Agathocles limbatus Stål, 1876

Salini, S. & Kment, Petr, 2021, The genera Agathocles and Surenus (Hemiptera: Heteroptera: Pentatomidae) tribal reassessment, redescription, new synonyms, and description of two new species, Zootaxa 4958 (1), pp. 510-559 : 524-529

Agathocles limbatus Stål, 1876

( Figs 1–8 , 16, 22 , 25–30 , 33 , 37–59 , 139 )

Agathocles limbatus Stål, 1876: 106 (original description). Holotype: ♂, Bangladesh, Silhet [= Sylhet] (NHRS:?lost).

Agathocles limbatus: Atkinson (1888) : 153 (redescription, distribution); Lethierry & Severin (1893): 183 (catalogue); Distant (1902): 199, fig. 121 (redescription; figures of habitat in dorsal view, and mesothorax, metathorax and abdomen in ventral view; distribution); Kirkaldy (1909): 147 (catalogue); Ahmad (1981): 52 (list); Zhang & Lin (1985): 21 –22 (record); Zhang & Lin (1986): 61 (record); Rider (1997): 402 (checklist); Rider et al. (2002): 136 (Chinese distribution); Rider (2006): 342 (catalogue); Li et al. (2008): 212 (checklist); Fan (2011): 43 (key to species); Hoàng & Ðãng (2013): 789 (record).

Agathocles fimbatus (incorrect subsequent spelling): Hua (2000): 171 (checklist).

Agathocles limbaatus (incorrect subsequent spelling): Zhang (1994): 33 (catalogue).

Agothocles [sic!] limbatus : Ahmad et al. (1974): 43 (list, host plant), 75: figs 63, 66, 88, 89 (drawings of habitus in dorsal view and meso- and metapleuron in ventral view).

Agathocles yunnanensis Zhang & Lin, 1984 in Lin & Zhang (1984): 268–270, figs 3, 4 (original description, line drawings of habitus in dorsal view and female terminalia in ventral view). Holotype: ♀, China, Yunnan, Cang-yuan County, 675 m a.s.l., 19.vii.1980 (IZAS). New junior subjective synonym.

Agathocles yunnanensis: Zhang et al. (1987) : 144 (key to genera and species); Yang & Sun (1991): 28 (type depository); Rider (1997): 402 (checklist); Hua (2000): 171 (checklist); Rider et al. (2002): 136 (Chinese distribution); Rider (2006): 342 (catalogue); Fan (2011): 43 (key to species), 44–45, figs 2.14a–f (redescription, drawings of male genitalia), 555–556, figs 32–33 (photos of habitus in dorsal and ventral view).

Agathooles [sic!] yunnanensis : Jiang (1985): 67 (checklist).

Type locality. Agathocles limbatus . Bangladesh, Silhet [= Sylhet], 24°54′N 91°52′E . Agathocles yunnanensis . China, Yunnan, Cangyuan Va Autonomous County, 23°09′N 99°14′E

Type material. Agathocles limbatus . Not examined. The description was based on single specimen (holotype) as it is mentioned in the original description: ‘Antennae exempli descripti …’ [= antennae of examined specimen …]. The holotype should be deposited in NHRS (cf. Stål 1876; Rider 2006), but it is not included in the list of the collection by Gustafsson (2006). Dávid Rédei (pers. comm.) confirmed to us that he did not locate the holotype in NHRS during his carefull search of the collection.

Agathocles yunnanensis . Holotype: ♀ ( Figs 3–5 ), China, Yunnan, Cang-yuan County, 675 m a.s.l., 19.vii.1980 ( IZAS). The photographs of the holotype were provided by courtesy of D. Rédei.

Material examined. CAMBODIA: Pailin env., [12°52′N 102°36′E], 200 m a.s.l., 16.–17.vi.2008, 1 ♂, S. Murzin lgt., P. Kment det. ( NMPC) . CHINA: Guangxi: 广NJkdfi.乡 550m 2000-VI-5 李rff. ḤflLj 物 ¶ [= Guangxi: Fangchenggang, 21°42′N 108°21′E, Tongzhong village , 550 m a.s.l., 5.vi.2000], 1 ♂, W.-Z. Li lgt., Z-H. Fan det. as A. yunnanensis, P. Kment revid. ( IZAS → NKUM) . Tibet: Chanyu [= Zayü county], Xiachayu [= Xiazayü], 28°29′16″N 97°01′22″E, vi.2017, 3 ♀, local collector (1 ♀ NMPC, 2 ♀ MRCS) . LAOS: Houa Phan Province: Ban Saleui, Phou Pan ( Mt. ), 20°12′N 104°01′E, 1300–1900 m a.s.l., 13.vi.–6.vii.2013, 3 ♂ 4 ♀, C. Holzschuh lgt., Acq. Nr. 2013-13, P. Kment det. (2 ♂ 3 ♀ NHMW, 1 ♂ 1 ♀ NHMW → NMPC) ; Ban Saluey → Phou Pane Mt. , 20°12– 13.5′N 103°59.5′– 104°01′E, 1340–1870 m a.s.l., primary mountain forest, individual collecting, 1.–24.vi.2012, 1 ♂, Vít Kubáň & Lao collectors lgt., P. Kment det. ( NMPC) ; Phu Pane Mt. , 20°12′N 103°50′E, 900–1400 m a.s.l., 10.–21.vi.2010, 1 ♀, S. Jakl & Lao collectors lgt., P. Kment det. ( NMPC) ; Phu Phan Mt. , iv.2016, 1 ♂, P. Kment det. ( MRCS) . Louang Phrabang Province: Ban Song Cha (5 km W), 20°33 –34′N 102°14′E, ± 1200 m a.s.l., 24.iv.–16.v.1999, 1 ♂, V. Kubáň lgt., P. Kment det. ( MMBC) . Phongsaly Province: Phongsaly env., 21°41‒42′N 102°6–8′E, ca. 1500 m a.s.l., 28.v.–30.vi.2003, 2 ♀, P. Pacholátko lgt., P. Kment det. ( MMBC) . THAILAND: Loei Province: Phu Kradung NP, 16°53′N 101°47′E, 1300 m a.s.l., 11.–15.v.1999, 1 ♀, D. Hauck lgt., P. Kment det. ( NHMW) . VIETNAM: Cao Bang Province: Phia Oac NP, 7 km N to village, [22°35′N 105°53′E], 28.v.2014, light trap, 2 ♂ 2 ♀ ( NMNS ENT 7399-646 , 668 , 877 , 879 ) , M. L. Jeng lgt., J.-F. Tsai det. as A. yunnanensis, P. Kment revid. ( NMNS) .

Redescription. Colouration ( Figs 1–4 ). Head above and below, thorax ventrally, abdominal tergites V (medially) and VI–VIII (completely), and sternites, and pygophore posteroventrally black; pronotum, scutellum and hemelytra dark brown with mosaic of smaller to larger, irregular, yellowish- to orange-ochraceous callosities (largest callosity on corium at the end of median furrow). Lateral margin of pronotum (narrowly), costal margin of corium anteriad of connexivum, posterolateral margin of metapleuron, and connexivum ventraly largely orangeochraceous (connexival segments along margins narrowly black). Most of femora, tibiae, tarsi, and antennomeres I to III dark brown. Distiflagellum (IV) and labium yellowish. Coxae, trochanters, bases of femora, and basal portion of pygophore yellowish brown. Membrane dark brown, translucent. Thoracic pleura (outside evaporatorium), especially on acetabula, with variable distribution of yellowish callosities. Peritremal surface ochraceous.Abdominal tergites I–IV completely and V laterally brownish. Round fields surrounding abdominal trichobothria very small, brown, not much contrasting with their black surroundings.

Integument and vestiture. Body dorsally matt, ventrally shining. Body above (including inner portion of corium) covered with dense, coarse, black punctures, disc of pronotum (posteromedially) and scutellum (anteriorly) with fine transverse wrinkles; head underside and thoracic sterna with dense, coarse punctures; surface of abdominal sternites smooth with fine punctures. Body mostly glabrous (see generic redescription).

Structure. Head ( Figs 6–8 ) slightly shorter than wide (across eyes), head length to width ratio 0.82–0.90. Mandibular plates with lateral margins strongly insinuate in front of eyes above small antenniferous tubercle, parallel-sided in about their midlength, and rather regularly rounded anteriad; apical incision in front of clypeus Vshaped; lateral margins of mandibular plates slightly raised, more prominent laterally than apically. Antennomeres from shortest to longest: I <IIa <IV <IIb <III. Scape (I) about reaching apex of head; basipedicellite IIa distinctly (1.70–2.66×) longer than scape (I), only slightly shorter than IIb (length of IIa: IIb ratio: 1.03–1.63×). Labiomeres from shortest to longest (n = 1): IV (1.32 mm) <I (1.52 mm) <III (1.96 mm) <II (2.21 mm). Labium reaching between mesocoxae or slightly beyond.

Pronotum ( Figs 1, 3 ). Lateral tooth on anterolateral angles larger, directed laterad. Lateral pronotal margin anteriorly only minutely crenulate. Humeri rectangular, slightly surpassing laterad over base of corium. Posterior margin slightly concave.

Scutellum ( Figs 1, 3 ). Disc of scutellum convex in prefrenal portion, its postfrenal portion flat, only slightly raised medially, apex narrowly rounded.

Hemelytra ( Figs 1, 3 ). Costal margin of corium slightly more insinuated basally.

Thorax. Ostiole large, oval, opening posteriad; median furrow short and wide, not apparent in apical half of peritreme length ( Fig. 16 ). Metasternum wide ( Fig. 24 ).

Male genitalia. Genital capsule ( Figs 37–41 ) nearly quadrangular in dorsal/ventral view, longer than wide. Ventral rim with double shallow insinuation submedially, posterolateral projections not prominent ( Fig. 41 ); ventral rim infolding wide, semi-circular, biconcave, slightly raised medially ( Figs 37–39 ). Ventral margin of genital capsule produced ventrally (see in lateral view, Figs 28 , 40 ). Superior processes of dorsal rim subquadrate ( Figs 37–39, 43 ). Parameral socket open ( Fig. 38 : ps). Proctiger trapezoid, laterally constricted about its midlength ( Figs 37, 42 ). Paramere ( Figs 44–53 ) with basal process (bp) foliaceous, roughly semicircular, apically with short row of long setae ( Figs 44, 47–48, 51 ); apical process (ap) triangular, more pointed apically ( Figs 44, 46–47, 49, 52–53 ); mesal angle of stem with an additional lamellate projection ( Figs 44, 46–47, 50–51 ). Phallus ( Figs 54–59 ): A pair of dorsal conjunctival processes, transversely placed and ending in sclerotised processes ( Figs 57, 59 ); median penial plates or processes of aedeagus (pa) fused basally, outer margins broadly and thickly sclerotized, encircling short, curved aedeagus ( Fig. 59 : ad).

Female genitalia ( Figs 29–30 , 33 ). Laterotergite VIII (lt 8) subtriangular. Laterotergites IX (lt9) widely rounded apically, not touching each other medially, segment X visible ( Figs 29–30 ). Valvifers VIII (vf8) quadrant with median margins juxtaposed and dorsal margins slightly convex ( Figs 29–30 ). Valvifers IX (vf9) fused, medially with impressed median fusion line. Ring sclerites drop-shaped, narrowly sclerotized ( Fig. 33 : rs). Spermatheca: dilation long, narrow in the examined specimen ( Fig. 33 : dil), distal invagination (‘sclerotized rod’; Fig. 33 : div) distinctly broadened distally, subparallel in most of its length, basally straight and slightly broadened, distal flange tightly attached to apical receptacle ( Fig. 33 : df); apical receptacle subhemispherical ( Figs 33 : ar) with one long projection directed proximad and far surpassing proximal flange (pf), and three short processes of various orientation, not surpassing distal flange.

Measurements. See Table 2.

Variation. Among the specimens examined there is distinct variability in the following characters: Colouration: Dorsal connexival segments orange-ochraceous with concolorous punctures ( Fig. 1 ) to dull ochraceous with dense black punctures ( Fig. 3 ); antennomeres I basally, IIa basally and in middle, III basally and apically, and tarsomeres I and II dorsally ochraceous; lateral margin of pronotum (including anterolateral angle or not) dark, concolorous with surrounding surface ( Fig. 4 ); small circles around trichobothria yellowish, more contrasting with surrounding surface; femora brown or blackish brown ( Figs 2, 4 ). Distribution of callosities on body, e.g. abdominal ventrites with yellowish callosities or completely black. Mandibular plates meeting in front of clypeus ( Fig. 6 ) or not, leaving apex of clypeus free ( Fig. 7 ). Length of antennomeres (see Table 2) and their ratios, from shortest to longest: I <IV <IIa <IIb <III (2 specimens), I <IIa <IIb <IV <III (1 specimen), I <IIa = IV <IIb <III (1 specimen), I <IIa <IV <III <IIb (2 specimens). Ventral rim infolding of genital capsule medially with slightly raised keel ( Fig. 37 ) or the keel is indistinct ( Fig. 38 ). Apical process and lamellate projection of mesal angle of paramere slightly different in shape (cf. Figs 44–46, 50, 52 versus 47–49, 51, 53).

Differential diagnosis. See key to species and Table 3. This species is easy to identify due to the wide yellow or orange stripe ventrally on connexivum.

Distribution ( Fig. 139 ). Bangladesh ( Stål 1876; Atkinson 1888; Distant 1902); Cambodia (new record); China: Fujian ( Zhang & Lin 1985: ‘Sha County, Jianyang’; Rider et al. 2002), Guangxi ( Fan 2011 [unpubl.]; new record), Jiangxi ( Zhang & Lin 1985: ‘Xunwu, Guizhumao’, 1986; Zhang 1994; Li et al. 2008), Tibet (new record); Yunnan ( Lin & Zhang 1984; Jiang 1985, both as A. yunnanensis ); India: Assam ( Distant 1902, without exact locality); Laos (new record); Thailand (new record); Vietnam ( Hoàng & Đăng 2013).

The Distant’s (1902) record from Assam may refer also to other Indian states later separated from the original territory of Assam.

Taxonomy. Zhang & Lin in Lin & Zhang (1984) described a new species, A. yunnanesis from Yunnan, distinguishing it from A. limbatus . The description was written in Chinese with only a short English summary. By the courtesy of D. Rédei, we provide here translation of the original description: ‘Female body length 23 mm, width 13 mm, humeral width 11 mm. Oval, ground colour blackish brown, with orange marmorated spots, finely and densely punctured. Mandibular plates longer than clypeus, anteriorly separated and forming a V-shaped interspace, lateral margin turned upwards; compound eyes black, ocelli pale yellowish. Antennomeres III, IV lacking; the bases and tips of the remaining antennomeres provided with a small protrusion on their outer sides, antennomeres I–IIb dark brown, bases and tips of each segments pale, lengths as 1.09, 3.00, 3.46 mm, antennomeres IIa–IIb pilose. Labium yellowish brown, reaching slightly before mesocoxae. Pronotum and basal half of scutellum with several irregular rugosities. Anterior margin of pronotum elevated at middle, anterior angle orange brown, tip sharp and directed outwards, anterolateral margin narrowly protruding, concave close to its middle; humeral angle slightly protruding, apex rounded, with crescent-shaped orange line, margin of callar area smooth, edge with several small, scattered, orange spots; pro- and mesosternum with low and narrow median carina, metasternum flat. Legs black, coxae, bases of femora, and tarsi yellowish brown. Scutellum with several small, marmorate, orange spots. Corium with several irregular orange markings, anterior margin marked with orange at base, slightly turned upwards; membrane blackish brown, not surpassing apex of abdomen. Connexivum exposed. Abdominal venter pitch-black, slightly shiny, densely and finely punctured, lateral margin of each segment with a broad orange band. Female terminalia in ventral view as in fig. 4.

Similar to A. limbatus , but the latter is somewhat smaller, not surpassing 21 mm in length, dorsum decorated with fine, marmorate, orange markings, ventrally also with such fine, marmorate, orange markings, all femora and tibiae yellowish brown, labium reaching between mid coxae.’ ( Lin & Zhang 1984).

The English abstract by Lin & Zhang (1984) says that the labium reaches ‘beyond the middle of coxae’, however, the Chinese text clearly states that it extends to the area slightly anteriad of mesocoxae (D. Rédei, pers. comm.).

All the characters distinguishing between A. limbatus and A. yunnanensis provided by Lin & Zhang (1984) and observed on the photographs of the holotype of A. yunnanensis fits within the variability of A. limbatus as observed in our study. Therefore, we consider A. yunnanensis as junior subjective synonym of A. limbatus . The observed differences in structure of pygophore and paramere seems rather minor and we prefer to treat A. limbatus as single though variable species.

Figures

FIGURE 139. Distribution of the Agathocles species. T—type locality;?—doubtful record. Orig. P. Kment.

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.

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.

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).

FIGURES 37–43. Male genital capsule of Agathocles limbatus Stål, 1876 (37, 40, 41—Laos: Phou Pane Mt.; 38, 42–43— Cambodia: Pailin env.; 39—China: Guangxi): 37—caudal view, not dissected (27×); 38–39—caudal view, dissected (27×); 40—lateral view (27×); 41—ventral view (19×); 42—proctiger (75×); 43—superior process of dorsal rim (110×). Lettering: pa –paramere, plp—posterolateral projection, ps—paramere socket, sp—superior process of dorsal rim, vif—ventral rim infolding, x—proctiger. Scale bars: 1 mm (37–41), 0.5 mm (42–43). Orig. P. Kment.

FIGURES 44–53. Paramere of Agathocles limbatus Stål, 1876 from Cambodia: Pailin env. (44–46, 50, 52) and China: Guangxi (47–49, 51, 53) (five different views, 70×). Lettering: ap—apical process of paramere, bp—basal process of paramere, map— projection of mesal angle of paramere stem. Scale bars: 0.5 mm. Orig. P. Kment.

FIGURES 54–59. Penis of Agathocles limbatus Stål, 1876 from Cambodia: Pailin env. (54–56—non inflated, 57–59—inflated): 54, 57—dorsal view; 55, 58—lateral view; 56—ventral view; 59—apical view. Lettering: ad—aedeagus, dlc—dorsolateral conjunctival process, mpp—median penial plate, pt—phallotheca. Scale bars: 1 mm. Orig. P. Kment.