Scutellera perplexa (Westwood, 1837)

Rédei, Dávid & Tsai, Jing-Fu, 2022, A revision of Scutellera (Hemiptera: Heteroptera: Scutelleridae), Zootaxa 5092 (1), pp. 1-40 : 17-24

Scutellera perplexa (Westwood, 1837)

( Figs. 18–23 , 34–37 , 44, 45 , 49–51 , 57–60 , 67, 68 )

Cimex nobilis (non Linnaeus, 1763): Fabricius (1775: 697; 1794: 80). Misidentification (Westwood 1837: 4, Dallas 1851: 18).

Tectocoris perplexa Westwood, 1837: 4 . Proposed for Cimex nobilis (non Linnaeus, 1763): Fabricius (1775: 697; 1794: 80). Syntypes: Ind[ia] Orient[ali] [= East Indies]; OXUM?, ZMUC? (not found).

Scutellera fasciata (non Panzer, 1798): Ahmad & Mushtaq (1977: 25, 30–37) and authors. Misidentification.

Cimex nobilis (misidentification): Sulzer (1776a: 113, 114) (redescription,distribution), Sulzer (1776b:pl. XI fig.c) (redescription, habitus), Fabricius ([1782]: 338) (diagnosis, distribution), Herbst (1784: 255) (redescription, habitus, distribution), Fabricius (1787: 280) (diagnosis), Gmelin (1790: 2128) (diagnosis, distribution), Fabricius (1794: 80) (diagnosis, distribution), Lichtenstein (1796: 102) (listed), Panzer (1798: 106) (redescription, habitus), Wolff (1801a: 49) (redescription, habitus, distribution), Wolff (1801b: 49) (redescription, habitus, distribution), Turton (1802: 615) (diagnosis, distribution).

Tetyra nobilis (misidentification): Fabricius (1803: 129) (diagnosis, distribution), Thunberg (1823: 5) (listed).

Tectocoris nobilis (misidentification): Hahn (1836: 24) (redescription, habitus, record).

Calliphara nobilis (misidentification): Germar (1839: 124) (redescription, distribution), Herrich-Schäffer (1839: 59) (in key), Herrich-Schäffer (1840: 73) (figures), Amyot & Serville (1843: 30) (redescription, distribution), Herrich-Schäffer (1853: 30) (listed, distribution).

Philia nobilis (misidentification): Schiødte (1843: 286) (diagnostic characters).

Scutellera nobilis (misidentification): Lamarck (1801: 293) (listed), Latreille ([1804a]: 164) (diagnosis), Latreille ([1804b]: 177) (diagnosis, distribution), Lamarck (1816: 491) (diagnosis, distribution), Latreille (1819: 442) (diagnosis, distribution), LePeletier & Serville (1828: 410) (listed), Burmeister (1835: 395) (diagnosis, distribution), Spinola (1837: 378) (diagnostic characters), Blanchard (1840: 158) (redescription, habitus, distribution), Dallas (1851: 18) (records), Dohrn (1859: 2) (catalogue, distribution), Dohrn (1860: 399) (listed), Vollenhoven (1863:11) (redescription, records, distribution, variability), Mayr (1866: 22) (record, variability), Walker (1867: 15) (records), Stål (1873: 14) (listed, records, distribution), Distant (1879: 44) (listed), Mason (1882: 43) (distribution), Atkinson (1884: 165) (distribution), Atkinson (1887: 161) (redescription, interspecific variability, records, distribution), Lethierry (1891: 142) (record), Cotes (1896: 82) (record, host plant), Distant (1901: 100) (record), Distant (1902: 51) (diagnostic characters, variability, distribution), Claybrooke (1903: 133) (listed), Nicéville (1903: 119) (figures of adult and preimaginal stages, host plant, economic importance, record, distribution), Schouteden (1904: 23) (catalogue, figure, distribution), Maxwell-Lefroy (1909a: 303) (records), Maxwell-Lefroy (1909b: 672) (habitus, figures of eggs and larvae, host plant), Distant (1918: 116) (synonymy), Bainbrigge Fletcher (1920: 250) (records, host plants), Singh-Pruthi (1925: 145) (male genitalia, figure), Reh (1932: 437) (host plant, phenology), Chatterjee (1934: 5, 28) (records, distribution, host plants), Hoffmann (1935: 31, 163, 181) (catalogue, distribution), Beeson (1941: 772) (host plants), Chandra (1953: 100) (records), Mathur et al. (1958: 88) (host plant, economic importance), Jande (1959: 215) (karyotype), Kumar (1965: 46) (male and female genitalia, figures), Baloch et al. (1968: 107) (host plant), Srivastava & Dogra (1969: 525) (anatomy), Takenouchi & Muramoto (1969: 10) (karyotype), Kavadia et al. (1971: 372) (host plant, economic importance, control), Nair (1975: 63, 101, 222) (host plants, economic importance), Ahmad & Mushtaq (1977: 25, 30) (habitus, figures, morphology), Mathur (1977: 9) (anatomy of nervous system), Ahmad et al. (1979: 30) (listed), Ueshima (1979: 75) (karyotype), Ahmad (1980: 134) (listed), Afzal et al. (1982: 378) (cladistics), Nuamah (1982: 16) (karyotype), Mall & Gupta (1983: 93) (physiology), Agarwal & Baijal (1984: 55) (figures, male genitalia), Datta et al. (1985: 21) (redescription, figure, distribution), Nair (1986: 71, 109, 221) (host plants, economic importance), Afzal & Sahibzada (1988: 254) (morphology), Ghosh et al. (1994: 495, 498) (listed, diagnosis, figure, records, distribution, habitat), Meshram & Garg (1999: 536) (record, host plant, economic importance), Hua (2000: 169) (listed, distribution), Chakraborty (2004: 169) (listed), Khokhar & Khokhar (2004: 218) (redescription, habitus, host plant, record), Manoharan et al. (2006: 67) (photo, records, host plants, economic importance, parasitoids), Ambika et al. (2007: 370) (host plant, economic importance, development, economic importance, control), Biswas & Bal (2007: 301, 304) (listed, diagnostic characters, record, distribution), Chandra (2008: 142, 152) (listed, diagnosis, record, distribution), Ghosh (2008: 417) (diagnosis, figure, photo, host plant, economic importance, control, distribution), Prabhakar et al. (2008: 84) (diagnostic characters, phenology, bionomics, economic importance, distribution), Kulkarni et al. (2009: 126) (record, host plant), Aland et al. (2010: 463) (listed, photo), Biswas & Bal (2010: 230, 241) (listed, diagnosis, record, distribution), Chandra et al. (2010: 43) (record), Divakara et al. (2010: 738) (host plant, economic importance), Kumar & Naidu (2010: 62) (listed), Patel & Borad (2010: 800) (host plant, control), Ranga Rao et al. (2010: 11) (diagnostic characters, photos, host plants, life history, economic importance), Sharma & Srivastava (2010: 1116) (host plant, economic importance, control), Rajmohana et al. (2011: 106) (parasitoid), Regupathy & Ayyasamy (2011: 169) (host plant, economic importance), Sharma & Srivastava (2011: 89) (host plant, economic importance), Tara et al. (2011: 223) (listed), Anitha & Varaprasad (2012: 190) (distribution, host plant, bionomics, economic importance, control), Chandra & Kushwaha (2012: 103) (record, photo), Chandra et al. (2012: 159) (listed), Terren et al. (2012: 226) (host plant, economic importance), Sanyal et al. (2012: 656) (distribution), Sharma & Srivastava (2012: 74) (host plant, economic importance, oviposition, development, growth), Wu et al. (2012: 25) (listed), Becker et al. (2013: 240) (host plant, economic importance), Datinton et al. (2013: 104) (host plant, economic importance), Francis et al. (2013: 400) (host plant, economic importance), Habou et al. (2013: 606, 608) (host plant, economic importance), López-Guillén et al. (2013: 4, 6, 15, 16) (host plant, economic importance, natural enemies), Martin et al. (2013: 2) (host plant, economic importance), Singh et al. (2013: 199) (host plant, economic importance), Alonso & Lezcano (2014: 8) (host plant, economic importance), Babu & Livingstone (2014: 1813) (mtDNA-COI sequences), Kumar & Singh (2014: 72) (host plant), Rashid et al. (2014: 45) (host plant, economic importance), Djimmy & Nacro (2015: 1525) (host plant, economic importance), Alamu et al. (2016: 82) (host plant, economic importance), Nerlekar & Rajmohana (2016: 8737) (host plant, economic importance, natural enemy), Nikam & More (2016: 210) (listed), Tekam et al. (2018: 962) (host plant, economic importance), Pathan et al. (2019: 408) (diagnostic characters, photos, record, host plant, economic importance), Neupane et al. (2021: 7) (host plant, economic importance).

Scutelera [inadvertent error] nobilis (misidentification): Ahmad et al. (1979: 47) (listed).

Scutellera noblis [inadvertent error] (misidentification): Ahmad et al. (1979: 35) (records, host plants, phenology, habitus).

Scutellera norrbilis [inadvertent error] (misidentification): Chakraborty et al. (1994: 473) (listed).

Scutellera nobiles [inadvertent error] (misidentification): Hedge (1995: 17) (diagnosis, distribution).

Callidea perplexa: Kirby (1891: 75) (record, distribution, variability).

Scutellera perplexa: Lethierry & Severin (1893: 269) (catalogue, distribution), Breddin (1909: 258) (record, comparison with S. brevirostris ), Kirkaldy (1909: 304, 380) (catalogue, distribution, host plants), Kirkaldy (1910: 109) (record), Bergroth (1915: 171) (listed), Hoffmann (1932a: 10) (listed), Tang (1935: 282) (catalogue, distribution), Ebeling (1950: 536, 554) (host plant, economic importance), Manna (1951: 6, 41) (record, host plant, karyotype), Manna (1958: 921) (karyotype), Stichel (1961: 728) (catalogue, distribution), Kumar (1962: 46, 53) (morphology, male and female genitalia, figures), Stichel (1962: 208) (catalogue, distribution), Sienkiewicz (1964: 114) (records), Baloch et al. (1968: 107) (host plant), Hsiao & Cheng (1977: 61) (in key, redescription, figure, photo, host plant, distribution), Nuamah (1982: 16) (karyotype), Wu (1984: 31) (distribution, host plants), Chen et al. (1985: 46) (host plants, distribution), Jiang (1985: 56) (distribution, host plants), Zhang et al. (1987: 117) (in key, host plants, distribution), Yang (1988: 101) (redescription, host plants, economic importance), Chen & Yang (1989: 46) (economic importance, distribution), Hua (1989: 43) (listed), Li et al. (1989: 38) (listed), Chen (1990: 154) (listed), Yang (1993: 211) (listed), Yu & Sun (1993: 77) (redescription, host plants, records), Zhang et al. (1994: 65) (distribution, host plants), Li et al. (1997: 44, 65, 144) (listed, host plants, distribution), Lin et al. (1999: 47) (redescription, habitus, distribution, host plants), Chen & Gu (2000: 53) (listed), Hua (2000: 169) (listed, distribution, host plants), Javahery et al. (2000: 491) (distribution, host plants, economic importance, control), Zhang & Chen (2003: 685) (listed, host plant), Göllner-Scheiding (2006: 200) (catalogue, distribution), Ou et al. (2006: 24) (listed), Shi & Xu (2006: 36) (host plant, in key), Wen & Yang (2007: 153) (listed, host plants), Yang (2007: 138) (host plants, economic importance), Chen et al. (2008: 106) (listed), Grazia et al. (2008: 936) (listed), Huang & Song (2008: 117) (records, host plant, economic importance), Parveen et al. (2010: 401) (redescription, photos, description, figures and photos of preimaginal stages, bionomics, phenology, development, host plant, economic importance), Tara & Sharma (2010a: 74) (diagnosis, photo, distribution, host plants, economic importance), Tara & Sharma (2010b: 266, 268) (host plant, diagnosis, economic importance), Sahai et al. (2011: 4684) (record, photos, description and photos of preimaginal stages, bionomics, phenology, development, host plant, economic importance), Tsai et al. (2011: 154) (diagnostic characters), Aland (2014: 90) (host plant, bionomics), Chandra et al. (2014: 200) (record, photo), Kumar & Singh (2014: 72) (host plant), Parveen et al. (2014: 237, 251) (record, morphology, figures), J.P. Singh et al. (2014: 265) (photo, record, host plant), K. Singh et al. (2014: 358) (photo, record, host plant, economic importance), Onkar & Nerlekar (2015: 179) (record, photo of 5th instar larvae, host plant), Singh & Kaur (2015: 90) (diagnosis, photos of adults and immatures, record, development, host plants, economic importance), Cai & Cui (2017: 622) (redescription, habitus, host plants, distribution), Sheikh & Iqbal (2017: 80) (diagnostic characters, photo, record, distribution), Jamwal et al. (2020: 154) (redescription and photos of adult and immatures, development, behaviour, host plant, economic importance, distribution), Neupane et al. (2021: 7) (host plant, economic importance).

Scutellera perplex [inadvertent error]: Yang (1934: 257) (diagnostic characters, figure, records, distribution), Yang (1962: 23, 28) (in key, redescription, host plant, records, distribution), Yang & Wu (1981: 74) (host plants, distribution), Zhang et al. (1985: 11) (record, distribution), Chen (1987: 127) (redescription, habitus, distribution), Shen (1993: 32) (listed, host plants), Shen et al. (2014: 329) (listed, distribution), Bhagat (2015: 132) (host plant, economic importance).

Scutellera perflexa [inadvertent error]: Goel (1972: 169) (unguitractor plate).

Scutellera preplexa [inadvertent error]: Ueshima (1979: 75) (karyotype).

Scutelleria [inadvertent error] perplexa: Hilgendorf & Goeden (1982: 149) (host plant, distribution).

Scutella [inadvertent error] perplexa: Zhang (1985: 42) (distribution), Zhang (1986: 74) (distribution), Chen & Yang (1988: 89) (listed).

Scutella [inadvertent error] perplex [inadvertent error]: Bhagat (2015: 129) (listed, distribution).

Scutllera [inadvertent error] perplexa: He et al. (2010: 14) (host plant, distribution).

Scutellera fasciata (misidentification): Mushtaq et al. (1975: 45) (preimaginal stages), Ahmad & Mushtaq (1977: 25, 30) (listed, habitus, figures, morphology), Ahmad & Moizuddin (1978: 95) (descriptions and figures of preimaginal stages, host plant, economic importance), Ahmad et al. (1979: 30, 35) (listed, record, host plant, phenology), Ahmad (1980: 134) (listed), Ahmad & Moizuddin (1980: 198) (anatomy), Moizuddin & Ahmad (1980: 19) (record, host plant, anatomy), Moizuddin & Ahmad (1981: 63) (anatomy), Afzal et al. (1982: 378) (cladistics), Javahery et al. (2000: 491) (host plant, economic importance).

Diagnosis. Highly similar to S. nepalensis , the morphology of the exoskeleton is virtually identical in the two species. The dorsal markings of S. perplexa are highly variable ( Figs. 18, 20, 22 ) but even in extreme cases the paired, obliquely transverse fasciae anteriad and posteriad of the middle of scutellum do not join the median vitta to form an extensive, confluent pattern as in S. nepalensis ( Figs. 1, 2 , 11, 12, 15 ). The two species are most reliably separated by the genitalia as it is explained under the Diagnosis of S. nepalensis .

Redescription. Colour. Dorsum bright metallic blue-green, with complex black markings ( Figs. 18, 20 ) which might be reduced or completely lacking ( Fig. 22 ); head uniformly metallic blue-green, without black markings except of a pair of narrow, linear, parallel vittae margining clypeus and extending to base of head; scape orange to red with a narrow black annulus apically, pedicel and flagellum black; labium orange to reddish, segment III darkened, segment IV black; pronotum broadly margined with orange or red laterally, cicatrices narrowly bordered by black, a broad median vitta, a pair of small, oval submedian spots and a pair of small humeral spots on posterior lobe black; scutellum with a median vitta from its base to about its basal half, gradually tapering posteriad, with a pair of rounded spots at posterior part of basal tumescence, a pair of oval submedian spots anteriad and another pair posteriad to middle, invariably separated from median vitta, a pair of small marginal spots at middle, and a large subapical spot black; exposed portion of fore wing black, its base marginally orange; venter of head, thorax and abdomen as described in S. nepalensis .

Morphology of the exoskeleton as in S. nepalensis .

External male genitalia ( Figs. 26, 27 , 34–37 , 44, 45 , 49–51 ). Genital capsule ( Figs. 26, 27 , 34 ) subrectangular, boadly transversely truncate posteriorly in dorsal view, minute submedian denticles on posterior margin broadly separated, lateral margin (ventral rim) not emarginate and lacking distinct tubercles; infolding of ventral rim conspicuously bulging laterally, without distinct ridge along meson. Paramere ( Figs. 35–37 ) similar to that of S. nepalensis but crown more strongly curved, its distal portion almost perpendicular to axis of stem. Phallus (repose: Figs. 44, 45 , 51 ; inflated: Figs. 49, 50 ): second conjunctival processes with a large, flap-like lateral lobe (cp-II 1) with its ventral portion provided with several minute denticles, and with a mesal lobe (cp-II 2) terminating in a relatively long, claw-shaped sclerotized process; third conjunctival processes (cp-III) long, gradually tapering towards apex but distalmost portion again broadened; distal portion of aedeagus s. str. and phallotreme narrow.

External female genitalia ( Figs. 57–60 , 67, 68 ). Ovipositor ( Figs. 57–60 ). Laterotergites VIII flat, not protruding posteriad; laterotergites IX rather broadly rounded distally, closely approaching each other or adjacent along midline, greatly covering sternite X and median part of fused valvifers IX. Gynatrium ( Fig. 67 ) with ring sclerites rather broadly separated from apex of anterolateral pouch proximally; a pair of small sclerites posteriad of spermathecal opening. Spermatheca: proximal duct slightly longer than distal duct and longitudinal diameter of dilation.

Measurements (in mm). Body length to apex of scutellum 16.0–20.5; length of head 3.50–3.75, width across eyes 3.75–4.30, interocular distance 2.60–3.05; lengths of scape 1.00–1.13: basipedicellite 0.73–0.88: distipedicellite 1.85–1.90: basiflagellum 2.30–2.60: distiflagellum 2.25–2.55; median length of pronotum 3.95–5.20, humeral width 6.80–8.65; length of scutellum 10.0–13.2, greatest width 5.95–7.70.

Intraspecific variability. The black markings of the dorsum are strongly variable, individuals lacking any markings ( Figs. 22, 23 ), others with extensive markings ( Fig. 18–21 ), and various transitional forms exist. Individuals with strongly reduced or absent black markings are most common in the Malabar Subregion and Sri Lanka.

Preimaginal stages. Descriptions, figures or photos of the egg batch and different larval instars were presented by Nicéville (1903), Ahmad & Moizuddin (1978) (misidentified as S. fasciata ), Parveen et al. (2010), J.P. Singh et al. (2014), K. Singh et al. (2014), Onkar & Nerlekar (2015), Singh & Kaur (2015) and Jamwal et al. (2020).

Karyotype. 10+XY ( Manna 1951, Jande 1959). Detailed description and figures were presented by Manna (1951).

Bionomics, economic importance. The species was recorded (partly as S. nobilis ) from the jujube species Ziziphus nummularia (Burm.f.) Wight & Arn. (as S. fasciata , misidentification) and Z. jujuba (Burm.f.) Wight & Arn. (both Rhamnaceae ), neem, Azadirachta indica A.Juss. ( Meliaceae ) ( Ahmad et al. 1979, as S. nobilis and S. fasciata , misidentification), and on Xanthium strumarium L. ( Asteraceae ) ( Baloch et al. 1968, as both S. nobilis and S. perplexa ) in Pakistan. In India it is common in sandal wood plantations on Santalum album L. ( Santalaceae ), Phyllanthus emblica L. and Ph. acidus (L.) Skeels (= Ph. distichus Müll.Arg. ) (both Phyllanthaceae ) ( Bainbrigge Fletcher 1920, Chatterjee 1934, Beeson 1941, Kavadia et al. 1971, Datta et al. 1985, Ghosh et al. 1994, Ghosh 2008, Aland 2014). It is a well-known pest of physic nut, Jatropha curcas L. ( Manoharan et al. 2006, Ambika et al. 2007, Prabhakar et al. 2008, Divakara et al. 2010, Parveen et al. 2010, Ranga Rao et al. 2010, Sharma & Srivastava 2010, Tara & Sharma 2010 a, Sahai et al. 2011, Singh et al. 2013, K. Singh et al. 2014, Jamwal et al. 2020 etc.); it probably also feeds on J. nana Dalzell & A.Gibson ( Kulkarni et al. 2009) (both Euphorbiaceae ). Further plants recorded as its host plants in India include Antidesma ghaesembilla Gaertn. ( Manna 1951) (Phyllanthaceae) , Emblica officinalis Gaertn. ( Meshram & Garg 1999, Pathan et al. 2019), ricinus (castor), Ricinus communis L. ( Nair 1975, 1986, Panwar 1995) (all Euphorbiaceae ), the jujube species Ziziphus mauritiana Lam. (Rhamnaceae) (J.P. Singh et al. 2014), citrus plants ( Ebeling 1950) and the curry tree, Murraya koenigii (L.) Spreng. ( Tara & Sharma 2010a, 2010b) (both Rutaceae ), Casearia tomentosa Roxb. (Flacourtiaceae) ( Cotes 1896, Chatterjee 1934), Dodonaea viscosa Jacq. (Sapindaceae) ( Chatterjee 1934, Beeson 1941), Buchanania cochinchinensis (Lour.) M.R. Almeida (Anacardiaceae) ( Onkar & Nerlekar 2015), grape vine, Vitis vinifera L. ( Vitaceae ) ( Nicéville 1903, Mathur et al. 1958, Nair 1975, 1986, Singh & Kaur 2015), cotton, Gossypium sp. (Malvaceae) ( Bainbrigge Fletcher 1920, Nair 1975, 1986, Khokhar & Khokhar 2004) and “wild melon” (unspecified species of Cucurbitaceae ) ( Chatterjee 1934). It was reported to damage “gooseberry” (probably Indian gooseberry, Phyllanthus emblica L., Phyllanthaceae ) ( Ranga Rao et al. 2010) as well, but the nature and extent of the damage it caused was not stated and is probably insignificant; a specimen collected on this plant in Andra Pradesh, India, was seen during the present study.

FIGURES 65–70. Scutellera spp. , ectodermal genital tract of female. Figs. 65, 66, S. nepalensis (Westwood, 1837) ; Figs. 67, 68, S. perplexa (Westwood, 1837) ; Figs. 69, 70 , S. spilogastra ( Walker, 1867) . Gynatrium and spermatheca, dorsal view ( Figs. 65, 67, 69); apical receptacle ( Figs. 66, 68, 70 ). Abbreviations: dil = dilation of spermatheca; dd = distal duct of spermatheca; fec = fecundation sclerite; gy = gynatrium; pd = proximal duct of spermatheca; rs = ring sclerite; v 8 = ventrite VIII; va 8, va 9 = valvulae VIII, IX. Scales in mm.

Host plants reported from China include Indian gooseberry ( Zhang et al. 1994); “mandarin orange”, probably referring to Citrus ponki (Hayata) hort. ex Tanaka and/or other hybrids and cultivars based on C. reticulata Blanco , and other unspecified citrus species ( Rutaceae ) ( Hsiao & Cheng 1977, Yang 1988, Shen 1993, Hua 2000); oilseed camellia, Camellia oleifera Abel (Theaceae) ( Chen et al. 1985, Yang 1988, Shen 1993, Yu & Sun 1993, Lin et al. 1999, Zhang & Chen 2003); tung tree, Vernicia fordii (Hemsl.) Airy Shaw (all Euphorbiaceae ) ( Zhang et al. 1987); olive, Olea europaea L. ( Oleaceae ) ( Zhang et al. 1987, Yang 2007); Eucalyptus camaldulensis Dehnh. ( Zhang et al. 1987) , E. urophylla S.T. Blake and its hybrid species with E. grandis W.Hill (all Myrtaceae ) ( Huang & Song 2008); and Alnus cremastogyne Burkill (Betulaceae) ( Zhang et al. 1987). Records from Yunnan pine, Pinus yunnanensis Franch. , and unspecified pine trees ( Pinaceae ) ( Chen et al. 1985, Zhang et al. 1987, Yang 1988, Shen 1993, Yu & Sun 1993, Shi & Xu 2006) are considered as questionable.

Detailed studies on the life history of this species were presented by Ambika et al. (2007), Parveen et al. (2010), Sahai et al. (2011) based on observations made on populations developing on physic nut in northern India and by Sharma & Srivastava (2012) and Jamwal et al. (2020) based on laboratory colonies. Their data are partly unclear and contradictory but apparently suggest that the species is bivoltine and follows the phenology of the plant that exhibits fruit production from midsummer to late autumn. Individuals are active all year round, with a peak in the monsoon season (June to August), but with reduced activity during the hottest and coldest months; they migrate to surrounding plants or weeds when the plants defoliate ( Parveen et al. 2010, Jamwal et al. 2020). Copulating pairs can be found from April to late November, but most abundantly from June to August. Oviposition apparently takes place mostly in late March ( Sahai et al. 2011) and in August ( Sahai et al. 2011) or September–October ( Parveen et al. 2010). Eggs are laid usually on the undersides of young leaves, more rarely on the stems, petioles or fruits of the host plant in a single batch forming two rows, containing about 26–52 (laboratory) or 37–71 (field) eggs. The fecundity is 76– 84 eggs / female ( Ambika et al. 2007, Jamwal et al. 2020). Eggs hatch after 5–7 days; first instars remain aggregated around the empty shells for about three days, then disperse, but the subsequent instars frequently exhibit aggregating behaviour too. The postembryonic development takes about 27 days ( Ambika et al. 2007, Parveen et al. 2010, Sahai et al. 2011, Sharma & Srivastava 2012, Jamwal et al. 2020). No published information is available about its life history in China.

It is a major pests of the physic nut in India. Adults and larvae mainly attack the flowers, young green fruits and fruit pedicels. Feeding on the inflorescence causes premature flower fall ( Prabhakar et al. 2008). Feeding on fruits cause punctures on the pericarp; strongly injured fruits will deform, dark necrotic spots appear around the place of the feeding, the pericarp will frequently rupture; the damage was studied in detailed and damaged fruits were illustrated by Sahai et al. (2011) and Jamwal et al. (2020). It causes deformed or hollowed seeds, decreases seed production ( Sahai et al. 2011, Jamwal et al. 2020) and reduces the weight of the pod ( Ambika et al. 2007) and the quality and quantity of the oil ( Parveen et al. 2010). Fruits on the periphery of the bunch are affected more severely. An indirect damage is inflicted by fungi attacking the fruits through the feeding punctures ( Parveen et al. 2010). Feeding on the leaf causes chlorotic or necrotic spots and premature leaf fall ( Jamwal et al. 2020).

The species inflicts serious damage on Murraya koenigii in northern India by feeding on the fruits, seeds and leaves ( Tara & Sharma 2010a, 2010b). It causes significant harm to citrus trees as well ( Ebeling 1950). Mass occurrence on the jujube species Ziziphus mauritiana , with individuals present on more than 75% of the trees in the plantation, resulting in serious damage was reported from Jharkhand, India (K. Singh et al. 2014). It is a minor pest of Ziziphus nummularia in Pakistan ( Ahmad & Moizuddin 1978), and of the grape vine ( Mathur et al. 1958, Singh & Kaur 2015), Phyllanthus acidus ( Kavadia et al. 1971) and Emblica officinalis ( Meshram & Garg 1999, Pathan et al. 2019) in India.

Yang (1962) claimed that it is a forestry pest in China but without providing any details. Slight damage caused by this species in plantations of Eucalyptus spp. was reported ( Huang & Song 2008) and it was listed among pests of mandarin orange ( Chen & Yang 1989), olive ( Yang 2007) and Yunnan pine ( Shi & Xu 2006) in southern China without further details. Li et al. (1997) listed it as minor pest of citrus crops in southern China. The record of an unidentified Scutellera sp. inflicting damage in physic nut plantations in Sichuan ( Zhou et al. 2008) probably pertains to this species.

Kavadia et al. (1971) found that carbaryl, malathione, lindane, and “BHC” (= hexachlorobenzene) are effective against the species in descending order; the latter two are now banned under the Stockholm Convention on Persistent Organic Pollutants. Ambika et al. (2007) and Sharma & Srivastava (2010) found that the species can effectively be controlled in physic nut plantations using lambda-cyhalothrin, imidacloprid, carbosulfan, monocrotophos or dichlorvos. Patel & Borad (2010) compared the efficacy of several plant-derived insecticides. Biological control might be perspectivic as the entomopathogenic fungi Beauveria bassiana (Bals.-Criv.) Vuill. and Metarhizium anisopliae (Metchnikoff) Sorokin (both Clavicipitaceae ) were found to be effective in reducing the population under laboratory conditions ( Ambika et al. 2007). The hymenopteran egg parasitoids Anastatus bangalorensis Mani & Kurian, 1953 (Eupelmidae) , Trissolcus jatrophae Rajmohana, Narendran & Manoharan, 2011 (Scelionidae) , and unidentified species of Eurytomidae and Pteromalidae , as well as tachinids also have role in controlling the populations of S. perplexa in India ( Manoharan et al. 2006, Rajmohana et al. 2011).

Remarks. Cimex nobilis Linnaeus, 1763 was listed by Fabricius (1775: 697, 1794: 80) together with two very brief and inadequate redescriptions which nevertheless greatly overlap therefore apparently pertain to the same biological species. In the second work Fabricius (1794) referred to illustrations of scutellerids by Stoll (1780: pl. I fig. 1), Schröter (1776: pl. I fig. 9) (both unnamed) and Sulzer (1776b: pl. XI fig. c) (as Cimex nobilis ) as putatively pertaining to Cimex nobilis . However, none of these illustrations actually show Cimex (now Calliphara ) nobilis . Westwood (1837) correctly pointed out the misidentification and proposed the name Tectocoris perplexa for “Nobilis. Fab. non Linn.”, presumably referring to Fabricius (1775, 1794). The name is accordingly to be treated as proposed by indication ( ICZN 1999, Art. 12.2.1), and its type material consisted of an unspecified number of specimen(s) of the Hope collection (Westwood 1837) plus specimens in the collection of Fabricius ( Hemiptera now deposited in ZMUC) and those of Stoll, Sulzer and Schröter (now all lost) ( ICZN 1999, Art. 72.4.2). No specimen which could be considered as a syntype could be located during a visit of us to OXUM where F.W. Hope’s collection, including most of O.J. Westwood’s Hemiptera types, are currently deposited; no specimen of this species was listed by Zimsen (1964); and Lars Vilhelmsen informed us that no specimen of this genus is present in Fabricius’s collection at ZMUC either. The species, however, is a distinctive one unquestionably identifiable from the above mentioned three illustrations of early authors, and except of a few evident misidentifications almost all subsequent authors identified the species accordingly. As there is a general agreement among the authors about the identity of this species, currently there is no exceptional need ( ICZN 1999, Art. 75.3) to designate a neotype. A redescription and illustrations are provided in the present work to facilitate identification of the species.

Although the misidentification of Cimex nobilis sensu Fabricius (1775, 1794 ) was already correctly pointed out by Westwood (1837), still several subsequent authors persisted in referring to the species as Scutellera nobilis , but with the authorship of J.C. Fabricius. This practice is incorrect and should be discontinued.

The photos presented by Shanker & Dhyani (2006) and Anitha & Varaprasad (2012) as S. nobilis are based on misidentification and pertain to Chrysocoris purpureus (Westwood, 1837) . A partial sequence of the mitochondrial gene of cytochrome oxidase subunit I (mtDNA-COI) of S. perplexa was provided by Babu & Livingstone (2014).

Distribution. Widely distributed in the Indian Subcontinent , Southern China and Indo-China, restricted to the zone of tropical and subtropical moist broadleaf forests ( Fig. 71 ). The records of S. fasciata (= S. nepalensis ) from Pakistan ( Ahmad & Moizuddin 1978, Ahmad et al. 1979) apparently pertain to this species (see discussion under S. nepalensis ). We provide first country records for Nepal, Vietnam and Laos. The records from Henan ( Hsiao & Cheng 1977, voucher specimen seen) and Shaanxi (Wen & Yang 2007) are perhaps based on mislabelled or migrating specimens, the presence of stable population of this species in these areas seems unlikely. Old specimens were seen from Palawan, the Malay Peninsula, Sumatra, and Java (in all cases without precise localities), but these are doubtful, almost certainly representing cases of mislabelling or at best accidental introductions.

PAKISTAN. Swat Distr.!; Islamabad!; Punjab: Lahore!, Gojra ( Maxwell-Lefroy 1909a, as nobilis ); Sind: Karachi ( Atkinson 1887, Distant 1902, Ahmad et al. 1979, all as nobilis ); North-West Frontier Province: Kaghan ( Ahmad et al. 1979, as noblis [sic]).— INDIA. Jammu and Kashmir: Ladakh!, Jammu, Kathua and Samba Districts ( Tara & Sharma 2010a, Jamwal et al. 2020); Himachal Pradesh!; Punjab: Panjab [= Punjab] ( Atkinson 1887, as nobilis ), Ludhiana ( Singh & Kaur 2015); Uttarakhand!; Haryana ( Khokhar & Khokhar 2004, as nobilis ); Delhi ( Parveen et al. 2010, 2014); Uttar Pradesh: Barkachha ( Sharma & Srivastava 2010, as nobilis ), Lucknow ( Sahai et al. 2011); Gujarat: Anand ( Patel & Borad 2010, as nobilis ), Vadodara ( Kumar & Naidu 2010, as nobilis ), Jagudan ( Pathan et al. 2019, as nobilis ); Madhya Pradesh: Churikhurd ( Meshram & Garg 1999, as nobilis ), Jabalpur Distr. ( Chandra 2008, as nobilis ; Sheikh & Iqbal 2017); Maharashtra: Mumbai!, Matheran!, Ahmednagar!, Pune ( Kulkarni et al. 2009, as nobilis ; Onkar & Nerlekar 2015), Kolhapur Distr. ( Aland et al. 2010, Nikam & More 2016, both as nobilis ); Goa!; Karnataka: Belagavi!, Srirangapatna!, Ablathi!, Bangalore [= Bengaluru] ( Distant 1902, as nobilis ), Fraserpet [= Kushalanagar], Kottur[u] ( Chatterjee 1934, as nobilis ); Kerala!; Tamil Nadu: several localities!, also Salem: Aiyur, Jawalagiri ( Chatterjee 1934, Chandra 1953, both as nobilis ); Puducherry!; Andhra Pradesh: Anakapalle!, Visakhapatnam ( Biswas & Bal 2007, as nobilis ); Chhattisgarh: Barnawapara Wildlife Sanctuary (Chandra & Kushwaha 2012, as nobilis ); Odisha (Orissa)!; Bihar: Pusa ( Chatterjee 1934, Chandra 1953, both as nobilis ); Jharkhand: Konbir!, Ranchi (J.P. Singh et al. 2014); West Bengal: Kolkata!, Gopaldhara!, Asansol ( Lethierry 1891, as nobilis ), S[h]ibpur ( Manna 1951); Sikkim!; Assam: Cachar [= Silchar] ( Atkinson 1887, as nobilis ); Meghalaya!; Manipur ( Distant 1902, as nobilis ).— SRI LANKA!— NEPAL!— BURMA ( MYANMAR). Mon State!; Tanintharyi Region!: Moulmein ( Dallas 1851, Distant 1901, 1902, all as nobilis ), Tavoy [= Dawei] ( Distant 1902, as nobilis ).— THAILAND. Chiang Rai Prov.!; Chiang Mai Prov.!— CHINA. Shaanxi?: Sangyuan National Nature Reserve? (Wen & Yang 2007); Henan?: Xinyang! [in error?], Xuchang?, Shangcheng? ( Yu & Sun 1993, Shen et al. 2014); Shanghai!; Sichuan!; Guizhou: Libo ( Wu 1984); Yunnan: all over the province!, several additional localities by Yang (1962), Hsiao & Cheng (1977), Jiang (1985), Zhang et al. (1987) etc.; Guangxi!; Hainan!; Guangdong!; Fujian: Fuzhou!, Jianyang!, Sanming ( Chen et al. 1985, Lin et al. 1999), Hua’an, Longhai, Changtai ( Huang & Song 2008); Macau ( Kirkaldy 1910).— VIETNAM. Ho Chi Minh City!, Hanoi [= Hà Nội] ( Claybrooke 1903, as nobilis ).

Specimens examined. PAKISTAN. Swat Distr.: Kalam, 6800 ft., 8.x.1966, leg. R. Traub (1 ♀ USNM); Islamabad: 20.xi.[19]75, from Zizyphus jujuba , leg. Mohsin (1 ♂ SDEI, 1 ♂ 1 ♀ USNM, all det. as S. fasciata by A.A. Khan, 1981), same but 17.xi.[19]75 (1 ♀ SDEI, det. as S. fasciata by A.A. Khan, 1981); Punjab: Lahore, 18. vi.[19]65, leg. Rahi [?] (1 ♂ USNM, det. as S. fasciata by I. Ahmad, 1975).— INDIA. Jammu and Kashmir: “Tibet: Ladak” [= Ladakh], coll. Felder (1 ♂ 2 ♀♀ RMNH), Jammu ( Tara & Sharma 2010a, 2010b); Himachal Pradesh: Sadhupul, nr. S[h]imla, 1250 m, 27.x.1978 (1 ♀ SEHU); Uttarakhand: Lansdowne Division, F.W.C. (2 ♀♀ BMNH), Sarda [= Sharda River?], F.W.C. (1 ♂ BMNH), Dehra Dun: Dobhalwalla [= Dobhalwala], 2304 ft., on Jatropha curcas , 4.xi.1945, leg. J.K. Uniyal, coll. J.C. Lutz 1961 (5 ♂♂ 6 ♀♀ USNM), Kumaon, Almora, H.G.C. (3 ♂♂ BMNH), Kumaon, Gori Valley, 4000 ft., H.G.C. (1 ♂ BMNH); Maharashtra: Bombay [= Mumbai] (1 ♀ HNHM, 1 ♀ ZSMC), same, coll. Bergroth (1 ♀ MZHF), same, coll. Leith> W.L. Distant, B.M.1911-383 (1 ♂ 2 ♀♀ BMNH), Matheran, 800 m, 8.viii.[1]902, leg. Bíró (1 ♀ HNHM), Achmednagar [= Ahmednagar], coll. Gebauer (2 ♂♂ 2 ♀♀ NHMW), ca. 30 km W of Patan, 17º22’N 73º54’E, 570 m, near river, 12.vi.2006, leg. Z. Kejval (1 ♂ MMBC); Goa: Mormugao, vi.1925, leg. J.C. Bridwell (2 ♂♂ 3 ♀♀ USNM), 30 km S of Margao, env. of Palolem, 0–20 m, 15º00.47’N 74º01.58’E, 14–21.viii.2002, leg. P. Šípek & M. Fikáček (1 ♂ 1 ♀ NMPC); Karnataka: Belgaum [= Belagavi], 88.148 (1 ♀ BMNH), Chikkaballapura, T.V. Campbell, ix.[19]14, B.M.1930-599 (1 ♂ BMNH), same but iii.1915 (1 ♀ BMNH), Seringapatam [= Srirangapatna] (1 ♀ HNHM), Ablathi, 800 m, 12.17ºN 76.06ºE, x.1984, leg. W. Lorenz (1 ♀ ZSMC); Kerala: Malabar, coll. W.L. Distant, B.M.1911-383 (1 ♀ BMNH), same, leg. Speyer, coll. G. Breddin (1 ♀ SDEI), S. Travancore, Thekkadi [= Thekkady], leg. Bourdillon, coll. G. Breddin (1 ♀ SDEI), Periyar Dam, Travancore, 6–10.v.[19]37, B.M.–C.M. Expedition to South India 1937 (1 ♀ BMNH), Trivandrum [= Thiruvananthapuram] Distr., Poonmudi [= Ponmudi], 900 m, iv–v.[19]71, leg. P. Susai Nathan (1 ♀ BPBM); Tamil Nadu: Nilgiri [Hills], leg. Hampson, coll. W.L. Distant, B.M.1911-383 (1 ♀ BMNH), Nilgiri Hills, 20.x.1983, leg. H. Kitahara (2 ♀♀ NSMT), Kodai Kanal [= Kodaikanal], T.V. Campbell, B.M.1930- 599 (2 ♂♂ BMNH), Madura[i]: Shembaganur, coll. H. Rolle (1 ♀ RMNH), Ana[i]mala[i] Hills, Cinchona [= Cinkona], 3500 ft., v.1960, leg. P. Susai Nathan (1 ♀ RMNH), Anaimalai, Top Slip, 550–800 m, 2.xii.1978 (1 ♂ 3 ♀♀ SEHU), Hasanur, Biligirirangan Hills, 29.iv.[19]37, B.M.–C.M. Expedition to South India 1937 (1 ♀ BMNH), Coimbatore, vii.[19]26, leg. P. Susai Nathan, coll. C.J. Drake 1956 (1 ♂ 1 ♀ USNM), same but x.1937, leg. P. Susai Nathan, coll. H.M. Harris [19]74 (1 ♂ USNM), same but 11.viii.1938 (1 ♀ USNM), same locality, vii.[19]46, leg. P. Susai Nathan (1 ♂ 1 ♀ NMPC), same but vii.[19]47 (2 ♂♂ 1 ♀ NMPC), same locality and collector, 4.iv.1963, B.M.1966-407 (1 ♀ BMNH), same locality and collector, 1400 ft. (1 ♀ RMNH), same but iv.1960 (3 ♀♀ RMNH), same but xi.1966 (1 ♂ 2 ♀♀ ZMAN > RMNH), same but x.1968 (1 ♂ 3 ♀♀ ZMAN > RMNH), Coimbatore Distr., Marudamalai Hills, 1800 ft., x.1969, leg. P. Susai Nathan (2 ♂♂ 2 ♀♀ ZMAN > RMNH), Walayar, 9.ix.1938, leg. P. Susai Nathan, coll. H.M. Harris [19]74 (1 ♂ USNM), Walayar Forest, 1000 ft., ix.1951, leg. P. Susai Nathan, coll. J.C. Lutz 1961 (3 ♀♀ USNM), same but vii.1952 (2 ♂♂ USNM), same but viii.1952 (2 ♂♂ 1 ♀ USNM), Shembaganur, leg. A. Heyne (1 ♀ SDEI), Madras (1 ♂ 1 ♀ RMNH, 1 ♂ USNM, 1 ♂ 1 ♀ ZSMC), same, coll. Signoret (1 ♀ NHMW), same, coll. G. Breddin (2 ♂♂ 1 ♀ SDEI), same but coll. Moore (1 ♀ RMNH); Puducherry: Pondichéry [= Pondicherry, Puducherry], coll. E. le Moult (1 ♀ RMNH), same but coll. Signoret (1 ♂ 1 ♀ NHMW), same but 1902, coll. Maindron (1 ♀ MNHN > IZAS), Kar[a]ikal, x.1968, leg. P. Susai Nathan (1 ♀ BPBM); Andhra Pradesh: Anakapalli [= Anakapalle], breeding on Phyllanthus emblicus [= emblica ], ii.1925, leg. J.R. Rao (1 ♂ 1 ♀ BMNH); Odisha (Orissa): Balasore, 21.vii.1915, leg. H. Lindsay (2 ♂♂ 1 ♀ BMNH); Jharkhand: Konbir (1 ♀ USNM), Kombir [= Konbir] (1 ♂ USNM); West Bengal: Calcutta [= Kolkata] (1 ♀ HNHM, 1 ♂ 1 ♀ USNM), same but coll. E.T. Atkinson, B.M.92-6 (1 ♀ BMNH), same but vi.1968, leg. D.W.P., coll. P. Roche (1 ♀ RMNH), Gopaldhara, Rungbong [= Rangbhang] Valley, H. Stevens, B.M.1922-307 (1 ♀ BMNH), Gopaldhara, 1916, H. Stevens (1 ♂ BMNH); Sikkim (1 ♀ HNHM); Meghalaya: Garo Hills, 1500-2500 ft., coll. W.L. Distant, B.M.1911-383 (1 ♂ BMNH), W Garo Hills, 25º30.7’N 90º13.9’E (WGS84), 600–800 m, 29–31.v.1996, leg. E. Jendek & O. Šauša (1 ♀ NHMW).— SRI LANKA. Trincomali [= Trincomalee], ix.1910 (2 ♀♀ MZHF), xii.1979, leg. P.J.J.H. Kuijten & F. Bouricius (4 ♀♀ RMNH), Trinconmale [sic, = Trincomalee], coll. Felder (1 ♀ RMNH), Tri[ncomalee] Distr.: China Bay, 0–100 ft., 27–31.i.1977, leg. K.V. Krombein, P. Fernando, D.W. Balasooriya & V. Gunawardane (1 ♀ USNM), Anuradhapura, leg. W. Horn (5 ♀♀ SDEI, 1 ♀ USNM), same, 1899 (3 ♂♂ 1 ♀ SDEI, 2 ♀♀ USNM), same locality, 19–21.xii.1910, leg. A. Luther (1 ♂ 1 ♀ MZHF), Kala Wewa, 12.ii.[18]96, leg. Madarász (2 ♂♂ HNHM), same but 14.ii.[18]96 (1 ♀ HNHM), same but 17.ii.[18]96 (1 ♀ HNHM), env. of Dambulla, 300 m, 19.iv–9.v.1991, leg. J. Kolibáč (1 ♀ ZSMC), Polonnaruwa, Girit[h]ale, 12.iii.1981, leg. M. Cofais (22 specimens MNHN), Wilpattu [National Park], Hunuwilagama, Wildlife Soc[iety] Bungalow, 200 ft., 10– 19.iii.1970, leg. Davis & Rowe (1 ♂ USNM), Kandy, leg. M. Löbell (1 ♂ NHMW), same but coll. E.T. Atkinson, B.M.92-6 (1 ♀ BMNH); same but xi.[18]97, coll. G.W. Kirkaldy, B.M.1912-513 (2 ♂♂ BMNH), Marawila, 50 km N of Colombo, 1970, leg. P. Kandulawa (1 ♂ 3 ♀♀ ZSMC), Colombo, coll. Nickerl (1 ♀ NMPC), Colombo Distr., Ratmalana Airport, 27.x.1973, leg. M. & B. Robinson (2 ♂♂ 2 ♀♀ USNM), Wellawaya, 3.i.[19]28, Imp. Bur. Ent., B.M.1930-197 (1 ♂ 1 ♀ BMNH), Inginiyagala, 2–4.vi.1975, leg. D.H. Messerschmith, G.L. Williams & P.B. Karunaratne (1 ♀ USNM), same but 1–5.vi.1975 (2 ♂♂ 1 ♀ USNM), Ham[bantota] Distr., Palatupana tank, 10–16 m, 6–7.x.1980, leg. K.V. Krombein, P.B. Karunaratne, T. Wijesinhe & V. Gunawardane (1 ♂ 2 ♀♀ USNM), Palatupana, WLNPS [= Wild Life and Nature Protection Society] Bungalow, 0–50 ft., 18–21.i.1979, Malaise trap, leg. K.V. Krombein, P.B. Karunaratne, S. Siriwardane & T. Gunawardane (1 ♀ USNM), Ceylon, coll. Green (3 ♂♂ 3 ♀♀ BMNH), same but coll. Staudinger (1 ♀ ZMHB), same but coll. E.A. Böttiker (1 ♀ ZMHB), same but leg. Baker (1 ♂ USNM), same but coll. Xántus (17 ♂♂ 18 ♀♀ HNHM), same but 1929, leg. W. Robinson (4 ♂♂ 7 ♀♀ USNM), same but leg. Xántus, coll. Nickerl (1 ♂ 1 ♀ NMPC), same but Novara Exp. (1 ♂ 2 ♀♀ NHMW), N. Ceylon, vi.1889, leg. H. Fruhstorfer (1 ♂ 1 ♀ ZMHB), same, coll. G. Breddin (1 ♂ SDEI), S. Ceylon, v.1889, leg. H. Fruhstorfer (2 ♂♂ 1 ♀ HNHM, 3 ♂♂ 1 ♀ ZMHB).— NEPAL. Gandaki Distr., Gorkha, 600–1500 m, 1.vi.1999, leg. Z. Andrš (1 ♂ MMBC), Gorkha Distr., Sudi-Samri Bazar, 1300–1650 m, 19.vi.1993, leg. J. & J. Probst (1 ♂ NHMW), 10 km NNW of Chehere, 27º39.29’N 85º42.69’E, 1478 m, 8.v.2000, dry forest, leg. A. Konstantinov, S. Lingafelter & M. Volkovitsh (1 ♂ USNM), Sun Khosi Tal [= Valley], 2150 m, 2.v.[19]62, leg. G. Ebert (1 ♂ 1 ♀ NMPC), Arun Valley, Mure, 2050 m, 2.vi.1992, leg. J. & J. Probst (1 ♂ NHMW).— BURMA ( MYANMAR). Mon State: Theinzeik, 1913, leg. P. Lolzeau (1 ♂ RMNH); Tanintharyi Region: Tenasserim [= Tanintharyi Region], B.M.95-188 (1 ♂ BMNH).— THAILAND. Chiang Rai Prov.: Wiang Pa Pao, 30.viii.1988 (1 ♀ NSMT); Chiang Mai Prov.: Nong Hoi, viii.1974, leg. P.J.J.H. Kuijten & F. Bouricius (3 ♂♂ 2 ♀♀ RMNH), San Pakia, 19.19ºN 98.50ºE, 1400 m, 1–15.v.1998, leg. V. Kubáň (1 ♀ MMBC); Mae Hong Son Prov.: env. of Sappong, 19°27’N 98°20’E, 1500 m, 7–12.v.1996, leg. J. Horák (1 ♂ ZJPC).— CHINA. Henan: Xinyang (1 ♀ NKUM); Shanghai (1 ♀ BMNH); Sichuan: Xichang, 1600–1700 m, 8.vii.1958, leg. S.M. Song (1 ♂ 1 ♀ IZAS), Dechang, vi.[19]58, from mandarin orange (1 ♀ NKUM), Yanyuan, Jinhe, 1250 m, 28.vi.1984, leg. C.F. Li (1 ♂ IZAS), Panzhihua, Jinjiang Distr., Madianhe, Mafeng village, 24.vii.2008, leg. J.H. Zhou (3 ♂♂ 6 ♀♀ NKUM), Yalung Tal [= valley of Yalong River] zur Huili und Yenyüen [= Yanyuan], 1200 m, 26.ix.1914, leg. H[andel]-Mazzetti (3 ♂♂ 4 ♀♀ NHMW); Yunnan: Lijiang, 26º50’N 100º24’E, 2524 m, 4.vi.2006, leg. Kremitovský (1 ♂ MMBC), Yongsheng, vii.1955 (4 ♂♂ 1 ♀ IZAS), Ta-Li-Fou [= Dali] (1 ♀ RMNH), Ta-Pin-Tze [= N of Dali], leg. R.P. Delavay (1 ♂ 2 ♀♀ RMNH), same, coll R. Oberthür 1899 (2 ♀♀ MNHN), Hang-Kia-Pin [= env. of Dali?], 1400 m (1 ♀ RMNH), Est de Tchin- Kiang [= E of Chengjiang], Lou-Fou-Tsouen (Ing-Ko-Tsoué), confl. Lou-Nan-Ho et Ta-Ken-Ho, 1905, Gervais (1 ♀ MNHN), Djo-Kou-La [= env. of Binchuan?], 1200 m, Coll. E. le Moult (3 ♀♀ RNMH), Yuanmo, viii.[19]80 (1 ♂ NKUM), Chuxiong, viii.[19]80 (1 ♂ NKUM), Magai [= Majie], NW of Yunnanfu [= Kunming], 10.ix.[19]14, leg. H[andel]-Mazzetti (1 ♂ 2 ♀♀ NHMW), Longling, 1060 m, 24.vi.1956, leg. B.S. Zhou (1 ♂ IZAS), Jingdong, 1200 m, 5.iii.1957, leg. A. Monchadsky (1 ♂ IZAS), same but 1400 m, 21.v.1956, leg. B.S. Zhou (1 ♀ IZAS), same but 1250 m, 25.v.1956, leg. Zagulyaev (1 ♀ IZAS), same but 27.v.1956, leg. Zagulyaev (1 ♀ IZAS), same but 28.v.1956, leg. V. Popov (1 ♂ 1 ♀ IZAS), same but leg. T.R. Huang (1 ♂ IZAS), same but leg. X.C. Yang (1 ♀ IZAS), same but 2.vi.1956, leg. Kryzhanovsky (3 ♂♂ 4 ♀♀ IZAS), same but 21.vi.1956, leg. Zagulyaev (1 ♂ IZAS), same but 24.vi.1956, leg. D.Y. Lin (1 ♂ IZAS), same but 1170 m, 23.vi.1956, leg. W. Zhang (1 ♀ IZAS), same but 29.vi.1956, leg. Kryzhanovsky (3 ♂♂ IZAS), same but 4.vii.1956, leg. Zagulyaev (1 ♀ IZAS), N of Jingdong, 28.v.1956, leg. Y. Zhao (1 ♂ 3 ♀♀ IZAS), same but 30.v.1956 (1 ♀ IZAS), env. of Jingdong, 1450 m, 23.iii.1957, leg. D. Panfilov (1 ♂ IZAS), road between Midu and Jingdong, 2.iii.1957, leg. D. Panfilov (6 ♂♂ 12 ♀♀ IZAS), Yuanjiang, Mt. Mandan, 770 m, 25.vii.2006, leg. Z.H. Fan (1 ♂ NKUM), Yuanjiang, 500 m, 16.v.1957, leg. S.Y. Wang (1 ♀ IZAS), 50 km SW of Mojiang, 900–1000 m, 30.iii.1955, leg. Bushchik (1 ♂ IZAS), Yuanyang, Nansha, 8.viii.1986, leg. H. Tao (1 ♀ SYSU), Pu’er, 1400 m, 19.iv.1955, leg. Y.F. Xue (1 ♀ IZAS), same but 21. iv.1955, leg. Y. Zhao (1 ♀ IZAS), same but 1080 m, 8.v.1956, leg. B.S. Zhou (1 ♂ IZAS), same but 1020 m, 10.v.1956 (1 ♀ IZAS), Lancang, 1200 m, 30.vii.1957, leg. L.C. Zang (1 ♀ IZAS), Lancang, Yuantian, 650–950 m, 26.xi.2001 (1 ♀ NKUM), Menghai, Mt. Nannuo, 1400 m, 4.iii.1957, leg. D.H. Liu (1 ♂ 1 ♀ IZAS), env. of Jinping, 900 m, 27.v.1956, leg. Panfilov (1 ♀ IZAS), “Ouest Yunnan, Kouang Si Hien” [unlocated], 2100 m (1 ♂ RMNH), “Ouest Yunnan, Ma-Chang” [unlocated], 1000 m (1 ♂ RMNH), “Sud-Yunnan, Tche-Ping-Tcheou” [unlocated] (1 ♀ RMNH), Yunnan, coll. E. le Moult (100+ ♂♂ 100+ ♀♀ RMNH), same but coll. A. David 1899 (1 ♀ MNHN); Guangxi: Tianlin, from niúgānguŏ [= Phyllanthus emblica ], leg. L.Z. Ming (1 ♂ NKUM), Ningming, 21.xii.1958, leg. Y.X. Chen (1 ♂ SYSU); Hainan: coll. W.L. Distant, B.M.1911-383 (1 ♂ BMNH), Lin-kao [= Lingao] Distr., 15 miles SE of Naam-fung [= Nanfeng], Ch’ung-mei [unlocated], 20–22.viii.1932, leg. F.K. To (1 ♂ SYSU), Jianfengling, Institute of Tropical Forestry, 19.xi.1983, leg. [illegible] Hu (1 ♂ SYSU); Guangdong: Henan [now Haizhu Distr.], 3.ix.1954, leg. X.R. Lin (1 ex. lacking abdomen SYSU); Fujian: Fuzhou, Xihu, 3.iii.1960, leg. S.Q. Jiang (1 ♀ IZAS), Jianyang, 21.vi.1965, leg. L.C. Wang (1 ♂ NKUM).— VIETNAM. Saigon [= Ho Chi Minh City], vi.1969, leg. R.J. Jae (1 ♂ USNM).— LAOS. Saravan Prov.: Wapikhamthong Prov. [now in Saravan Prov.], Khong Sedone [= Khongxedon], 3.viii.1965, native collector (1 ♂ BPBM), same locality, Wapi, 17.viii.1965, native collector (1 ♂ BPBM), same but 30.iv.1967 (1 ♀ BPBM), same but 15.vi.1967 (1 ♂ BPBM); Champasak Prov.: Sedone Prov. [now in Champasak Prov.], Paksong, 15.vii.1967, native collector (1 ♂ BPBM).— PHILIPPINES? Palawan [in error?]: coll. Staudinger (1 ♂ ZMHB).— MALAYSIA? Malacca [in error?], coll. Noualhier 1898 (1 ♂ MNHN > IZAS).— INDONESIA? Sumatra [in error?] (2 ♂♂ NHMW); Java [in error?] (2 ♂♂ 6 ♀♀ RMNH), same, coll. Fitch (1 ♂ USNM).

Figures

FIGURES 18–23. Scutellera perplexa (Westwood, 1837). Figs. 18, 19, a female from Thailand: Nong Hoi (RMNH); Figs. 20, 21, a male from India: Himachal Pradesh, Sadhupul, near Simla (SEHU); Figs. 22, 23, a female from Sri Lanka: Uppuveli (RMNH). Dorsal (Figs. 18, 20, 22) and vental (Figs. 19, 21, 23) views. Scales in mm.

FIGURES 30–41. Scutellera spp., external male genitalia. Figs. 30–33, S. nepalensis (Westwood, 1837); Figs. 34–37, S. perplexa (Westwood, 1837); Figs. 38–41, S. spilogastra (Walker, 1867). Genital capsule, dorsal view (Figs. 30, 34, 38); right paramere, three different aspects (Figs. 31–33, 35–37, 39–41). Abbreviations: dsp = dorsal setal patch; mp = median projection of cuplike sclerite; vsp = ventral setal patch. Scales in mm. Figs. 30–33 from Tsai et al. (2011).

FIGURES 42–47. Scutellera spp., phallus (in repose). Figs. 42, 43, S. nepalensis (Westwood, 1837); Figs. 44, 45, S. perplexa (Westwood, 1837); Figs. 46, 47, S. spilogastra (Walker, 1867). Left lateral view (Figs. 42, 44, 46); posteroventral view (Figs. 43, 45, 47).Abbreviations: aed = aedeagus s. str.; cp-II 1, cp-II 2 = lobes of second conjunctival process; cp-III = third conjunctival process. Arrows in Figs. 42, 44 and 46 show aspects of Figs. 43, 45 and 47, respectively. Scales in mm. Figs. 42–43 after Tsai et al. (2011).

FIGURES 48–52. Scutellera spp., phallus. Figs. 4, S. nepalensis (Westwood, 1837), distal portion, ventral view; Fig. 49, S. perplexa (Westwood, 1837), inflated, lateral view; Fig. 50, same, posteroventral view; Fig. 51, same, distal portion, ventral view; Fig. 52, S. spilogastra (Walker, 1867), distal portion, ventral view. Abbreviations as in Figs. 42 and 43. Arrow in Figs. 49 shows aspect of Fig. 50. Scales in mm. Fig. 48 after Tsai et al. (2011).

FIGURES 53–64. Scutellera spp., female terminalia and ovipositor. Figs. 53–56, S. nepalensis (Westwood, 1837); Figs. 57–60, S. perplexa (Westwood, 1837); Figs. 61–64, S. spilogastra (Walker, 1867). Terminalia, posterior view (Figs. 53, 57, 61); same, posterodorsal view (Figs. 54, 58, 62); same, ventral view (Figs. 55, 59, 63); ovipositor and sternum X, posterior view. Abbreviations: ga = gonangulum; lt8, lt9 = laterotergites VIII, IX; = laterotergite IX; s10 = sternum X; t8 = tergite VIII; va8, va9 = valvulae VIII, IX; vf8, vf9 = valvifers VIII, IX. Scales in mm.

FIGURES 65–70. Scutellera spp., ectodermal genital tract of female. Figs. 65, 66, S. nepalensis (Westwood, 1837); Figs. 67,

FIGURES 1–10. Lectotypes of Tectocoris nepalensis Westwood, 1837, Callidea spilogastra Walker, 1867, and Scutellera brevirostris Breddin, 1909, with their labels. Figs. 1, T. nepalensis, dorsal view; Fig. 2, same, lateral view; Fig. 3, same, ventral view; Fig. 4, same, labels; Fig. 5, C. spilogastra, dorsal view; Fig. 6, same, ventral view; Fig. 7, same, labels; Fig. 8, S. brevirostris, dorsal view; Fig. 9, same, ventral view; Fig. 10, same, labels. Scales in mm. © OXUM (Figs. 1–4), © BMNH (Figs. 5–7), © SDEI (Figs. 8–10).

FIGURES 11–17. Lectotypes of Calliphara amethystina Germar, 1839 and Scutellera lanius Stål, 1854 and their labels. Fig. 11, C. amethystina, dorsal view; Fig. 12, same, lateral view; Fig. 13, same, ventral view; Fig. 14, same, labels; Fig. 15, S. lanius, dorsal view; Fig. 16, same, ventral view; Fig. 17, labels. Scales in mm. © ZMHB (Figs. 11–14), © NHRS, courtesy of G. Lindberg (Figs. 15–17).

FIGURES 24–29. Scutellera spp., terminalia of males. Figs. 24, 25, S. nepalensis (Westwood, 1837); Figs. 26, 27, S. perplexa (Westwood, 1837); Figs. 28, 29, S. spilogastra (Walker, 1867). Lateral (Figs. 24, 26, 28) and ventral (Figs. 25, 27, 29) views. Abbreviations: fw = fore wing; gc = genital capsule; sc = scutellum. Scales in mm.

FIGURES 71–72. Geographical distribution of Scutellera spp. Fig. 71: S. perplexa (Westwood, 1837). Fig. 72: Circles = S. nepalensis nepalensis (Westwood, 1837); squares = S. nepalensis amethystina (Germar, 1839); triangles = S. spilogastra (Walker, 1867). Solid markers represent specimens examined during the present study, empty markers unverified literature data.