Tijdschrift voor Entomologie 161 (2018) 11–24
Two new species of the mirine plant bug genus Cheilocapsus Kirkaldy (Heteroptera: Miridae: Mirinae) from Japanese Ryukyus and Taiwan Tomohide Yasunaga
The fauna of the Asian mirine plant bug genus Cheilocapsus Kirkaldy, 1902 (Mirinae: Mirini) in the Japanese Ryukyu islands and Taiwan is updated, based mainly on new morphological features. Four species are now recognized, with descriptions of two new species, Cheilocapsus maius sp. n. (from Taiwan) and C. martius sp. n. (Ishigaki and Iriomote Islands of the Ryukyus). An annotated checklist including all known congeners and a key to the four species are provided. The phylogenetic relationship between Cheilocapsus and Pantilius Curtis is argued on the basis of their detailed structures and zoogeographical distribution pattern. Keywords: Heteroptera; Miridae; Cheilocapsus; new species; Japan; Ryukyus; Taiwan; phylogenetic relationship Tomohide Yasunaga, Research Associate, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA. [email protected]
Introduction Reevaluation of detailed structures with the aid of The plant bug genus Cheilocapsus was proposed by a Tabletop SEM for some Asian mirine plant bugs Kirkaldy (1902) to accommodate a single Burmese revealed misidentifications ofCheilocapsus specimens species, C. flavomarginatus Kirkaldy, from the north- from Ishigaki and Iriomote Islands of the Japanese eastern montane region (possibly Shan District of Ryukyus as well as from Taiwan. Some of these current Myanmar Union), neighboring Yunnan Prov- specimens (5 ♂, 2 ♀) are paratypes of Parapantilius ince of SW China. Reuter (1903) established a genus flavomarginatus (= C. miyamotoi). Comparison of Parapantilius for a single Chinese taxon P. thibetanus biometrics and structures of the male and female Reuter, and subsequently two species were added genitalia unequivocally clarified that these specimens from the Ryukyus, Japan (P. flavomarginatus Miya- actually represented either of two new species de- moto & Yasunaga, 1989) and Taiwan (P. taiwanicus scribed herein. Yasunaga, 1994). However, Yasunaga & Kerzhner The present work revises Cheilocapsus from the (1998) considered Cheilocapsus and Parapantilius Japanese Ryukyus and Taiwan, and recognizes four congeneric, and provided the substitute name C. mi- species, including two new species, C. maius sp. n. yamotoi Yasunaga for P. flavomarginatus as the revised (Yaeyama-group) and C. martius sp. n. (Taiwan). combination resulted in a secondary junior hom- The latter, previously identified as C. miyamotoi, is onym of Kirkaldy’s preoccupied flavomarginatus. Liu now considered sympatric with C. taiwanicus. The & Wang (2001) further described two species from genus Cheilocapsus is rediagnosed, based on micro- continental China, so that currently Cheilocapsus is scopic surface structures and genitalic morphology. represented by five species, althoughYasunaga (2011) The phylogenetic relationship of Cheilocapsus to the suspected one of the two Chinese species may be con- Palearctic genus Pantilius Curtis (previously suggest- specific with the Burmese type species. ed by Yasunaga 2011) is reviewed and discussed, and
Tijdschrift voor Entomologie 161: 11–24, Table 1, Figs 1–65. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 15 December 2018. DOI 10.1163/22119434-00002071 Downloaded from Brill.com09/27/2021 12:24:15AM via free access
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Figs 1–9. Habitus images of Cheilocapsus species in dorsal (1, 4, 7), ventral (2, 5, 8) and lateral (3, 6, 9) view (2, 5, 7 each with shagreened base of mesocoxa in left lateral view). – 1–3, C. martius, holotype male; 4–6, C. martius, paratype female; 7, C. maius, holotype male; 8, C. maius, paratype female; 9, C. miyamotoi, paratype female from Amami-Oshima Island. a dark spot on the epimeron (Figs 6, 9); green, olive diagnostic characters were provided by Yasunaga or brown median hemelytron (but forewing without (1994, 2011). reddish tinge); a series of minute callosities on lateral base of mesocoxa (presumably a stridulatory device Distribution or simply toughened structure for coxal base, cf. From northern Indochina via southern China to Figs 2, 5, 7, 25, 31); more or less spotted ventral Japan (Ryukyus) and Taiwan. surface of metafemur (Figs 15–18); five similar en- dosomal lobes; roundly sclerotized margin of female Biology genital chamber; enlarged, thick-rimmed sclero- For the Japanese and Taiwanese species, one gen- tized rings; and small interramal lobes. Additional eration per year is assumed. The egg appears to
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Figs 10–18. Cheilocapsus miyamotoi (10, 13, 17), C. maius (11, 15), C. martius (12, 16) and C. taiwanicus (14, 18). – 10, live adult female (Amami-Oshima Island); 11–14, dorsal habitus of anterior body; 15–18, left metafemur, ventral surface.
hibernate and hatch in early spring. The adults are Discussion limited to spring (mostly between March and May). Yasunaga (1994, 2011) posited that Cheilocapsus Most specimens I have examined were captured by most probably has the closest relationship to Pan- sweep-netting branches or inflorescences of ever- tilius Curtis (cf. Figs 19, 20) as they share the tough, green broadleaved trees; however, no information is coreid-like body, more or less tumid frons, thick- currently available on the immature form or breed- ened antennal segments I and II, and presence of ing host. Collection records (Zheng et al. 2004) sug- at least three homologous lobes on the endosoma. gest the adults of two continental Chinese species My recent examinations by SEM have revealed ad- inhabiting the temperate climate zone, C. maculipes ditional characters possessed by both of them, such and C. nigrescens, to appear in mid summer (July and as partly shagreened surface structure of head (cf. August). Figs 33 vs. 36); minutely pruinosed or shagreened Downloaded from Brill.com09/27/2021 12:24:15AM via free access
Figs 19–20. Adult female of Pantilius spp. – 19, P. tunicatus (Hyogo Pref., SW Honshu, Japan, taken by T.B. Shishido); 20, P. hayashii, both images showing the same individual (Tochigi Pref., Honshu, by S. Maehara). – Scale bars ≈ 2.0 mm.
hemelytron with short, reclining setae (Figs 34 vs. being difficult to observe using a stereoscopic micro- 37); general shape of scent-efferent system and scope (as in Figs 2, 4, 5), are currently presumed to metathoracic spiracle (Figs 25, 29 vs. 40); and wide, represent a stridulatory plectrum, possibly with the somewhat rhombic parempodia (Figs 26 vs. 38). On roughened ventral edge of the epimeron sensu Schuh the other hand, Cheilocapsus is distinct from Pan- (1984) who reported a similar structure in a few tilius in having the following differential characters: genera of phyline plant bugs, or maybe simply the dorsum without reddish tinge; eye larger; mesal sul- toughened structure for the coxal base. This struc- cus on vertex wider and shallower, with short setae; ture is not present in Pantilius (cf. Fig. 40) but is in frons less bulbous; lateral carina of the pronotum; some species of other mirine genera (e.g., Adelphoco- areolar veins on forewing membrane rounded pos- risella Miyamoto & Yasunaga, Eurystylus Stål, Pachy- teriorly (Figs 35 vs. 44); and base of mesocoxa with lygus Yasunaga) (Yasunaga unpublished observation). callosities (Fig. 25). Incidentally, the minute callosi- Cheilocapsus and Pantilius are now considered to ties or bumps on the base of the mesocoxa observed constitute a monophyletic group within the tribe in five species ofCheilocapsus (C. maius sp. n. (Figs 7, Mirini, based on the above morphological evidence. 29), C. martius sp. n. (Figs 2, 5, 25), C. miyamotoi The two genera have different habitat require- (Fig. 31), C. taiwanicus and C. thibetanus), despite ments with Cheilocapsus composed of thermophilic Downloaded from Brill.com09/27/2021 12:24:15AM via free access
Figs 21–32. Scanning electron micrographs for Cheilocapsus martius (21–27), C. maius (28, 29), C. thibetanus (30) and C. miyamotoi (31, 32). – 21, 22, 24, 28, 30, 31, head and pronotum; 23, pronotum, scutellum and base of hemelytron; 25, 29, thoracic pleura; 26, protarsus; 27, 32, metatarsus. Abbreviations: cw: claw; pp: parempodia.
members occurring principally in the Asian warm eastern Eurasia as the occurrence of all other related temperate and subtropical zones rich in evergreen taxa is restricted to Asia. Recently, Kim et al. (2016) broadleaf vegetation, whereas all species of Pantilius reported P. hayashii Miyamoto & Yasunaga, 1989 are known to inhabit deciduous forests in colder cli- from Korea. The identity of the Korean population mate zones in the Palearctic Region and Himalayas needs further verification, because the dorsal color (Yasunaga 1992). Pantilius tunicatus (F.) (Fig. 19) is pattern, pronotal shape and male endosoma figured the only species of the two related genera widespread by Kim et al. (2016) do not exactly fit those exhib- in the Palearctic Region (Yasunaga 1992, Schuh ited by the Japanese population. 2002–2014). In eastern Asia, this mirid was re- In the Japanese Ryukyu islands, Cheilocapsus mar- corded only from China (Ningxia Hui Autonomous tius and C. miyamotoi are allopatric (Fig. 65), whereas Region, N35.77 E106.18, see Zheng et al. 2004), the two Taiwanese species, C. maius and C. taiwani- Japan (recently introduced to Osaka area, south- cus, were found to co-occur in the Puli area of Nan- western Honshu, see Yasunaga et al., 2018) and tou (central montane part of Taiwan). Speciation of Russian Primorsky Territory (Kerzhner 1988). The the two Japanese species is most probably attribut- primal origin of P. tunicatus is still perplexing, but able to isolation onto the respective island-groups this species is presumed to have originated from after the Pleistocene (cf. Kimura 2002). However, Downloaded from Brill.com09/27/2021 12:24:15AM via free access
Figs 33–44. Scanning electron micrographs for Cheilocapsus martius (33–35), Pantilius hayashii (36–41) and P. tunicatus (42–44). – 33, 36, head, with sulcus on vertex; 34, 37, vestiture on corium; 35, 44, cuneus and forewing membrane; 38, pretarsal structure of mesotarsus; 39, head and thoracic pleura; 40 scent-efferent system; 41, meta- tarsus; 42 head and pronotum; 43 pronotum, scutellum and anterior hemelytron. Abbreviations: cw: claw; pp: parempodia. further field investigation of their ecology is required – Head width across eyes obviously less than to elucidate the interspecific relationship between length of antennal segments I or III; epimeron the two sympatric Taiwanese species. with a well-defined dark spot (Fig. 9) ���������� 2 2. Pronotum and scutellum lacking dark spots at base of setae (Fig. 14); dark spots on ventral metafemur sparsely distributed (Fig. 18); Tai- Key to Cheilocapsus species from Japan wan ������������������������C. taiwanicus (Yasunaga) and Taiwan – Pronotum and scutellum with dark spots at 1. Head width across eyes almost equal to length base of setae (Figs 11–13); ventral surface of of antennal segment I and greater than length metafemur with densely distributed, clear dark of segment III; dark spot on epimeron weak spots (Figs 15–17) ��������������������������������������� 3 or faint (Figs 3, 6); known from Ishigaki and 3. Basal 1/3 of antennal segment III yellowish Iriomote Islands of the Ryukyus �������������������� (or whitish) brown; scutellum with distinct, ������������������������������������������� C. martius sp. n. paired dark maculae subapically (Figs 10, 13); Downloaded from Brill.com09/27/2021 12:24:15AM via free access
Figs 45–48. Male genitalia of Cheilocapsus maius. – 45, pygophore in left lateral view; 46, left paramere; 47, right paramere; 48, endosoma. Scale bars 0.5 mm. Abbreviations as mentioned in Materials and methods section.
metafemoral spots sparse or faint at basal 1/4 Type material. Holotype, ♂: Taiwan, Nantou (Fig. 17); Amami-Oshima and Okinawa Is- Hsien, Puli Township, Mt. Hewang-shan (= current lands �������������������������C. miyamotoi Yasunaga Habon, near Wushe), N23.92 E120.97, 20 May – Basal half of antennal segment III pale; apical 1990, S. Gotoh (AMNH_PBI 00380602) (NMNS). dark maculae on scutellum faint or obliterated Paratypes: Taiwan, 1 ♂, Kagi Hsien, Chiayi County, (Figs 8, 11); metafemoral spots uniformly dis- Fenchihu, N23.50 E120.69, 12 Apr 1964, S. Miya- tributed (Fig. 15); Taiwan ������ C. maius sp. n. moto (AMNH_PBI 00380601) (TYCN) (paratype of Parapantilius flavomarginatus Miyamoto & Yasu- naga = C. miyamotoi Yasunaga); 1 ♂, 3 ♀, same data as for holotype (00380603–00380606) (TYCN); 1 Cheilocapsus maius Yasunaga, sp. n. ♀, Puli Township, Mt. Kanto-shan, 11 May 1987, S. Figs 7–8, 28–29, 15, 45–51, 65 Gotoh (TYCN, without USIs).
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Figs 49–55. Female genitalia. – 49–51, Cheilocapsus maius; 52–55, C. miyamotoi. 49, 52, posterior wall; 50, 53, bursa copulatrix; 51, 54, valvula II; 55, valvula I. Scale bars 0.2 mm. Abbreviations as mentioned in Materials and methods section.
Diagnosis Description Recognized by the characters mentioned in the ge- Body generally pale smoky brown, elongate, parallel- neric diagnosis and key. Externally most similar to C. sided; dorsum weakly shining, with sparsely distrib- miyamotoi; distinguished by densely and uniformly uted, golden, sericeous setae mainly on clavus and distributed dark spots on ventral surface of metafe- corium. Head pale brown, almost smooth; mesal sul- mur (Fig. 15), endosoma with more developed apical cus on vertex faintly infuscate, rather short. Antenna sclerite of median lobe and weakly curved spicules reddish brown; segment I darkened at base and later- (Fig. 48), and narrower interramal lobe (Fig. 49), in ally, obviously longer than head width across eyes; addition to its allopatric distribution in Taiwan. segment II with fuscous apical half, longer than
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Figs 56–57. Male genitalia (aedeagus) – 56, Cheilocapsus martius; 57, C. miyamotoi. Scale bars 0.5 mm. Abbrevia- tions as mentioned in Materials and methods section. metafemur; two distal segments dark brown; basal paramere semi-circularly curved, with moderately half of segment III and base of segment IV yellow- elongate hypophysis (Fig. 46); right paramere with ish brown. Labium shiny yellowish brown, reaching basally thickened sensory lobe and rather developed but not surpassing base of mesocoxa; apical 1/3 of hypophysis (Fig. 47). Endosomal spicules (Fig. 48, segment IV infuscate. Pronotum shining, sometimes SP1 and SP2) elongate, not strongly hooked apicad; tinged with olive, shallowly and transversely rugose, process of median lobe (ML) narrow, conical. with dark, small spots at base of dark, simple setae; Female genitalia (Figs 49–51): interramal lobe small scent-efferent system as in Fig. 29; apical paired dark (Fig. 49); sclerotized rings ovoid, somewhat separat- maculae on scutellum obscure. Hemelytron coffee ed each other at middle (Fig. 50). brown, somewhat matte, with yellowish brown exo- corium and cuneus; apical 1/5 of cuneus darkened; Measurements membrane smoky brown, with dark brown veins As in Table 1. and a pale, translucent spot posterior to large areolar cell. Legs including coxae pale brown; each femur with dark small spots; ventral surface of metafemur Etymology uniformly spotted; base of metatibia and all tarso- From Latin maius (=May), referring to the appear- meres III darkened. Male genitalia (Figs 45–48): Left ance of this new species in May. Downloaded from Brill.com09/27/2021 12:24:15AM via free access
Figs 58–64. Male (58–60) and female (61–64) genitalia of Cheilocapsus martius. – 58, pygophore; 59, left paramere; 60, right paramere; 61 posterior wall, dorsal margin; 62, bursa copulatrix; 63, valvula I; 64 valvula II. – Scale bars 0.2 mm. – Abbreviations as mentioned in Materials and methods section.
Biology flowers of Styrax japonica, 7 Mar 2002, T. Yasunaga Unknown. (AMNH_PBI 00380608) (TYCN).
Diagnosis Cheilocapsus martius Yasunaga, sp. n. Readily distinguished from congeners by the short Figs 1–6, 12, 15, 21–27, 33–35, 56, 58–65 antennal segments I and III each shorter than the head width across eyes, small, faint dark spot on Type material. Holotype, ♂: Japan, Ryukyus, the epimeron (Figs 5, 6), narrowly infuscate apex of Yaeyama-group, Iriomote Island, Mombanare the cuneus (Figs 1, 4), rather slender spicules and tri- near Otomi, N24.298 E123.866, 26 Mar 1991, angular median lobe on the endosoma (Fig. 56) and M. Hayashi (AMNH_PBI 00380607) (AMNH). small, circular interramal lobe (Fig. 61), in addition Paratype: Japan, 1 ♀, Ryukyus, Yaeyama-group, Ishi to the isolated distribution on Ishigaki and Iriomote gaki Island, Shiramizu, N24.4186 E124.1623, on Islands (Fig. 65). Downloaded from Brill.com09/27/2021 12:24:15AM via free access
Fig. 65. Map showing distribution of Cheilocapsus species.
Description or less tinged with red; scent-efferent system as in Body generally brown, subparallel-sided, compara- Fig. 25, with posterior corner not produced; scutel- tively small in size (less than 10 mm in total length); lum with paired, apical dark brown maculae. Hem- dorsum weakly shining, with sparsely distributed, elytron dark brown, somewhat matte, with yellow silvery, sericeous setae. Head pale brown, partly exocorium, cuneus and anterior half of clavus; apex slightly tinged with red, with silvery, short, upright of cuneus narrowly darkened; membrane pale grayish setae; base of mesal sulcus on vertex removed from brown, with pale posteromedial macula, dark brown anterior margin of pronotal collar, with several short, veins and a pale, translucent spot posterior to large stiff setae (Fig. 33). Antenna pale brown; segment I areolar cell. Legs including coxae pale brown; each reddish brown, darkened at base and apex and lat- femur with sparsely distributed, dark small spots ex- erally, almost equal in length to head width across cept on base; ventral surface of metafemur sparsely eyes; segment II with dark reddish brown apical 1/4, spotted; base of metatibia and apical half of each tar- longer than metafemur; two distal segments brown; somere III darkened; pretarsal structures as in Figs basal 3/5–2/3 of segment III and base of segment 25 and 26. Male genitalia (Figs 56, 58–60): Hypoph- IV yellowish brown. Labium shiny pale brown, ysis of left paramere rather short, somewhat inflated partly tinged with red, reaching base of mesocoxa; subapically (Fig. 59); right paramere almost straight apex of segment IV dark reddish brown. Pronotum and uniform, with small hypophysis (Fig. 60). Endo- yellowish brown, shining, somewhat darkened pos- somal spiculi (Fig. 56, SP1 and SP2) rather slender, teriorly; weakly and transversely rugose, with dark, not strongly hooked apicad; process of median lobe small spots at base of dark, simple setae; calli more (ML) triangular. Female genitalia (Figs 61–64, not Downloaded from Brill.com09/27/2021 12:24:15AM via free access
completely sclerotized because of teneral specimen): Interramal lobe small, circular (Fig. 61).
Measurements As in Table 1. metatarsus I:II:III 10:15:16 — 10:13:14 10:12:15 10:11:15 10:13:16 4:5:6 7:9:10
Etymology From Latin martius (=March), as this new species has tarsus 0.88 — 0.98 0.93 0.91 0.83 0.93 0.91 been collected only in March. 0
tibia 5.34 — 5.51 5.83 5.73 5.39 4.85 4.9 Biology An adult male was collected from an inflorescence of Styrax japonica Siebold & Zucc. (Styracaceae), Length metaleg femur 3.92 — 3.87 4.43 4.17 4.04 3.68 3.77 although the breeding host is yet to be confirmed.
0 0 Acknowledgements Labium length 2.79 2.72 2.94 — 2.96 2.84 2.5 2.7 Special thanks are due to Ex-Prof. Leyi Zheng (Nan- kai University, Tianjin, China), the late Mr. Shin
.00 Gotoh (Tanabe City, Wakayama, Japan) and Prof. IV 0.86 — — 1 0.98 — 0.81 0.88 Em. Masami Hayashi (Saitama University, Urawa, Japan) for providing invaluable specimens. Dr. Jing- fu Tsai (NMNS) kindly suggested correct locality III 1.52 — 1.64 1.91 1.72 1.84 1.23 1.25 names in Taiwan, and Mr. Satoshi Maehara (Tochigi, Japan) and Mr. Takayuki B. Shishido (Hyogo, Japan) provided images of Pantilius species. I am also grateful II 4.24 4.36 4.41 4.41 4.66 5.07 3.65 3.55 to Nagasaki West High School (Super-Science High School program, biology, Mr. Tetsuya Nagashima) and to Mr. Daihei Terada (CSR Division, Hi- Length antennal segments I 1.72 1.69 1.74 1.96 1.96 1.76 1.31 1.38 tachi High-Technologies Corporation, Tokyo) for generously providing access to the scanning electron
microscope. Thanks are extended to Dr. Michael D. . Schwartz (Agriculture & Agri-Food Canada, Cana- Max Max width 3.21 3.19 3.41 3.58 3.5 3.58 3.23 3.53 dian National Collection of Insects, Ottawa) and Dr. Dan A. Polhemus (associate editor, TvE) for review- ing and improving the manuscript. Cheilocapsus width 2.84 2.89 2.99 3.16 3.21 3.11 2.77 3.04
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