Helopeltis, SIGNORET, 1858

Namyatova, Anna A. & Cassis, Gerasimos, 2016, Systematic revision and phylogeny of the plant bug tribe Monaloniini (Insecta: Heteroptera: Miridae: Bryocorinae) of the world, Zoological Journal of the Linnean Society 176 (1), pp. 36-136 : 96-97

HELOPELTIS SIGNORET

Figures 9 , 12F , 13A, H , 15M–Q , 19C, D , 22

Helopeltis Signoret, 1858: 502 (gen. nov.; type species Helopeltis antonii Signoret, 1858 by monotypy); Walker, 1873: 165 (cat., syn.); Atkinson, 1890a: 51 (cat.); Atkinson, 1890b: 175 (descr.); Watt & Mann, 1898: 247 (bibliography, history, biology, etc.); Distant, 1904b: 439 (descr.); Kirkaldy, 1906: 134 (list.); Mann, 1907: 277 (descr., disc.); Reuter, 1910: 153 (cat.); Poppius, 1911: 39 (key to spp.); Poppius, 1912: 175 (key to gen., descr., key to spp.); Bergroth, 1922: 54 (list); Ghesquière, 1922: 281 (disc., key to spp.); China, 1944: 144 (key to gen.); Ghesquière & Carayon: 1948: 59 (note); Carayon & Delattre, 1948: 185 (note, key to spp.); Villiers, 1952: 191 (descr.); Carvalho, 1952: 59 (cat,); Carvalho, 1955: 38 (key to gen.); Carvalho, 1957: 133 (cat.); Schmitz, 1968: 1 (disc., descr., diag.; key to gen., key to spp.); Steyskal: 1973: 206 (correction); Lavabre, 1977a: 51 (descr., disc.); Lavabre, 1977b: 107 (note); Carvalho, 1981: 39 (key to gen.); Stonedahl, 1991: 465 (descr., diag., key to spp.); Cassis & Gross, 1995: 142 (cat.); Schuh, 1995: 511 (cat.); Schuh, 2002 –2013 (cat.); Namyatova et al., in press (phylogeny).

Aspicellus Costa, 1864: 146 (gen, nov.; type species Aspicellus podagricus A. Costa, 1864 by monotypy, synonymized by Walker, 1873: 165); Schuh, 1995: 512 (cat.); Schuh, 2002 –2013 (cat.).

Diagnosis: Helopeltis belongs to the Monalonion - complex (see discussion for tribe), and can be separated from other genera in this complex by: long, undivided spinelike projection on scutellum ( Fig. 12F ); antenna distinctly longer than body; femora curved (as in fig. 18A in Namyatova et al., in press); collar flat; base of clypeus delimited with depression; metepimeron with rounded or rectangular outgrowth (as in Fig. 13A ); claw with apical tooth (as in fig. 10D in Namyatova & Cassis, 2013b); place of attachment of ductus seminis on phallobase only slightly to left of midline ( Fig. 15M ); DLP with two sclerotized bands ( Fig. 19C ).

Redescription (partly based on Stonedahl, 1991): Male: Body length 5–8 mm. COLORATION ( Fig. 9 ). Variable, from pale brown to dark brown or almost black, sometimes reddish, often with paler or darker markings. TEXTURE. Body mostly smooth, without punctures and tubercles; vertex without flattened areas; semicircular depression between scutellum and mesoscutum absent; striations on scutellum laterally absent; only small depression on anterior angle of pronotum present (as in fig. 9H in Namyatova & Cassis, 2013b). VESTITURE. Setae on dorsum and thoracic pleura absent; very short, pale or dark simple suberect setae present on clypeus, labium, antenna, apices of femora, tibiae, tarsi, genital segment and often on scutellar process; ASI sometimes almost without setae; setae on clypeus sometimes flattened; tibiae regularly setose; small black spinules on femora absent, spinules on tibia apically absent or present. STRUCTURE. Head. Distance between eye and pronotum slightly shorter or subequal to eye diameter (as in Fig. 10J ); occipital region delimited with shallow depression; longitudinal depression distinct, as long as or longer than eye diameter, sometimes extending on frons; eyes not stylate, in line with contour of head, c. 0.2× as long as head width (as in Fig. 10J ); distance between antennal fossae subequal or slightly longer than antennal fossa diameter; frons straight, without ridges, outgrowths or longitudinal depression; anterior view of head c. 1.6– 1.9× wide as high; eye height c. 1–2× as long as distance between eye and apex of clypeus; antennal fossa oval, only slightly shorter than eye height, not raised, inferior margin placed above inferior margin of eye; base of clypeus placed slightly below than or near inferior margin of antennal fossa delimited with depression; head swollen or almost flat in lateral view; gula c. 1.5–2.5× as long as buccula, convex. Labium. Length varying from reaching posterior margin of mesosternum to reaching abdominal LSII; LSI c. 3–4× as long as wide; LSII c. 4–6× as long as wide, subequal to or slightly longer than LSI; LSIII c. 3–6× subequal to or slightly longer than LSII; LSIV c. 5–8× as long as wide, c. 1.2–1.5× as long as segment III. Antenna. Twice as long as body, rarely only 1.5× as long as body ( H. cinchonae ); ASI c. 2–2.5× as long as head width, rarely subequal to head width ( H. cinchonae ), swollen apically (as in fig. 8I in Namyatova et al., in press); ASII c. 1.6–3.5× as long as ASI, c. 2.2–3.2× as long as head and pronotum combined; AS III c. 0.7– 1× as long as ASII; ASIV c. 0.3–0.5× as long as ASIII, ASII–IV filiform. Thorax. Collar flat, delimited posteriorly; calli separated (as in Fig. 10J ), flat, almost indistinct; depression delimiting calli posteriorly absent; humeral angles of pronotum rounded, not dilated (as in Fig. 10J ); posterior margin of pronotum straight or concave; scutellum swollen, with elongate spinelike process with expanded apex ( Fig. 12F ); scutellum distinctly rounded apically, without ridge or medial depression; metepimeron narrow, c. 3–4× as high as long, rounded or with small subrectangular outgrowth ( Fig. 13A ); metasternum rounded posteriorly, without medial projection on to abdominal segment II (fig. 17A in Namyatova et al., in press). Hemelytron. Costal margin straight; hemelytra tapering posteriorly; margins of claval commissure c. 3× as long as scutellum, curved; R + M distinct, reaching posterior margin of corium; medial fracture subparallel to R + M; corium without swelling posteriorly; cuneus c. 4–7× as long as wide, c. 1.2–1.4× as long as pronotum, medial margin distinctly concave (as in fig. 13C in Namyatova et al., in press); membrane cell c. 1.6–3× as long as pronotum, rounded or acute apically; auxiliary vein absent or short present; distance between cell and apex of membrane c. 0.2–0.3× as long as cell. Legs. Forecoxae separated (as in fig. 17B in Namyatova et al., in press); femora swollen apically, with additional swelling medially, distinctly curved ( Fig. 13H ); foretibia longer than head and pronotum combined; swellings on tibiae absent; segment I of hind tarsus distinctly longer than segments II and III each, segment II slightly shorter than segment III; claw broadly rounded, tooth subdivided into basal and subapical parts ( Fig. 13L ). Genitalia ( Fig. 15M– Q ). Genital capsule slightly longer than wide, without outgrowth(s); ventral margin of genital capsule shortened anteriorly; left paramere r-shaped or only slightly curved, c. 3× as long as right paramere; phallobase sclerite of primary gonopore subtriangular, often with outgrowth in front of ductus seminis attachment place; ductus seminis longer than phallotheca length, with coils, forming wide tube, without sclerotization basally or apically, attached to phallobase on left-hand side or almost medially; sclerotized part of phallotheca broad occupying entire dorsal side, rounded apically, without outgrowth(s) or ridges; endosoma usually with lobal sclerite and fields of small spicules, sometimes endosoma without sclerotization (see also illustrations of Stonedahl, 1991).

Female: Body length 6–9 mm. Similar to male, but coloration sometimes paler and body larger ( Fig. 9 ). Genitalia ( Fig. 19C, D ). DLP mostly membranous with two sclerotized bands, sometimes fused; DLP often with medial ridge and sclerotization around it, sometimes without ridge ( H. cinchonae ); striations absent or present only at base of lateral oviducts; lateral oviducts attached at halfway of DLP, removed from each other, placed close to lateral margins of DLP; spermathecal gland placed in posterior half, slightly below midline or close to posterior margin, not shifted right or left; posterior wall of bursa copulatrix covered with small tubercles and with pair of spinose lobes; base of second valvula with bifurcate outgrowth or straight ( H. cinchonae ); posterior wall of bursa copulatrix membranous, not bearing sclerites around vulva (see also illustrations from Stonedahl, 1991).

Distribution: Widely distributed in South-East Asia, Pacific Islands, Australia ( Fig. 22 ).

Host plants: The host plants are known for many species of Helopeltis ; some Helopeltis species are serious pests of cocoa, tea and cashew (see Stonedahl, 1991 for details).

Figures

Figure 9. Habitus photographs. Monalonion dissimulatum ♂ AMNH_PBI 19564, ♀ AMNH_PBI 5258; Physophoropterella bondroiti ♂ AMNH_PBI 400348, ♀ AMNH_PBI 5037; Physophoroptera mirabilis ♂ AMNH_PBI 5070, ♀ AMNH_PBI 5083; Arthriticus eugeniae ♂ AMNH_BPI 19573, ♀ AMNH_PBI 19575; Ragwelellus suspectus ♂ AMNH_PBI 34153, ♀ AMNH_PBI 34154; Ragwelellus vittatus ♂ AMNH_PBI 34152, ♀ AMNH_PBI 45821; Schuhirandella fulva ♂ PT AMNH_PBI 400345, ♀ PT AMNH_PBI 19576; Helopeltis clavifer ♂ AMNH_PBI 40416, ♀ AMNH_PBI 202016; Afropeltis corbisieri ♂ AMNH_PBI 5133, ♀ AMNH_PBI 5216.

Figure 10. Scanning electron micrographs. Head and pronotum, dorsal view. A, Mansoniella nitida ♀ AMNH_PBI 46067; B, Poppiusia leroyi ♀ AMNH_PBI 5838; C, Chamus bellus ♀ AMNH_PBI 5223; D, Lycidocoris mimeticus ♀ AMNH_PBI 5043; E, Physophoropterella bondroiti ♂ AMNH_PBI 19110; G, Odoniellia reuteri ♂ AMNH_PBI 19090; H, Volkelius carvalhoi ♀ PT AMNH_PBI 19630; I, Yangambia vesiculata ♀ AMNH_PBI 19084; J, Ragwelellus suspectus ♂ AMNH_PBI 19629. Head, dorsal view. F, Sahlbergella tai ♀ AMNH_PBI 5106. Head, anterior view. K, Parachamus bellus ♀ AMNH_ PBI 5837. Head, lateral view. L, Poppiusia leroyi ♀ AMNH_PBI 5838. Frons, dorsal view. M, Poppiusia leroyi ♀ AMNH_PBI 5838.

Figure 12. Scanning electron micrographs. Scutellum, lateral view. A, Bryocoropsis soror ♂ AMNH_PBI 5115; B, Sahlbergella singularis ♂ AMNH_PBI 19053; C, Physophoroptera mirabilis ♀ AMNH_PBI 20202; D, Physophoropterella bondroiti ♂ AMNH_PBI 19112; E, Afropeltis lalandei ♂ AMNH_PBI 5272; F, Helopeltis pellucida sex unknown AMNH_PBI 19628.

Figure 13. Scanning electron micrographs. Metepimeron. A, Helopletis pellucida sex unknown AMNH_PBI 19628; B, Afropeltis hyalospilosus ♂ AMNH_PBI 5137; C, Poppiusia leroyi ♀ AMNH_PBI 5838; D, Pachypeltis reuteri ♂ AMNH_PBI 45980; E, Odoniella reuteri ♂ AMNH_PBI 19194. Setae on hemelytron. F, Boxia khayae ♂ AMNH_PBI 5065. Fore- and middle femora. G, Physophoropterella bondroiti ♂ AMNH_PBI 19110. Hind femur. H, Helopeltis pellucida sex unknown AMNH_PBI 19628. Hind tarsus. I, Physophoropterella bondroiti ♂ AMNH_PBI 19110. Claw, lateral view. J, Chamus sp. sex unknown, not databased; K, Pachypeltis brevirostris sp. nov. sex unknown, not databased; L, Helopeltis bradyi sex unknown, not databased.

Figure 15. Male genitalia. Dorsal view. Bryocoropsis laticollis var. infuscata. AMNH_PBI 18942. A, aedeagus; B, right paramere; C, left paramere. AMNH_PBI 5108. D, genital capsule. Bryocoropsis soror AMNH_PBI 19072. E, aedeagus; F, right paramere; G, left paramere; H, genital capsule. Eupachypeltis flavicornis LT AMNH_PBI 19128. I, aedeagus; J, right paramere; K, left paramere; L, genital capsule. Helopeltis clavifer AMNH_PBI 34191. M, aedeagus; N, right parameres; O, left paramere; P, genital capsule. Dimia inexpectata PT AMNH_PBI 271336. R, aedeagus; S, right paramere; T, left paramere; U, genital capsule. Lycidocoris mimericus AMNH_PBI 5066. V, aedeagus; W, right paramere; X, left paramere; Y, genital capsule. Lycidocoris thoracicus no USI. Z, aedeagus; AA, right paramere; AB, left paramere; AC, genital capsule. Lateral view. Helopeltis clavifer AMNH_PBI 34191. Q, genital capsule. The smaller scale is for genital capsule, the larger scale is for aedeagi and parameres.

Figure 19. Female genitalia, bursa copulatrix. Eupachypletis pilosus AMNH_PBI 46068. A, dorsal labiate plate; B, posterior wall. Helopletis clavifer AMNH_PBI 74192. C, dorsal labiate; D, posterior wall. Lycidocoris modestus AMNH_PBI 5099. E, dorsal labiate plate. Lycidocoris mimeticus AMNH_PBI 5086. F, dorsal labiate plate; G, posterior wall. Mansoniella nitida AMNH_PBI 45970. H, dorsal labiate plate; I, posterior wall. Lycidocoris thoracicus AMNH_PBI 5096. G, dorsal labiate plate; K, posterior wall. Mansoniella sassafri AMNH_PBI 19067. L, dorsal labiate plate; M, posterior wall. Monalonion sp. AMNH_PBI 19632. N, dorsal labiate plate. Monaloion dissimulatum no USI. O, posterior wall. Odoniella rubra AMNH_PBI 18952. P, dorsal labiate plate; Q, posterior wall. Rhopaliceschatus quadrimaculatus AMNH_PBI 20523. R, dorsal labiate plate.

Figure 22. Distribution maps of Afropeltis, Arculanus, Arthriticus, Boxia, Chamus, Eupachypeltis, Helopeltis, Mansoniella, Physophoropterella, Pseudodoniella and Sahlbergella.