Neocentrocnemis stali (Reuter, 1881)

Rédei, Dávid & Tsai, Jing-Fu, 2011, The assassin bug subfamilies Centrocnemidinae and Holoptilinae in Taiwan (Hemiptera: Heteroptera: Reduviidae), Acta Entomologica Musei Nationalis Pragae 51 (2), pp. 411-442 : 414-422

Neocentrocnemis stali (Reuter, 1881)

( Figs. 1–40 )

Centrocnemis stali [ as Ståli ] Reuter, 1881: 65. LECTOTYPE ( MILLER 1956: 267, by use of ‘holotype’) (♀): India: Darjeeling ; NHRS!

Centrocnemis formosana Matsumura, 1913: 161 . HOLOTYPE (♀): ‘Formosa’ [= Taiwan]:‘Arikan’ [= Likang, Pingtung County]; EIHU ( TOMOKUNI & CAI 2002). Erroneously synonymized with C. deyrollii Signoret, 1852 by ESAKI (1926): 165. Syn. nov.

Centrocnemis deyrollii (non Signoret, 1852): BERGROTH (1914): 364. Misidentification.

Neocentrocnemis baudoni Dispons, 1965: 91 . HOLOTYPE (♀): Laos; MNHN! Syn. nov.

References. LETHIERRY & SEVERIN (1896): 96 (catalogue); DISTANT (1904): 246 (redescription, records); PAIVA (1919): 367 (record); MATSUMURA (1930): 182 ( formosana , redescription, record, habitus); MATSUMURA (1931): 1207 ( formosana , redescription, record, habitus); ESAKI (1932): 1655 ( deyrollei , redescription, habitus); KATO (1933): [plate 22] ( deyrollei , distribution, colour photo); CHINA (1940): 252 (listed); HOFFMANN (1944): 17 (record, distribution); MILLER (1956): 251 ( formosana , new combination, type material, redescription, figures, records), 265 (type material, redescription, figures, records); DISPONS (1965): 94 (in key, distribution); HSIAO (1974): 321 ( stali and formosana , in key); HSIAO & REN (1981): 414 ( stali and formosana , redescription, figures, photos); REN (1985): 178 (redescription, habitus, habitat, distribution); MALDONADO CAPRILES (1990): 16 ( formosana and stali , catalogue); BISWAS et al. (2004): 370, 379 (listed, record, distribution); PUTSHKOV & PUTSHKOV (1996): 148 ( formosana , catalogue); HUA (2000): 209 ( formosana and stali , listed, distribution); TOMOKUNI & CAI (2002): 102 (type material of formosana ); LIN (2003): 128 ( formosana , listed, distribution), 129 ( stali , listed, distribution); YANG (2003): 129 ( formosana , male genitalia); ISHIKAWA (2005): 26 ( formosana , photo); AMBROSE (2006): 2395 (listed, distribution); PUTSHKOV & PLUOT- SIGWALT (2008): 152 ( baudoni , type material); WEIRAUCH et al. (2011): 141 (molecular genetics).

Type material examined. Centrocnemis stali Reuter, 1881 . LECTOTYPE (♀): ‘3 \ 55’ [light red square, printed + handwritten], ‘Darjee- \ ling.’ [printed], ‘ Staudinger. ’ [printed], ‘ Ståli Reut. \ Typ.’ [Reuter’s handwriting], ‘Typus’ [red square with black frame, printed], ‘ Neocentrocnemis \ stali (Reut.) \ N.C.E. Miller det.1955’ [Miller’s handwriting + printed], ‘NHRS-GULI \000000152’ [printed]; deposited in NHRS ( Figs. 1–3 ). – Neocentrocnemis baudoni Dispons, 1965 . HOLOTYPE (♀): ‘M. Baudon \ Laos \ S. W 63’ [handwritten], ‘Type ♀ ’ [handwritten in red], ‘ Neocentrocnemis \ baudoni DISPONS’ [handwritten], ‘TYPE’ [red square, printed], ‘ Museum Paris \ MNHN ( EH) \ 1573 ’ [printed]; deposited in MNHN ( Figs. 4–7 ).

Specimens examined. TAIWAN: TAICHUNG COUNTY: Baxianshan National Forest Recreation Area , 13–15.vii.2006, leg. J. F. Tsai (1 ♂ 1 ♀, 1 L4, NCHU) ; same locality, 8.vii.2010, leg. T.J. Hsieh (1 ♂, NCHU) ; Guguan , 12.iv.2001, leg. Q.Z. Huang (1 ♀, NCHU) . NANTOU COUNTY: Howangshan , 10.ix.2000, leg. C.C. Lo (2 ♀♀, NMNS, ENT 4463-123 , ENT 4463-927 ) ; Huisun Forest Station , 31.vii.2008, leg. Y.M. Weng (1 ♀, NCHU) ; ‘ ChipChip’ [= Jiji], ii.1909, leg. H. Sauter (1 ♂ 1 ♀, HNHM) ; Nanshanxi , 6.vii.2008, leg. H.Y. Lin (1 ♀, NCHU) ; ‘ Polisha’ [= Puli], xii.1908, leg. H. Sauter (1 ♂ [identified as ‘ Centrocnemis deyrollei Sign. ’ by G. Horváth], 2 ♀♀, HNHM) ; ‘ Hori (Puli, Polisia [= Polisha])’ [= Puli], 800 m, 23.viii.[19]47, leg. L. Gressitt (1 ♀ [identified as ‘ Neocentrocnemis formosana (Mats.) ’ by T.Y. Hsiao, photographed by HSIAO & REN (1981: plate 57, fig. 571)], NKUC) ; ‘ Fuhosho’ [= Wucheng], iv.1909, leg. H. Sauter (1 ♂, HNHM) ; same locality and collector, vii.1909 (1 ♂ 5 ♀♀, HNHM) ; same locality and collector, viii.1909 (1 ♂, HNHM) ; same locality and collector, ix.1909 (8 ♂♂ 9 ♀♀, HNHM) ; same locality and collector, x.1909 (2 ♀♀, HNHM) . CHIAYI COUNTY: ‘ Chikutoge’ [= Jhuci], V. 1909, leg. H. Sauter (1 ♀, HNHM) ; Shanmei , 600 m, 23.v.1977, leg. J. & S. Klapperich (1 L5, coll. E. Heiss, Innsbruck) . KAOHSIUNG COUNTY: ‘ Kosempo’ [= Jiasian], iv.1909, leg. H. Sauter (1 ♀, HNHM) ; same locality and collector, vii.1909 (1 ♂ 1 ♀, HNHM) ; same locality and collector, ix.1909, leg. H. Sauter (1 ♀, HNHM) ; same locality and collector, x.1909 (1 ♀, HNHM) ; same locality and collector, 7.vii.1911 (1 ♀ [identified as ‘ Centrocnemis deyrollii Sign. ’ by E. Bergroth, as ‘ Neocentrocnemis formosana (Mats.) ’ by N. C.E. Miller, 1955], DEIC). COUNTY UNCERTAIN: ‘ Formosa’, leg. H. Sauter (1 ♂, HNHM) . CHINA: HAINAN: Jianfengling , 8.iv.1983, leg. M. B. Gu (1 ♂, NKUC) . VIETNAM. BẮC KẠN PROV.: Ba B ể National Park, near headquarters, from bark and litter, 1.v.2007, leg. G. Csorba ( HNHM). HÀ TÂY PROV.: Mt. Bavi , 800–1000 m, vii.1941, leg. A. de Cooman (1 ♂ [photographed by HSIAO & REN (1981: plate 57, fig. 570)], Musée Heude> NKUC). HÒA BÌNH PROV.: Hòa Bình, leg. A. de Cooman (1 ♂ [identified as ‘ Centrocnemis deyrollei Sign. ’ by T.Y. Hsiao, 1966], 2 ♀♀, NKUC) .

Diagnosis. Recognized within the genus by the combination of the following characters: body relatively small, total length 19–24 mm; flattened humeral projections of the pronotum relatively short, their apical triangular processes short and wide; disk of posterior lobe of pronotum with 1+1 short and stout spine-like tubercles; corium of fore wing gray with whitish venation, membrane greatly smoky gray, basal angle as well as an apical spot in each membranal cells blackish; posterolateral angle of connexival segments II–III each with 2 processes, that of segments IV–VI each with a single process ( Figs. 17–18 ). For faciliting the recognition of the species, the male and female genitalia are described and illustrated in detail.

Description of male genitalia. Genital capsule ( Figs. 8–9 ) elongate, dorsally membranous, anterior and posterior apertures not separated by distinct dorsal sclerotized bridge-like portion therefore seemingly confluent; superoposterior margin with a broad triangular median projection and 1+1 broad lateral projections of variable shape ( Figs. 10–11 ); sternite IX with a partly sclerotized, mobile cuplike sclerite within genital cavity surrounding apical portion of phallus in repose ( Fig. 14 : cs). Parameres ( Figs. 12–13 , 19–20 ) symmetrical, apical portions strongly curved, hook-like, crossing each other in resting position ( Fig. 8 ). Phallus ( Fig. 14 , 21–28 ): articulatory apparatus ( Fig. 22 : aa; Fig. 15 ) small, restricted to the extreme base of the phallus; basal plates ( Fig. 15 : bp [dotted]) short, U-shaped, longitudinal arms thickened around middle; support bridge complex ( Fig. 15 : sbc) with a thick ponticulus transversalis and with 2+2 capitate processes of lateral position, continued in 1+1 greatly elongate, narrow support bridge prolongations ( Fig. 15 : sbp) narrowly separated at their base but fused at most of their length, bulbously thickened apically; ductus seminis could not be traced; phallosoma with a large, membranous basal aula ( Fig. 22 : ba) provided with a pedicel ( Fig. 22 : ped); phallotheca ( Figs. 23 , 25–26 ) short, simple, tubular, apically broadly rounded, with 1+1 dorsal ( Figs. 23 , 25 : dsp) and 1+1 ventral ( Figs. 23 , 25 : vsp) narrow, band-like sclerotized plates, dorsal ones arched and widely separated, ventral ones parallel and adjacent; phallotheca mouth ( Fig. 25 : phm) broad; endosoma ( Figs. 24 , 27–28 ) tubular, membranous, impossible to inflate because of rigid sclerotized elements in its wall, with processes as follows: 1+1 struts subdivided into two portions: basal portions ( Figs. 24 , 27–28 : strb) thick, stout, apically tumid, provided with a ventroapical finger-like projection, apical portions ( Figs. 24 , 26–28 : stra) whip-like, thick basally, strongly narrowing towards apex, apical four-fifth thin, protruding from phallotheca; 1+1 membranous dorsolateral projections ( Figs. 24 , 27–28 : dlp); with a dorsal sclerite provided with 1+1 lobe-like protrusions laterally ( Fig. 24 , 26–28 : dsc); with 1+1 elongate, plate-like lateral sclerites ( Figs. 24 , 26–28 : lsc); 1+1 small and thick basal sclerites ( Fig. 24 : bsc) at the base of the basal portions of the struts.

Description of female genitalia. Segment VIII. Tergite VIII ( Fig. 31 : t 8) short, transversal, nearly horizontal, lacking spiracles; with 1+1 obtuse longitudinal ridges, the broad, obliquely declivous portions laterad of these ridges are possibly homologous with laterotergites VIII. Valvifer VIII (= first valvifer, first gonocoxa) ( Figs. 29, 31–32 , 35, 37–39 : vf 8) large, plate-like. Valvula VIII (= first valvula, first gonapophysis) ( Figs. 31–32 , 35, 37–39 : va 8): external portion narrow, separated from the ipsilateral valvifer VIII by a narrow suture, its median margin membranous; median portion membranous, provided with dense, long pilosity. Segment IX. Valvifer IX (= second valvifer, second gonocoxa) ( Fig. 36–37 : vf 9) rod-like, articulated with the ipsilateral valvula IX (= second valvula, second gonapophysis) ( Figs. 36–37 : va 9), latter greatly membranous, median surface with dense, long pilosity, lateral wall provided with a plate-like, elongate, apically truncate sclerite. Gonoplac (= styloid, = ‘valvula 3’ of authors; the individualized apical portion of valvifer IX, cf. SNODGRASS 1935, SCUDDER 1957, ŠTYS 1959) ( Figs. 29, 31–32 , 36–37, 39–40 : gpc) thick, sclerotized dorsally, apically rounded with a small marginal tooth projecting ventrad, greatly membranous ventrally; connected to its contralateral counterpart by a membrane except of apical portion. Ectodermal genital tracts. Gynatrium (= vagina) ( Fig. 33 : gy) sheath-like, broad, abruptly narrowed apically; extreme apex of gynatrium with 1+1 simple, thread-like, apically strongly tortuous pseudospermathecae (= lateral spermathecae) ( Fig. 33 : psth); spermatheca (= ‘vermiform gland’) (cf. DAVIS 1966, WYGODZINSKY 1966) ( Fig. 33 : sth; Fig. 34 ) simple, tubular, basal half thicker, with cross-striate wall, apical half thin, with simple wall; a common oviduct ( Fig. 33 : cod) broad, its short basal portion strongly narrowed.

Distribution. This is a widely distributed species occurring all over the continental South and Southeast Asia. It is recorded as new to Vietnam. The southeastern border of its area should be clarified as we have seen specimens from Sumatra probably belonging to this species. — Pakistan ( DISPONS 1965); India: Assam: Khasi Mts. ( DISTANT 1904), Meghalaya: ‘Tura’ ( PAIVA 1919), ‘Sikhim’ [= Sikkim] ( DISTANT 1904), West Bengal: Darjeeling!; Sri Lanka ( HSIAO & REN 1981); Bangladesh: Sylhet ( MILLER 1956); Myanmar ( Burma): ‘N. Khasia’ ( HOFFMANN 1944); Laos: Luang Prabang Prov., ‘Lak Mune’ ( HOFFMANN 1944); Vietnam: Bắc Kạn Prov.!, Hòa Bình Prov.!; China: Hainan!; Taiwan!.

Biology. This species is generally rare in collections, however, apparently it is relatively frequent in Taiwan. Specimens in Taiwan and Vietnam were observed to hide in tree holes, bark crevices, or on the bark of various trees (J. F. Tsai, pers. observ.; R. J. Chen, G. Csorba, pers. comm.). A specimen was observed to feed on a female of Aegus laevicollis formosae Bates, 1866 ( Coleoptera : Lucanidae ) (J. F. Tsai, pers. observ.).

Synonymy. Centrocnemis formosana was described by MATSUMURA (1913) based on a single female holotype from Taiwan; subsequently the species occurring in Taiwan was cited under this name in the literature. The species was transferred to Neocentrocnemis by MILLER (1956). All material from Taiwan seen by us was conspecific, and the specimens fully agree with the original description and illustrations, and the several redescriptions and illustrations ( MATSUMURA 1930, 1931; ESAKI 1932 [as deyrollei ]; MILLER 1956; HSIAO & REN 1981; ISHIKAWA 2005) of N. formosana . The lectotype (a female) of N. stali and several additional specimens of the same appearance collected in Vietnam, China: Hainan Island, and Taiwan were examined. Virtually no external difference could be observed among them. Specimens from Taiwan have usually conspicuously short marginal processes of the abdomen ( Fig. 17 ), whilst specimens from the mainland of Asia can have similar, much longer ( Fig. 18 ), or intermediate spines; these differences are of no diagnostic value. Similarly, the 1+1 short transverse carinae of the posterior lobe of the pronotum, used as a diagnostic character for separating the two species by MILLER (1956: 246) and HSIAO & REN (1981: 414), show strong intraspecific variability even among long series collected at the same locality, and certainly are of no taxonomic value. The genitalia of several males from Taiwan ( Figs. 8–15 , 22–28 ) and two males from northern Vietnam ( Figs. 19–21 ) were examined and compared. Specimens from Taiwan showed only small intraspecific variability; the phallus of one of the males from Vietnam was identical with specimens from Taiwan, the other was slightly different in the shape of the sclerotized elements of the wall of the endosoma ( Fig. 21 ). Females from Taiwan and the Asian mainland are indistinguishable from each other and the female lectotype. As a consequence, N. formosana is hereby sunk as synonym of N. stali . Based on re-examination of the holotype, N. baudoni is also synonymized with the latter.

Examining various Oriental specimens belonging to Neocentrocnemis and a few holotypes of taxa described by N. C. E. Miller, we are convinced that the descriptions and illustrations of this author are insufficient for acceptable and adequate species definition. By a careful study of his species descriptions and identification keys it is clear that his species are based mostly on insignificant and obscure differences, and probably many of them are also synonyms of N. stali . This problem is not examined further now.

BERGROTH (1914: 364) reported Centrocnemis deyrollii Signoret, 1852 from Taiwan: ‘Fuhosho’ (currently Wucheng). Re-examination of a specimen studied by Bergroth, deposited in DEIC (based on photos received from Stephan Blank) concluded that it belongs to N. stali , just like several other specimens collected at the same locality by the same collector which were directly examined by us. ESAKI (1926) synonymized C. formosana with C. deyrollii and later he provided a redescription of the species under the latter name ( ESAKI 1932). As it is clear from the text and habitus illustration of the latter work, this is in fact misidentification of C. stali . No specimens of Centrocnemis Signoret, 1852 were examined during the present study from Taiwan, therefore the genus Centrocnemis and the species C. deyrollii are deleted from the checklist of the Reduviidae of this country.

Figures

Figs. 1–7. Neocentrocnemis spp., type specimens and their labels. 1–3 – N. stali (Reuter, 1881), lectotype; 4–7 – N. baudoni Dispons, holotype.Scales in mm; labels not to scale. (Figs. 1–3: © NRMS, photographed by G. Lindberg; Figs. 4–7: © MNHN, photographed by L. Fauvre; published with permission).

Figs.8-16. Neocentrocnemis stali (Reuter, 1881) (Taiwan), male genitalia. 8 – genital capsule, dorsal view; 9 – same, lateral view (right paramere omitted); 10–11 – superoposterior margin of genital capsule of two different specimens, posteroventral view; 12–13 – right paramere, two different aspects; 14 – phallus with cuplike sclerite, lateral view; 15 – articulatory apparatus; 16 – pedicel and support bridge prolongation, cross-section around middle of pedicel. Lettering: bp = basal plates; cs = cuplike sclerite; ism 9–10 = inter-sternal membrane between segments IX–X; sbc = support bridge complex; sbp = support bridge prolongation. Scale bars in mm.

Figs.17-21. Neocentrocnemis stali (Reuter,1881),diagnostic characters.17 – right margin of abdomen (female,Taiwan); 18 – same (female, Vietnam); 19 – right paramere (Vietnam); 20 – same, another specimen (Vietnam); 21 – apical portion of the phallus, lateral view (same specimen as in Fig. 19). Lettering as in Fig. 24. Scale bars in mm.

Figs. 22–24. Neocentrocnemis stali (Reuter, 1881). 22 – phallus, lateral view; 23 – phallotheca, lateral view; 24 – endosoma, lateral view (phallotheca wall removed). Lettering: aa = articulatory apparatus; ba = basal aula; bsc = basal sclerite; dlp = dorsolateral projection of endosoma; dsc = dorsal sclerite of endosoma; dsp = dorsal sclerotized plate of phallotheca; lsc = lateral sclerite of endosoma; ped = pedicel; phs = phallosoma; phth = phallotheca; stra = apical portion of strut; strb = basal portion of strut; vsp = ventral sclerotized plate of phallotheca. Scale bars in mm.

Figs. 25–28. Neocentrocnemis stali (Reuter, 1881), apical portion of the phallus. 25 – dorsal view; 26 – ventral view; 27 – dorsal view, phallotheca removed; 28 – ventral view, phallotheca removed. Lettering as in Figs. 22–24, furthermore ensw = wall of endosoma; phm = phallotheca mouth; phthw = wall of phallotheca. Scale bars in mm.

Figs. 29–34. Neocentrocnemis stali (Reuter, 1881), female genitalia. 29 – apex of abdomen, posterior view; 30 – same, lateral view; 31 – same, dorsal view, ovipositor slightly opened; 32 – same, ventral view; 33 – gynatrium; 34 – spermatheca. Lettering: cod = common oviduct; gpc = gonoplac; gy = gynatrium; psth = pseudospermatheca; sth = spermatheca; t9 = tergite IX; va8 = valvula VIII; vf 8 = valvifer VIII. Scale bars in mm.

Figs. 35–40. Neocentrocnemis stali (Reuter, 1881), female genitalia. 35 – apex of abdomen, lateral view, ovipositor slightly opened; 36 – same, right valvifer VIII and valvula VIII removed; 37 – same, posterior view, ovipositor opened, left gonoplac removed; 38 – right valvifer VIII and valvula VIII, inner surface; 39 – left half of ovipositor, ventral view, schematical; 40 – ovipositor in ventral view, valvifer VIII and valvula VIII removed. Lettering as in Figs. 29–32, furthermore: va9 = valvula IX; vf9 = valvifer IX; X = segment X. Scale bar in mm.