The genera Agathocles and Surenus (Hemiptera: Heteroptera: Pentatomidae) tribal reassessment, redescription, new synonyms, and description of two new species Author Salini, S. National Bureau of Agricultural Insect Resources, Bangalore, 560024, India Author Kment, Petr 0000-0002-7026-5691 Department of Entomology, National Museum, Cirkusová 1740, CZ- 193 00 Praha 9 - Horní Počernice, Czech Republic sigara @ post. cz; https: // orcid. org / 0000 - 0002 - 7026 - 5691 sigara@post.cz text Zootaxa 2021 2021-04-14 4958 1 510 559 journal article 7213 10.11646/zootaxa.4958.1.31 a1df68e3-0e4c-4010-9556-cb15d51a5499 1175-5326 4692369 9E63F8E2-1CCA-4F58-9019-39AE7A3D0640 Agathocles Stål, 1876 Agathocles Stål, 1876: 69 (key to genera), 106 (description of type species). Type species: Agathocles limbatus Stål, 1876 , by monotypy. Agathocles : Atkinson (1888) : 152 (redescription); Lethierry & Severin (1893) : 183 (catalogue); Distant (1900) : 173 (comparison with Fernelius ); Distant (1902) : 196 (key to genera), 198 (redescription); Kirkaldy (1909) : 147 (catalogue); Ahmad et al. (1974): 88, 89 (figure legend); Ahmad (1981) : 16 (key to genera), 43 (list); Rider (1997) : 402 (checklist); Rider (2006) : 342 (catalogue); Fan (2011) : 15 (key to genera), 43 (diagnosis, key to Chinese species); Rider et al. (2018) : 80 , 96–97 (comments on tribal placement). Agathooles (incorrect subsequent spelling): Jiang (1985) : 67 (checklist). Agothocles (incorrect subsequent spelling): Ahmad et al. (1974): 43 (list, host plant). Surenus Distant, 1901a: 101 (list), 106–107 (original description, systematic position). Type species: Surenus normalis Distant, 1901a , by monotypy. New junior subjective synonym. Surenus : Distant (1902) : 110 (key to genera), 116–117 (description repeated); Bergroth (1908) : 154 (catalogue); Kirkaldy (1909) : 195 (catalogue); Memon et al. (2011) : 1151 –1155, 1157, 1162–1163 (phylogenetic analysis, key to the genera); Salini & Viraktamath (2015) : 10 , 22, 51, 59 (key to tribes and genera, checklist for south Indian Pentatomidae , illustration); Arnold & Arnold (2017) : 13 (faunistic record); Rider et al. (2018) : 80 , 86 (comments on tribal placement). Redescription. Colouration . Body above dark brown to black, underside black, sometimes either dorsally or ventrally with yellow pattern; membrane brownish; compound eyes silver or dark brown; ocelli yellowish or reddish; rostrum yellowish; antennae and legs always paler, yellowish to brown. Abdominal spiracles concolorous with surrounding surface of ventrites. Integument and vestiture. Body above matt or slightly shining with only clavus and corium matt; ventral side shining except of matt evaporatorium. Body above and below covered with dense, coarse punctures; posterior portion of pronotal disc with very coarse punctures; punctures on scutellum partly confluent to short transverse wrinkles; abdominal sternites with finer punctures. Body glabrous, only antennomeres I to IV with short, erect, brown setae, antennomeres III and IV in addition with short, semierect, golden setae. Labium with short, sparse golden setae, femora with short sparse brown setae, tibiae with dense, short, erect setae, tarsi dorsally with comparatively long, semierect sparse setae and short, dense mat of golden setae ventrally. Genital capsule along ventral and lateral rim and ventral rim infolding with dense, semierect, golden brown setae ( Fig. 37 ). External female genitalia (valvifers VIII on dorsal side and on mesial margins, valvifers IX and laterotergite IX) with sparse golden setae. Structure. Body robust, 13–24 mm long, widely oval, widest in posterior half (across abdominal segment III), biconvex, more strongly vaulted ventrally. Head ( Figs 6–13 ) slightly shorter than wide across compound eyes (head length: width ratio 0.73–0.99); dorsum of head more or less flat, lateral margins concave in front of eyes, more or less reflexed upwards including apex of mandibular plates. Mandibular plates explanate, wider and longer than clypeus, each of them separately rounded apically, meeting ( Fig. 6 ) or nearly meeting ( Figs 7, 9, 10 ) in front of clypeus, leaving narrow V-shaped cleft anteriad of clypeus apex, or not meeting ( Fig. 12 ); clypeus correspondingly narrowing and anteriorly closed ( Fig. 6 ), nearly closed ( Figs 7, 9, 10 ), or apically free ( Fig. 12 ). Compound eyes moderately large, rounded, protruding out of the head outline in most of their width ( Figs 6–7, 9–10, 12 ). Ocelli small, situated posteriad of compound eyes, close to anterior margin of pronotum. Round, smooth, unpunctured spot present between compound eye and ocellus ( Figs 6–7, 9–10, 12 ). Postgenae behind eyes narrow. Base of antennae including antenniferous tubercle partly ( Figs 6–7 ) to completely visible ( Fig. 12 ) from above. Antennae 5-segmented, slender. Scape (I) cylindrical, shortest and stoutest; basipedicellite (IIa) shortest; antennomeres IIa to IV slender and uniformly cylindrical. Bucculae ( Fig. 8 ) short, low, anteriorly obtusangulate, with small tooth-like projection anteroventrally, posteriorly widely rounded, leaving apex of labiomere I free. Labrum ( Figs 11, 13 ) flat and narrow, reaching base of labiomere II. Lengths of labiomeres from shortest to longest: IV ≤ I <III <II. Labiomere I moderately stout, short, nearly reaching or slightly surpassing posterior margin of bucculae ( Figs 11, 13 ); apex of labium reaching between mesocoxae to metacoxae. Pronotum ( Figs 1 , 64 , 88 , 108, 110 ). Anterior pronotal margin medially incised, concave, forming arcuate to quadrangular concavity receiving the postocular portion of head ( Figs 6, 10 , 64 ); anterolateral angles truncate behind eyes, each laterally with one minute, laterally directed denticle ( Figs 6, 10, 12 ); anterolateral margins sharp but not emarginate, slightly reflexed and minutely to distinctly crenulate ( Figs 64 , 88 ) in anterior half, smooth in posterior half, shallowly concave in middle. Humeri rectangulate to slightly obtusangulate with small tubercle present sublaterally; posterolateral margin of pronotum straight or slightly convex, directed obliquely towards anterolateral angles of scutellum, posterolateral angles of pronotum widely rounded; posterior margin of pronotum slightly concave to nearly straight. Anterior part of pronotal disc (in front of humeri) declivent and flat; posterior part of pronotal disc slightly convex ( Fig. 111 ). Scutellum ( Figs 1, 3 , 88 , 108 ) triangular, longer than wide at base, lateral margins nearly straight or slightly convex at frenal portion, slightly incised at apices of frena (in about two thirds of scutellar length); apex of scutellum parabolic, not surpassing anterodistal angles of corium, narrowly to more broadly rounded. Disc of scutellum anteriorly convex, posteriorly flat, in some species with saddle-like concavity in front of frenal apices. Hemelytra ( Figs 1, 3 , 88 , 108 ). Clavus narrowly triangular, reaching only two thirds of scutellar length. Corium laterally more or less arcuate, anterodistal angles acutangulate, narrowly rounded apically, much surpassing apex of scutellum, reaching about middle of segment V. Membrane translucent, infumate, widely rounded apically, reaching or slightly surpassing apex of abdomen. Membrane bearing 7‒12 prominent and simple veins ( Fig. 65 ). Thoracic pleuron and sternum. Mesosternum with narrow, median longitudinal carina ( Fig. 26 : mec). Metasternum with lateral margins raised, narrow ( Fig. 23 ) to wide, flat ( Figs 24–25 : mts). Ostiole ( Figs 14–19 : o) situated between outer margins of meso- and metacetabulum, elongate oval, directed posterolaterad, accompanied with short and narrow, slightly elevated, spout-shaped peritreme, reaching about middle of metapleuron width; median furrow distinct, at least in basal half of peritreme; periostiolar depression shallow. Evaporatorium very large ( Figs 14, 15 , 68 ), covering most of metapleuron (except its posterior and lateral areas), prolonged medially between meso- and metacoxae, laterally delimited by low longitudinal ridge; on mesopleuron covering its posterior half (mesepimeron), but leaving smooth ca. triangular spot in middle of lateral margin ( Fig. 20 : sms). Gyrification developed, fine and shallow ( Figs 14, 21 ). Metathoracic spiracle wide, long and narrow, visible in ventral view ( Figs 14–15, 21 : sp), protected by anastomosing, reticulate mycoid filter processes ( Fig. 22 ). Legs ( Figs 2 , 89 , 109 ). Femora cylindrical, unarmed, rounded in cross section, slightly longer and thicker than appropriate tibiae. All tibiae shallowly sulcate on dorsal surface. All tarsi with tarsomere II shortest, I longest, nearly as long as II and III combined; all tarsomeres dorsally regularly rounded, not grooved. FIGURES 1–5. Agathocles limbatus Stål, 1876 . 1–2—habitus of male from Laos: Ban Saluey → Phou Pane Mt. (22.2 mm). 3–5—habitus and labels of holotype of A. yunnanensis Zhang & Lin, 1984 , syn. nov. , female, China: Yunnan: Cang-yuan (scale bar 1 mm). Orig. P. Kment (1–2), D. Rédei (3–5). FIGURES 6–13. Head. 6–8— Agathocles limbatus Stål, 1876: 6 —dorsal view, Vietnam: Phia Oac NP (magnification 20×); 7— dorsal view, Thailand: Phu Kradung NP (20×); 8—lateral view, Laos: Phou Pane Mt. (30×). 9— A. normalis ( Distant, 1901a ) , male, dorsal view, Laos: Phou Pane Mt. (23× mm). 10–11— A. flavipes sp. nov. , female, dorsal (10, 30×) and ventral (11, 40×) view, India: Maharashtra: 70 km N of Pune. 12–13— A. joceliae sp. nov. : 12—holotype, male, dorsal view, Malaysia: Kelantan (25×); 13—female, ventral view, Malaysia: Kelantan: Kampong Sungai Om (27×). Lettering: at—antenniferous tubercle, bu— buccula, cl—clypeus, la—labium, lr—labrum, mp—mandibular plate, oc—ocellus. Scale bars: 1 mm. Orig. P. Kment. FIGURES 14–22. External scent efferent system of metathoracic scent gland. 14— Agathocles normalis (Distant, 1901) , male, Laos: Phou Pane Mt. (23×). 15— A. joceliae sp. nov. , female, Malaysia: Kelantan: Kampong Timor. 16–19—ostiole and peritreme: 16— A. limbatus Stål, 1876 (90×); 17— A. flavipes sp. nov. (100×); 18— A. joceliae sp. nov. (75×); 19— A. normalis (Distant, 1901) (85×). 20— A. joceliae sp. nov. , evaporatorium laterally on mesopleuron (75×). 21— A. normalis , metathoracic spiracle (42×). 22— A. limbatus , detail of metathoracic spiracle with reticulate filter processes (320×). Lettering: ev—evaporatorium, mf—median furrow, o—ostiole, pe—peritreme, sms—smooth spot on mesopleuron, sp—spiracle. Scale bars: 1 mm (14, 15, 21), 0.5 mm (16–20), 0.1 mm (22). Orig. P. Kment. FIGURES 23–28. Morphology of thorax and abdomen. 23–25—metasternum: 23— Agathocles flavipes sp. nov. (50×); 24— A. joceliae sp. nov. (55×); 25— A. limbatus Stål, 1876 (37×). 26–28— A. limbatus : 26—mesosternum (50×); 27—spiracle and trichobothria on abdominal segment VI (170×); 28—apex of abdomen and genital capsule, lateral view (35×). Lettering: cx2— mesocoxa, cx3—metacoxa, gc—genital capsule, la2‒4—labiomere II‒IV; msc—mesosternal carina; mts—metasternum; sp— abdominal spiracle; tr—trichobothrium. Scale bars: 1 mm (23–26, 28), 0.3 mm (27). Orig. P. Kment. Pregenital abdomen slightly wider than pronotum across humeri ( Figs 1 , 88 , 108 ). Connexivum wide, exposed laterally and slightly lamellate ( Fig. 1 ); posterolateral angles of each laterotergite III–VII about rectangulate, only slightly protruding from outline of the abdomen. Abdominal venter uniformly convex, neither grooved, nor keeled medially; ventrite III without spinose projection or depression medially ( Figs 2 , 89 , 109 ). Spiracle on ventrite II covered by metapleuron or its posterior margin more or less exposed. Each ventrite III–VII laterally with spiracle and pair of trichobothria situated posterolaterad of spiracle ( Figs 27–28 ); inner trichobothrium situated in and outer trichobothrium laterad of the imaginary spiracular line. FIGURES 29–32. External female genitalia. 29–30— Agathocles limbatus Stål, 1876 (29—27×); 31— A. flavipes sp. nov. ; 32— A. joceliae sp. nov. Lettering: lt8–9—laterotergites VIII–IX, t8—tergite VIII, vf8–9—valvifers VIII–IX, x—segment X. Scale bars: 1 mm. Orig. P. Kment. Male genitalia. Genital capsule ( Figs 37–41 , 69–74 , 90–94 , 113–116 ) nearly quadrangular in dorsal/ventral view, longer than wide. Dorsal rim ( Figs 72 , 92 : dr) deeply and concavely excavated with median emargination. Ventral rim undulated, slightly ( Figs 41 , 73 ) to deeply ( Figs 93 , 115 ) incised medially; ventral rim infolding unmodified ( Fig. 91 ), medially with short median process perpendicular to ventral rim ( Figs 71, 74 ), or with wide depression ( Figs 37–39 , 113–114 ). Superior processes of dorsal rim usually present ( Figs 37–39, 43 , 90–91, 96 , 113–114, 117 ), in A. flavipes sp. nov. absent. Parameral socket open ( Figs 38 , 91 , 114 : ps) or in A. flavipes closed, kidney-shaped ( Figs 71, 74 : ps). Paramere ( Figs 44–53 , 75–77 , 84–87 , 97–101 , 118–124 ) with main axis of parameral crown Cshaped in lateral view; basal process (bp) foliaceous, roughly semicircular, apically with row of few to many long setae ( Figs 44, 47–48, 51 , 75–76 , 97, 101 , 118, 121 ); stem (st) short and broad; apical process (ap) hook-shaped, apically more or less obtuse ( Figs 76 , 84–87 , 97–99 , 118, 122 ) or triangular, more pointed apically ( Figs 44, 46–47, 49, 52–53 ), always provided with an elongate patch of unpigmented, scaled texture ( Fig. 124 ); mesal angle of stem with or without an additional projection variable in shape ( Figs 44, 46–47, 50–51 , 84–87 ). Phallus ( Figs 54–59 , 78– 80 , 82–83 , 102–107 ): Articulatory apparatus composed of basal plate (bpt) and support bridge complex surrounded by intersegmental membrane (ism); 1+1 capitate processes (cp) connected to support bridge complex (sbc) by means of dorsal connectives (dc) ( Figs 82–83 ). Phallotheca short, barrel shaped, constricted at both ends, broadest in middle. A pair of dorsal conjunctival processes, transversely placed and ending in sclerotised processes ( Figs 57, 59 , 78–79 , 105–107 ); median penial plates fused basally, outer margins broadly and thickly sclerotized, encircling short, curved aedeagus ( Figs 57 , 80 , 105 : ad). Female genitalia ( Figs 29–36 ). Laterotergite VIII subtriangular. Laterotergite IX broadest preapically, gradually narrowing apicad, apically more or less rounded, medially touching each other or not ( Figs 29–32 , 34 ). Valvifers VIII quadrant with median margins juxtaposed and dorsal margins slightly concave to slightly convex ( Figs 29–32 , 34 ). Valvifers IX fused medially with median fusion line well apparent or not. Spermathecal opening associated with a small annular sclerite which has a pair of oblique projections surrounding spermathecal opening laterally, and a larger, arch-shaped sclerite surrounding the previous sclerite ( Figs 33, 36 : so); ring sclerites present, unsclerotized ( Fig. 36 : rs) or slightly sclerotized ( Figs 33, 34 : rs). Spermatheca: proximal duct ( Fig. 33 : pd) thicker and longer than distal duct; dilation long ( Figs 33, 35, 36 : dil), distal invagination (‘sclerotized rod’) distinctly broadened distally ( Figs 33, 35 : div), subparallel in most of its length, basally either straight and broadened ( Figs 33, 35 ) or thin and strongly curved ( Fig. 36 ); distal duct ( Figs 33, 35, 36 : dd) short and thin, much shorter than distal invagination; proximal flange ( Figs 33, 35, 36 : pf) narrower than distal flange; intermediate part of spermatheca narrow, distal flange either separated ( Fig. 35 ) or tightly attached to apical receptacle ( Figs 33, 36 : df); apical receptacle subglobular ( Fig. 35 ) or subhemispherical ( Figs 33, 36 : ar), with one long projection directed proximad, not reaching to surpassing proximal flange, and one to four short processes of various orientation, not surpassing distal flange. FIGURES 33–36. External and internal female genitalia. 33— Agathocles limbatus Stål, 1876 ; 34–35— A. flavipes sp. nov. ; 36— A. joceliae sp. nov. Lettering: ar—apical receptacle, dd—distal duct of spermatheca, df—dorsal flange, dil—spermathecal dilation, div—distal invagination of spermathecal duct, ip—intermediate part, pd—proximal duct of spermatheca, pf—proximal flange, rs—ring sclerite, so—spermathecal opening. Scale bars: 1 mm. Orig. P. Kment (33, 36), Salini (34–35). Differential diagnosis. The genus Agathocles is well recognizable with the following diagnostic characters: large (body length 13–24 mm ), with robust body, widest across abdominal segment III; head only slightly broader than long with mandibular plates explanate and distinctly longer than clypeus, meeting or not in front of it; anterolateral margins of pronotum crenulate to slightly serrate; mesosternum carinate; labium reaching mesocoxae or slightly beyond, at most to metacoxae; all femora unarmed; peritreme spout-shaped, well developed; and abdominal sternites convex regularly without any tubercles or furrows. Among SE Asian genera of Caystrini , Agathocles is most similar to Valescus (also having anterolateral margins of pronotum crenulate; 10–12 mm ) and Praetextatus (anterolateral margins of pronotum smooth, 9–13 mm )—both are smaller, their scape does not reach apex of head, and mesosternum is furrowed (cf. Distant 1902 ; Hsiao et al. 1977 ; Ahmad & Afzal 1979 ; Chen 1983 ; Ahmad & Kamaluddin 1989 ; Ahmad & Zahid 2006b ; Zahid 2006 ; Fan 2011 ). The enigmatic genus Paramecocoris (body length 6 linien = 13.17 mm ), known only from its original description, differs in having the head longer than wide and basipedicellite (IIa) as long as antennomeres III and IV combined ( Fieber 1851 ; Distant 1902 ; Zahid 2006 ). The remaining genera differ in having anterolateral margins of pronotum smooth and heads distinctly wider than long with mandibular plates not explanate, continually narrowing in front of eyes, as long as or only slightly longer than clypeus, clypeus apically free in Heissocoris , Neoriazocoris , Parodius and Riazocoris ( Distant 1902 ; Ahmad & Afzal 1979 ; Ahmad & Kamaluddin 1989 ; Zahid 2006 ; Ahmad & Zahid 2012 ) or closed by mandibular plates in Caystrus and Hippotiscus (e.g., Distant 1902 ; Ahmad & Zahid 2006a ; Zahid 2006 ; Zahid & Ahmad 2011 ). The Rolstoniellini genera Exithemus and Critheus differs markedly by laterally prolonged and narrowing, rugashaped peritreme, while species of Rolstoniellus differs by conspicuously produced humeral angles of pronotum ( Distant 1902 ). Etymology. Agathocles (= ἈγαθΟΚλῆς ) is a Greek name, e.g. of several ancient Greek philosophers and rulers. The name is derived from agathos (= ἀγαθός ) meaning good and kleos (= ΚλέΟς ) meaning glory; masculine. Neither the etymology nor the gender of the name Surenus were explicitly given in the original description. As we failed to trace it in the Latin or Greek dictionaries, the name must be considered masculine according to the Article 30.2.4 of ICZN (1999) . The name is also a hemihomonym with the plant genus Surenus (family Meliaceae ). Key to species of Agathocles 1 Small species (body length 13.6–15.5 mm ). Body dorsally and ventrally with uniform dark brown colouration with yellow callosities instead of patterns ( Figs 60–63 ). Head shorter (head length: width ratio 0.73), continually narrowing in front of eyes ( Fig. 10 ). Scape (I) surpassing apex of head about half of its length ( Fig. 64 ). Basipedicellite (IIa) short, about half length of distipedicellite (IIb). Genital capsule lacking superior processes of dorsal rim; ventral rim infolding with distinct median process ( Figs 71, 74 ). Apical process of paramere hook-shaped, mesal angle with additional projection ( Figs 75–77 , 84–87 )....................................................................................... A. flavipes sp. nov. – Large species ( 18.8–24.2 mm ). Body dorsally and ventrally with dark brown to blackish colouration with yellow patterns on scutellum or connexivum. Head longer (head length: width ratio 0.79–0.90), parallel-sided anteapically. Basipedicellite (IIa) long, only slightly shorter than distipedicellite (IIb). Genital capsule with superior processes of dorsal rim developed; ventral rim infolding without median process..................................................................... 2 2 Connexivum ventrally, sometimes also dorsally, yellow to orange; scutellum unicolorous except very small and scattered yellowish callosities ( Figs 1–4 ). Scape reaching about apex of head ( Fig. 6 ). Genital capsule with subquadrate superior processes of dorsal rim ( Figs 37–39, 43 ); ventral rim shallowly bisinuate in ventral view ( Fig. 41 ); ventral rim infolding with large semicircular impression medially ( Figs 37–39 ); ventral rim produced ventrally (see in lateral view, Fig. 40 ). Apical process of paramere triangular, mesal angle with additional projection ( Figs 44, 46–47, 49, 52–53 ). Body length 20.61–24.15 mm ................................................................................................................................................................................... A. limbatus Stål, 1876 – Connexivum dark, concolorous with surrounding abdominal terga and sterna; scutellum with yellowish spot on apex ( Figs 88 , 108, 110 ). Ventral rim of genital capsule with single, deep incision medially ( Figs 93 , 115 ), not produced ventrally. Apical process of paramere hook-shaped, mesal angle without additional projection ( Figs 97–101 , 118–123 )................... 3 3 Scutellum with small, yellowish, punctate spot on apex ( Figs 108, 110 ). Lateral pronotal margin anteriorly minutely crenulate. Scape about reaching apex of head ( Fig. 108 ). Genital capsule with small, subtriangular superior processes of dorsal rim ( Figs 113–114, 117 ); ventral rim infolding with large crescent-shaped impression medially ( Figs 113–114 ). Basal process of paramere oriented more vertically, bearing row of many setae ( Figs 118–123 ). Body length 18.80–20.00 mm.......................................................................................... A. normalis ( Distant, 1901a ) – Scutellum with large yellowish impunctate spot on apex ( Fig. 88 ). Lateral pronotal margin anteriorly slightly serrate. Scape surpassing apex of head about half of its length ( Figs 12 , 88 ). Genital capsule with crescent-shaped superior processes of dorsal rim ( Figs 90–91, 96 ); ventral rim sharp, without apparent infolding ( Figs 90–91 ). Basal process of paramere oriented more horizontally, bearing group of few setae ( Figs 97–101 ). Body length 18.90–20.98 mm ................ A. joceliae sp. nov.