Amblycara gladiatoria ( Stål, 1876 )

Tsai, Jing-Fu & Rédei, Dávid, 2014, A revision of the genus Amblycara (Hemiptera: Heteroptera: Pentatomidae), Acta Entomologica Musei Nationalis Pragae 54 (1), pp. 133-155 : 138-147

Amblycara gladiatoria ( Stål, 1876)

( Figs 1, 3 , 5–6 , 9–13 , 17–20 , 25–28 , 33–34 , 37–38 , 41–42 )

Abeona gladiatoria Stål, 1876: 102 . SYNTYPE (S): ♀: “ India orientalis” [= East Indies ]; NHMW. Amblycara gladiatoris: LIN & ZHANG (1985) : 7; HUA (2000): 171. Incorrect subsequent spelling.

References. ATKINSON (1888): 146 (English translation of original description, distribution); LETHIERRY & SEVERIN (1893): 179 (catalogue, distribution); DISTANT (1902): 217 (redescription, habitus, ¿gures, distribution); DISTANT (1908a): 420 (record); DISTANT (1908b): 127 (record); KIRKALDY (1909): 140 (catalogue, distribution), 381 (record); DISTANT (1918): 139 (listed, distribution); ESAKI (1937): 89 (redescription, record); TAKARA (1957): 31 (record, distribution); BLACK (1968): 562 (record); LIN & ZHANG (1985): 7, 10 ( gladiatoris , record); MIYAMOTO & YASUNAGA (1989): 184 (listed, distribution); CASSIS & GROSS (2002):524 (catalogue, distribution, biology); ISHIKAWA (1999): 24 (distribution, record, photo); HUA (2000): 171 ( gladiatoris , listed, distribution); HAYASHI (2002): 147 (distribution); RIDER et al. (2002): 136 (listed, distribution); RIDER (2006): 326 (catalogue, distribution); TAKAI & ISHIKAWA (2012): 488 (redescription, distribution, records, photos of adult, egg batch, 4th and 5th instar larvae).

Type material examined. LECTOTYPE (present designation): ♀, “Ostindien [handwritten] \ coll. Signoret [printed]”, “gladiator [handwritten] \ det. Stal [printed]”; left antenna, right antennal segments IIa–IV, labial segments II–IV, right fore leg, tibia and tarsus of left mid leg lacking; deposited in NHMW ( Figs 5–7 ).

Additional specimens examined. JAPAN: Okinawa Is., date unknown, leg. Sakaguchi (1 ♀, SEHU). – SRI LAN- KA: “Ceylon”, Kandy , coll. H. Rolle (1 J [ Figs 1, 3 , 9–13 , 17–20 , 25–27 ], det. G. Horváth, HNHM) ; “Ceylon”, leg. [E.E.] Green, coll. W.L. Distant 1911-383 (1 J, BMNH) . – INDIA: KERALA: Kardamon Hills, 50 km NW of Pathanamthitta,near Pambaiyar river , 77°05ƍE 9°25ƍN, 6–9.v.1994, leg. Z. Kejval (2JJ 1♀, ZJPC) – LAOS: Louang Phrabang Prov., Muang Ngoy , 20°43ƍN 102°41ƍE, 500 m, 22.iv.1999, leg.V.KubáĖ(1J, det. P.Kment 2007, MMBC) ; Attapeu Prov., Bolavens Plateau , bridge ca. 4 km E Tad Katamtok, 15°07.8ƍN 106°40.1ƍE, 260 m, 11–12.v.2010, at light, leg J. Hájek (1 J, det. P. Kment, NMPC) . – CAMBODIA: Siem Reap, Angkor Thom , 26.v.2003, light trap, leg. J. Constant, K. Smets & P. Grootaert (1 ♀, ISNB) . – MALAYSIA: Malay Penin [sula], Kuala Lumpur, ex coll. Agri. Dept., ex F[ederated] M[alay] S[tates] Museum, B.M. 1955-354 (1 J, BMNH) . – PHILIPPINES: BALABAC: Dalawan Bay , 5.x.1961 (1 ♀, det. G.M. Black, ZMUC, H. Enghoff pers. comm.) . – INDONESIA: “Dutch New Guinea”, Humboldt Bay Dist., Bewani Mts. , ix.1937, W. Stüber, B.M. 1938-177 (1 ♀, BMNH) . – PAPUA NEW GUINEA: NE Wau, 1750 m, 5.ii.1966, leg. J. & M. Sedlacek (1 J, DARC, D.A. Rider pers. comm.). – AUSTRA- LIA: QUEENSLAND: Mt. Garnet, 15–19.i.1989, leg. S. Lamond (1 ♀, JEEC, J.E. Eger pers. comm.) ; Queensland, unspeci¿ed locality, leg. F.P. Dodd, 1904-284 (1 ♀, BMNH) . – NEW CALEDONIA: Farino, 31.iii.[20]01, leg. I. Salesne (1 J, IANC, D.A. Rider pers. comm.) . – FIJI: Viti Levu , SW-Suva, 22.x.1985, leg. G.F. Bornemissza (1 ♀ [ Figs 28 , 33–34 , 37–38 ], HNHM) ; Viti Levu, Nandarivatu , 16–20.xii.1968, leg. H.S. & G.S. Robinson, B.M. 1969-516 (1 ♀, BMNH) .

Diagnosis. Distinguished from the only known congener, A. innocens sp. nov., by the humeral angle being produced into a long, anterolaterally directed process ( Figs 1, 3 ) and by the genitalia of both sexes. A detailed comparison of the diagnostic characters of the two species is provided in Table 1.

Redescription. Macropterous male and female.

Colour, integument and vestiture. Dorsum yellowish brown to light brown, with dense, dark punctation; head (lateral margin of mandibular plates and apical margin of clypeus) narrowly black; antenna stramineous, apical fourths of segments IIb and III and apical third of segment IV black; anterior portion of lateral margin of pronotum with a conspicuous, broad, stramineous callosity bordered by an irregular series of densely arranged, confluent, brown punctures, posterior portion of lateral margin narrowly margined with black and usually also with more or less strong metallic lustre; lateral margin of scutellum with a pair of small, elongate, longitudinally directed black spots subbasally; basal fourth of anterior (= external) margin of fore wing thickened, callose, stramineous; puncturation of exocorium denser and ¿ner than that of endocorium; membrane colourless, with a brown patch apically; legs stramineous, impunctate; connexivum brown to black, puncturation partly confluent, each segment with lateral margin broadly stramineous, impunctate, denticle of posterolateral angle black; venter stramineous, preocular portion of head with a short, narrow longitudinal vitta dorsad of antenniferous tubercle, a small black spot at dorsal extremity of supracoxal clefts; abdominal spiracles III–VII narrowly surrounded by brown or black. Body glabrous; short, ¿ne, semierect or erect, inconspicuous hairs present on antenna, labium, legs, and terminalia; integument rather smooth, except of inconspicuous rugosity on dorsum of head; venter mostly impunctate, smooth.

Structure mainly as in generic description. Pronotum 3.6–3.7 times as broad across humeri as its median length, humeral angle produced into a long process directed slightly anterolaterad.

Pregenital abdomen. Connexivum narrower than in A. innocens sp. nov. (width of each laterotergites of segments III–VII smaller) ( Fig. 33 ) and more broadly covered by the fore wing. Male: Posterior margin of sternite VII relatively narrowly emarginate medially ( Fig. 17 ). Female: Posterior margin of sternite VII broadly concave along its whole width ( Fig. 28 ).

External male genitalia. Genital capsule ( Figs 9, 12–13 ) relatively small in relation to pregenital abdomen, convex, posterior margin broadly concave, with a pair of distinct posterolateral projections ( Fig. 9 : plp) directed posterolaterad, infolding of ventral rim provided with a pair of strongly protruding submedian tumescences ( Fig. 9 : smt); dorsal sclerite ( Figs 9, 12 : dsc; Figs 10–11 ) relatively large, kidney-shaped. Paramere ( Figs 18–20 ) with subtriangular basal process (sensory lobe) ( Fig. 20 : blp), apical process ( Fig. 20 : app) obliquely angled apicad of basal lobe and again around its middle, apical portion slightly narrowed towards apex, provided with an elongate patch of pigmented, scaled texture, obtuse apically. Phallus ( Figs 25–27 ): basal plates ( Fig. 26 : bp) broadly laminate, subquadrate and relatively short in lateral view; phallotheca ( Fig. 26 : phth) barrel-shaped, broadest around middle, base and apex constricted; second conjunctival process ( Fig. 26 : cp-II) with lateral part forming a fully membranous lobe, inner part heavily sclerotized and those of the contralateral processes fused along midline (“median penial plates”, Fig. 26 : mpp); endophallic duct ( Fig. 25 : end) nearly straight, abruptly bent subapically, apical portion directed posterodorsad, ensheathed within aedeagus ( Fig. 26 : aed; Fig. 27 ); phallotreme (opening of the aedeagus) relatively broad ( Fig. 27 ).

External female genitalia. Laterotergites VIII ( Fig. 28 : lt 8) broadly rounded, exposed portion conspicuously smaller than that of A. innocens sp. nov.; visible portion of valvifers VIII ( Fig. 28 : vf 8) subtriangular, posterior margins oblique, laterotergites IX ( Fig. 28 : lt 9) surpassing posterior margin of tergite VIII ( Fig. 28 : t 8). Gynatrium ( Fig. 37 : gy; Fig. 39 ) simple, saccular; spermathecal opening associated with a small annular sclerite which has a pair of oblique projections surrounding spermathecal opening laterally, and a larger, archshaped sclerite surrounding the previous sclerite; ring sclerites could not be traced in the single examined female. Spermatheca: proximal duct ( Fig. 37 : pd) short, somewhat shorter than gynatrium, narrow; dilation long ( Fig. 37 : dil), apex of dilation reaching middle of sternite V within abdomen ( Fig. 34 ); basal cross-striate portion restricted to about basal 1/5 of total length of distal invagination (“sclerotized rod”), distinctly broadened distally ( Fig. 38 ); distal invagination ( Fig. 37 : div) subparallel in most of its length, weakly and gradually broadened subbasally, then relatively strongly narrowed to its base; distal duct ( Fig. 37 : dd) much thinner than proximal duct, longer than half of length of distal invagination; intermediate part of spermatheca rather narrow, broadened distally, proximal flange much smaller than distal flange, apical receptacle subglobular, with three long projections directed proximad, all of them far surpassing proximal flange, one of them even far surpassing distal flange ( Fig. 42 ).

Measurements (n = 3 JJ 2 ♀♀) (in mm). Body length 17.0–21.6, length of head 3.50–3.80, width across eyes 3.65–3.99, interocular distance 2.04–2.45, lengths of antennal segments I: IIa: IIb: III: IV as 0.98–1.10: 2.05–2.40: 2.47–2.96: 3.25–3.80: 3.00–3.35, median length of pronotum 3.45–3.68, width across apices of humeral processes 12.5–13.9, median length of scutellum 7.10–8.07, greatest width 5.90–6.64, greatest width of abdomen 10.4–11.9.

Immatures. Photographs of an egg batch and 4th and 5th instar larvae were presented by KOHNO & TAKAI (2003) and TAKAI & ISHIKAWA (2012).

Habitat and bionomics. All available observations were made in the Ryukyu Archipelago of Japan (Ishigaki and Miyako Islands). Larvae of A. gladiatoria were recorded to feed on fruits of the evergreen tree Drypetes matsumurae (Koidz.) Kaneh. ( Putranjivaceae , formerly Euphorbiaceae ) in July, and adults on fruits of Antidesma pentandrum Merr. ( Phyllanthaceae , formerly Euphorbiaceae ) in early October ( KOHNO & HAYASHI 2003, TAKAI & ISHIKAWA 2012). Both host plants are native in the Ryukyu Archipelago and commonly occur in coral-reef habitats.

Of the above mentioned two host plants, D. matsumurae is a species endemic to the Ryukyu Archipelago. The species was not included in the monograph of Euphorbiaceae (sensu lato) of Taiwan by HSIEH et al. (1993). It was listed, however, from southern Taiwan by LI (1994) without any further information; this record is apparently erroneous and probably pertains to D. littoralis Merr. which is rather common in the Hengchun Peninsula. Antidesma pentandrum is restricted to the Ryukyus, the northern Philippines, and southern and eastern parts of Taiwan ( LI 1963, HSIEH et al. 1993, LI 1994). Since both of the two reported host plants are narrowly distributed, A. gladiatoria obviously feeds on different, so far unknown host plants in other parts of its area of distribution. Both Drypetes and Antidesma are species-rich genera widely distributed in the Oriental Region ( LI 1994), also represented by several species in New Guinea, northern Australia ( AIRY SHAW 1980), New Caledonia ( MCPHERSON & SCHMID 1991) and Fiji ( SMITH 1981); at least part of the included species, perhaps also members of other related genera or families of Malpighiales could potentially be consumed by A. gladiatoria . Most species of Drypetes and Antidesma are commonly found in damp understorey, in riverine forests, swamps, frequently on seashores; coastal forests are perhaps the preferred habitat of A. gladiatoria .

The only paper providing data on the bionomics of A. gladiatoria is that of KOHNO & HAYASHI (2003) based on observations in uplifted coral-reef forests of Ishigaki Island in early June of 2002. During this period aggregations of several larvae of 3rd–5th instars were observed to feed on fruits of the host plant at about 3–4 m above the ground. Not a single individual was found after July, according to the authors adults presumably left the plant soon after emergence and migrated to A. pentandrum . Two observed egg batches contained 81 and 58 eggs, respectively.

Note. The original description of A. gladiatoria does not state the number of specimens available to the author, and it cannot be excluded that the description was based on more than one specimen. The specimen illustrated in Figs 5–6 was apparently ¿gured by DISTANT (1902: 217, ¿g. 137), although with a modi¿ed position of legs and antennae. This author referred to the specimen as “a typical one”, making it clear by the use of an inde¿nite article that he considered that the original description was possibly based on more than one specimen. Because of this the above expression cannot be considered the equivalent of the term lectotype, and therefore Distant’s act cannot be considered a valid ¿xation of lectotype under the International Code of Zoological Nomenclature ( ICZN 1999, Articles 74.5 and 74.6). As a consequence, the identity of the species is ¿xed in the present paper by designating (presumably) the same specimen ( Figs 5–7 ) as lectotype.

Distribution. SRI LANKA: Kandy! INDIA: KERALA: Kardamon Hills! (new record). CHINA: YUNNAN: Jingdong ( LIN & ZHANG 1985). JAPAN: TOKARA ISLANDS: Akuseki Is. ( HAYASHI 2002, KOHNO & HAYASHI 2003, TAKAI & ISHIKAWA 2012); RY Nj KY Nj ISLANDS: Okinawa Is.!; SAKISHIMA ISLANDS: Miyako Is.; YAEYAMA ISLANDS: Ishigaki Is., Iriomote Is., Yonaguni Is. ( ESAKI 1937, TAKARA 1957, MIYAMOTO & YASUNAGA 1989, ISHIKAWA 1999, HAYASHI 2002; KOHNO & HAYASHI 2003, TAKAI & ISHIKAWA 2012). LAOS: ATTAPEU PROV.: Muang Ngoy! (new record). CAMBODIA: Siem Reap, Angkor Thom! (new record). MALAYSIA: MALAY PENINSULA! (new record). PHILIPPINES: BALABAC IS.: Dalawan Bay! ( BLACK 1968). INDONESIA: PAPUA: Bewani Mts.! (new record). PAPUA NEW GUINEA: Wau! (new record). AUSTRALIA: QUEENSLAND!: Mt. Garnet!, Kuranda ( CASSIS & GROSS 2002). NEW CALEDONIA: Farino! (new record). FIJI: VITI LEVU! (new record).

The species has a broad distribution area from Sri Lanka apparently all over South and Southeast Asia and the Malay Archipelago to northern Australia and Fiji ( Fig. 8 ). However, it is rarely captured and only a few sporadical records exist, except for the Ryukyu Archipelago of Japan where it was recorded from several islands. We present the ¿rst record from Laos, Cambodia, Malaysia, New Guinea (both Indonesian and Papuan parts), New Caledonia and Fiji (partly based on personal communication of D. A. Rider). The record from Balabac Is. ( BLACK

1968) was veri¿ed by H. Enghoff by comparing the voucher specimen to the illustrations of the present paper. The source of the record from Queensland: Kuranda ( CASSIS & GROSS 2002) is unknown, but a specimen from an unspeci¿ed locality in Queensland deposited in BMNH was seen ; its collector, F. P. Dodd collected extensively around Kuranda, therefore the specimen potentially comes from the same region. Another female from the nearby Mt. Garnet was communicated by J. E. Eger. Papers listing the species from India or “E. India ” apparently refer to the original description, however, the type locality, “ India orientalis” (= “ East Indies ”) pertains to a much broader area comprising practically the whole Oriental Region. Based on personal communication of Z. Jindra we present the ¿rst con¿rmed record from India .

Figures

Figs 1–4. Males of Amblycara in dorsal (1–2) and ventral view (3–4). 1, 3 – A. gladiatoria (Stål, 1876); 2, 4 – A. innocens sp. nov. (holotype).

Figs 5–7. Lectotype of Amblycara gladiatoria (Stål, 1876). 5 – dorsal view; 6 – ventral view; 7 – labels. Scales in mm. ©NHMW.

Figs 9–16. Amblycara gladiatoria (Stål, 1876) (9–13) and A. innocens sp. nov. (14–16), genital capsules. 9 and 14 – dorsal view; 10–11 – dorsal sclerite, two different views; 12 and 15 – posterodorsal view; 13 and 16 – lateral view. Scales in mm. Lettering: dsc = dorsal sclerite; smt = submedian tumescence; plp = posterolateral projection.

Figs 17–24. Amblycara gladiatoria (Stål, 1876) (17–20) and A. innocens sp. nov. (21–24), pregenital abdomen and paramere of male. 17 and 21 – abdominal sternites VI and VII, ventral view; 18–20 and 22–24 – right paramere, different views. Scales in mm. Lettering: app = apical process; blp = basal process (sensory lobe); s7 = sternite VII.

Figs 25–32. Amblycara gladiatoria (Stål, 1876) (25–28) and A. innocens sp. nov. (29–32), phallus and female terminalia. 25 and 29 – phallus, lateral view; 26 – same, dorsal view; 30 – same, articulatory apparatus omitted; 27 and 31 – apex of phallus, posteroventral view; 28 and 32 – terminalia of female, most exposed (posteroventral) view. Scales in mm. Lettering: aed = aedeagus; bp = basal plates; cp-II = second conjunctival process; end = endophallic duct; erp = erection fluid pump; hi = hinge; lt8–lt9 = laterotergites VIII and IX; mpp = median penial plate; phth = phallotheca; res = endophallic reservoir; s7 = sternite VII; sb = support bridge; sbc = support bridge complex; sut = support tube; t8 = tergite VIII; vf8–vf9 = valvifers VIII and IX; X = segment X.

Figs 33–36. Amblycara gladiatoria (Stål, 1876) (33–34) and A. innocens sp. nov. (35–36), abdomen of female. 33 and 35 – dorsal view; 34 and 36 – ventral view. Scales in mm. Lettering: mfu = median furrow; sth = spermatheca; s5–s7 = sternites V–VII.

Figs 37–40. Amblycara gladiatoria (Stål, 1876) (37–38) and A. innocens sp. nov.(39–40), female ectodermal genital tracts. 37 and 39 – gynatrium and spermatheca; 38 and 40 – basal portion of dilation of spermatheca with distal invagination (“sclerotized rod”), longitudinal optical section.Scales in mm.Lettering:dd = distal duct of spermatheca; dil = spermathecal dilation; div = distal invagination of spermathecal duct; gy = gynatrium; pd = proximal duct of spermatheca. Arrow shows possible artefactual distortion of apical portion (Fig. 39) and basal portion (Fig. 40) of distal invagination of spermathecal duct.

Figs 41–44. Amblycara gladiatoria (Stål, 1876) (41–42) and A. innocens sp. nov.(43–44), female ectodermal genital tracts. 41 and 43 – gynatrium; 42 and 44 – intermediate part and apical receptacle of spermatheca. Scales in mm.

Fig. 8. Distribution of Amblycara gladiatoria (Stål, 1876) (circles) and A. innocens sp. nov. (squares). Filled symbols represent records based on specimens examined by us or veri¿ed literature records, open symbols represent unveri¿ed literature data.