 6). ♂, 3 ♀ (TYCN); Aomori City, Tashirotai, 12 Aug Description. Body elongate oval, medium-sized; basic 1989 & 13 Aug 1990, T. Ichita, 2 ♂ (TYCN); coloration orange-red, sanguineous, or whitish pink in Aomori City, Kansuizawa, 8 Sep 1992, T. Ichita, 1 ♂ teneral specimens; dorsal surface matte, with uni- (TYCN); Aomori City, Sukayu, 21 Aug 1989, T. formly distributed, brown, semierect setae and rather Ichita, 1 ♂ (TYCN); Aomori City, Hibakozawa, 6 sparsely distributed, sericeous, reclining setae. Head Sep 1992, T. Ichita, 1 ♂ (TYCN); Towadako-machi, often widely yellowish brown, mixed with reddish Tsuta, 29 Aug 1989, 1 ♂, 1 ♀ (TYCN); Yamagata portions. Antennal segment I reddish brown, speckled Pref.: Fujishima, 6 Sep 1954, H. Hasegawa, 1 ♀ with small, yellow spots; segment II yellowish brown, (NIAES); Yamagata City, Zaou, Dokko-numa, 10 sometimes tinged with red; segments III and IV Aug 1991, K. Watanabe, 2 ♀ (TYCN); Oe Town, brown. Labium shiny reddish brown, reaching Tanosawa, 20 Jul 1994, Y. Nakatani, 1 ♀ (NIAES); abdominal sternum VI; apical half of segment IV Nishikawa, Shizu, 6 Sep 1997, S. Sakurai, 1 ♀ darkened. Pronotum with shiny, paler calli; collar (TYCN); Fukushima Pref.: Ozawa, 17 Aug 1988, about as thick as antennal segment I; scutellum usually M. Hayashi, 1 ♀ (TYCN); Kawairi, 18 Aug 1988, with paler corners; pleura and all coxae pale brown, M. Hayashi, 1 ♂ (TYCN). Kanto Region: Tochigi partly with red-orange marks and/or spots. Hemelytron Pref., Mt. Yamizosan, light trap, 23 Aug 1993, K. nearly matte, almost uniformly bright orange-red or Konishi, 1 ♂ (TYCN); Tochigi Pref., Chugu-shi, 20 sanguineous, in teneral specimens speckled with pale, Jul 2004, S. Maehara, 1 ♂ (TYCN); Tochigi Pref., semitransparent spots. Leg pale reddish brown; all Nikko, Kiyotaki, 20 Aug 2012, S. Maehara, 1 ♂ femora and metatibia more or less speckled with pale, (TYCN); Tokyo, Okutama, Nippara, 21 Sep 1976, small spots; tibial spines reddish brown, prominent. M. Tomokuni, 2 ♀ (NSMT); Tokyo, Mt. Takao, 12 Abdomen shiny reddish brown. Male genitalia as in Jul 1961, K. Mizusawa, 2 ♂ (NIAES); Kanagawa Figs 59, 75. Pygophore with a strong projection above Pref., Izu, Amagi-toge–Haccho-ike, 29 Sep 1980, base of left paramere; right paramere with many M. Tomokuni, 3 ♀ (NSMT). Koushin’etsu Region: notches (Fig. 59); endosoma with long comb-shaped Niigata Pref., Nishikawa, Shizu, 6 Sep 1997, S. sclerite that is notched apically (Fig. 75). Female geni- Sakurai, 1 ♀ (TYCN); Nagano Pref.: Kamikochi, 10 talia as in Fig. 76. Sclerotized ring rather thick- Sep 1951, H. Hasegawa, 1 ♂ (NIAES); Nagano rimmed, narrowed inward. Pref., Sarukura-Hakuba, 16 Sep 1966, H. Hasegawa, Measurements. ♂/♀: Total length of body 5.7– 1 ♀ (NIAES); Yamanashi Pref., Masutomi, 18 Jul 6.2/6.1–7.1; head width including eyes 0.96– 1994, K. Yoshizawa, 1 ♀ (TYCN). Kinki Region: 0.99/1.01–1.03; vertex width 0.32–0.34/0.40; same data as for holotype, 3 ♂ (TYCN); same local- lengths of antennal segments I–IV 1.23–1.42, 2.76– ity, 18 Aug 1996, S. Gotoh, 2 ♂ (TYCN); Nara 2.96, 1.26–1.42, 1.23–1.42/1.29–1.54, 2.79–3.33, Pref., Kami-kitayama, light trap, 17 Aug 1997, Y. 1.44–1.57, 1.26–1.57; labial length 2.76–2.80/3.01– Nakatani, 2 ♂ (NIAES); Nara Pref., Totsugawa, Mt. 3.08; mesal length of pronotum including collar Obakodake, 3 Aug 1995, Y. Nakatani, 2 ♂ (NIAES); 0.92–0.99/1.04–1.14; basal width of pronotum

Downloaded from Brill.com10/06/2021 05:01:49PM via free access

36 Tijdschrift voor Entomologie, volume 158, 2015

1.56–1.70/1.81–1.85; maximum width across hem- 2001, 2 ♂ (TYCN); Tokushima Pref., Kitou-son, 15 elytron 2.03–2.06/2.33–2.40; and lengths of metafe- Aug 1998, M. Takai, 8 ♂, 1 ♀ (TYCN). Kyushu, mur, tibia and tarsus 3.22–3.33, 4.61–5.23, Fukuoka Pref., Mt. Hikosan (Hikosan Biol. Lab., 0.58–0.62/3.38–3.69, 5.19–5.85, 0.64–0.68. Kyushu Univ.), N33.482046, E130.908898, light Etymology. Named for Izanami, a Japanese mytho- trap, 10–11 Aug 1985, N. Koda, 1 ♂ (TYCN); logical goddess in the Shinto worship and spouse of Kumamoto Pref., Izumi-mura, Mt. Yamaingiri, Izanagi; a noun in genitive case. 1,100 m, light trap, 26–27 Jun 1992, T. Yasunaga, 3 Biology. Almost all known specimens were collected ♀ (TYCN); Kumamoto Pref., Izumi-mura, Momiki, with light traps. No other information on biology is light trap, 6 Aug 1991, T. Yasunaga, 1 ♂ (TYCN). available. A univoltine life cycle is assumed for this Additional material examined. Hokkaido, Sapporo City, new species. Hokkaido Univ. Campus, on Salix, 14−30 Sep 2001, T. Ogita, 3 ♂, 2 ♀ (TYCN). Shikoku, Kochi Pref., Monobe, Phytocoris kerzhnerianus Yasunaga & Schwartz, Nishikuma, light trap, 10 Oct 1998, T. Befu, 7 ♀ (TYCN). n. sp. Kyushu, Oita Pref., Mt. Kurodake, 15 Sep 1985, Figs 23, 33–34, 60, 78, 95–98 S. Nomura, 1 ♀ (TYCN). Diagnosis. Recognized by its medium size; generally Type material. Holotype ♂: Japan, Honshu, Kumano brownish or castaneous body with matte, more or area: Wakayama Pref., Hatenashi Mts., Mt. Ando, less mottled dorsum; not significantly infuscate 1,100 m alt., light trap, 18 Aug 1996, S. Gotoh posterior pronotum; widely pale castaneous scutel- (AMNH_PBI 00380370) (AMNH). Paratypes: lum; usually almost darkened clavus (Figs 23, 33); Japan, Honshu, Tohoku Region: Aomori Pref., and developed comb-shaped sclerite on endosoma Aomori City, Nyuunai, 2 Aug 1987, T. Ichita, 1 ♂ (Fig. 78). Most related to P. shabliovskii Kerzhner, (TYCN); Yamagata Pref., Murayama City, Hayama, 1978 (Figs 52, 53), currently known only from the light trap, 11 Sep 1993, K. Watanabe, 1 ♂ (TYCN). Russian Far East (South Primorsky Territory); this Kohshin’etsu Region: Niigata Pref., Myoukou, new species can be distinguished from it by the Tsubame Hot Spa, light trap, 14 Sep 1996, S. larger size, less mottled dorsum, not significantly Sakurai, 13 ♂, 1 ♀ (TYCN); Niigata Pref., darkened posterior part of the pronotum, and Myokokogen Town, Hikosa Waterfall, 7 Aug 1997, different shape of the male genitalia (cf. Figs 77, x Sueyoshi, 1 ♂ (TYCN); Nagano Pref., Kamikochi, 9 vs. 78, y). Sep 1972, K. Arito, 1 ♂ (NIAES). Hokuriku Region: Description. Body elongate, medium-sized; basic col- Ishikawa Pref., Mt. Hakusan, Nakahanba, 1,300– oration brown or castaneous; dorsal surface matte, 1,500 m, 24 Aug 1991, I. Togashi, 1 ♂ (TYCN); castaneous to dark brown, less mottled, with three Ishikawa Pref., Zenjodo, 17 Aug 1991, I. Togashi, 1 types of vestiture: almost uniformly distributed, seri- ♂ (TYCN). Kanto Region: Tokyo, Chichibu, Futase- ceous, reclining setae (1) and brown, simple, erect or dam, 10 Aug 1982, H. Hasegawa, 1 ♂ (NIAES). semierect setae (2), and partly distributed, dark, Kinki Region: Wakayama Pref., same data as for semierect setae (3). Head pale brown, sometimes holotype, 2 ♂ (TYCN); Wakayama Pref., Mt. with darker pattern. Antenna brown to dark brown; Kohyasan, Juglans mandshrica, 17 Sep 1998, S. segment I creamy yellow, mottled with castaneous Gotoh, 1 ♂ (TYCN); Nara Pref., Tenkawa Village, marks or spots; base of segment II, and base and Gyojyagaeri, Stewartia monadelpha Sieb. et Zucc., 15 extreme apex of segment III yellowish brown. Aug 1996, Y. Nakatani, 2 ♂ (TYCN); same locality Labium shiny pale brown, reaching abdominal ster- and collector, light trap, 2 Aug 1996, 2 ♂, 1 ♀ num VI or VII; apical half of segment IV chestnut (NIAES); Nara Pref., Kamikitayama, Mt. Wasamata, brown. Pronotum castaneous, usually less mottled; 11 Aug 1994, N. Hirai, 5 ♂, 2 ♀ (TYCN); same collar about as thick as antennal segment I; scutel- locality, light trap, 20 Aug 1997, Y. Nakatani, 1 ♀ lum almost uniformly brown or pale brown; pleura (NIAES); same locality, light trap, 24−25 Jul 1992, widely chestnut brown; ostiolar peritreme and all Y. Nakatani, 1 ♂ (NIAES). Chugoku Region: coxae yellowish brown. Hemelytron varying from Hiroshima Pref., Geihoku (=current Kitahiroshima) brown to dark brown, irregularly speckled with pale, Town, Yawata, Chojabaru, light trap, 10−11 July semitransparent portions; cuneus usually castaneous 1994, K. Yoshizawa, 51 ♂ (AMNH, TYCN). with pale base; membrane smoky brown, with api- Shikoku, Ehime/Kochi Prefs., Jiho Pass (900−1,000 cally whitish veins. All femora and tibiae variously m), flowers of Swida controversa (Helms. ex Prain) mottled (brown and pale); tibial spines reddish Soják, 6 Aug 1993, T. Yasunaga, 1 ♂ (TYCN); Kochi brown, prominent; all tarsi castaneous. Abdomen Pref., Tengu-Kogen (Plateau), light trap, 31 Aug shiny chestnut brown; ventral surface often mottled 1998, T. Befu, 2 ♂ (TYCN); Kochi Pref., Hongawa, in female. Male genitalia as in Figs 60, 78, q, y. Teragawa, light trap, 17 Jul 1999, M. Takai, 2 ♂ Female genitalia as in Figs 95−98. Sclerotized ring (TYCN); Kochi Pref., Hongawa, Yosakoi-toge, 7 Jul elongate-ovoid, narrowed. Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 37

Figs 77–80. Male endosoma of Phytocoris species. – 77, P. shabliovskii; 78, P. kerzhnerianus; 79, P. longipennis; 80, P. pallidicollis.

Measurements. ♂/♀: Total length of body 5.5– Biology. Since most specimens were collected using 6.3/6.2–7.7; head width including eyes 0.99– light traps, biological information is unknown. A 1.05/0.96–1.08; vertex width 0.27–0.28/0.36–0.43; few adults were captured from the inflorescence of lengths of antennal segments I–IV 1.23–1.27, 2.89– Stewartia monadelpha Sieb. et Zucc. (Theaceae) and 2.96, 1.32–1.50, 1.17–1.33/1.26–1.67, 2.77–3.60, Swida controversa (Helms. ex Prain) Soják 1.50–1.85, 1.29–1.54; labial length 2.46–2.65/2.76– (Cornaceae), and leaves and branches of Salix sp. 3.17; mesal length of pronotum including collar These data may indicate that this new species is 0.89–0.93/0.95–1.17; basal width of pronotum ­associated with broadleaf angiosperms rather than 1.53–1.57/1.72–1.94; maximum width across conifers. Collection records suggest the adults appear ­hemelytron 1.90–2.10/2.27–2.46; and lengths of from July to October. metafemur, tibia and tarsus 3.07–3.17, 4.92–4.96, 0.58–0.62/3.38–4.00, 5.13–5.85, 0.62–0.65. Phytocoris longipennis Flor Etymology. Named in honor of the late Dr. I. M. Figs 17, 24–25, 62, 79, 99–100 Kerzhner (ZMAS), who greatly improved knowledge about the heteropteran faunas of the Russian Far Phytocoris longipennis Flor, 1861: 593, 601 (n. sp.); East; an adjective. Schuh, 1995: 890 (cat.); Kerzhner & Josifov,

Downloaded from Brill.com10/06/2021 05:01:49PM via free access

38 Tijdschrift voor Entomologie, volume 158, 2015

Figs 81–86. Male genitalia of Japanese Phytocoris species. – 81–83, P. ohataensis; 84–86, P. minakatai.

1999: 157 (cat.); Yasunaga, 2001: 255 (diag.); maximum width across hemelytron 2.0–2.3 mm Wachmann et al., 2004: 107 (diag.); Zheng et (♂)/2.3–2.5 mm (♀). There are some more species al., 2004: 495 (diag.); Aukema et al., 2013: similar to P. longipennis in western Palearctic Region, 227 (cat.). but many authors have provided diagnostic charac- ters, redescriptions and/or figures for this wide- Diagnosis. Recognized by its rather large size; long spread, Trans-Palearctic species (Kerzhner 1988, antenna, labium and leg; antennal segment III lon- Wachmann et al. 2004, Wagner 1974, Wagner & ger than basal width of pronotum; pale brown pro- Weber 1964, Yasunaga 2001). notum with obscurely darkened posterior part; and a Biology. According to several authors, this univoltine dark, oblique, slash-shaped mark at posterior corium mirid is found on various deciduous broadleaf trees, (Figs 24, 25). In Japan, most similar to P. pallidicollis, such as Alnus and Betula (Betulaceae), Quercus from which P. longipennis can be distinguished by the (Fagaceae), Populus and Salix (Salicaceae), Crataegus, characters in the key (couplet 10), smaller right para- Malus, Prunus, and Sorbus (Rosaceae), and Tilia mere (Fig. 62), and shorter endosomal comb-shaped (Tiliaceae). Predation on aphids, small lepidopteran sclerite (cf. Figs 79, r vs. 80, p). Male genitalia as in larvae, or mites is occasionally observed. Yasunaga Figs 62, 79; female genitalia as in Figs 99, 100. (2001) briefly documented the molt of final-instar Total body length 6.6–7.3 mm (♂)/6.9–7.5 mm (♀); immatures found on ferns in deciduous forest floor Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 39 habitat that emerged during observation (Fig. 17). Diagnosis. Recognized by its medium to large size; The adults appear from late July to early September generally dark brown body (Fig. 22); widely matte, in Hokkaido, Japan. less mottled dorsum; short labium (Figs 36–37); Material examined. Japan: Hokkaido, Kushiro: widely darkened pronotum (Fig. 35); and shape of Takkobu, on Salix, 30 Aug 1995, T. Yasunaga, 1 ♂ the male and female genitalia. Most closely related to (TYCN); Shibecha Town, Kami-Isobunnai, 30 Aug P. ohataensis, from which this new species can be dis- 1995, Y. Todo, 1 ♀ (TYCN). Horokanai: Moshiri, on tinguished by the characters in the key (couplet 8), Ulmus, Salix & Alnus, N44.364352, E142.261536, 2–7 larger and more lanceolate comb-shaped sclerite of Aug 1994, T. Yasunaga et al., 2 ♂, 4 ♀ (TYCN). the endosoma (Figs 86, 87), and much narrowed, Kamikawa: Taisetsu Mts., Tenninkyo, on Salix, 11 Aug elongate sclerotized rings that are mesally contiguous 1994, Y. Todo, 1 ♀ (TYCN); Higashikawa Town, to each other (Figs 88–90). Nokanan, on Alnus, 7 Aug 1999, T. Yasunaga, 1 ♀ Description. Body elongate oval, parallel-sided, com- (TYCN). Tokachi: Obihiro City, Iwanai-senkyo, on paratively large; basic coloration dark brown to fus- Salix, 25 Aug 1995, Y. Todo, 2 ♀ (TYCN); Rikubetsu cous; dorsal surface matte, partly mottled, with City, Shotoshibetsu, 24 Jul 1984, M. Tomokuni, 3 ♀ uniformly distributed, dark, simple, semierect setae (NSMT). Finland: Lojo, 20 Aug 1918, H. Lindb. and rather sparsely distributed, sericeous, reclining (Lindberg), 1 ♀ (TYCN). Russia: Moscow area, setae. Head brown, shining, irregularly speckled with Yamburgsk u., Serjozhino, 21 Aug 1896, (V. L.) Bianki, darker maculae; eyes large; frons somewhat striolate. Phytocoris longipennis Flor det. by Bianchi, 1 ♀ (TYCN). Antenna dark brown; segment I speckled with pale, Sankt Petersburg, Universitateskaya, Botanical Garden, small spots; extreme bases of segments III and IV yel- on Sorbus, 18–19 Aug 2002, T. Yasunaga, 1 ♀ (TYCN). lowish brown. Labium shiny chocolate brown, slightly S. Primorsky, Khasanskij Dist., Kedrovaya Pad’ Nature exceeding apex of metacoxa, not exceeding abdominal Reserve, I. M. Kerzhner, 27 Jul 1982, 1 ♂ & 28 Aug sternum V. Pronotum weakly shining, usually widely 1982, 2 ♀ (TYCN); Ussurisk Reserve, west border, 14 Jul dark brown, with narrowly pale posterior margin; calli 1993, T. Yasunaga, 1 ♂ 1 ♀. irregularly pale brown; collar about as thick as antennal segment I; scutellum shining, dark brown, somewhat Phytocoris minakatai Yasunaga & Schwartz, arched, with yellowish brown apex; pleura rather shin- n. sp. ing, widely chestnut brown except for creamy yellow ventral margins and ostiolar peritreme. Hemelytron Figs 22, 35–37, 84–87, 90 weakly shining, widely dark brown, speckled with pale Type material. Holotype ♂: Japan, Honshu, Kinki semitransparent portions; apex of corium usually with Region: Kumano area, Wakayama Pref., Doro- a pale, semicircular macula; cuneus dark reddish valley, Tamakiguchi, N33.8955 E135.8784, 15 Jun brown, with a pale, comma-shaped spot at basal inner 1993, S. Gotoh (AMNH_PBI 00380371) corner; membrane smoky brown, with a small, semi- (AMNH). Paratypes: Japan, Honshu, Kinki Region: transparent portion near apex of cuneus. All coxae pale Wakayama Pref., Tanabe City, Nakahechi, Hyoze, brown, sometimes narrowly darkened apically. Leg on Pinus densiflora, 1 Jun 1997, T. Yasunaga & chestnut brown; all femora speckled with pale spots as S. Gotoh, 1 ♂, 1 ♀ (TYCN); Wakayama Pref., same in antennal segment I; each tibia with one or two yel- data as for holotype, 1 ♂, 1 ♀ (TYCN); Wakayama lowish annulation; tibial spines pale brown, promi- Pref., Tanabe City, Ohtoh, Tsubakio-toge, nent; tarsi brown to dark brown. Abdomen shiny N33.7982, E135.6658, 600 m, 30 May 1998, chestnut brown; ventral median part more or less S. Gotoh, 2 ♂ (TYCN); Wakayama Pref., Tanabe mottled. Male genitalia as in Figs 84–87. Comb- City, Mt. Toragamine, N33.8550, E135.4646, on shaped sclerite large, rather lanceolate (Fig. 86). Female Pinus densiflora, 7 Jun 1997, T. Yasunaga & genitalia as in Figs 88–90. Sclerotized rings elongate S. Gotoh, 4 ♀ (TYCN); Nara Pref., Otoh Village, ovoid, mesally contiguous to each other. Akadani, 15 Jul 1995, T. Hirowatari, 2 ♀ (TYCN). Measurements. ♂/♀: Total length of body Hokuriku Region: Ishikawa Pref., Suzu City, Mt. 6.4–6.6/7.3–7.8; head width including eyes Nekogatake, light trap, 8 Jul 1993, Y. Nakatani, 3 ♂ 1.15–1.20/1.16–1.22; vertex width 0.26–0.30/0.36– (NIAES, TYCN). Kanto Region: Tokyo, Hibara 0.40; lengths of antennal segments I–IV 1.26–1.42, Village, 22 Jun 1990, N. Nishimura, 1 ♂ (TYCN). 3.35–3.39, 1.56–1.63, 1.30–1.36/1.53–1.56, 3.38– Shikoku, Kochi Pref., Kuroson, 24 Jun 1995, M. 3.40, 1.59–1.74, 1.23–1.25; labial length 3.07– Takai, 2 ♂ (TYCN); Tengu Plateau, light trap, 30 3.23/3.32–3.39; mesal length of pronotum including Jun 2001, T. Befu, 1 ♂ (TYCN). Kyushu, collar 1.01–1.08/1.16–1.23; basal width of prono- Kumamoto Pref., Izumi-mura, Mt. Yamaingiri, tum 1.81–1.85/2.06–2.13; maximum width across 1,100 m, light trap, 26–27 Jun 1992, T. Yasunaga, 1 hemelytron 2.12–2.22/2.39–2.59; and lengths of ♂ (TYCN); Yakushima Island, Shiratani-Unsuikyo, metafemur, tibia and tarsus 3.07–3.54, 4.92–5.23, 13 May 1983, K. Konishi, 1 ♂ (TYCN). 0.73–0.80/3.69–4.00, 5.59–5.85, 0.72–0.87. Downloaded from Brill.com10/06/2021 05:01:49PM via free access

40 Tijdschrift voor Entomologie, volume 158, 2015

Figs 87–94. Male endosoma (87, 91) and female bursa copulatrix of Japanese Phytocoris species. – 87–90, P. mi- nakatai; 91–94, P. ohataensis.

Etymology. Named in honor of Kumagusu Minakata teneral adults were found. The adults of (1867–1941), who is the most outstanding Japanese P. minakatai (from mid-May to early July) appear naturalist and enthusiastically investigated organ- obviously earlier than those of its sister species, isms in Nanki area, including type locality of this P. ohataensis (see below). new species; a noun in genitive case. The late Mr. S. Gotoh who collected many specimens of this new Phytocoris miyamotoi Yasunaga & Schwartz, n. sp. species used to respect Sir Minakata in particular. Figs 3–5, 41–42, 55, 66, 70 Biology. This new species is assumed to be associated Type material. Holotype ♂: Japan, Honshu, Kinki with Pinus densiflora Sieb. & Zucc., on which some Region, Kumano, Wakayama Pref., Tanabe City, Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 41

Figs 95–102. Female bursa copulatrix of Japanese Phytocoris species. – 95–98, P. kerzhnerianus; 99, 100, P. lon- gipennis; 101, 102, P. pallidicollis.

Ohtoh, Tsubakio-toge, N33.7982, E135.6658, 600 m, on Abies sachalinensis (Fr. Schmidt) Mast., 21 Aug 18 Aug 1998, S. Gotoh (AMNH_PBI 00380372) 1997, T. Yasunaga, 11 ♂, 1 ♀ (TYCN). Hiyama: (AMNH). Paratypes: Japan, Hokkaido, Kushiro: Kaminokuni, 18 Aug 1994, K. Yoshizawa, 1 ♀ Shibecha, Futatsuyama, 28 Aug–5 Sep 1994, (TYCN). Honshu, Kanto Region: Tochigi Pref., K. Ijima, 1 ♂, 2 ♀ (TYCN). Kamikawa: Horokanai, Nikko, Dorobu, 20 Aug 2007, S. Maehara, 2 ♂ Moshiri-Shirakaba, flowers ofKalopanax septemlobus, (TYCN); Tokyo, Ome City, Mt. Mitake, 27 Aug 2−7 Aug 1994, T. Yasunaga, 1 ♂ (TYCN); Taisetsu 1960, T. Maenami, 1 ♂, 2 ♀ (NIAES). Tokai Region: Lake, Picea jezoensis, 14 Aug 2000, T. Yasunaga Shizuoka Pref., Atami, 300 m, 17 Aug 1996, Y. (TYCN); Mts. Taisetsu, Mt. Asahidake, 200−800 m, Sawada, 2 ♀ (TYCN). Kinki Region: Osaka Pref., 9−11 Aug 1994, T. Yasunaga, 2 ♂, 1 ♀ (TYCN). Kishiwada City, Mt. Katsuragi, 16 Jul 1993, Y. Tokachi: Obihiro City, Iwanaisenkyo, 25 Aug 1995, Nakatani, 1 ♂ (NIAES); Wakayama Pref., same data Y. Todo, 1 ♂ (TYCN). Ishikari: Sapporo City, as for holotype, 2 ♂, 1 ♀ (TYCN). Shikoku, Kochi Ainosato, Picea abies & P. pungens, 2 Aug–6 Sep Pref., Befu, Monobe, light trap, 6 Aug 1994, M. 2001, T. Ogita, 21 ♂, 13 ♀ (TYCN). Shiribeshi: Takai, 2 ♂, 1 ♀ (TYCN); same locality, light trap, 5 Yuni, 25 Aug 1990, S. Kudo, 1 ♀ (TYCN). Oshima: Sep 1997, T. Yasunaga & M. Takai, 1 ♂ (TYCN); Yakumo Town, Nodaoi Park, N42.2083, E140.3717, same locality, 11 Sep 1998, M. Takai, 1 ♂ (TYCN); Downloaded from Brill.com10/06/2021 05:01:49PM via free access

42 Tijdschrift voor Entomologie, volume 158, 2015

Tokushima Pref., Kitou-son, 15 Aug 1998, M. Takai, including collar 0.68–0.76/0.71–0.86; basal width 1 ♀ (TYCN); Tokushima Pref., Mt. Tsurugi, 8 Aug of pronotum 1.25–1.30/1.22–1.40; maximum 1998, M. Takai, 1 ♂ (TYCN). width across hemelytron 1.45–1.70/1.47–1.67; and Diagnosis. Recognized by its small size; dark brown lengths of metafemur, tibia and tarsus 2.49–2.55, to brown basic coloration; comparatively shiny dor- 3.55–4.02, 0.58–0.62/2.57–2.82, 3.57–4.02, sum (Figs 3–5); very long labium (Fig. 41); some- 0.56–0.62. times pale or reddish scutellum (Figs 3, 4); less Etymology. Named in honor of the late Dr. Syoiti mottled hemelytron; often reddish cuneus (Figs 4, Miyamoto who greatly contributed to improving 42); and small, medially membranous comb-shaped knowledge on the Asian Heteroptera; a noun in gen- sclerite (Fig. 66). Closely allied to P. amateras, judg- itive case. ing from the general appearance and shape of the Biology. Records confirm that this new species inhab- female genitalia; however, P. miyamotoi can be distin- its pinaceous conifers broadly, Abies sachalinensis F. guished from P. amateras by the generally darker Schmidt, Pinus densiflora Sieb. & Zucc., Picea basic coloration, weakly shining dorsum, left param- jezoensis (Sieb. & Zucc.) Carrière, Picea glehnii ere sensory lobe lacking a pointed process (cf. Figs 54 (F. Schmidt) Mast., etc. Some populations are obvi- vs. 55), and different shape of the endosoma (cf. ously associated with pinaceous trees introduced for Figs 64 vs. 66). landscaping (e.g., Picea abies (L.) H. Karst., P. pun- Description. Body small; dorsal surface compara- gens Engelm.), and this mirid is gradually expanding tively shining, partly mottled, with uniformly dis- its habitat into artificial zones, such as parks, gar- tributed, dark, simple, semierect setae and sericeous, dens, and university campuses. A few individuals reclining setae. Head pale chestnut brown; maxillary were collected from the inflorescence of deciduous and mandibular plates and apical part of clypeus broadleaf angiosperms, Kalopanax septemlobus somewhat darkened. Antenna dark brown; segment (Thunb.) Koidz. (Araliaceae) and Syringa reticulate I speckled with pale, small spots. Labium shiny dark (Blume) H. Hara (Oleaceae). The adults are occa- brown, somewhat tinged with red, very long, exceed- sionally attracted to light. A univoltine life cycle is ing abdominal segment VII or sometimes reaching assumed with newly emerged adults appearing in genital segment. Pronotum sometimes pale anteri- July in southern Japan and August in northern areas. orly; collar often pale brown, about as thick as or slightly thicker than antennal segment I; scutellum Phytocoris nowickyi Fieber almost uniformly brown or pale brown, sparsely, Figs 13–15, 56, 67, 71 shallowly and transversely rugose; pleura shiny, Phytocoris nowickyi Fieber, 1870: 261 (n. sp.); Schuh, widely chestnut brown except for creamy yellow ven- 1995: 895 (cat.); Kerzhner & Josifov, 1999: tral margins and ostiolar peritreme. Hemelytron 152 (cat., in subgenus Ktenocoris Wagner, rather shining, sometimes partly or widely pale 1954); Yasunaga, 2001: 256 (diag.); brown, not significantly mottled; cuneus sometimes Wachmann et al., 2004: 108 (diag.); Zheng et widely sanguineous, usually with pale, semitranspar- al., 2004: 513 (diag.); Aukema et al., 2013: ent base; membrane smoky brown, with distally pale 226 (cat., in Ktenocoris Wagner). veins and a narrow, pale portion along apical half of cuneus. All coxae and legs chestnut brown (reddish Diagnosis. Recognized by its moderate size; always brown in teneral specimens); each femur and tibia brachypterous female; reddish or sanguineous basic more or less mottled with pale brown spots; tibial coloration; and long antennal segment III. Male spines brown, long; all tarsi brown, except for apical genitalia as in Figs 56, 67; female genitalia as in 71. half of each tarsomere III usually darkened. Abdomen Total body length 6.4–6.8 mm (♂)/6.0–6.2 mm (♀); almost entirely chestnut brown, sometimes pale maximum width across hemelytron 1.8–2.0 mm brown. Male genitalia as in Figs 55, 66. Generally (♂)/2.0–2.2 mm (♀). Detailed diagnostic small in size; sensory lobe of left paramere lacking a characters including the male genitalia and/or pho- pointed process; comb-shaped sclerite widely mem- tographic images are provided by quite a few authors branous except for notches. Female genitalia as in for this widespread species (Wagner 1954, 1974, Fig. 70. Similar in general shape to P. amateras but Wagner & Weber 1964, Yasunaga, 2001, Wachmann sclerotized ring smaller. et al. 2004, Zheng et al. 2004). Measurements. ♂/♀: Total length of body 4.4– Biology. Both adults and immatures are found on 5.1/4.5–5.2; head width including eyes 0.85– various herbs (mostly Asteraceae and Fabaceae) and 0.91/0.79–0.89; vertex width 0.24–0.25/0.30–0.43; some deciduous broadleaf trees, such as willows lengths of antennal segments I–IV 0.78–0.91, (Salix spp.) (Kerzhner 1988, Yasunaga 2001, 2.35–2.63, 1.10–1.21, 0.82–0.89/0.83–0.98, Wachmann et al. 2004). 2.10–2.36, 1.07–1.13, 0.90–0.98; labial length Remarks. The previous records of P. nowickyi from 2.20–2.36/2.27–2.65; mesal length of pronotum southwestern Japan (Honshu west of Kanto area, Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 43

Shikoku and Kyushu) are erroneous, possibly confused thick as antennal segment II; scutellum pale brown, with P. izanamiae n. sp. which has similar reddish col- yellowish brown, pale olive or creamy yellow, some- oration. Phytocoris nowickyi is now known to occur what pruinose or roughened, weakly arched, often only in cold temperate and boreal zones in Japan. with a pair of dark, symmetrical spots or maculae; Material examined. 77 specimens collected between Aug pleura usually widely darkened, except for creamy 1 and Aug 28 from the following localities (NIAES, yellow ventral margins and ostiolar peritreme. NSMT, TYCN): Japan: Hokkaido: Kamikawa: Hemelytron partly shining, very variable in color- Asahikawa City, Etanbetsu; Mts. Taisetsu. Kushiro: ation, variously mottled with greenish, yellowish, Kushiro Marsh, Sarubobira; Kushiro, Miyama. Tokachi: and/or reddish maculae or spots, usually with a Hiroo, Oshirabetsu; Churui, Horokayanto; Ashoro; shiny, pale, squared or circular mark across cuneal Obihiro City. Hidaka: Atsuma, Uryu. Ishikari: Sapporo fracture; cuneus sometimes partly tinged with red; City; Tobetsu, Aoyama; Atsuta, Hattari. Iburi: membrane smoky brown, with pale veins. All Tomakomai City, Takaoka. Shiribeshi: Otaru City. Soya: coxae and trochanters shiny pale brown; metacoxa Wakkanai City; Teshio, Sarobetsu. Oshima: Toyoura. sometimes darkened basally. Leg chestnut brown, Hiyama: Yunotai. Honshu: Aomori Pref., Mutsu City, reddish brown or brown; femora and tibiae vari- Sekine; Tochigi Pref., Yunishigawa & Nikko, Senjogahara; ously speckled with pale spots or maculae; each Nagano Pref., Shioziri City, Mt. Takabotchi-yama; tibia usually with two or three, yellowish annula- Nagano Pref., Japan Alps, Mt. Kiso-komagatake, Mt. tions; tibial spines brown, prominent. Abdomen Norikura & M. Nyugasa; Yamanashi Pref., Kiyosato. weakly shining, usually dark brown or brown, China: Manchuria (= current Heilongjiang Prov.), sometimes tinged with green or olive; ventral sur- Mutanchiang (= Mudanjiang City). face often paler medially. Male genitalia as in Figs 81–83, 91. Comb-shaped sclerite rather ovoid. Phytocoris ohataensis Linnavuori Female genitalia as in Figs 92–94. Sclerotized rings Figs 18–21, 44–50, 81–83, 91–94 elongate ovoid, curved, medially separated from each other. Phytocoris ohataensis Linnavuori, 1963: 74 (n. sp.); Measurements. / : Total length of body 6.1– Schuh, 1995: 896 (cat.); Kerzhner & Josifov, ♂ ♀ 7.3/6.8–8.0; head width including eyes 1.16– 1999: 158 (cat.); Yasunaga, 2001: 256 (diag.). 1.23/1.07–1.23; vertex width 0.24–0.30/0.43–0.47; Phytocoris scotinus Kerzhner, 1977: 8 (n. sp.); Schuh, lengths of antennal segments I–IV 1.38–1.45, 1995: 906 (cat.); Kerzhner & Josifov, 1999: 3.29–3.69, 1.66–1.85, 1.29–1.45/1.53–1.85, 159 (cat.); Yasunaga, 2001: 256 (diag.) n. syn. 3.38–3.69, 1.75–1.97, 1.35–1.54; labial length Diagnosis. Recognized by its comparatively large size; 3.22–3.51/3.38–3.76; mesal length of pronotum generally pale (often creamy or greenish) brown including collar 0.98–1.05/1.04–1.54; basal width basic coloration; matte, variously mottled dorsum; of pronotum 1.66–1.76/1.72–2.10; maximum long labium; bicolorous pronotum with pale ante- width across hemelytron 1.93–2.40/2.24–2.59; rior part; squared or circular, shiny pale mark across and lengths of metafemur, tibia and tarsus 3.38– cuneal fracture; often reddish cuneus; small, oval 3.69, 5.32–5.85, 0.73–0.83/3.38–4.56, 5.96–6.83, comb-shaped sclerite on endosoma (Fig. 83); and 0.86–0.93. elongate-ovoid sclerotized rings that are Biology. This species is found predominantly on medially separated from each other (Figs 92–94). conifers. Sometimes numerous specimens are Very variable species in coloration and size; each attracted to UV light traps. The adults appear from individual possessing a unique design, or color pat- late July to mid-October, later than its closest rela- tern (cf. Figs 18–21, 44–50). tive, P. minakatai n. sp. The immature forms of Redescription. Body elongate oval; basic coloration P. ohataensis are yet to be found although the adults brown but sometimes widely greenish; dorsal sur- were collected from the following plant species: Abies face matte, partly weakly shining, with uniformly sachalinensis F. Schmidt, Larix leptolepis (Sieb. et distributed, dark, simple, semierect setae and seri- Zucc.) Gordon, Picea glehnii (F. Schmidt) Mast., ceous, reclining setae. Head shining, pale brown, P. abies (L.) H. Karst. and P. pungens Engelm. sometimes irregularly darkened; frons sometimes (Pinaceae), Salix spp. (Salicaceae), and Rhus javanica striolate; eye large; vertex narrow. Antenna chestnut L. (Anacardiaceae). This is the most commonly dis- brown; segment I speckled with pale, small spots; tributed mirid in Japan, whereas its habitats in the extreme bases of segments II and III yellowish southwestern parts (Kinki, Chugoku, Shikoku and brown. Labium shining, pale brown, brown or Kyushu regions) are liable to be restricted to the chocolate brown, long, reaching abdominal ster- mountains or hills. num VI or VII. Pronotum usually darkened poste- Remarks. This species and P. scotinus Kerzhner have riorly, with pale posterior margin; calli usually pale, been considered independent species, distinguished sometimes speckled; collar usually pale, about as by their different color patterns. Individuals having Downloaded from Brill.com10/06/2021 05:01:49PM via free access

44 Tijdschrift voor Entomologie, volume 158, 2015 brownish basic coloration with narrowly darkened Phytocoris pallidicollis Kerzhner posterior pronotum (Figs 19–21, 46, 47) were Figs 16, 26–28, 51, 63, 80, 101–102 regarded previously as P. scotinus whereas specimens Phytocoris pallidicollis Kerzhner, 1977: 10 (n. sp.); with often greenish basic coloration with widely Schuh, 1995: 897 (cat.); Kerzhner & Josifov, infuscate posterior pronotum (Figs 18, 44, 45, 50) 1999: 158 (cat.); Yasunaga, 2001: 256 (diag.). were considered to be P. ohataensis. However, detailed comparison of both male and female genitalia in Diagnosis. Recognized by its comparatively large size; more than 40 specimens (from Japan, Korea, and generally pale (often creamy) brown basic coloration; Russian Far East) suggests that what has been limited darkened patterns on dorsum (Figs 26–28); regarded as P. scotinus is unequivocally conspecific almost entirely pale scutellum; pale base of each with P. ohataensis. No significant difference is recog- femur (Fig. 51); broad right paramere (Fig. 63); and nized in the genitalia, although among the examined developed, elongate comb-shaped sclerite of endo- specimens conspicuous intraspecific variation is soma (Fig. 80, p). Very similar to exhibited in coloration and size. P. longipennis, from which it can be distinguished by Material examined. Holotype ♀: Japan, Honshu, Ohata, the shorter antennal segment III which is shorter Mutsu (= Aomori Pref.), 12 Aug 1957 (AMNH). Holotype than the basal width of the pronotum, apical part of (P. scotinus Kerzhner, jr. syn.) ♂: Japan, S. Kuril Islands, the corium without a slash-like mark (Figs 26–28), Kunashiri (Kunashir) Is., Vulkan Mendeleeva, on Picea and more elongate endosomal comb-shaped sclerite glehnii, 11 Aug 1973, I. M. Kerzhner (ZMAS). Paratypes (cf. Figs 79, r vs. 80, p). (P. scotinus Kerzhner, jr. syn.): S. Kuril Islands, Kunashiri Redescription. Body elongate oval; basic coloration (Kunashir) Is., same data as for the holotype, 1 ♂ (TYCN). whitish or creamy brown; dorsal surface matte, usu- Russia: S. Sakhalin: Novoaleksandrovsk, 25 Aug 1973, ally speckled with dark patterns on posterior hemely- Ermolenko, 1 ♀ (TYCN). tron, with uniformly distributed, sericeous, reclining In addition to the type-series, more than 500 setae and rather sparsely or regionally distributed, specimens collected between July 22 and October 11 dark, semierect setae. Head pale brown; maxillary from the following localities (AMNH, NIAES, plate sometimes partly darkened. Antenna pale NSMT & TYCN): Japan: Hokkaido, Rishiri Island, brown; segment I speckled with dark brown macu- Oniwaki; Kamikawa, Kushiro, Horokanai, Ishikari, lae; apical part of segment II more or less darkened; Iburi, Shiribeshi, Oshima, Tokachi and Hiyama segments III and IV brown or dark brown. Labium areas (almost all parts from coasts to subboreal zones, shiny pale brown, long, reaching abdominal sternum up to 800 m, including parks, gardens and cam- VI or VII; apical part of segment IV widely reddish puses). Honshu, Tohoku Region: Aomori & brown. Pronotum unicolorous pale (posterior part Yamagata Prefs. widely; Iwate Pref., Mt. Hayachine; rarely obscured), with a pair of bundles of dark setae Akita Pref., Akita City; Miyagi Pref., Mt. Zaou. near posterior margin; scutellum and pleura almost Koushin’etsu Region: Niigata Pref.: Tsubame immaculate; ostiolar peritreme creamy yellow. Hot Spa; Tsunan Town; Myokoh-kogen Town; Hemelytron widely pale, less mottled; dark maculae Gifu/Nagano Prefs.: Mt. Nishi-hotaka, 2,100m or spots present mostly along inner margins of alt.; Nagano Pref.: Kaida Village, Kamikochi, and corium and cuneus; apical half of cuneus usually Matsumoto City. Kanto Region: Gunma Pref., widely darkened; membrane pale grayish brown, Ozagahara; Tokyo, Okutama, Nippara; Tokyo, speckled with pale, semitransparent spots. All coxae Mt. Takao; Saitama Pref., Okuchichibu. Kinki pale brown; apical 4/5−1/2 of each femur speckled Region: Nara Pref.: Ohdaigahara. Kumano area: with brown maculae or spots (Fig. 51); each tibia Wakayama Pref., Maenokawa Forest, Hatenashi usually with three dark annulations; tibial spines red- Mts., Mt. Ando (1,100 m), Mt. Oto-san, and moun- dish brown, prominent; all tarsi brown. Abdomen tains around Tanabe City. Shikoku, Ehime Pref.: pale or creamy brown; male pygophore usually shiny Omogo & Ishizuchi mountain areas. Kochi castaneous. Male genitalia as in Figs 63, 80. Right Pref.: Monobe, Nishikuma; Kuroson; Tengu-kogen paramere broad, with distinct row of notches (plateau); Tosashimizu, Imanoyama. Tokushima (Fig. 63); comb-shaped sclerite longest among Pref.: Mt. Tsurugi; Kitoh-son; Naka-gun, Mt. Japanese congeners (Fig. 80, p). Female genitalia as Takashiro (1,450 m). Kyushu, Fukuoka Pref., in Figs 101, 102. Bursa copulatrix rather widened. Mt. Hikosan; Saga Pref., Mt. Sefuri; Nagasaki Measurements. ♂/♀: Total length of body 6.3–6.6/ Pref., Isahaya City, Mt. Gokahara (1,050 m); 7.0–7.5; head width including eyes 1.04–1.11/1.01– Kumamoto Pref., Izumi-mura; Miyazaki Pref., 1.05; vertex width 0.27–0.30/0.36–0.40; lengths of Shiiba-mura. Korea: Gyeongsangbuk-do, Yeongju- antennal segments I–IV 1.26–1.48, 3.13–3.23, si, Huibangsa; Gangwon-do, Hoengseong, Mt. 1.50–1.54, 1.23–1.42/1.63–1.73, 3.38–3.45, 1.59– Cheongtae. Russia: S. Primorskij Krai, Khasanskij 1.79, 1.26–1.42; labial length 2.79–2.83/2.99– Dist., Ryazanovka; Ussury area, Ussurijsk Reserve. 3.14; mesal length of pronotum including collar Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 45

0.92–0.96/1.04–1.11; basal width of pronotum (Figs 6–8); long labium (Fig. 43); and distinctly 1.72–1.79/1.81–1.88; maximum width across hem- notched sensory lobe of left paramere (Fig. 57). Most elytron 2.06–2.13/2.42–2.46; and lengths of meta- similar to P. miyamotoi, from which this new species femur, tibia and tarsus 3.32–3.42, 5.22–5.26, can be distinguished by the distinctly mottled head, 0.64–0.67/3.69–4.00, 5.53–5.94, 0.67–0.71. pronotum and hemelytron, and spinose sensory lobe Biology. This species is assumed to be associated pre- of the left paramere (cf. Figs 55 vs. 57), in addition dominantly with deciduous trees, and the newly to being allopatric. emerged, teneral adults have been found from Alnus, Description. Body elongate oval, subparallel-sided, Juglans, Salix, and Tilia. The immature forms were small; basic coloration chestnut brown; dorsum collected from a Japanese lime, Tilia japonica (Miq.) weakly shining, variously mottled with pale brown Simonk. The population density of P. pallidicollis is maculae, with uniformly distributed, sericeous, generally low, except for broadleaf forests in reclining setae and rather sparsely distributed, dark, Hokkaido, where several deciduous trees often yield semierect setae. Head pale brown, with irregular a few specimens when sweeping the leaves and dark and reddish pattern. Antenna chocolate brown; branches during the day. segment I with several pale spots; extreme bases of Material examined. Holotype ♂: Japan, S. Kuril Islands, segments II and III yellowish brown. Labium shiny Kunashiri (Kunashir) Is., Dubovoe, on Betula, 31 Aug brown, partly reddish, very long, exceeding abdomi- 1973, I. M. Kerzhner (ZMAS). Paratypes: Japan, S. Kuril nal sternum VII. Pronotum weakly shining, mottled, Islands, same data as for the holotype, 1 ♂ (TYCN). widely darkened posteriorly except for pale posterior Russia, S. Sakhalin, Novoaleksandrovsk, 10 Aug 1973, margin; calli with several dark spots; collar mottled, Kuporosov, 1 ♀ (TYCN). about as thick as antennal segment II; scutellum In addition to the type-series, 39 specimens col- rather flat, pale brown, with variable, symmetrical, lected between July 6 and September 8 from the fol- dark maculae or spots; pleura widely chocolate lowing localities (NSMT, TYCN): Japan: Hokkaido, brown except for yellow ventral margins; propleuron Kamikawa: Mts. Taisetsu, Mt. Asahidake (200−800 medially with a yellow stripe that is sometimes con- m); Horokanai, Moshiri-Shirakaba. Tokachi: Ashoro, tinuing via episternum to epimeron; ostiolar perit- Kami-Ashoro; Obihiro City, Inada. Ishikari: Sapporo reme creamy yellow. Hemelytron matte, dark brown, City, Hokkaido University Campus; Jozankey; variously mottled with pale brown and/or reddish Tobetsu, Aoyama. Shiribeshi: Niseko, Mt. Chisenupuri. patterns; apex of corium with a pale, shiny, triangu- Honshu, Tochigi Pref., Nikko & Yunishigawa. lar mark; basal 1/3 of cuneus reddish brown, with a Shikoku, Kochi Pref., Nishikuma & Kuroson. Kyushu, dark spot at inner basal corner; membrane pale Kumamoto Pref., Izumi-mura, Momiki. smoky brown, with distally pale veins. All coxae and trochanters pale brown; bases of meso- and meta- Phytocoris protobothrops Yasunaga & Schwartz, coxae reddish or dark brown. Leg chocolate brown, n. sp. partly tinged with red; all femora and tibiae more or Figs 6–8, 43, 57, 68, 72 less speckled with yellowish brown maculae and/or spots; basal 1/4 of metafemur pale; tibial spines dark Type material. Holotype ♂: Japan, Ryukyus, Okinawa reddish brown, prominent. Abdomen almost entirely Is., Kunigami Village, Okuni Pass, 25 Jun 1999, dark reddish brown. Male genitalia as in Figs 57, 68. T. Yasunaga & M. Takai (AMNH_PBI 00380373) Pygophore with swell at base of each paramere; sen- (AMNH). Paratypes: Japan, Ryukyus, Amami- sory lobe of left paramere with four distinct spines Oshima Is., Uken Village, 19 May 1999, K. Takahashi, (Fig. 57); endosoma with crescent comb-shaped 1 (TYCN). Okinawa Is., Kunigami Village, Yona, ♂ sclerite and rounded LBS (Fig. 68). Female genitalia light trap, N26.763155, E128.215979, T. Yasunaga, 1 as in Fig. 72. Sclerotized ring narrow but widened. (TYCN); Kunigami Village, Yona, 19 Oct 1987, ♂ Measurements. / : Total length of body 4.2– M. Tomokuni, 1 (NSMT); Kunigami Village, Mt. ♂ ♀ ♀ 5.4/4.6–5.3; head width including eyes 0.95– Yonahadake, on rotten log, N26.7777, E128.2209, 4 1.06/0.93–0.98; vertex width 0.28–0.35/0.39–0.42; Oct 2002, T. Yasunaga, 1 (TYCN); Kunigami ♂ lengths of antennal segments I–IV 0.85–1.01, 2.42– Village, Mt. Yonahadake, 20 Oct 1987, M. Tomokuni, 2.75, 1.02–1.11, 1.10–1.16/0.95–0.98, 2.25–2.38, 1 , 1 (NSMT); Kunigami Village, Hiji River, ♂ ♀ 1.00–1.03, 1.02–1.03; labial length 2.08– N26.7085, E128.1965, 30 Mar 1999, M. Takai, 1 ♂ 2.26/2.27–2.36; mesal length of pronotum includ- (TYCN); Nago City, Uehara, 7 Apr 1991, M. Hayashi ing collar 0.73–0.84/0.80–0.84; basal width of et al., 4 (TYCN); Nago City, Oo’ura, 4 Apr 1990, ♀ pronotum 1.22–1.52/1.42–1.50; maximum width M. Hayashi et al., 2 , 1 (TYCN). ♂ ♀ across hemelytron 1.51–1.96/1.71–1.87; and Diagnosis. Recognized by its small size; chestnut lengths of metafemur, tibia and tarsus 2.20–2.82, brown basic coloration; rather shiny dorsum vari- 3.57–4.22, 0.49–0.59/2.64–2.77, 3.99–4.17, ously speckled with pale and/or reddish portions 0.53–0.59. Downloaded from Brill.com10/06/2021 05:01:49PM via free access

46 Tijdschrift voor Entomologie, volume 158, 2015

Etymology. Named from the generic name of a pit Fieber, F.X., 1870. Dodecas Neuer Gattungen und Neuer viper serpent, Protobothrops flavoviridis (Hallowell), Arten Europäischer Hemiptera. – Verhandlungen der which is almost exactly sympatric with this new Kaiserlich-Königlichen Zoologisch-Botanischen mirid species and has a similar mottled pattern on Gesellschaft in Wien 20: 243–264, pls. 5, 6. the trunk; original meaning in Greek: protos (= first, Flor, G., 1861. Die Rhynchoten Livlands in systematischer primary) + bothros (= pit, hole) + ops or opos (= face, Folge Beschrieben. Band II. – C. Schulz, Dorpat, countenance). 638 pp. Kerzhner, I.M., 1977. New and little-known species of Biology. Most specimens were collected using UV light traps. A teneral adult was found on a rotten log Heteroptera from the Far East of the USSR. – Trudy Zoolologicheskogo Instituta Akademiya Nauk SSSR on the floor of well-preserved, subtropical broadleaf 62(1976): 6–35. (In Russian) forest, but its breeding host is unknown. A bivoltine Kerzhner, I.M., 1988. Infraorder Cimicomorpha. 21. life cycle is assumed for P. protobothrops, as newly Family Miridae (Capsidae). – In: Ler, P.A. (Ed.), emerged adults were collected in both April and Opredelitel’ nasekomykh Dal’nego Vostoka SSSR October. Generally, the population density of this (Keys to the identification of insects of the Soviet Far new species is low. East), Vol. 2: Homoptera and Heteroptera), pp. 778– 857. Nauka, Leningrad, USSR. (In Russian) Kerzhner, I.M. & M. Josifov, 1999. Miridae Hahn, 1833. Acknowledgements – In: Aukema, B. & C. Rieger (Eds), Catalogue of the Special thanks are due to the late Drs S. Miyamoto Heteroptera of the Palearctic Region, Vol. 3, (Fukuoka, Japan) and I. M. Kerzhner (ZMAS), and Cimicomorpha II, pp. 1–576. The Netherlands the late Mr S. Gotoh (Tanabe, Wakayama Pref.), Entomological Society, Amsterdam, Netherlands. who had been positively supporting this study with Linnavuori, R.E., 1963. Contribution to the Miridae providing valuable suggestions and specimens but so fauna of the Far East III. – Annales Entomologici Fennici 29: 73–82. regretfully passed away before completion of this Miyamoto, S. & T. Yasunaga, 1993. A new genus of the paper. We are also much indebted to the following mirid bug, Adelphocorisella (Heteroptera, Miridae) with individuals or institutions for offering or loaning two new species from Japan. – Proceedings of the materials: Dr R.T. Schuh (AMNH); Dr T.J. Henry Japanese Society of Systematic Zoology 49: 47–52. (Systematic Entomology Laboratory, USDA, Schuh, R.T., 1995. Plant Bugs of the World (Insecta: Washington, D.C.); Prof. S.H. Lee (Seoul National Heteroptera: Miridae). Systematic Catalog, University, Korea); Dr M. Webb (Natural History Distributions, Host List and Bibliography. – The New Museum, London); Messrs T. Kawasawa, M. York Entomological Society, New York, NY, USA, xii + Kawamura, M. Takai, T. Befu and I. Yamashita 1329 pp. (Kochi Pref.); Mr S. Sakurai (Niigata Pref.); Emer. Schuh, R.T., 2002–2014. On-line Systematic Catalog of Prof. M. Hayashi (Saitama Univ.); Dr K. Takahashi Plant Bugs (Insecta: Heteroptera: Miridae). Available (Tsukuba, Ibaraki Pref.); Drs M. Tomokuni and S. online at http://research.amnh.org/pbi/catalog/ Nomura (NSMT); Dr Y. Nakatani (NIAES); Prof. T. (accessed on 1 August 2015). Hirowatari and Dr S. Kamitani (Kyushu Univ.); Stonedahl, G.M., 1988. Revision of the mirine genus Prof. K. Konishi (Ehime Univ.); Dr K. Yamada Phytocoris Fallén (Heteroptera: Miridae) for western (TKPM); Dr K. Yoshizawa (Hokkaido Univ.); Mr B. North America. – Bulletin of the American Museum of Shishido (Himeji, Hyogo Pref.); and R.K. Duwal Natural History 188: 1–257. and many pupils of T. Yasunaga. Mr M. Takai was Wachmann, E., A. Melber & J. Deckert, 2004. Wanzen. kind enough to provide quite a few vivid images Band 2. – Goecke & Avers, Keltern, Germany, 278 (Figs 2, 4–6, 8, 9–12, 18, 21, 23, 27–28). pp. Wagner, E., 1954. Eine Beitrag zur Systematik der Gattung Phytocoris Fall. (Hem. Heteropt. Miridae). – Nachrichten des Naturwissenschaftlichen Museums References der Stadt Aschaffenburg 42: 1–44, taf. I–VI. Aukema, B., C. Rieger & R. Wolfgang (Eds), 2013. Wagner, E., 1968. Die Untergattung Leptophytocoris Catalogue of the Heteroptera of the Palearctic Region, (Hemiptera, Heteroptera, Miridae). – Reichenbachia Vol. 6, supplement. – The Netherlands Entomological 10: 103–111. Society, Amsterdam, Netherlands, xxiv + 629 pp. Wagner, E., 1974. Die Miridae Hahn, 1831, des Cassis, G., 2008. The Lattinova complex of austromirine Mittelmeerraumes und der Makaronesischen Inseln plant bugs (Hemiptera: Heteroptera: Miridae: (Hemiptera, Heteroptera), Teil. 1. – Entomologische Orthotylinae). – Proceedings of the Entomological Abhandlungen 37 Suppl., iii + 484 pp. Society of Washington 110: 845–939. Wagner, E. & H.H. Weber, 1964. Héteroptères Miridae. – Fallén, C.F., 1814. Specimen novam Hemiptera dispo- In: Faune de France 67: 1–592. Librairie de la Faculte nendi methodum exhibens. – Lundae, 26 pp. des Sciences, Paris, France.

Downloaded from Brill.com10/06/2021 05:01:49PM via free access

Yasunaga and Schwartz: Phytocoris of Japan 47

Westwood, J.O., 1840. Synopsis of the Genera of British Yasunaga, T., 2001. Family Miridae Hahn, plant bugs. – Insects. – Longman, Orme, Brown, Green, and In: Yasunaga T., M. Takai & T. Kawasawa (Eds), A Longmans, London, UK, 158 pp. Field Guide to Japanese Bugs II, pp. 1–96, 111–351. Yasunaga, T., 1990a. A revision of the genus Adelphocoris Zenkoku Noson Kyoiku Kyokai Publ. Co. Ltd., Tokyo, Reuter (Heteroptera, Miridae) from Japan, Part I. – Japan. (In Japanese) Japanese Journal of Entomology 58: 606–618. Yasunaga, T. & M.D. Schwartz, 2007. Revision of the mir- Yasunaga, T., 1990b. A revision of the genus Adelphocoris ine plant bug genus Philostephanus Distant and allies Reuter (Heteroptera, Miridae) from Japan, Part II. – (Heteroptera: Miridae: Mirinae: Mirini). – Tijdschrift Japanese Journal of Entomology 58: 725–733. voor Entomologie 150: 100–180. Yasunaga, T., 1997. Revision of the mirine genus Zheng, L.Y., N. Lu, G. Liu & B. Xu, 2004. Hemiptera, Creontiades Distant and allies from Japan (Heteroptera, Miridae, Mirinae. Fauna Sinica, Insecta, Vol. 33. – Miridae). Part I. True members of Creontiades. – Science Press, Beijing, China, xix+797 pp., 8 pls. (In Japanese Journal of Entomology 65: 541–555. Chinese, with English keys and descriptions of new Yasunaga, T., 1998. Revision of the mirine genus taxa) Creontiades Distant and allies from Japan (Heteroptera: Miridae). Part III. Neomegacoelum gen. n. and exotic Received: February 15, 2015 new taxa, new synonymy and new combinations. – Accepted: May 18, 2015 Entomological Science 1: 63–70.

Downloaded from Brill.com10/06/2021 05:01:49PM via free access