Tijdschrift voor Entomologie 159 (2016) 89–96

Further records of the plant bug subfamily Isometopinae from Thailand (Heteroptera: ), with descriptions of three new species Tomohide Yasunaga, Jomsurang Duangthisan, Kazutaka Yamada & Taksin Artchawakom

Three new species, Isometopus chaiyaphum, Paloniella microchelys, and Myiomma phuvasae, of the plant bug subfamily Isometopinae Fieber are described; additional records for I. siamensis Yasunaga, Yamada & Artchawakom are documented. The habitat of P. microchelys is reported. A checklist of the isometopine fauna of Thailand is provided. Keywords: Heteroptera; Miridae; Isometopinae; new species; Thailand Tomohide Yasunaga*, Research Associate, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA. [email protected] Jomsurang Duangthisan, Collection, Entomology & Zoology Group, Plant Protection Research and Development Office, Department of Agriculture, Chatuchak, Bangkok 10900, Thailand. [email protected] Kazutaka Yamada, Tokushima Prefectural Museum, Bunka-no-Mori Park, Mukôterayama, Hachiman-chô, Tokushima 770–8070, Japan. yamada [email protected] Taksin Artchawakom, Sakaerat Environmental Research Station (SERS), Sakaerat Biosphere Reserve, Thailand Institute of Scientific & Technological Research, Ministry of Science Technology, Wang Nam Khieo, Nakhon Ratchasima 30370, Thailand. [email protected]

Introduction (1904, 1911) and Poppius (1913, 1915), have been The plant bug subfamily Isometopinae Fieber is augmented mainly by Lin (2004) and Lin & Yang composed of very small mirids with a pair of ocelli, (2004) for Taiwan, Ren (1991) and Ren & Yang and together with the subfamily Psallopinae Schuh, (1988) for continental China, Yasunaga (2001, 2005) putatively constitutes the most primitive Miridae. for Japan, and Yasunaga & Duwal (2006) for Nepal. Isometopines are distributed globally, but the major- However, the Isometopinae of Indochina, known to ity of species occur in the tropics, subtropics, and be one of the most species-rich regions in the world, warm temperate climate zones. is currently represented only by Isometopus feanus The Isometopinae currently comprise approxi- Distant, 1904 (Myanmar) and I. siamensis Yasunaga, mately 50 nominal species in Asia (Schuh 1995, Yamada & Artchawakom, 2013 (Thailand). 2002−2014, Kerzhner & Josifov 1999). The Asian The habits of isometopines are poorly known faunas, historically known from the works of Distant because of their unique habitat preference for

Tijdschrift voor Entomologie 159: 89–96, Figs 1–16. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 1 November 2016. DOI 10.1163/22119434-15902003 *Corresponding author Downloaded from Brill.com10/02/2021 09:50:00AM via free access

90 Tijdschrift voor Entomologie, volume 159, 2016 tree trunks (often under bark) or branches, which necessary literature are now available (Schuh, 1995, make collecting the bugs with a sweep net difficult 2002–2014, Kerzhner & Josifov 1999). (Wheeler 2001, Yasunaga 2005). In addition, most isometopines are assumed to be diurnal, with UV light traps attracting a limited number of specimens Checklist of Thai Isometopinae presumably from nearby habitats (Yasunaga 2001, Family Miridae Hahn 2005); therefore, few individuals come to light be- Subfamily Isometopinae Fieber cause isometopine population densities are inher- Tribe Isometopini, subtribe Isometopina ently low in most habitats, particularly in tropical Genus Isometopus Fieber, 1860 or well-preserved forests. As a result, quite a few Isometopus chaiyaphum Yasunaga, Duan- species have been described from the holotype only. thisan & Yamada, sp. n. — Distribution: Biological information on tropical isometopines is Thailand (Chaiyaphum). hardly available, with many species in tropical and Isometopus siamensis Yasunaga, Yamada & subtropical zones remaining undescribed. Artchawakom, 2013 — Thailand (Nakhon During our continuing field investigations in Ratchasima). Thailand, one undescribed specimen each from the Genus Paloniella Poppius, 1915 genera Isometopus, Paloniella, and Myiomma were Paloniella microchelys Yasunaga, Duan- recently discovered (two by uv light traps and one thisan & Yamada, sp. n. — Thailand from the trunk of a broadleaf tree). Herein we de- (Nakhon Ratchasima). scribe these as the new species, I. chaiyaphum, M. Tribe Myiommini Bergroth, subtribe Myiom-­ phuvasae, and P. microchelys. Additionally the habitat mina of P. microchelys is briefly documented,Isometopus Genus Myiomma Puton, 1872 siamensis is diagnosed, and a checklist of the Isome- Myiomma phuvasae Yasunaga, Duanthisan topinae from Thailand, following the classification & Yamada, sp. n. — Thailand (Nakhon system of Herczek (1993), is provided. Ratchasima).

Materials and methods Taxonomy Almost all specimens examined were collected Subfamily Isometopinae Fieber at ‘Sakaerat Environmental Research Station, Sakaerat Biosphere Reserve, Ministry of Science Diagnosis. Members of this subfamily are easily rec- and ­Technology, Nakhon Ratchasima Provinces, ognized by the presence of a pair of ocelli between com- ­Thailand’ cited in the text simply as ‘SERS’ (http:// pound eyes. General shape and size (total length 2−8 www.tistr.or.th/sakaerat/index_en.php). All holo- mm) variable, especially form of the head, compound types are deposited in the Insect Collection, Ento- eyes, antennae and pronotum (see Herczek 1993). mology & Zoology Group, Plant Protection Re- Discussion. Although the present work follows a search and Development Office, Department of classification system proposed for isometopine taxa Agriculture, Bangkok, ­Thailand (DOAT). Matrix by Herczek (1993), we encourage a revised supra- code labels were attached to all type specimens, generic classification. In our opinion many unde- which uniquely identify each specimen, and are scribed taxa, especially from the tropics and sub- referred to as ‘unique specimen identifiers’ (USIs). tropics, cannot be accommodated by the current The USI codes [e.g., AMNH_PBI 000123] com- system (Yasunaga unpublished data). In general, the prise an institution and project code (AMNH_PBI) isometopine pretarsal and male genitalic structures and a unique number (000123). These data were are fundamentally similar to those exhibited in the digitized on the ­ Easy Capture (former- Psallopinae and some members of the Cylapinae ly the Planetary Biodiversity Inventory) database (Yasunaga et al. 2010, Herczek & Popov 2014). A maintained by the ­American ­Museum of Natural broader survey of these and other character systems, ­History, New York, USA (http://research.amnh.org/ as well as documentation of undescribed taxa, is re- pbi/), are ­incorporated in http:// www.discoverlife. quired to substantiate a relationship between these org, and are also searchable on the ‘­Heteroptera three subfamilies and provide an updated suprage- Species Pages’ (http://research.amnh.org/pbi/hetero neric classification system. pteraspeciespage/). All the species of the Isometopinae are presumed Digital images of live individuals were taken by to be predators, with some species having a close TY with a Canon EOS Kiss digital camera body + association with coccids (Wheeler & Henry 1978, Olympus OM-System. All measurements are in Wheeler 2001, Yasunaga 2001, 2005). Ren (1987) millimeters (mm). Synonymic lists for known taxa reported a continental Chinese species, Isometopus are omitted, as comprehensive catalogs covering the citri Ren, found on citrus trees heavily infested by a Downloaded from Brill.com10/02/2021 09:50:00AM via free access

Yasunaga et al.: New species of Isometopinae from Thailand 91

Figs 1–6. Isometopines in Thailand, habitat and habitus images. — 1, an Indian ash broadleaf, Lannea coroman- delica, habitat of Paloniella microchelys; 2, P. microchelys, holotype female, on bark of L. coromandelica; 3, ditto; 4, Myiomma phuvasae, holotype female; 5, Isometopus chaiyaphum, holotype male; 6, I. siamensis, male. scale, Unaspis sp. (Diaspididae), the assumed prey of able refuge from the predators of these isometopine the isometopine. On Ishigaki Island of the Ryuky- plant bugs. us, a subtropical island of Japan, Yasunaga (2005) A similar habitat preference was confirmed in the reported six species of the Isometopinae inhabit- psallopine, Psallops fulvioides Yasunaga & Yamada, ing (or sometimes co-occurring on) the ash tree, which might support a close relationship between Fraxinus griffithii C.B. Clarke (Oleaceae). There are the Isometopinae and the Psallopinae (Yasunaga et al. usually many half-detached fragments and cracks 2010). Incidentally, Wolski & Henry (2015) down- on the ash bark surface, which may provide suit- graded the Psallopinae to a tribal level and placed Downloaded from Brill.com10/02/2021 09:50:00AM via free access

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Figs 7–12 Habitus images of isometopines in Thailand. — 7, 8, Isometopus chaiyaphum, holotype male (AMNH_ PBI 00380407); 9, 10, Paloniella microchelys, holotype female (00380408); 11, 12, Myiomma phuvasae, holotype female (00380409). it in the Cylapinae. As pointed out by Herczec et al. Genus Isometopus Fieber (2015), this proposal, inclusion of Psallopinae in Isometopus chaiyaphum Yasunaga, Duanthisan & Cylapinae at the rank of tribe, seems to require care- Yamada sp. n. ful consideration. The identity and relationships of Figs 5–8, 15, 16 the Psallopinae are still poorly demonstrated, and their subfamilial status may be compromised by a Type material. Holotype, ♂: Thailand: Chai- more comprehensive phylogenetic assessment of the yaphum, Chulabhom Dam, UV light trap, 760−780 Isometopinae and Cylapinae (Cassis & Schuh 2012). m, 16°32–33’N, 101°38–39’E, 16–18 Apr 2013, T. Therefore, a definitive treatment of the Psallopinae Yasunaga et al. (AMNH_PBI 00380407) (DOAT). (or Psallopini in Cylapinae) is beyond the scope of Diagnosis. Recognized by its bicolored, oval this study. We currently encourage much broader body; pale yellow-brown, translucent hemelytron surveys of characters as well as their biology, to cor- and membrane; fuscous pronotum with pale pos- rectly ascertain a lineage between isometopines, psal- terolateral corner; and fuscous scutellum with an lopines and cylapines. apical, creamy yellow,Downloaded V-shaped from mark Brill.com10/02/2021 at apex. Most 09:50:00AM via free access

Yasunaga et al.: New species of Isometopinae from Thailand 93

Figs 13–16. Male genital segment (pygophore) of Isometopus spp. in Thailand. — 13, 14, I. siamensis, holotype (AMNH_PBI 00379601); 15, 16, I. chaiyaphum, holotype (00380407). similar in general coloration and shape to I. nagarjun weakly shining, roughly punctate, with pale lateral Yasunaga & Duwal, 2006 from Kathmandu, Nepal; carina and posterolateral corner; posterior margin distinguished by its flat posterior margin of prono- of pronotum flat, not projecting; scutellum shiny tum not projecting mesally, larger apical pale mark fuscous, with a creamy yellow, V-shaped mark from on the scutellum, and uniformly pale, translucent median part to apex; pleura widely shiny reddish hemelytron without infuscate paracuneus and base brown. Wings pale, translucent; hemelytron mi- of cuneus. nutely punctate; cuneus with an obscure spot along inner margin. Pro- and mesofemora each with a brown ring at apex; metafemur with a circular spot Description ventrally. Abdomen entirely shiny dark brown. Male Male. Body oval, widely fuscous with pale, translu- genitalia as in Figs 15, 16. Hypophysis of left para- cent wings and creamy yellow legs; dorsal surface mere sharply curved at middle; right paramere with weakly shining, with uniformly distributed, simple, short, rather broad hypophysis. Endosoma almost pale, semierect setae. Head weakly flattened in front; totally membranous. compound eye a little produced anteriad; vertex Female. Unknown. darkened medially around ocelli; buccula and clyp- eus shiny fuscous. Antenna creamy yellow; segments Measurements. Male: Total body length 2.05; III and IV slightly darker. Labium shiny yellowish length apex of clypeus to cuneal fracture 1.33; width brown, long, reaching genital segment; apical 2/3 of of head across compound eyes 0.55; head height segment IV reddish brown. Pronotum dark brown, in frontal view 0.36;Downloaded width from of Brill.com10/02/2021vertex 0.17; lengths 09:50:00AM via free access

94 Tijdschrift voor Entomologie, volume 159, 2016 of antennal segments I–IV: 0.09, 0.43, 0.30, 0.16; sparsely punctate, with uniformly distributed, very length of labium 1.05; mesal pronotal length 0.30; short setae. Head tinged with maroon, strongly flat- basal pronotal width 1.02; maximum width across tened anteriad, shallowly and irregularly punctate, hemelytron 1.15; and length of metafemur, tibia and somewhat broadened ventrad, narrowly carinate tarsus 0.67, 0.67, 0.22. basally; compound eye small, flattened; ocellus con- Etymology. Named for its type locality, Chiyaphum tiguous to compound eye and basal carina. Antenna Province; a noun in apposition. brown, short; segment I dark brown; median part of Biology. Unknown; the adult male was attracted to segment II broadly and basal 1/4 of segment III yel- a UV light trap. lowish brown; apical brown part of segment II a little swollen. Labium shiny chocolate brown, reaching Isometopus siamensis Yasunaga, Yamada & apex of metacoxa. Pronotum anteriorly concealed by Artchawakom, 2013 head and compound eyes, transversely rugose, nar- Figs 6, 13, 14 rowly margined laterally; mesoscutum and scutellum Diagnosis. Recognized by its uniformly fuscous roughly and transversely rugose. Hemelytron slanting general coloration; deeply punctate dorsum; rather laterally; cuneus wider than long; membrane grayish large compound eye; elongate left paramere with brown. Legs generally short; apex of each femur, base circularly curved, apically tapered hypophysis; and and apex of each tibia and most of tarsi brown. broad base of right paramere. See Yasunaga et al. Male: Unknown. (2013) for detailed description. Measurements. Female: Total body length 1.84; Biology. The adults are occasionally attracted to length apex of clypeus to cuneal fracture 1.27; light. Collection records suggest this isometopine width of head across compound eyes 0.63 (maxi- has at least two generations per year. mum width in front 0.69); head height in frontal Material examined. 1 ♂, Thailand, Nakhon Rat- view 0.47; width of vertex 0.17; lengths of antennal chasima Prov.: SERS, 14°30’27”N, 101°55’39”E, segments I–IV: 0.08, 0.38, 0.26, 0.13; length of la- 410 m alt., light trap, 3 Jun 2012, T. Yasunaga bium 0.86; mesal pronotal length 0.33; basal prono- (holotype, AMNH_PBI 00379601) (DOAT); 1 ♀, tal width 0.97; maximum width across hemelytron same data, except for date 2 Jun 2012 (00379602) 1.24; and length of metafemur, tibia and tarsus 0.50, (TYCN); 1 ♂, SERS, in guestroom, 14°30’39”N, 0.57, 0.18. 101°55’52”E, 385 m, 13 Dec 2015 (01:50 a.m.), T. Etymology. From Greek, mikros (= tiny) combined Yasunaga (TYCN, without USI label). with chelys (= tortoise), referring to the general shape of this new species reminiscent of a tortoise. Genus Paloniella Poppius Biology. A single female adult was found on the bark Paloniella microchelys Yasunaga, Duanthisan & (40 cm high from ground) of an Indian ash tree, Lan- Yamada sp. n. nea coromandelica (Houtt.) Merr. (Anacardiaceae), Figs 1–3, 9, 10 width about 25 cm diameter at breast height (Fig. 1). Several ant workers and small spiders were observed Type material: Holotype ♀: Thailand: Nakhon on the same trunk, but no suitable prey was evident. Ratchasima, SERS, Lannea coromandelica (Houtt.) Merr. (Anacardiaceae), on bark, 390 m alt., 14°30’38.38”N, 101°55’55.88”E, 11 Dec 2015 Genus Myiomma Puton (14:30 p.m.), T. Yasunaga (AMNH_PBI 00380408) Myiomma phuvasae Yasunaga, Duanthisan & Ya- (DOAT). mada, sp. n. Diagnosis. Recognized by its ovoid, tortoise-shaped Figs 4, 11, 12 body; shiny dorsum that is minutely and sparsely Type material. Holotype ♀, Thailand, Nakhon punctate; very short vestiture; extremely flattened head Ratchasima, SERS, 14°30’27”N, 101°55’39”E, 410 including compound eyes slightly broadened ventrad; m alt., UV light trap, 10 Dec 2015, T. Yasunaga and narrow membrane. Similar in overall appearance (AMNH_PBI 00380409) (DOAT). to P. parallela Yasunaga & Hayashi, 2002, known only from Iriomote Island, Japan; distinguished by Diagnosis. Recognized by its oval, somewhat flat- its smaller size, generally fuscous coloration, minute tened body; conspicuously reddish lateral margins of and sparse punctures on the dorsum, and higher head, dorsum; comparatively small head; short antennal which is somewhat broadened ventrad. segments III and IV; uniformly punctate pronotum and hemelytron; and broadened exocorium. Most similar to M. zhengi Lin & Yang, 2004; readily dis- Description tinguished by its more ovate body shape and con- Female: Body almost totally fuscous, oval, tortoise- spicuously reddish dorsum, and uniformly punctate shaped; dorsal surface shiny, minutely, shallowly and pronotum and hemelytron. Downloaded from Brill.com10/02/2021 09:50:00AM via free access

Yasunaga et al.: New species of Isometopinae from Thailand 95

Description Distant, W.L., 1911. The Fauna of British India, Including Ceylon and Burma. Rhynchota. — Vol. V Heteroptera: Female: Body ovate, broadened posteriorly in dorsal Appendix, xii+362 pp. Taylor & Francis, London. view; dorsal surface fuscous, with conspicuous, red Herczek, A., 1993. Systematic position of Isometopinae lateral margins from pronotum to cuneus, shallowly Fieb. (Miridae, Heteroptera) and their interrelation- and roughly punctate, with uniformly distributed, ships. — Prace Naukowe Uniwersytetu Slaskiego w simple, pale, reclining setae. Head including com- Katowicach 1357: 1–86. pound eyes hemispheric, comparatively small. An- Herczek, A., & Y.A. Popov, 2014. New psallopinous plant tenna whitish brown, short; segment II broadened, bugs (: Heteroptera, Miridae, Psallopinae) with apex sanguineous; segments III and IV very from the New Hebrides and Nigeria. — Zootaxa 3878: short. Labium pale brown, slightly exceeding meta- 366–378. coxa; apical part of segment IV broadly darkened. Herczek, A., Y.A. Popov & J. Gorczyca, 2015. First new Pronotum dark maroon with red margin; scutellum fossil plant bugs of the genus Psallops Usinger, 1946 dark maroon; hemelytron widely dark maroon, with (Hemiptera: Heteroptera, Miridae, Psallopinae) from broadly red exocorium and cuneus; membrane pale the Eocene Baltic amber. — Zootaxa 4052: 495–500. smoky brown. Legs almost totally pale brown; apices Kerzhner, I.M. and M. Josifov, 1999. Miridae Hahn, 1833. of pro- and mesofemora narrowly sanguineous; api- — In: B. Aukema & C. Rieger (Eds), Catalogue of the cal half of metafemora tinged with red. Abdomen Heteroptera of the Palearctic Region, Vol. 3, Cimico- shiny reddish brown; sterna VIII and IX pale brown. morpha ii, pp. 1–576. Netherlands Entomological So- Male: Unknown. ciety, Amsterdam, Netherlands. Lin, C.S., 2004. Seven new species of Isometopinae (He- Measurements. Female: Total body length 2.28; miptera: Miridae) from Taiwan. — Formosan Ento- length apex of clypeus to cuneal fracture 1.52; width mologist 24: 317−326. of head across compound eyes 0.48; head height in Lin, C.S., & C.T. Yang, 2004. Isometopinae (Hemiptera: frontal view 0.42; width of vertex 0.09 (maximum in- Miridae) from Taiwan. — Formosan Entomologist 24: terocular space); lengths of antennal segments I–IV: 27–42. 0.07, 0.46, 0.11, 0.10; length of labium 0.91; me- Poppius, B., 1913. Zur Kenntnis der Miriden, Isometop- sal pronotal length 0.29; basal pronotal width 0.98; iden, Anthocoriden, Nabiden und Schizopteriden Cey- maximum width across hemelytron 1.30; and length lon’s. — Entomologisk Tidskrift 34: 239–260. of metafemur, tibia and tarsus 0.76, 0.86, 0.19. Poppius, B., 1915. H. Sauter’s Formosa-Ausbeute: Nabi- Etymology. Named to honor Phuvasa Chanon- dae, Anthocoridae, Termatophylidae, Miridae, Isome- muang (SERS), who positively supports our research topidae und Ceratocombidae (Hemiptera). — Archiv activities; a noun in genitive case. für Naturgeschichte 80A(8): 1−80 (1914). Ren, S.Z., 1987. New species and a newly recorded genus Biology. Unknown; a female adult was collected by of Isometopidae from China (Hemiptera: Heterop- UV light trap. tera). — Acta Zootaxonomica Sinica 12: 398−403 (in Chinese with English summary). Acknowledgements Ren, S.Z., 1991. A new species of Isometopus Fieber from Yunnan, China (Heteroptera: Isometopidae). — Acta We are much indebted to Dr Charuwat Taekul Zootaxonomica Sinica 16: 204−206 (in Chinese with (DOAT), Ms Phuvasa Chanonmuang (SERS), and English summary). Drs Andrzej Wolski and Milosz Mazur (Opole Univer- Ren, S.Z. & C.K. Yang, 1988. New genus and new spe- sity, Poland) for supporting our collection of specimens cies of Isometopidae from China (Hemiptera–Heter- used in this paper. Dr Michael D. Schwartz (Agricul- optera). — Entomotaxonomia 10: 75−82 (in Chinese ture & Agri-Food Canada, Ottawa, Ontario) kindly re- with English summary). viewed the manuscript and offered some necessary ref- Schuh, R.T., 1995. Plant Bugs of the World (Insecta: erences. Thanks are extended to Dr Alfred G. Wheeler Heteroptera: Miridae). Systematic Catalog, Distribu- (Clemson University, SC, USA) and Dr. Dan Polhe- tions, Host List and Bibliography. — The New York mus (associate editor, TvE) for reviewing the manu- Entomological Society, New York, NY, USA, xii + 1329 script and providing useful comments and suggestions. pp. Schuh, R.T., 2002–2014. On-line Systematic Catalog of Plant Bugs (Insecta: Heteroptera: Miridae). — http:// References research.amnh.org/pbi/catalog/ Cassis, G. & R.T. Schuh, 2012. Systematics, biodiversity, Wheeler, A.G. jr, 2001. Biology of the Plant Bugs (He- biogeography, and host associations of the Miridae miptera: Miridae), Pests, Predators, Opportunists. — (Insecta: Hemiptera: Heteroptera: ). — Cornell University Press, Ithaca, NY, USA & London, Annual Review of Entomology 57: 377–404. UK, xv + 507 pp. Distant, W.L., 1904. The Fauna of British India, Includ- Wheeler, A.G. jr & T.J. Henry, 1978. Isometopinae ing Ceylon and Burma. Rhynchota. — Vol. 2, part 2, (Hemiptera: Miridae) in Pennsylvania: Biology and pp. 243–503. Taylor & Francis, London. descriptions of fifth instars, with observations of Downloaded from Brill.com10/02/2021 09:50:00AM via free access

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predation on obscure scale. — Annals of the Entomo- Yasunaga, T., & R.K. Duwal, 2006. First record of the logical Society of America 71: 607–614. plant bug subfamily Isometopinae (Heteroptera, Miri- Wolski, A. & T.J. Henry, 2015. Review and a new sub- dae) from Nepal, with description of a new genus and family placement of the plant bug genus Isometocoris five new species. — Biogeography 8: 55−61. Carvalho and Sailer, 1954 (Hemiptera: Heteroptera: Yasunaga, T., K. Yamada & T. Artchawakom, 2010. First Miridae), with the description of a new species from record of the plant bug subfamily Psallopinae (Heter- Brazil. — Proceedings of the Entomological Society of optera: Miridae) from Thailand, with descriptions of Washington 117: 407−408. new species and immature forms. — Tijdschrift voor Yasunaga, T., 2001. Family Miridae Hahn, plant bugs. Entomologie 153: 91–98. — In: T. Yasunaga, M. Takai & T. Kawasawa (Eds), A Yasunaga, T., K. Yamada & T. Artchawakom, 2013. A Field Guide to Japanese Bugs ii, pp. 1–96, 111–351. new species of Isometopus Fieber, the first record of Zenkoku Noson Kyoiku Kyokai, Publishing Co. Ltd., Isometopinae (Heteroptera: Miridae) from Thailand. Tokyo, Japan (in Japanese). — Zootaxa 3599: 197–200. Yasunaga, T., 2005. Isometopine plant bugs (Heteroptera: Miridae) preferably inhabiting Fraxinus griffithii on Ishigaki Island of the Ryukyus, Japan. — Tijdschrift Received: January 25, 2016 voor Entomologie 148: 341−349. Accepted: July 23, 2016

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