Physopelta (Neophysopelta) gutta gutta ( Burmeister, 1834 )

Souma, Jun & Ishikawa, Tadashi, 2021, A taxonomic review of the genus Physopelta (Hemiptera: Heteroptera: Largidae) from Japan, Korea, and Taiwan, with a new record of Ph. parviceps from China, Zootaxa 4951 (3), pp. 461-491 : 465-467

Physopelta (Neophysopelta) gutta gutta ( Burmeister, 1834)

( Figs. 1A, B , 3A , 4A , 5A , 6A , 7A , 8A , 9A , 10A , 11A , 12A , 13A , 14A )

Lygaeus ( Pyrrhocoris Fall. , Platynotus Schill. ) gutta Burmeister, 1834: 30 . Syntype (s): macropterous ♂, Philippines: Manila env; ZMHB ( Stehlík 2013: 528).

Pyrrhocoris gutta: Burmeister (1835: 285) (new combination).

Physopelta bimaculata Stål 1855: 186 . Syntype (s): macropterous sex unknown; Indonesia: Java; NHRS ( Stehlík 2013: 528).

Synonymized by Stål (1861: 195).

Physopelta gutta: Stål (1861: 195) (new combination).

References. Walker (1873: 18) (distribution); Scott (1874: 291) (distribution); Esaki (1926: 157) (distribution); Hussey (1929: 30) (distribution); Blöte (1931: 99) (distribution); Kato (1940: 704) (distribution); Esaki (1952: 230) (distribution); Takara (1957: 54) (distribution); Miyamoto (1965a: 89) (monograph); Sawada & Watanabe (1969: 7) (distribution); Watanabe & Sohma (1972: 8) (distribution); Yamaguchi et al. (1973: 90) (distribution); Japanese Society of Applied Entomology and Zoology (1980: 24) (pest); Tomokuni (1981: 108) (distribution); Urata (1981a: 429) (distribution, as Physopelta guttata ); Urata (1981b: 738) (distribution); Ahmad & Abbas (1987: 137) (distribution); Higuchi & Sato (1988: 84) (distribution); Hatada (1991: 28) (distribution); Lee & Kwon (1991: 50) (distribution); Yasunaga et al. (1993: 197) (monograph); Ehira (1996: 53) (distribution); Tomokuni et al. (2000: 43) (distribution); Kerzhner (2001: 246) (checklist: Palaearctic); Kwon et al. (2001: 300) (checklist: Korea); Hayashi (2002: 144) (distribution); Tomokuni & Ishikawa (2002: 175) (distribution); Hiromori (2003: 75) (distribution); Umeya & Okada (2003: 546) (biology); Stehlík (2004: 2) (distribution); Yang (2003: 200) (male genitalia, as Physopelta guttata ); Japanese Society of Applied Entomology and Zoology (2006: 59) (pest); Tomokuni (2005: 403) (distribution); Tomokuni & Hayashi (2006: 298) (distribution); Tomokuni (2006: 352) (distribution); Stehlík (2007: 117) (distribution); Yang (2007: 6) (male genitalia, as Physopelta guttata ); Miyamoto (2008: 155) (monograph); Komatsu et al. (2009: 82) (distribution); Rédei et al. (2009: 8) (monograph); Yazaki (2009: 32) (distribution); Kohno et al. (2012: 400) (monograph); Yano et al. (2012: 92) (distribution); Aukema et al. (2013: 401) (checklist: Palaearctic); Ito (2013: 42) (distribution); Stehlík (2013: 528) (monograph); Zheng & Lin (2013: 148) (distribution); Kanai & Moriyama (2014: 39) (distribution); Nozaki et al. (2015: 30) (distribution); Ishikawa (2016: 477) (checklist: Japan); Komatsu (2016: 37) (distribution); Nozaki et al. (2016: 89) (distribution); Kanai (2017: 16) (distribution); Ahn et al. (2018: 203) (distribution); Hayashi & Kadowaki (2018: 313) (distribution); Souma et al. (2019: 71) (distribution); Ito et al. (2020: 116) (distribution); Okuda (2020: 47) (distribution).

Material examined. Non-types , a total of 1,492 specimens (spec.) of macropterous morph deposited in ELKU, KUM and TUA from the following localities: JAPAN: Honshu: Tochigi Pref. (1 spec.) ; Ibaraki Pref. (1 spec.) ; Saitama Pref. (12 spec.) ; Chiba Pref. (2 spec.) ; Tokyo Metropolis (23 spec.) ; Kanagawa Pref. (223 spec.) ; Shizuoka Pref. (14 spec.) ; Aichi Pref. (3 spec.) ; Wakayama Pref. (10 spec.) ; Tottori Pref. (1 spec.) ; Hiroshima Pref. (3 spec.) ; Yamaguchi Pref. (2 spec.) . Izu Islands : Oshima Is. (7 spec.) ; Miyake Is. (37 spec.) ; Mikura Is. (15 spec.) ; Kozu Is. (6 spec.) ; Hachijo Is. (22 spec.) ; Aogashima Is (9 spec.) . Shikoku : Ehime Pref. (4 spec.) ; Kochi Pref. (2 spec.) . Kyushu : Fukuoka Pref. (34 spec.) ; Saga Pref. (3 spec.) ; Nagasaki Pref. (3 spec.) ; Oita Pref. (2 spec.) ; Kumamoto Pref. (56 spec.) ; Kagoshima Pref. (22 spec.) . Iki Island (6 spec.) . Tsushima Island (17 spec.) . Danjo Islands : Onna Is. (3 spec.) . Koshiki Islands : Kamikoshiki Is. (3 spec.) ; Shimokoshiki Is. (12 spec.) . Ryukyu Islands: Osumi Islands : Yaku Is. (5 spec.) . Kuro Is. (18 spec.) . Tokara Islands : Kuchinoshima Is. (1 spec.); Taira Is. (1 spec.) . Amami Islands : Amami-Oshima Is. (31 spec.) ; Tokunoshima Is. (11 spec.) ; Okinoerabu Is. (1 spec.) ; Yoron Is. (1 spec.) . Okinawa Islands : Okinawa-Honto Is. (108 spec.) ; Iheya Is. (17 spec.) ; Noho Is. (1 spec.) ; Izena Is. (1 spec.) ; Ie Is. (3 spec.) ; Yabuchi Is. (1 spec.) ; Tonaki Is. (2 spec.) ; Kume Is. (1 spec.) ; Tokashiki Is. (1 spec.) . Yaeyama Islands : Ishigaki Is. (7 spec.) ; Kohama Is. (5 spec.) ; Iriomote Is. (95 spec.) ; Yonaguni Is. (9 spec.) . KOREA: Korean Peninsula: (1 spec.) ; Chejudo Island (1 spec.) . TAIWAN: main island (610 spec) .

Diagnosis. Recognized among other species of Physopelta by a combination of the following characters: only macropterous morph known; body 3 times as long as maximum width across fore wings ( Fig. 1A, B ); calli and disc of pronotum brown ( Fig. 5A ); scutellum dark brown, without pale callosities; membrane of fore wing dark brown ( Fig. 6A ); abdominal sternite orange with black markings on sutures ( Figs. 3A , 4A ); compound eye more than 0.3 times as wide as vertex in dorsal view; antennomere I shorter than antennomere II; antennomere II nearly clavate; calli in male convex; punctures of scutellum sparser in central part than in marginal part, smaller than punctures of pronotum; anterior margins of fore wings nearly parallel to each other in rest; procoxal projection straight, more than 1.5 times as long as its maximum width ( Fig. 7A ); protrochanteral wrinkles present; male profemur more than 2 times as wide as mesofemur at widest part of each; male protibia lacking tooth at apex, with a single row of denticles throughout its length ventrally; peritreme of scent gland ostiole crescent-shaped, protruding posterolaterad ( Figs. 8A , 10A ); infolding of ventral rim of genital capsule less convex in middle part ( Fig. 9A ); outer margin of endophallic reservoir outgrowth not emarginate in apical part ( Fig. 11A ); stem of paramere not emarginate in basal part ( Fig. 12A ); crown of paramere at apex convex in posterior view; and inner margins of ring sclerites nearly parallel to each other ( Fig. 13A ).

Description of genitalia. Genital capsule ( Fig. 9A ) spherical, semicircular in ventral view, smooth on surface; infolding of ventral rim less convex in middle part. Phallus ( Fig. 11A ) oblong; capitate process membranous; basal plate and phallotheca coriaceous; conjunctiva with two pairs of partly sclerotized conjunctival appendages; endophallic reservoir with a pair of outgrowths; outer margin of outgrowth not emarginate in apical part. Paramere ( Fig. 12A ) longer than its maximum width across crown; stem not emarginate in basal part; crown at apex not convex in posterior view.

Female terminalia ( Fig. 13A ) triangular in anterior view; valvulae VIII and IX combining to form an ovipositor, fused with valvifers VIII and IX, respectively; ring sclerites coriaceous in marginal part, membranous in central part, protruding inward, with inner margins nearly parallel to each other. Spermatheca ( Fig. 14A ) membranous; apical receptacle spherical; intermediate part in width uniform; spermathecal duct apically widened.

Remarks. Physopelta (Neophysopelta) gutta is divided into two subspecies, namely, Ph. (N.) g. gutta ( Burmeister, 1834) and Ph. (N.) g. famelica Stål, 1863, and the eastern Asian population corresponds to the nominotypical subspecies ( Stehlík 2013).

Distribution. Japan: Oshima-Oshima Island, Honshu, Izu Islands (Oshima Is, Toshima Island, Niijima Island, Miyake Island, Mikura Island, Kozu Island, Hachijo Island, Aogashima Island), Sado Island., Oki Islands, Shikoku, Kyushu, Iki Island, Tsushima Island, Goto Islands, Danjo Islands (Otoko Island, Onna Island), Amakusa Islands (Shimo Island), Koshiki Islands (Kamikoshiki Island, Shimokoshiki Island), Ryukyu Islands (Yaku Island, Kuchinoerabu Island, Kuro Island, Kuchinoshima Island, Nakanoshima Island, Taira Island, Akuseki Island, Takara Island, Amami-Oshima Island, Tokunoshima Island, Okinoerabu Island, Yoron Island, Okinawa-Honto Island, Iheya Island, Noho Island, Izena Island, Aka Island, Ie Island, Yabuchi Island, Tonaki Island, Kume Island, Tokashiki Island, Iotorishima Island, Ishigaki Island, Kohama Island, Iriomote Island, Yonaguni Island) ( Scott 1874; Blöte 1931; Yasunaga et al. 1993; Kohno et al. 2012; present study, etc.); Korea (southern part of Korean Peninsula, Chejudo Island) ( Lee & Kwon 1991; Kwon et al. 2001; Kerzhner 2001; Ahn et al. 2018; present study); Taiwan (main island) ( Esaki 1926; Blöte 1931; Kerzhner 2001; Rédei et al. 2009; Stehlík 2013; present study); Afghanistan ( Stehlík 2013); Bangladesh ( Walker 1873); Bhutan ( Stehlík 2013); Brunei ( Stehlík 2013); Cambodia ( Stehlík 2013); China ( Stehlík 2013); India ( Stehlík 2007, 2013); Indonesia ( Blöte 1931); Laos ( Stehlík 2013); Malaysia ( Stehlík 2013); Myanmar ( Stehlik 2013); Nepal ( Stehlík 2004, 2013); Pakistan ( Stehlík 2013); Philippines ( Stehlík 2013); Singapore ( Stehlík 2013); Sri Lanka ( Stehlík 2013); Thailand ( Stehlík 2013); Vietnam ( Stehlík 2013).

Host plant. In Japan, Mallotus japonicus (Euphorbiaceae) is a host plant for Physopelta (Neophysopelta) gutta gutta because nymphs have been observed to feed on the seeds in the field, whereas adults feed on various flowers and fruits from five plant families other than Euphorbiaceae , namely, Acer spp. ( Sapindaceae ), Amygdalus persica L. ( Rosaceae ), Callistemon sp. ( Myrtaceae ), Citrus spp. ( Rutaceae ), Fortunella spp. ( Rutaceae ), Poncirus trifoliata (L.) Raf. ( Rutaceae ), Prunus domestica L. ( Rosaceae ), and P. salicina Lindl. (Miyamoto 1965; Japanese Society of Applied Entomology and Zoology 1980, 2006; Yasunaga et al. 1993; Umeya & Okada; 2003; Miyamoto 2008; Kohno et al. 2012; Komatsu 2016; present study).

This species is found on various woody angiosperms belonging to Rosaceae , Rutaceae , and Sapindaceae in Korea (cf. Kwon et al. 2001) and on Mallotus spp. and Poaceae spp. in Taiwan ( Zheng & Lin 2013).

Biology. Adults and nymphs are attracted to artificial light (Miyamoto 1965; Tomokuni 1993; Miyamoto 2008; Kohno et al. 2012; present study).

In Japan proper, adults are collected in almost all seasons ( Yasunaga et al. 1993; Kohno et al. 2012; Nozaki et al. 2016; Okuda 2020; present study, etc.), suggesting that the overwintering stage is adult.

Teneral adults were observed from August to September in Japan proper (present study). Nymphs were observed from August to September in Japan proper ( Kohno et al. 2012; present study). In Japan proper, host plant of Physopelta (Neophysopelta) gutta gutta , Mallotus japonicus blooms from June to July, and mature seed is fallen to ground surface from September to November ( Tomikawa et al. 2013). Due to one season blooming of its host plant and emergence time of teneral adult and nymph, Ph. (N.) gutta gutta could be univoltinism in Japan proper. Number of generations in other distribution areas is unknown.

Figures

FIGURE 1. Habitus images of four Physopelta (Neophysopelta) species, dorsal view: Ph. (N.) gutta gutta, male (A) and female (B) from Japan proper; Ph. (N.) fusciscutellata sp. nov., male from the main island of Taiwan (C) and female from the Ryukyu Islands, Japan (D); Ph. (N.) quadriguttata, male (E, G) and female (F) from the main island of Taiwan; Ph. (N.) lutaspidata sp. nov., male from the main island of Taiwan (H). Arrows mark antenna with antennomeral oligomery and malformed calli. Scale bar: 5.0 mm.

FIGURE 3. Habitus images of five Physopelta (Neophysopelta) species, lateral view: Ph. (N.) gutta gutta, female from Japan proper (A); Ph. (N.) fusciscutellata sp. nov., female from the Ryukyu Islands, Japan (B); Ph. (N.) lutaspidata sp. nov., male from the main island of Taiwan (C); Ph. (N.) parviceps, male from Japan proper (D) and female from the Ryukyu Islands (E); Ph. (N.) quadriguttata, male from the main island of Taiwan (F). Scale bar: 5.0 mm.

FIGURE 4. Habitus images of five Physopelta (Neophysopelta) species, ventral view: Ph. (N.) gutta gutta, female from Japan proper (A); Ph. (N.) fusciscutellata sp. nov., female from the Ryukyu Islands, Japan (B); Ph. (N.) lutaspidata sp. nov., male from the main island of Taiwan (C); Ph. (N.) parviceps, male from Japan proper (D) and female from the Ryukyu Islands (E); Ph. (N.) quadriguttata, male from the main island of Taiwan (F). Scale bar: 5.0 mm.

FIGURE 5. Male pronotum and scutellum of five Physopelta (Neophysopelta) species, dorsal view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); macropterous morph of Ph. (N.) parviceps from Japan proper (C, D); macropterous morph of Ph. (N.) parviceps from the Ryukyu Islands, Japan (E); brachypterous morph of Ph. (N.) parviceps from Japan proper (F); Ph. (N.) fusciscutellata sp. nov. (G); Ph. (N.) quadriguttata (H). Arrows mark malformed calli. Scale bars: 1.0 mm.

FIGURE 6. Left fore wing of five macropterous species of Physopelta (Neophysopelta), dorsal view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Scale bar: 1.0 mm.

FIGURE 7. Procoxa of five Physopelta (Neophysopelta) species, ventral view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Abbreviation: pp = procoxal projection. Scale bars: 0.2 mm.

FIGURE 8. Left scent gland ostiole of five Physopelta (Neophysopelta) species, ventral view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Abbreviation: pe = peritreme. Scale bars: 0.1 mm.

FIGURE 9. Genital capsule of five Physopelta (Neophysopelta) species, posterior view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Abbreviations: vif = infolding of ventral rim; cr = crown of paramere. Scale bars: 0.5 mm.

FIGURE 10. Left scent gland ostiole of five Physopelta (Neophysopelta) species, lateral view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Abbreviation: pe = peritreme. Scale bars: 0.2 mm.

FIGURE 11. Phallus of five Physopelta (Neophysopelta) species, dorsal view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Abbreviations: ad = aedeagus; bb = basal plate bridge; bp = basal plate; co = conjunctiva; cp = capitate process; dc = dorsal connective; ds = ductus seminis; ed = endophallic duct; eo = endophallic reservoir outgrowth; ep = erection fluid pump; er = endophallic reservoir; pt = phallotheca; 1ca, 2ca = first and second conjunctival process. Scale bars: 0.5 mm.

FIGURE 12. Left paramere of five Physopelta (Neophysopelta) species, dorsal view: Ph. (N.) gutta gutta (A); Ph. (N.) lutaspidata sp. nov. (B); Ph. (N.) parviceps from Japan proper (C); Ph. (N.) parviceps from the Ryukyu Islands, Japan (D); Ph. (N.) fusciscutellata sp. nov. (E); Ph. (N.) quadriguttata (F). Abbreviations: ba = basal apodeme; cr = crown; ne = neck; st = stem. Scale bars: 0.1 mm.

FIGURE 13. Female terminalia of four Physopelta (Neophysopelta) species, anterior view: Ph. (N.) gutta gutta (A); Ph. (N.) parviceps from Japan proper (B); Ph. (N.) parviceps from the Ryukyu Islands, Japan (C); Ph. (N.) fusciscutellata sp. nov. (D); Ph. (N.) quadriguttata (E). Abbreviations: go = gonangulum; lt9 = laterotergite IX; mt8 = mediotergite VIII; ra8, ra9 = ramus of valvula XIII, IX; rs = ring sclerite; va8, va9 = valvulae VIII, IX; vf8, vf9 = valvifers VIII, IX. Scale bars: 0.5 mm.

FIGURE 14. Spermatheca of four Physopelta (Neophysopelta) species: Ph. (N.) gutta gutta (A); Ph. (N.) parviceps from Japan proper (B); Ph. (N.) parviceps from the Ryukyu Islands, Japan (C); Ph. (N.) fusciscutellata sp. nov. (D); Ph. (N.) quadriguttata (E). Abbreviations: ar = apical receptacle; ip = intermediate part; sd = spermathecal duct. Scale bars: 0.1 mm.