Tijdschrift voor Entomologie 160 (2017) 25–40
Three new species of Nesidiocoris Kirkaldy from Japan, with a taxonomic review of the tribe Dicyphini for eastern Asia (Heteroptera: Miridae: Bryocorinae) Tomohide Yasunaga
The fauna of the bryocorine plant bug tribe Dicyphini in eastern Asia (including Japan, Korea, Russian Far East and Taiwan) is reviewed, with emphasis on the genus Nesidiocoris Kirkaldy, which is rediagnosed and discussed. Twelve species in six genera are now recognized. Three new species of Nesidiocoris are described from Japan: Nesidiocoris nozakianus sp. n., N. okinawanus sp. n. and N. simotukensis sp. n. Nesidiocoris poppiusi (Carvalho) is proposed as a junior synonym of N. tenuis, and N. plebejus (Poppius) is transferred to Singhalesia China & Carvalho. An annotated checklist and a key to genera and species are also provided to aid in appreciating the east Asian dicyphine fauna. Keywords: Miridae; Dicyphini; Nesidiocoris; new species; new combination; new synonymy Tomohide Yasunaga, Research Associate, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA. [email protected]
Introduction Nesidiocoris Kirkaldy are present in Japan; (2) a Plant bugs belonging to the tribe Dicyphini are well Taiwanese species, N. poppiusi (Carvalho), is in known to be predominantly predaceous among the all likelihood synonymous with the cosmopolitan highly herbivorous subfamily Bryocorinae, and some N. tenuis (R euter); and (3) the current generic place- of them have attracted significant attention as ef- ment of N. plebejus (Poppius) is erroneous. fective components in integrated pest management The present work is therefore organized to re- (IPM) programs against economically important view the east Asian taxa of Dicyphini, particularly pests (Wheeler 2001, Alomar et al. 2006, Sánchez & treating the genus Nesidiocoris which is in need of Lacasa 2008, Calvo et al. 2016). The Dicyphini cur- further taxonomic revision. Three Japanese species, rently contain approximately 200 species in 16 valid Nesidiocoris nozakianus, N. okinawanus and N. simo- genera, most of which were described from the Ethi- tukensis, are described as new. The genus Nesidiocoris opian and Neotropical Regions (Schuh 2002–2013); and N. tenuis (Reuter), assumed adventive to eastern however, a series of recent surveys by the author and Asia, are rediagnosed and discussed. Nesidiocoris pop- his colleagues have revealed that more than a few un- piusi (Carvalho) is proposed as a junior synonym of described taxa still remain in eastern Asia. N. tenuis, and N. plebejus (Poppius) is removed from The dicyphine faunas of Japan and adjacent re- Nesidiocoris and transferred to Singhalesia China & gions were documented by Yasunaga (2000, 2001), Carvalho. An annotated checklist, with updated in- Neimorovets (2006), and Vinokurov & Golub formation on distribution and/or biology, and a key (2007). Nonetheless, the present work has revealed to genera and species are provided to facilitate iden- that (1) three undescribed species of the genus tification of the east Asian dicyphine taxa.
Tijdschrift voor Entomologie 160: 25–40, Figs 1–66. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 21 July 2017. DOI 10.1163/22119434-16001002 Downloaded from Brill.com10/01/2021 07:03:11PM via free access
Materials and methods on Ternstroemia gymnanthera (Wight et Arn.) Almost all specimens examined in this study were Bedd. (Pentaphylacaceae) (Yasunaga & Yamada collected by the author and his colleagues, and are 2014); recently, the mirid was also found on deposited in the American Museum of Natural His- Euonymus japonicus Thunb. (Celastraceae) tory, New York, USA (AMNH); Natural History in an urbanized zone in Daito City, Osaka Museum, London (BMNH); National Institute of (N34.705, E135.626), according to informa- Agro-Environmental Sciences, Tsukuba, Japan tion on a webpage (http://osaka-insecta.sakura. (NIAES); Tokushima Prefectural Museum, Tokushi- ne.jp/shieldbug/d-kasumikamemushi_1.html). ma, Japan (TKPM); T. Nozaki Collection, Fukuoka, Because C. virgula can use various broadleaf Japan (TNCF); and T. Yasunaga Collection, Naga- angiosperms (of at least twelve plant families) saki, Japan (TYCN), unless otherwise stated. for completing nymphal development and fur- Matrix code labels were attached to the holotypes thermore is regarded to be parthenogenetic (cf. and several representative specimens (from different Wheeler 2001, Henry 2012), the introduced localities); these labels uniquely identify each speci- populations of C. virgula may have strong po- men, and are referred to as ‘unique specimen iden- tential for dispersal and colonization. No male tifiers’ (USIs). The USI codes [e.g., AMNH_PBI adult has hitherto been captured in Japan. 000123] comprise an institution and project code Genus Cyrtopeltis Fieber, 1860 (AMNH_PBI) and a unique number (.000123). Cy. miyamotoi (Yasunaga, 2000) (Figs 12, 37) — These data were digitized on the Arthropod Easy Japan (Hokkaido, Honshu, Shikoku, Kyushu), Capture (formerly the Planetary Biodiversity Inven- Korea. This species is externally most similar to tory) database maintained by the AMNH (http:// the European C. geniculata Fieber; both species research.amnh.org/pbi/) and are also searchable are, as far as known, geographically separated on the ‘Heteroptera Species Pages’ (http://research from each other. The breeding host of C. mi- .amnh.org/pbi/heteropteraspeciespage/). yamotoi was confirmed to be Rosa multiflora Measurements are given in millimeters (mm). L. (Rosaceae), whereas C. geniculata was re- The synonymic list for known taxa is in most cases ported to be associated with fabaceous hosts omitted, as comprehensive catalogs and databases (Schuh 2002–2013). are available (Schuh 1995, 2002–2013 online cata- Cy. nakatanii Yasunaga, 2000 (Figs 14, 38) — log, Kerzhner & Josifov 1999). Terminology of the Japan (Honshu). This bug is generally tinged genitalia mainly follows Namyatova et al. (2016) with vivid green when alive and inhabits a rasp- and Yasunaga & Ishikawa (2016). The following ab- berry, Rubus hirsutus Thunb. (Rosaceae), which breviations are used in the text and figures for the is assumed to be its breeding host (Maehara, genitalic structures: pers. obs.). Male genitalia: HP: hypophysis; DP: dorsal Cy. rufobrunnea Lee and Kerzhner, 1995 (Figs 13, (pygophoral) process; PL: primary lobe; PT: phal- 45) — Japan (Honshu, Kyushu), Korea. This lotheca; SB: sensory lobe; SD: seminal duct; SL: unique maroon-colored species is known to be secondary lobe; VP: ventral (pygophoral) process. associated with raspberries, Rubus spp. Female genitalia: DLP: dorsal labiate plate; SCR: Genus Dicyphus Fieber, 1858 sclerotized ring. D. incognitus Neimorovets, 2006 — Russia (Primorsky: Partizansk). Neimorovets (2006) documented the plant association of this mirid Annotated checklist of Dicyphini in with Circaea alpina L. (Onagraceae). Genus Nesidiocoris Kirkaldy, 1902 eastern Asia N. nozakianus Yasunaga, sp. n. — Japan (Kyushu: Tribe Dicyphini Reuter, 1883 [= subtribe Dicyphina Nagasaki Pref.). sensu Schuh (2002–2013), see Namyatova et al. N. okinawanus Yasunaga, sp. n. — Japan (2016)] (Ryukyus: Okinawa & Ishigaki Islands). Genus Campyloneura Fieber, 1858 N. simotukensis Yasunaga, sp. n. — Japan C. virgula (Herrich-Schaeffer, 1835) (Fig. 18) — (Honshu: Tochigi Pref.). Japan (Osaka, Tokyo), Holarctic Region, Afri- N. tenuis (Reuter, 1895) [= N. poppiusi (Carvalho, ca; populations in Japan and the United States 1958) syn. n.] — Japan (southwest of cen- are evidently not indigenous (possibly intro- tral Honshu, almost whole area including duced from Europe) (cf. Henry 2012, Yasunaga Ryukyus), Korea, Taiwan, China, Russia [Ker- & Yamada 2014). This exotic mirid was first zhner (1988) mentioned a record from Russia collected in 2003 at Yoyogi Park in central To- (Amur) was due to accidental introduction]; kyo Metropolitan (N35.670, E139.695), where widespread almost globally within the tropics, both adults and immature forms were confirmed subtropics, and warm temperate climate zones. Downloaded from Brill.com10/01/2021 07:03:11PM via free access
Figs 1–11. Habitus images for Nesidiocoris species. — 1–4, N. nozakianus sp. n., females on nuts of Vernicia cordata (Thunb.) Airy Shaw in Nagasaki; 5–7, N. caesar on Solanum carolinense L. in Kathmandu (5 and 7 lower females, 6 and 7 upper males); 8, N. okinawanus sp. n., female on Ishigaki Island (courtesy of Mr M. Takai); 9, N. tenuis, male in Nakhon Ratchasima, Thailand; 10, same, female in Siem Reap, Thailand; 11, N. orientalis (Poppius) [= N. poppiusi Carvalho], holotype male (courtesy of Dr M.L. Chan and Dr J.F. Tsai).
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Figs 12–18. Habitus images for dicyphine (non-Nesidiocoris) species. — 12, Cyrtopeltis miyamotoi, female; 13, Cy. rufobrunnea, female; 14, Cy. nakatanii, holotype male; 15, Tupiocoris annulifer, male; 16, an undetermined species of Singhalesia from Thailand; 17, female specimen probably representing S. obscuricornis (from Panay, Philip- pines); 18, Campyloneura virgula, female (from Tokyo). Scale bars 2.0 mm. Images 12, 13 courtesy of Mr M. Takai.
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Genus Singhalesia China & Carvalho, 1952 membranous, bilobate, sometimes with significantly S. obscuricornis (Poppius, 1915) (Figs 17, 36) — melanized lobal sclerites; bursa copulatrix thick- Philippines (Visayas: Panay Island), Taiwan. walled, with concentric wrinkles; posterior wall More than fifty specimens I collected on usually simple, less sclerotized. Further diagnostic Panay Island of the Philippines in May, 1994 characters and synapomorphies are presented and are identical to S. obscuricornis, originally de- discussed in detail by Cassis (1984) and Namyatova scribed from Taiwan. Images of the holotype et al. (2016). female can be seen on a webpage (http://twin- Discussion. The Asian dicyphine members are eas- secttype.nmns.edu.tw/specimen/?id=NMNS- ily recognized by the above-mentioned characters. SDEI-00158). The Philippine male has the Within five bryocorine tribes suggested by the latest short, C-shaped left paramere (Fig. 36) quite classification (Namyatova et al. 2016), only Dicy- similar to that of the type species of the genus, phini are predominantly composed of predaceous S. indica (Poppius) currently known from Sri species, whereas the majority of the species in the Lanka and Iran (Schuh 2002–2013). other tribes are phytophagous and host-plant- S. plebejus (Poppius, 1915) n. comb. [transferred specific (Stonedahl 1988, Cassis & Schuh 2012, from Nesidiocoris] — Taiwan. This species was Konstantinov & Knyshov 2015, Yasunaga & described by Poppius (1915) under the ge- Ishikawa 2016, Yasunaga et al. 2016). Several spe- nus Engytatus Reuter, composed of 28 species cies (e.g., Macrolophus caliginosus, Nesidiocoris tenuis) mainly known from the Neotropical Region. are commercially mass-propagated as biocontrol Subsequently, Cassis (1984) placed it in Nesidi- agents against whiteflies, thrips, aphids or spider- ocoris. The males are not yet known, and its ac- mites (Takai & Yasunaga 2001). However, little is curate generic position remains unclear (images currently known about the life history of ‘genuinely of the holotype female available on http://twin- indigenous’ Asian dicyphines, which require fur- secttype.nmns.edu.tw/specimen/?id=NMNS ther investigation. Incidentally, Wheeler & Krim- -HNHM-00051). I recently obtained some mel (2015) emphasized that dicyphines specialize specimens from Thailand similar to plebejus. on sticky plants that have glandular trichomes pro- The Thai species (presumed to be undescribed, ducing adhesive, viscous exudates. This theory can cf. Fig. 16) has a tiny body (2.4 mm) as in Pop- be applicable to some Asian species of Dicyphini; pius (1915) and a C-shaped rather than an many of the plant species mentioned in this paper elongate left paramere; this facies corresponds have more or less sticky leaves and stems. Nesidi- with the diagnostic characters of Singhalesia. ocoris caesar (on a wild Solanaceae plant in Nepal) Based on current evidence, it seems best to and an undetermined species of Nesidiocoris (on place plebejus in Singhalesia. Accordingly, a new Muntingia calabura L., Tiliaceae, in Thailand) scav- combination is herein proposed. enge on cadav ers of small dipterans trapped by the Genus Tupiocoris China & Carvalho, 1952 viscous hairs on the leaf and stem surfaces (Yasunaga T. annulifer (Lindberg, 1927) (Figs 15, 39, 44) — pers. obs.). Japan (Hokkaido, Chishima Islands), Russia (Far East, S. Siberia west to Altai). The breeding Key to east Asian genera and species of hosts of this boreal dicyphine were confirmed to be raspberries (e.g., Rubus chamaemorus L., Dicyphini R. idaeus L., R. phoenicolasius Maxim.) (Kerzh- [Two Taiwanese species of Singhalesia, each known ner 1988, Yasunaga 2001). only from a single, partly faded holotype female, are excluded from this key, but at the generic level of recognition, they are usually separable from other Taxonomy taxa by their notably small size (less than 2.5 mm Dicyphini Reuter in total length); see above checklist and Figs 16, 17] 1. Body elongate, more than 5.0 mm in total Diagnosis. The plant bugs belonging to this tribe length; antennal segment III 2/3 as long as II; are defined primarily by the following diagnos- currently known only from Russian Primorsky tic characters: Body noticeably slender, elongate; Territory ���������������������������Dicyphus incognitus dorsum usually smooth, impunctate, with uni- – Body shorter than 4.6 mm or much small- formly distributed, simple vestiture; ostiolar peri- er; antennal segment III about as long treme flat or weak Figs( 40–44); meta-tarsomere I as, slightly shorter or obviously longer than distinctly shorter than segment II; tarsomere III II ���������������������������������������������������������������� 2 shorter than segment II; pygophore greatly asym- 2. Pronotum, scutellum and clavus almost con- metrical, with distinct processes (Figs 34–39); left colorous and immaculate, uniformly pale paramere sometimes excessively elongate and right green (Fig. 12), brown-castaneous (Fig. 14) or paramere reduced or vestigial; endosoma broadly reddish (Fig.Downloaded 13) ����������������������� from Brill.com10/01/20213 [Cyrtopeltis 07:03:11PM] via free access
– Pronotum, scutellum and clavus somewhere – Body longer than 3.6 mm; antennal seg- with darkened, spotted, striped or more color- ment I white (Fig. 25), sometimes slightly ful pattern ���������������������������������������������������� 5 obscured medially (Fig. 3); antennal segment 3. Base of head (neck) yellow brown medially, III obviously longer than II; scutellum usually contrastingly blackened laterally behind eyes with broad, mesal stripe (Fig. 4) �������������������� (Fig. 12); extreme bases (knees) of pro- and �������������������������������������� N. nozakianus sp. n. mesotibiae clearly darkened �������������������������� ��������������������������������������Cyrtopeltis miyamotoi – Base of head concolorous; bases of pro- and Nesidiocoris Kirkaldy mesotibiae almost uniformly pale ���������������� 4 4. Dorsum almost uniformly pale brown Diagnosis (Fig. 14, but fresh specimens more green- Distinguished from other dicyphine genera by the ish); antennal segment III much longer than following combination of characters: small to moder- II ����������������������������������������������Cy. nakatanii ate size (total length 2.7–4.0 mm); pale yellow-green – Dorsum almost uniformly red-maroon or to dark olive brown general coloration; pale simple sanguineous (Fig. 13); antennal segment III setae uniformly distributed on pronotum, scutellum almost equal in length to II ���������������������������� and hemelytra; smooth, head more or less rounded in ����������������������������������������������Cy. rufobrunnea front; relatively large eyes removed from pronotum 5. Head with long neck convergent towards base for a distance subequal to thickness of pronotal col- (Fig. 15); eye distinctly removed from anterior lar; always pale base and apex of antennal segment I; margin of pronotum (distance between poste- almost linear antennal segment II as long as or shorter rior margin of eye and pronotal collar greater than segment III; weakly tumid scutellum; narrow, than length of eye in dorsal view); scutellum triangular evaporative area of metathoracic scent almost uniformly fuscous ������������������������������ efferent system (Figs 40–42); usually narrowly ����������������������������������������Tupiocoris annulifer darkened base (knee) of each tibia; deeply excavated – Head with short, parallel neck; eye rather pygophore with distinct dorsal (DP) and ventral contiguous to anterior margin of pronotum processes (VP) (Figs 34–36, 47, 51); noticeably (distance between posterior margin of eye and elongate, sharply curved left paramere (Figs 48, pronotal collar less than length of eye in dorsal 53, 62); and disk-shaped bursa copulatrix (Figs 50, view); scutellum pale laterally (or with a pair 55, 65). of pale spots as in Fig. 32) ���������������������������� 6 6. Apical part (including apex) of antennal Distribution segment I fuscous; scutellum and basal half Nearly throughout the Old World (except for of cuneus bright yellow (Fig. 18); base of N. tenuis adventive to New World); speciose in metatibia pale ���������������Campyloneura virgula tropics and subtropics. – Antennal segment I (even if its median part darkened) always with pale or whitish apex; scutellum creamy white, sometimes darkened Discussion mesally; cuneus without bright yellow tint; ex- This Old World genus is currently composed of treme base of metatibia (at knee) narrowly but 26 species (Schuh 2002–2013 and present study). clearly darkened ���������������������7 [Nesidiocoris] Nineteen of them were originally described from the 7. Pronotum broadly (Figs 8, 19) or at least later- Ethiopian Region (including N. tenuis); N. tabaci ally darkened (Figs 20, 22) �������������������������� 8 (Froggatt) is restricted to Australia; and six are – Pronotum almost uniformly pale, often with a Asian species [three new species described herein short, longitudinal stripe between calli �������� 9 as well as N. caesar (Ballard) and N. cruentatus 8. Vertex and frons pale (Fig. 22); apex of (Ballard) from India, and N. pulchricornis (Pop- metafemur darkened (Fig. 8); known pius) from Java]. I also confirmed at least three from subtropical Ryukyu Islands �������������������� undetermined congeners from Indochina and ������������������������Nesidiocoris okinawanus sp. n. the Philippines. However, N. cruentatus appears – Head uniformly shiny fuscous (Figs 19, 20); to be a member of Singhalesia and needs further metafemur whitish brown; known from verification. temperate central Honshu ������������������������������ Currently, four species of Nesidiocoris are rec- �������������������������������������N. simotukensis sp. n. ognized to occur in eastern Asia (temperate and 9. Body shorter than 3.2 mm; antennal segment subtropical climate zones); however, east Asian N. I darkened except extreme base and apex pale; tenuis is considered to have been introduced from antennal segment III almost equal in length other zoogeographical regions of the Old World, as to II; scutellum broadly pale except apex more supposed for the New World and Australian popula- or less darkened (Figs 9, 11) ������������ N. tenuis tions (cf. Wheeler & HenryDownloaded 1992 from). Brill.com10/01/2021 07:03:11PM via free access
Nesidiocoris nozakianus Yasunaga sp. n. Male genitalia (Figs 59–63, 66): Pygophore some- Figs 1–4, 24, 25, 30, 40, 59–66 what expanded (cf. Fig. 24), with basally widened and blunt-tipped DP (Figs 59–61, 66); left para- Type material. Holotype ♂: Japan, Kyushu, Na- mere sharply bent, with basally tumid sensory lobe gasaki City, Konoura-Natsui Town, N32.873763, and noticeably elongate hypophysis (Figs 62, 66); E129.682715, 70 m alt., on Vernicia cordata, 16 endosoma with primary lobe (PL) uniformly mela- Sep 2016, T. Nozaki (AMNH_PBI 00380554) nized apically and secondary lobe (SL) developed (NIAES). Paratypes: Japan, Kyushu, same data as (Fig. 63). Female genitalia (Figs 64, 65): Ovipositor for holotype, 7 ♂, 4 ♀ (AMNH, TKPM, TNCF, (valvula II) elongate, moderately curved like a sword TYCN); same locality and plant, 20 Sep 2016, T. subapically (Fig. 64); bursa copulatrix with some- Yasunaga, 1 ♂, 2 ♀ (TYCN). what squared dorsal labiate plate (Fig. 65); sclero- Diagnosis. Recognized by its comparatively large tized ring reduced. size among Asian congeners (Fig. 4); pale antennal Measurements. ♂/♀: Total body length 3.6–3.8/3.6– segment I sometimes weakly obscured medially; 3.9; width of head across eyes 0.56–0.57/0.55–0.58; largely pale head with darkened clypeus and pale width of vertex 0.16–0.18/0.19–0.20; lengths of neck (Fig. 3); dark, short, longitudinal stripe be- antennal segments I–IV: 0.28–0.35, 1.06–1.13, tween pronotal calli; pale orange brown pronotal 1.10–1.18, 0.43–0.48/0.30–0.35, 0.87–0.95, 0.98– disc, mesoscutum and propleuron (Figs 4, 24, 25); 1.07, 0.41–0.50; length of labium 1.34–1.35/1.36– creamy yellow scutellum with dark, mesal stripe; 1.52; mesal pronotal length including collar whitish, semitransparent hemelytra with narrowly 0.55–0.57/0.51–0.59; basal pronotal width 0.83– but continuously infuscate apical and inner margin 0.88/0.79–0.92; maximum width across hemelytra of corium; and rather rounded posterior corner of 0.95–1.09/0.98–1.11; lengths of metafemur, tibia membrane vein (Fig. 30). Most closely related and and tarsus 1.40–1.52, 2.09–2.17, 0.72–0.75/1.33– externally very similar to N. caesar known from India 1.38, 1.90–2.02, 0.68–0.70. and Nepal (cf. Fig. 4 vs. Fig. 5), from which this new Etymology. Named after Mr Tatsuya Nozaki, discov- species can be distinguished by the characters men- erer of this unique mirid; latinized as an adjective. tioned in the discussion below. Biology. The adults, including teneral individu- Description. Body elongate, nearly parallel-sided; als, were found on a tung-oil tree, Vernicia cordata dorsal surface shining, with uniformly distributed, (Thunb.) Airy Shaw (Euphorbiaceae); however, silky, short, semierect setae. Head shiny whitish nothing is known about its life cycle. brown, weakly tinged with orange; eyes compara- Discussion. As mentioned above, N. nozakianus [N] tively large; more than apical half of clypeus dark- is in all likelihood sister to N. caesar [C] known ened. Antennae brown; segment I white, sometimes from the Indian subcontinent, but the following weakly obscured medially; extreme base and apex of characters (1–8) are particularly useful in distin- segment II whitish; median part of segment II and guishing between [N] and [C]: (1) vertex in fresh base of segment III slightly pale; segment II longer specimen almost uniformly pale orange-brown as in than basal width of pronotum, obviously shorter Figs 3, 4 [N]/tinged with light blue as in Figs 5, 6 than segment III. Labium shiny creamy brown, [C]; (2) base of head (neck) pale or weakly obscured reaching apex of mesocoxa; more than apical half [N]/distinctly darkened [C]; (3) antennal segment of segment IV dark brown. Pronotum shining, pale II obviously shorter than III [N]/about as long as orange brown, shallowly and finely punctate; calli or or slightly shorter than II [C]; (4) small areolar cell anterior lobe greenish, with a narrow, dark stripe at on forewing membrane very tiny as in Fig. 34 [N]/ midline; collar whitish brown, about as thick as an- rather wide (Fig. 35) [C]; (5) extreme bases (knees) tennal segment I; mesoscutum pale orange-brown, of pro- and mesotibiae not darkened [N]/more or polished; scutellum creamy white, broadly darkened less darkened [C]; (6) pygophore narrower (Figs 56, along midline; thoracic pleurites largely shiny pale 57) [N]/wider (Figs 59–61) [C]; (7) endosoma with orange-brown; ostiolar peritreme pale brown, nar- uniformly melanized PL and developed SL (Fig. 63) row. Hemelytra generally shiny whitish, semitrans- [N]/with apically melanized PL and smaller SL parent; inner and posterior margin of corium and (Fig. 58) [C]; and (8) additional differences shown apex of cuneus narrowly fuscous; membrane pale in Fig. 66 (cf. shape of pygophoral processes and left smoky brown, semitransparent, with mesal, obscure, paramere). rectangular macula subapically; venation on mem- Next to these morphological characters, each brane as in Fig. 30, with small areolar cell tiny. Coxae species appears to have a dissimilar host preference; and legs whitish brown; extreme base of metatibia N. nozakianus was observed to be associated with (knee) infuscate; apical half of each tarsomere III euphorbiaceaeous Vernicia cordata, whereas N. cae- dark brown. Abdomen generally shiny pale green; sar was found on Cucurbitaceae (e.g., bottle gourd, left paramere tinged with orange in fresh specimen. 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Figs 19–29. Habitus images for Nesidiocoris species. — 19, N. simotukensis, holotype male in dorsal view; 20, same, female; 21, same, male in ventral view; 22, N. okinawanus, holotype female; 23, same, in left lateral view; 24, N. nozakianus, male; 25, same, female; 26, N. caesar, male (from Kathmandu, Nepal, AMNH_PBI 00380564); 27, same, female (00380565); 28, N. tenuis, male from Ishigaki Island, Japan; 29, same (holotype of N. orientalis [=N. poppiusi] from Taiwan. Image courtesy of Dr M.L. Chan and Dr J.F. Tsai).
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Figs 30–44. Forewing membrane (30–33), terminal abdominal segment of male (34–39) and metathoracic scent efferent system (40–44) of dicyphines. — 30, 40, Nesidiocoris nozakianus; 31, 41, N. caesar; 32, 35, N. tenuis from Ishigaki Island; 33, 34, 42, N. simotukensis; 36, species identical to S. obscuricornis, from Panay Island, Philippines; 37, Cyrtopeltis miyamotoi; 38, Cy. nakatanii; 39, 44, Tupiocoris annulifer; 43, Cy. rufobrunnea.
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Figs 45–55. Male and female (50, 55) genitalia of Nesidiocoris tenuis (45−50) and N. simotukensis (51−55). — 45– 47, pygophore with left paramere in right lateral (45), ventral (46) and left lateral (47) view; 48, 53, left paramere; 49, 54, endosoma; 50, 55, bursa copulatrix; 51, 52, apical part of pygophore in left (51) and right (52) lateral view. Scale bars 0.2 mm. Abbreviations as mentioned in Materials and methods section.
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Figs 56–65. Male and female (64, 65) genitalia of Nesidiocoris caesar (45−50) and N. nozakianus (59−65). — 56, 57, 59, 60, 61, pygophore; 58, 63, endosoma; 62, left paramere; 64, 2nd valvula (ovipositor); 65, bursa copulatrix. Scale bars 0.2 mm. Abbreviations as mentioned in Materials and methods section.
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Fig. 66. Schematic images of characters in the male genitalia distinguishing between Nesidiocoris caesar (right) and N. nozakianus (left).
Discussion. This is a typical omnivorous plant bug to attract and propagate the mirid. Recent experi- now known from almost all zoogeographical regions. ments demonstrated that populations of insecticide- Several workers presumed Nesidiocoris tenuis is na- resistant pests, Thrips palmi Karny and Bemisia tive to somewhere in the Old World (cf. Wheeler tabaci (Gennadius), were successfully decreased after & Henry 1992). It indeed is not easy to trace the releasing a combination of predators, Nesidiocoris geographic origin of this globally widespread species. tenuis and Amblyseius swirskii Athias-Henriot (Acari: Nonetheless, the native habitat is presumed to be ei- Phytoseiidae) in eggplant greenhouses (Yanagita ther the Ethiopian or Oriental Region where most et al., 2016). However, when prey densities are of its congeners occur (Schuh 2002–2013). Wheeler sparse, this zoophytophagous mirid may injure crops & Henry (1992) suggested that N. tenuis has colo- (Takai & Yasunaga 2001, Rurallib 2016). nized several island groups and has been transported Poppius (1915) described Dicyphus orientalis from widely by commerce. Taiwan, based on a single male specimen; as the spe- In Japan, N. tenuis is mainly utilized to control cific name was a junior primary homonym, Carvalho whiteflies Bemisia( spp., Aleyrodidae) and thrips (1958) provided a new name, poppiusi, and placed (Thrips spp., Thysanoptera) in greenhouses produc- it in Nesidiocoris. I recently accessed clear digital ing cucumber, eggplant and tomato (Rurallib 2016). images of the holotype (Figs 11, 29; see National To introduce live individuals (mostly immature Museum of Natural Science 2017), and this taxon forms) of Nesidiocoris tenuis to such vegetables, plant- was confirmed to be conspecific with N. tenuis. ing Sesamum indicum L. (Pedaliaceae) and Cleome Accordingly, N. poppiusi (Carvalho) is proposed as a hassleriana Chodat (Cleomaceae) is recommended junior synonym of N. tenuis (Reuter). Downloaded from Brill.com10/01/2021 07:03:11PM via free access
Acknowledgements Kerzhner, I.M., 1988. Sem. Miridae (Capsidae)—Slepnjaki. I particularly thank the following individuals or in- Opredelitel’ nasekomykh Dal’nego Vostoka SSSR [Keys to the insects of the Soviet Far East], Vol. 2. — Nauka, stitutions for providing invaluable material, helping Leningrad, USSR, pp. 778–857. [in Russian] with my fieldwork and/or sharing necessary informa- Kerzhner, I.M. & Josifov, M., 1999. Miridae Hahn, tion: the late Mr S. Gotoh (Tanabe, Wakayama Pref., 1833. — In: B. Aukema & C. Rieger (Eds), Catalogue Japan); Dr P.K. Shrestha (Kathmandu, Nepal); Dr T. of the Heteroptera of the Palearctic Region, Vol. 3, Artchawakom and Dr C. Phuvasa (Thailand Insti- Cimicomorpha II, pp. 1–576. The Netherlands Ento- tute of Science & Technology); Dr M. Rut (Surana- mological Society, Amsterdam, The Netherlands. ree University of Technology, Thailand); Mr M. Konstantinov, F.V. & A.A. Knyshov, 2015. The tribe Bryo- Takai and Mr T. Befu (Kochi Pref., Japan); Dr Pyone corini (Insecta: Heteroptera: Miridae: Bryocorinae): Pyone Kyi, Dr Mu Mu Thein, Dr Khin Nyunt Yee, Phylogeny, description of a new genus, and adaptive Mr Zayar Soe and Mr Shine Shane Naing (Plant radiation on ferns. — Zoological Journal of the Lin- Protection Division, Department of Agriculture, nean Society 175: 441–472. Yangon, Myanmar); Dr K. Takahashi (Ushiku City, Namyatova, A.A., F.V. Konstantinov & G. Cassis, 2016. Ibaraki Pref.), Dr Y. Nakatani (NIAES), Dr K. Ya- Phylogeny and systematics of the subfamily Bryocori- mada (TKPM); Mr T. Nozaki (Fukuoka City); Mr nae based on morphology with emphasis on the tribe S. Maehara (Tochigi Pref.); Dr R.K. Duwal (Kyushu Dicyphini sensu Schuh, 1976. — Systematic Entomol- University, Fukuoka); Dr H.M. Yeshwanth (Banga- ogy 41: 3–40. lore, India); and JICA (Japan International Coop- National Museum of Natural Science, 2017. Integrated eration Agency) Cambodia, Myanmar, Nepal and insect types database of Taiwanese species. http:// Thai Offices. Mr M. Takai kindly provided images twinsecttype.nmns.edu.tw/specimens/. [accessed of live individuals (Figs 8, 12, 13), and Dr M.L. 30.iv.2017] Neimorovets, V.V., 2006. A new species of the genus Dicy- Chan and Dr J.F. Tsai (National Museum of Natu- phus Fieber, 1858 (Heteroptera: Miridae) from the Far ral Science, Taiwan) permitted me to use images in East of Russia and notes on taxonomical position of this paper (Figs 11, 29). Thanks are extended to Dr Cyrtopeltis miyamotoi (Yasunaga, 2000) comb. n. — A.G. Wheeler (Clemson University, SC, USA) and Russian Entomological Journal 15: 131–132. Dr D.A. Polhemus (Associate editor of TVE) for re- [in Russian] viewing the manuscript with valuable comments and Poppius, B., 1915. H. 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