Zootaxa 4382 (2): 299–320 http://www.mapress.com/j/zt/ Copyright © 2018 Magnolia Press ISSN 1175-5326 (print edition) Article ZOOTAXA ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4382.2.5 http://zoobank.org/urn:lsid:zoobank.org:pub:3F5277F2-9FA5-4004-8BFF-4A1627AC6C07 The genus Alphocoris in the Indomalaya (Hemiptera: Heteroptera: Scutelleridae) DÁVID RÉDEI1,2, JING-FU TSAI3 & ZDENĚK JINDRA4 1 Institute of Entomology, College of Life Sciences, Nankai University, Weijin Road 94, 300071 Tianjin, China. E-mail: david.redei@gmail.com 2 Department of Zoology, Hungarian Natural History Museum, H-1088 Budapest, Baross u. 13, Hungary 3 Department of Biology, National Museum of Natural Science, 1 Kuan-Chian Rd., Taichung, Taiwan. E-mail: jingfu.tsai@gmail.com 4 Department of Plant Protection, Faculty of Agrobiology, Food and Natural Resources, Czech University of Agriculture, CZ-165 21 Praha 6—Suchdol, Czech Republic. E-mail: palomena@seznam.cz Abstract The Indomalayan species of the genus Alphocoris Germar, 1839 (Hemiptera: Heteroptera: Scutelleridae: Odontotarsinae: Odontotarsini) are revised. Three species, A. caudatus Rédei, Tsai & Jindra, sp. nov. (India: Maharashtra), A. naso Rédei & Tsai, sp. nov. (India: Goa), and A. asper Rédei, Tsai & Jindra, sp. nov. (India: Goa and Maharashtra), are recognized. The type material of A. lixoides Germar, 1839 (type locality: Senegal) is documented; previous records of this species from India and Pakistan are considered as based on misidentifications, the species is restricted to the Afrotropical Region. Key words: Hemiptera, Scutelleridae, taxonomy, new species, South Asia, India, Afrotropical Region Introduction The genus Alphocoris Germar, 1939 (Hemiptera: Heteroptera: Scutelleridae: Odontotarsinae: Odontotarsini) currently contains 9 described species distributed in the Afrotropical Region, the West Palaearctic (North Africa and the Middle East) and the Indomalaya (Pakistan, western India) (Schouteden 1903, 1904, Kirkaldy 1909, Bergevin 1931, Linnavuori 1975, 1984, Hoberlandt 1997, Göllner-Scheiding 2006). Several of the described species have never been reexamined since their original description and are in need of a taxonomic revision. The Afrotropical fauna was monographed by Schouteden (1903); an additional new species was described by Linnavuori (1975). The Palaearctic fauna, particularly the species of the Middle East, has remained greatly unrevised, described species were treated and new species were described by Bergevin (1931), Vidal (1949), Stichel (1960b), Putshkov (1965), Linnavuori (1984), Hoberlandt (1997), and others. A single species, A. lixoides Germar, 1839 (type locality: Senegal) was recorded from India by Dallas (1851), from Pakistan by Ahmad & Mustaq (1977), and from both countries by various subsequent authors. The present paper offers a revision of the Indomalayan species of Alphocoris. Based on the reexamination of the type material of A. lixoides it is demonstrated that previous records of this species from South Asia are erroneous. Three new species, all from India, are described. This paper forms a part of the ongoing revision of Indomalayan Scutelleridae by the first and second authors (Tsai & Rédei 2009, 2010, Rédei & Tsai 2014, 2016). Material and methods External structures and genitalia were examined using stereoscopic microscopes (Leica M205C, Zeiss Discovery V8), drawings were made with the aid of a camera lucida. Genitalia of both sexes were dissected after careful heating in hypertonic KOH solution, stained with Chlorazole Black E if necessary. Measurements were taken using a micrometer eyepiece. Digital photographs were taken with a Nikon D90 camera equipped with AF-S Micro Nikkor 60mm f/2.8G ED macro lens. Accepted by D. Rider: 11 Dec. 2017; published: 21 Feb. 2018 299 Label data are given in verbatim transcription. Backslash (\) indicates separation between lines on the label, comments on label data are provided in square brackets. Printed text (pr) is not indicated except if the preceding or following text is handwritten (hw). The sign ‘>’ between the original and final depository indicates specimen transfer between two collections. Specimens examined by us are marked by exclamatory point (!); doubtful literature records by question mark (?). Morphological terminology follows Tsai et al. (2011), Tsai & Rédei (2015), and Rédei (2017). Abbreviations for depositories: BMNH HNHM NMNS NMPC ZJPC ZMHB Natural History Museum, London, United Kingdom Hungarian Natural History Museum, Budapest, Hungary National Museum of Natural Science, Taichung, Taiwan National Museum, Prague, Czech Republic Zdeněk Jindra Personal Collection, currently deposited at the Department of Plant Protection, Czech University of Agriculture, Prague, Czech Republic Naturkundemuseum Berlin (former Zoologisches Museum der Humboldt-Universität), Germany Taxonomy Alphocoris Germar, 1839 Alphocoris Germar, 1839: 8, 58. Type species by subsequent designation (Distant 1902: 67): Alphocoris lixoides Germar, 1839. Sphenaspis Jakovlev, 1877: 72. Type species by monotypy: Sphenaspis curculionoides Jakovlev, 1877. Synonymized by Puton (1886: 6). Alphocoris: Herrich-Schäffer (1839: 20) (listed), Agassiz (1843: 2) (listed), Amyot & Serville (1843: 43) (redescription), Dallas (1851: 5, 41) (in key, listed), Herrich-Schäffer (1851: 293, 333) (in key, listed), Herrich-Schäffer (1853: 8) (listed), Dohrn (1859: 4) (catalogue), Stål (1865: 35, 60) (in key, redescription, fauna of Africa), Mayr (1866: 13) (in key), Stål (1866: 257) (listed), Walker (1867: 59) (listed), Stål (1873: 26) (in key, synopsis), Puton (1875: 7) (catalogue, Europe), Puton (1886: 6) (catalogue, Palaearctic), Atkinson (1887: 186) (redescription, fauna of India), Lethierry & Severin (1893: 40) (catalogue), Puton (1899: 6) (catalogue, Palaearctic), Distant (1902: 67) (distribution, fauna of British India, Ceylon and Burma), Kirkaldy (1903a: 214) (listed), Kirkaldy (1903b: 233) (listed), Schouteden (1903: 80) (redescription, Afrotropical fauna), Schouteden (1904: 71, 82) (in key, redescription, catalogue), Oshanin (1906: 33) (catalogue, Palaearctic), Kirkaldy (1909: XXXIV, 263, 268) (type species, distribution, catalogue), Schouteden (1909: 35) (fauna of Belgian Congo), Oshanin (1910: 10) (catalogue, Palaearctic), Oshanin (1912: 4) (catalogue, Palaearctic), Hesse (1925: 8) (fauna of Southwest Africa), Vidal (1949: 54, 61) (in key, fauna of the Mediterranean), Villiers (1952: 59) (diagnosis, distribution, selected African species), Priesner & Alfieri (1953: 11) (in key), Stichel (1960a: 444) (in key), Stichel (1960b: 451) (fauna of West Palaearctic), Stichel (1961: 731) (catalogue, Palaearctic), Stichel (1962: 211) (catalogue, Palaearctic), Putshkov (1965: 81, 82) (in key, fauna of Central Asia), Fuente (1973: 245) (in key), Ahmad et al. (1979: 27, 31, 32) (in key, listed), Ahmad (1980: 124) (distribution in Pakistan), Cassis & Vanags (2006: 290) (listed), Gadalla & Fadl (2000: 143, 145) (in key, redescription, fauna of Egypt), Göllner-Scheiding (2006: 203) (catalogue, Palaearctic), Czaja (2007: 311, 312) (hind wing venation), Tsai et al. (2011: 60, 63) (distribution), Czaja (2012: 61) (wing coupling structures), Czaja (2013a: 354, 363) (fore wing microtrichia), Czaja (2013b: 106, 117) (fore wing), Parveen & Gaur (2015: 169, 170, 174) (listed, in key, diagnosis), Czaja (2016a: 373) (diagnostic characters). Diagnosis. Conspicuously elongate, subcylindrical species differing from all other genera of Odontotarsini in their densely tomentose integument and the strongly broadened anterior portion of clypeus more or less protruding anteriad of mandibular plates. The monotypic Nearctic genus Fokkeria Schouteden, 1904, and the monotypic Afrotropical genus Rhinolaetia Schouteden, 1965, are both somewhat similar to Alphocoris in general habitus and the similarly tomentose integument, but members of both of these genera have a conspicuously different clypeus (in Fokkeria it is not conspicuously broadened anteriorly, in Rhinolaetia it is strongly tapering anteriad) and a posteriorly broadly rounded scutellum. Redescription. Body small to medium (about 6–12 mm), elongate, narrow, subcylindrical, strongly convex dorsally, less convex ventrally; dorsal outline of body almost contiguous between base of pronotum and base of scutellum in lateral view. Body surface and vestiture. Whole surface of body and legs covered by dense, tomentose 300 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. pubescence; antennae with very fine, inconspicuous, semierect pilosity; dorsum and venter distinctly punctured, punctures occasionally large and coarse. Head strongly convex, more or less declivous anteriorly; apical portion of clypeus far surpassing mandibular plates, conspicuously broadened and tumescent; lateral margin of mandibular plates obtuse or with a distinct carina; buccula high, anteriorly obtusely truncate (as in Figs. 20, 23, 26) or produced into an elongate projection; ocelli small, situated close to level of posterior margin of eyes. Antenna short, five-segmented, apex of scape far remote from apex of mandibular plates, basipedicellite at least slightly longer than distipedicellite; basi- and distiflagellum weakly flattened. Labium reaching between middle to hind coxae, segment II distinctly longer than combined length of segments III–IV. Pronotum rather elongate, less than two times as long as its greatest width; anterior collar indistinguishable; calli weakly demarcated; lateral margin straight to weakly convex, obtuse; humeral angle rounded, not produced; posterolateral angle widely rounded; posterior margin slightly convex, narrower than base of scutellum. Scutellum elongate, lateral margins of anterior portion subparallel or somewhat convex, posterior portion strongly tapering, apex reaching or surpassing apex of abdomen, rounded, truncate or excised; concealing exocorium except of its extreme base and whole of endocorium; basal tumescence indistinguishable. Thoracic pleura and sterna. Anterior explanate part of proepisternum slightly surpassing base of head, rounded; metathoracic scent gland ostiole and associated efferent structures lacking; thoracic sternites deeply furrowed mesally. Legs short, femora thick, tibiae cylindrical. Pregenital abdomen elongate, more or less tapering towards apex especially in males; lateral margin obtuse; posterolateral angles of ventrites unarmed or produced into minute denticles; ventrite VII produced posteriad, completely concealing genital capsule from below; openings of first dorso-abdominal scent gland (= DAG 1) situated about halfway between tergite IV and openings of DAG 2 or closer lateral margin of tergite IV than to opening of DAG 2; stridulatory vittae and androconial glands absent. Male genitalia. Genital capsule subquadrate in dorsal view; posterior opening directed dorsad, completely concealed by apical portion of scutellum; cuplike sclerite with a pair of strong suspensory apodemes and sublateral processes, but lacking median projection. Paramere fully sclerotized, with a stick-like stem and a finger-shaped crown. Phallus symmetrical, basal aula wide, support bridge prolongation lacking, phallotheca weakly sclerotized, with a single median thecal process apically; conjunctiva loosely eversible, with a flexible, thickened, pad-like invagination distally, tightly enclosing endophallic reservoir; conjunctiva with a single pair of processes (cp-II), subdivided into two or three branches, dorsal branch (cp-II1) membraneous, wrinkled, ventral branches (cp-II2, cp-II3) sclerotized, provided with numerous minute denticles; aedeagus membranous, receiving heavily sclerotized endophallic duct of single lumen, distal part narrowed and slightly invaginated; ductus seminis connecting to endophallic reservoir basoventrally. Female genitalia. Terminalia posterior, almost vertical; gonangulum indistinct; valvulae with sclerotized rami or completely membranous; gynatrium simple, with a pair of anterolateral pouches, with or without fecundation sclerite; proximal portion of spermatheca (probably homologous with proximal duct) broad, saccular, thin-walled, dilation thick-walled but without tubular projections or inner lamellae; distal duct very short; apical receptacle with broad, curved base and globose apex. Bionomics. Species of Alphocoris usually occur in arid, steppic habitats and they probably feed on grasses. Diversity and distribution. The genus contains 9 described species; by describing three additional species this number is increased to 12 now (Table 1). Nevertheless, several of the previously described species are in need of revision. All species described by earlier authors are restricted to the Afrotropical Region and the West Palaearctic (North Africa and the Middle East); the three new species occur in the western part of India. Key to the species of Alphocoris occurring in the Indomalaya 1(2) 2(1) Distance from anterior margin of eye to apex of mandibular plate about 2.75 times as long as length of eye in dorsal view, apical, protruding portion of clypeus shorter and broader (Figs. 19–20); dorsum with colourless, shallow punctures intermixed with a few dark punctures which never form dark longitudinal vittae (Figs. 1, 4); posterior, produced portion of scutellum more strongly tapering, apically narrowly and almost transversely truncate between a pair of minute lateral denticles (♂) or narrowly angularly excised (♀) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. caudatus Rédei, Tsai & Jindra, sp. nov. Distance from anterior margin of eye to apex of mandibular plate at most about 2 times as long as length of eye in dorsal view, apical, protruding portion of clypeus relatively long and narrow (Figs. 22–23, 25–26); dorsum with several densely distributed, coarse, black punctures forming more or less distinct dark longitudinal vittae (Figs. 7, 10, 13); posterior, produced portion of THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 301 3(4) 4(3) scutellum short, apex different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Converging lateral margins of posterior third of scutellum straight, terminal part relatively broadly, angularly excised apically (Figs. 7, 10); posterolateral angles of abdominal ventrites III–IV not produced . . . . . . . . . . . . . A. naso Rédei & Tsai, sp. nov. Lateral margins of posterior third of scutellum concave, apex relatively broad, angularly transversely truncate (Figs. 13, 16); posterolateral angles of abdominal ventrites III–IV each produced into a minute but distinct spine exposed laterad of scutellum in dorsal aspect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. asper Rédei, Tsai & Jindra, sp. nov. Alphocoris caudatus Rédei, Tsai & Jindra, sp. nov. Figs. 1–6, 19–21, 28–29, 34–36, 44, 51–54, 59–62 Type material. Holotype: ♂, “INDIA occ., Maharashtra state \ 7–11 October 2005 \ Mulshi env., 40 km W of Pune \ F. Kantner lgt.”; mounted on card, genital capsule removed and glued to the same card separately but otherwise intact (ZJPC). Paratypes: same label as in holotype (3 ♂♂ 1 ♀ ZJPC, 2 ♂♂ HNHM, 1 ♂ NMNS); “India W, 30.ix.-2.x.2005 \ Maharastra state \ 40 km W of Pune \ Mulshi env., J. BEZDĚK lgt.” (1 ♀ NMPC) Diagnosis. Readily recognized among all congeners by its rather uniformly coloured dorsum lacking dark longitudinal vittae (Figs. 1, 4), and the posterior portion of the scutellum and abdomen being strongly constricted in both sexes (Figs. 1–6) but particularly in males (Fig. 21). Description. Colour, integument and vestiture. Dorsum rather uniformly stramineous with contrasting black markings on head but without dark longitudinal vittae on scutellum, subshining; head stramineous to yellow, anterior part of clypeus, a longitudinal vitta on mandibular plate laterally anteriad of eyes, mesal margin of mandibular plates together with adjacent lateral margins of clypeus continued in a pair of broad submedian vittae extending to base of head, postocular portion laterally, and whole ventral surface of head black; yellow areas of head with a few scattered, colourless, inconspicuous punctures, black areas densely and coarsely punctured; scape and pedicel of antenna dark yellowish or brown, flagellum rather dark brown, but occasionally whole antenna suffused with dark brown except of base of scape and basipedicellite; labium stramineous, segments III and IV suffused with dark brown; pronotum, scutellum, and exposed portions of fore wings rather uniformly stramineous, with small, not conspicuous dark patches formed by confluent dark punctures as follows: an irregular longitudinal streak laterad and a small dot immediately mesad of cicatrices, and a pair of submedian, longitudinal, anteriorly indistinctly delimited vittae on posterior fourth to fifth of scutellum; pronotum and scutellum densely and rather uniformly and regularly punctured, but with a pair of unpunctured, slightly elevated, somewhat callose surfaces between cicatrices and anterior margin of pronotum, furthermore midline of scutellum and a pair of submedian longitudinal bands in posterior third of scutellum also unpunctured, with very fine and shallow punctures between and around them; middle and posterior portions of scutellum also frequently with a few irregularly scattered, very large, black punctures; prothoracic hypomeron (= deflexed marginal portion of pronotum) stramineous, thoracic pleurites greatly black but paler laterally, coarsely punctured; coxae and trochanters dark brown, suffused with black, femora and tibiae black, tarsi brown, suffused with black towards apex; ground colour of abdominal venter stramineous but densely covered by coarse, greatly confluent black punctures therefore pale ground colour only visible as a broad vitta along lateral margin, a median vitta usually interrupted at intersegmental sutures, and pale interspaces between punctures on disk; outer laterotergites stramineous, broadly suffused with black mesally, inner laterotergites and mediotergites black. Dense, tomentose pubescence of dorsum was lacking in the examined material (probably it was rubbed off), but its remnants were present on some specimens particularly on anterior portion of head, ventral surface of body, and legs; scape and basipedicellite nearly glabrous, distipedicellite with a few scattered, fine hairs, flagellum with fine, dense, adpressed to semierect pilosity. Structure. Body elongate, about 2.7 times as long as its humeral width (♂, ♀), more strongly constricted posteriorly than A. naso sp. nov. and A. asper sp. nov. therefore appearing more gracile. Head (Figs. 19–20) elongate subtriangular, about as wide as long, width across eyes about 1.35 (♂) / 1.25–1.3 (♀) times as long as interocular distance; eyes relatively small, semiglobose, strongly protruding laterally; distance from anterior margin of eye to level of apex of mandibular plate measured along longitudinal axis of body about 2.75 times as long as length of eye in dorsal view; apical, protruding portion of clypeus relatively short, less than one third as long as distance from anterior margin of eye to level of apex of mandibular plate measured along longitudinal axis of body; mandibular plates with a weak, obtuse longitudinal carina laterally anteriad of ventral margin of eye; apex of buccula closely approaching apex of clypeus. Labium reaching bases of hind coxae. Thorax. Pronotum 1.4–1.45 302 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. times as broad as its median length; lateral margins smooth in their whole length, straight anteriorly, then gradually transform to broadly rounded humeri. Scutellum 1.55–1.65 times as long as its greatest width, posterior portion strongly tapering with converging lateral margins straight, apex conspicuously narrow (distance between paired protruding terminal angles shorter than width of anterior portion of clypeus) and considerably surpassing apex of abdomen, terminal part apically narrowly and almost transversely truncate between a pair of minute lateral denticles (♂) or narrowly angularly excised (♀). Pregenital abdomen more strongly tapering apically than in A. naso sp. nov. and A. asper sp. nov., ventrite VII of male conspicuously elongate (Fig. 21), posterolateral angles of abdominal ventrites III–IV not produced. External male genitalia. Genital capsule (Figs. 28–29, 34) highly elongate oval in posterior view (Fig. 34), ventral rim broadly convex in dorsal view (Fig. 28), posterior portion relatively strongly produced (Fig. 29). Paramere as in Figs. 35–36. Phallus (Figs. 44, 51): second pair of conjunctival processes (cp-II) subdivided into 2+2 branches, dorsal branch (Fig. 44: cp-II1) membranous, greatly enlarged, surpassing apex of aedeagus; ventral branch (Fig. 44: cp-II2) sclerotized, fin-shaped, broad; aedeagus weakly dilated subapically, distalmost portion narrow, tubular. External female genitalia. Terminalia as in Figs. 52–53. Ovipositor: dorsomesal margin of laterotergite IX (Figs. 53, 59: lt9) deeply emarginate around postgenital abdomen; valvifer IX (Fig. 59: vf9) with a relatively small, oval median portion prolonged into a pair of elongate, obliquely transverse rod-like arms joining rami of valvulae IX without moveable articulation; valvulae VIII (Figs. 59: va8) and IX well developed, each with distinct sclerotized rami (Fig. 59: ra8), rami of valvulae VIII weakly sclerotized towards its base, distinct, sclerotized gonangulum lacking. Inner ectodermal genital tracts: gynatrium saccular, elongate, provided with a pair of long anterolateral pouches (Fig. 60: algp); ring sclerites could not be traced in the single examined specimen; base of spermatheca associated with a broad, plate-like fecundation sclerite (Fig. 60: fec). Spermatheca (Figs. 61–62): proximal duct (Figs. 61–62: pd) greatly broadened, thin-walled, saccular, joining to gynatrium through a broad opening; dilation (Figs. 61–62: dil) thick-walled, outer surface irregularly wrinkled, capable of invagination into proximal duct as shown in Fig. 64 for A. naso sp. nov.; distal duct (Figs. 61–62: dd) very short and thin, membranous, dorsoventrally flattened; intermediate part (Fig. 54) with a small proximal flange, lacking distal flange, thin-walled portion (flexible zone) occupying its distal two-thirds, septum and fretum could not be traced in the single examined specimen; apical receptacle (Fig. 54) with a cup-shaped, strongly curved proximal portion and a voluminous, globose apical portion. Measurements (in mm; N = 3 ♂♂ / 2 ♀♀). Body length 8.00–8.72 / 8.82–9.10; length of head 1.62–1.75 / 1.75–1.86, width across eyes 1.75–1.76 / 1.81–1.91, interocular distance 1.28–1.32 / 1.43–1.47; lengths of scape 0.37–0.44 / 0.43–0.44 : basipedicellite 0.32–0.39 / 0.39–0.43 : distipedicellite 0.25–0.27 / 0.27–0.29 : basiflagellum 0.43–0.44 / 0.49 : distiflagellum 0.62–0.63 / 0.68–0.69; median length of pronotum 2.08–2.16 / 2.21–2.35, humeral width 2.93–3.04 / 3.23–3.35; median length of scutellum 4.73–5.00 / 5.20–5.39, greatest width 2.88–3.12 / 3.33–3.37. Etymology. The specific epithet caudatus, -a, -um (meaning ‘caudate’, ‘having or provided with a tail’) is an adjective formed from the Latin noun cauda (‘tail’) by adding the suffix -atus; it refers to the posteriorly strongly produced and tapering scutellum. Distribution. Only known from the type locality near Pune in Maharashtra State, western coastal region of India. Alphocoris naso Rédei & Tsai, sp. nov. Figs. 7–12, 22–24, 30–31, 37–40, 45, 47–48, 55–56, 63–64 Type material. Holotype: ♂, “India, Molem, Goa, \ 120 m” [hw], “No.116. netted \ 21.II.1980. \ leg. Topál” [hw]; mounted on card, intact (HNHM). Paratypes: labels as in holotype (1 ♀ NMNS); “INDIA Molem, \ Goa, 120m” [pr], “No.116. netted \ 21.II.1980. \ leg. Topál” [hw] (1 ♂ 2 ♀♀ HNHM). Diagnosis. Similar to A. asper sp. nov. in size and habitus, but it can be distinguished from the latter species by the less coarse punctation of the dorsum (Figs. 7, 10) and the straight lateral margins of the posterior third of scutellum (Fig. 7). The male genitalia (Figs. 30–31, 37–40, 45, 47–48) and female terminalia (Figs. 55–56) offer several diagnostic characters for separation of the two species. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 303 Description. Colour, integument and vestiture. Ground colour of dorsum stramineous with contrasting black markings on head and with more or less distinct dark longitudinal vittae on scutellum formed by dark punctures, dull to weakly shining; head stramineous to yellow, with black markings on anterior and lateral part of clypeus, margins of mandibular plate, submedially between eyes, postocular portion, and ventral surface of head more or less as in A. caudatus; antenna rather uniformly stramineous; labium stramineous, segments III and IV suffused with dark brown; pronotum, scutellum, and exposed portions of fore wings stramineous, with two pairs of more or less distinct, broad, longitudinal bands on pronotum as well as a pair of discal and a pair of marginal bands on scutellum formed by dark but not confluent punctures, punctures forming these bands are confluent and form dark patches immediately mesad and laterad of cicatrices on pronotum and bilaterally before apex of scutellum; dark punctures forming above described longitudinal bands large, rather coarse, greatly confluent in male, smaller and isolated in female; paler interspaces between dark longitudinal bands more finely and sparsely punctured; midline of scutellum unpunctured and surrounded by several large, coarse, black punctures at two sides (♂) or finely punctured with a few scattered larger and coarser punctures (♀); prothoracic hypomeron and thoracic pleurites apparently as in A. caudatus sp. nov., but could not be studied in detail due to dense, tomentose pubescence; legs black, tarsi stramineous; abdominal venter as in A. caudatus sp. nov. but dark colour surrounding punctures more extensive and confluent therefore median vitta irregular and indistinct, and disk only with a few pale interspaces between punctures; outer laterotergites stramineous, broadly suffused with black mesally, inner laterotergites and mediotergites black. Dense, tomentose pubescence of dorsum was greatly rubbed off from the specimens examined but its remnants were present on all specimens dorsally and ventrally; scape and basipedicellite nearly glabrous, distipedicellite with scattered, fine hairs, flagellum with fine, dense, adpressed to semierect pilosity. Structure. Body elongate, about 2.6 times as long as its humeral width (♂, ♀), less strongly constricted posteriorly than in A. caudatus sp. nov. Head (Figs. 22–23) elongate subtriangular, about as wide as long, width across eyes about 1.45 (♂, ♀) times as long as interocular distance; eyes larger than in A. caudatus sp. nov., semiglobose, strongly protruding laterally; distance from anterior margin of eye to level of apex of mandibular plate measured along longitudinal axis of body about 1.65 times as long as length of eye in dorsal view; apical, protruding portion of clypeus long, about half as long as distance from anterior margin of eye to level of apex of mandibular plate measured along longitudinal axis of body; mandibular plates with a strong longitudinal carina laterally anteriad of ventral margin of eye, visible in dorsal view (Fig. 22: arrow). Labium surpassing middle or reaching bases of hind coxae. Thorax. Pronotum 1.45–1.5 times as broad as its median length; lateral margins smooth, very slightly concave posteriad of level of calli particularly in female. Scutellum 1.45–1.5 times as long as its greatest width, posterior portion less strongly tapering than in A. caudatus sp. nov., with converging lateral margins straight, distance between paired protruding terminal angles longer than width of anterior portion of clypeus, slightly surpassing apex of abdomen, terminal part angularly excised apically (♂, ♀). Pregenital abdomen broader, less strongly tapering apically than in A. caudatus sp. nov., ventrite VII of male relatively short and broad (Fig. 24), posterolateral angles of ventrites III–IV not produced. External male genitalia. Genital capsule (Figs. 30–31, 37–38) broadly oval in posterior view (Fig. 37), ventral rim very slightly convex, nearly straight in dorsal view (Fig. 30), posterior portion weakly produced (Fig. 31). Paramere as in Figs. 39–40. Phallus (Figs. 45, 47–48) with second pair of conjunctival processes (cp-II) subdivided into 3+3 branches, dorsal branch (Fig. 47: cp-II1) membranous, relatively short, ventral branches (Figs. 45, 47: cp-II2, cp-II3) sclerotized, elongate, apex slightly dilated, club-like; conjunctival pad (Fig. 47: cpd) rounded around endophallic reservoir; aedeagus (Fig. 45: aed) strongly dilated subapically, distalmost portion short, relatively broad, sclerotized. External female genitalia. Terminalia as in Figs. 55–56. Ovipositor: dorsomesal margin of laterotergite IX obliquely truncate around postgenital abdomen; valvifer IX with a relatively small, oval median portion prolonged into a pair of short, obliquely transverse rod-like arms terminating in a rounded tip; valvulae VIII greatly membranous, apical lobe small, rami VIII only recognizable in their distal portion but weakly sclerotized even there, proximal portion completely desclerotized and indiscernible; valvulae IX (Figs. 63: va9) completely membranous, lacking traceable sclerotized rami; distinct, sclerotized gonangulum lacking. Inner ectodermal genital tracts (Fig. 64): gynatrium similar to the condition found in A. caudatus sp. nov. but anterolateral pouches shorter; ring sclerites could not be traced in the single examined specimen; distinct fecundation sclerite lacking, base of spermathecal duct associated with a fecundation channel traceable as a membranous fold. Spermatheca highly similar to the condition found in A. caudatus sp. nov. 304 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. FIGURES 1–6. Alphocoris caudatus sp. nov. 1–3, a male paratype; 4–6, a female paratype. Dorsal (1, 4), lateral (2, 5) and ventral (3–6) views. Scale in mm. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 305 FIGURES 7–12. Alphocoris naso sp. nov. 7–9, male holotype; 10–12, a female paratype. Dorsal (7, 10), lateral (8, 11) and ventral (9–12) views. Scale in mm. 306 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. Measurements (in mm; N = 2 ♂♂ / 2 ♀♀). Body length 7.15–7.30 / 8.25–7.90; length of head 1.72–1.75 / 1.75–1.83, width across eyes 1.72–1.78 / 1.80–1.83, interocular distance 1.18–1.20 / 1.23–1.25; lengths of scape 0.39–0.42 / 0.42–0.44 : basipedicellite 0.29–0.31 / 0.29–0.31 : distipedicellite 0.20–0.27 / 0.25–0.27 : basiflagellum 0.39–0.44 / 0.40–0.43 : distiflagellum 0.59–0.65 / 0.56–0.60; median length of pronotum 1.84–1.86 / 2.05–2.12, humeral width 2.74–2.80 / 3.05–3.12; median length of scutellum 4.12–4.16 / 4.30–4.60, greatest width 2.74–2.76 / 3.00–3.10. Etymology. The specific epithet naso is the ablative singular of the Latin noun nasus ‘nose’, meaning ‘the one with the nose’ (i.e. “bignose”), referring to the anteriorly greatly protruding clypeus; to be treated as noun in apposition, ending not to be changed. Distribution. Only known from the type locality in Goa State, western coastal region of India. Alphocoris asper Rédei, Tsai & Jindra, sp. nov. Figs. 13–18, 25–27, 32–33, 41–43, 46, 49–50, 57–58 Alphocoris lixoides (non Germar, 1839): Dallas (1851: 41) and subsequent authors. Misidentification. Alphocoris lixoides (misidentification): Dallas (1851: 41) (record), Walker (1867: 59) (record), Atkinson (1884: 166) (distribution), Atkinson (1887: 187) (redescription, distribution), Distant (1902: 67) (redescription, habitus, figures, distribution), Maxwell-Lefroy (1909: 673) (habitus). Type material. Holotype: ♂, “INDIA OR. \ RAMANDOROG \ KATONA 1919”; mounted on card, intact (HNHM). Paratypes: same label as in holotype (1 ♂ HNHM); “INDIA occ., Maharashtra state \ 3–6 October 2005, Wai env. \ 70 km S of Pune, F. Kantner lgt.” (1 ♂ 1 ♀ ZJPC). Diagnosis. Similar to A. naso sp. nov. in size and habitus, but it can be distinguished from the latter species by the conspicuously coarse punctation of the dorsum and the concave lateral margins of the posterior third of scutellum (Fig. 13). The male genitalia (Figs. 29–30, 38–40, 43, 51–52) and female terminalia (Figs. 57–58) offer several diagnostic characters for separation of the two species. Description. Colour, integument and vestiture as in A. naso sp. nov. but with the following differences: punctures of dorsum conspicuously coarser and deeper; midline of dorsum lacking distinct unpunctured, smooth band; dark longitudinal bands on pronotum and scutellum formed by dark punctures obscure, rather indistinctly delimited; abdominal venter almost continuously dark except of lateral margins and a narrow median vitta interrupted at intersegmental sutures. Structure. Body elongate, about 2.7 times as long as its humeral width, similar in shape to A. naso sp. nov. but less strongly constricted posteriorly. Head (Figs. 25–26) similar to that of A. naso sp. nov., about as wide as long, width across eyes about 1.55 times as long as interocular distance; distance from anterior margin of eye to level of apex of mandibular plate measured along longitudinal axis of body about 2.0 times as long as length of eye in dorsal view; apical, protruding portion of clypeus about half as long as distance from anterior margin of eye to level of apex of mandibular plate measured along longitudinal axis of body; mandibular plates with a strong longitudinal carina laterally anteriad of ventral margin of eye, visible in dorsal view. Labium slightly surpassing apices of hind coxae. Thorax. Pronotum about 1.5 times as broad as its median length; lateral margins very finely granulate in their anterior halves, slightly concave posteriad of level of calli. Scutellum about 1.5 times as long as its greatest width, posterior portion less strongly tapering than in A. naso sp. nov., with converging lateral margins distinctly concave, apex broad, transversely truncate, distance between paired protruding terminal angles more than two times as long as width of anterior portion of clypeus, not surpassing apex of abdomen (occasionally leaving extreme apex of abdomen exposed in dorsal view). Pregenital abdomen broader, less strongly tapering apically than in A. caudatus sp. nov., ventrite VII of male relatively short and broad (Fig. 27), posterolateral angles of ventrites III–IV each produced into a minute but distinct spine exposed laterad of scutellum in dorsal aspect. External male genitalia. Genital capsule (Figs. 32–33, 41) with ventral rim distinctly convex in dorsal view (Fig. 32), posterior portion weakly produced (Fig. 33). Paramere as in Figs. 42–43. Phallus (Figs. 46, 49–50) with second pair of conjunctival processes (cp-II) subdivided into 3+3 branches, dorsal branch (cp-II1) membranous, relatively short, ventral branches (cp-II2, cp-II3) sclerotized, elongate, apex slightly dilated, club-like; conjunctival pad (Fig. 49: cpd) much broader than in A. naso sp. nov.; aedeagus moderately dilated subapically, distalmost portion elongate, relatively narrow, membranous. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 307 FIGURES 13–18. Alphocoris asper sp. nov. 13–15, male holotype; 16–18, female paratype. Dorsal (13, 16), lateral (14, 17) and ventral (15, 18) views. Scale in mm. External female genitalia. Terminalia as in Figs. 57–58. Ovipositor: dorsomesal margin of laterotergite IX obliquely truncate around postgenital abdomen; valvulae and valvifers of ovipositor much similar to the condition found in A. naso sp. nov.: valvulae VII as in A. naso sp. nov.; valvulae IX membranous, but rami IX can be traced as a very fine, thin, weakly sclerotized and pigmented line along anterior margin of valvulae IX; distinct, 308 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. sclerotized gonangulum lacking. Inner ectodermal genital tracts: gynatrium and spermatheca highly similar to the condition found in A. naso sp. nov. therefore not illustrated. Measurements (in mm; N = 3 ♂♂ / 1 ♀). Body length 7.00–7.74 / 7.60; length of head 1.68–1.72 / 1.69, width across eyes 1.68–1.76 / 1.70, interocular distance 1.08–1.23 / 1.16; lengths of scape 0.32–0.39 / 0.38 : basipedicellite 0.27–0.34 / 0.32 : distipedicellite 0.24–0.25 / 0.26 : basiflagellum 0.39–0.44 / 0.42: distiflagellum 0.55–0.59 / 0.56 ; median length of pronotum 1.70–1.86 / 1.81, humeral width 2.60–2.84 / 2.80; median length of scutellum 3.92–4.41 / 4.13, greatest width 2.56–2.84 / 2.80. Discussion. The species illustrated by Distant (1902: 67, fig. 30) and Maxwell-Lefroy (1909: plate LXXII: fig. 10) as A. lixoides was likely A. asper sp. nov., but its identity cannot be unambiguously elucidated; no specimens from the localities listed by Distant (1902) (Bombay, Khandala) were found in the BMNH where W.L. Distant’s collection is currently deposited. FIGURES 19–17. Alphocoris spp., diagnostic characters. 19–21, A. caudatus sp. nov.; 22–24, A. naso sp. nov.; 25–27, A. asper sp. nov. Head, dorsal view (19, 22, 25); same, lateral view (20, 23, 26); posterior part of abdomen of male with tip of scutellum, ventral view (21, 24, 27). Lettering: scu = scutellum. Scales in mm. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 309 FIGURES 28–33. Alphocoris spp., genital capsules. 28–29, A. caudatus sp. nov.; 30–31, A. naso sp. nov.; 32–33, A. asper sp. nov. Dorsal view, left paramere removed (28, 30, 32); lateral view (29, 31, 33). Scales in mm. Etymology. The specific epithet is the Latin adjective asper, -a, -um (‘rough’, ‘coarse’), given in allusion to the coarsely punctured dorsum. Distribution. Only known from Ramandorog [= Ramdurg], Goa State, and Wai, Maharashtra State, of western India. Previous record of A. lixoides from Maharashtra (Bombay, Khandala) (Distant 1902) possibly at least partly pertain to this species. Alphocoris lixoides Germar, 1839 Figs. 65–72 Alphocoris lixoides Germar, 1839: 59. Syntypes: Senegallia [= Senegal]; ZMHB! Alphocoris lixoides: Amyot & Serville (1843: 44) (diagnosis, distribution), Dohrn (1859: 4) (catalogue, distribution), Stål (1873: 26) (in key, diagnostic characters, distribution), Lethierry & Severin (1893: 40) (catalogue, distribution), Schouteden (1903: 81) (in key, redescription, distribution), Schouteden (1904: 83) (catalogue, distribution), Schouteden (1905: 6) (record), Kirkaldy (1909: 269) (catalogue, distribution), Villiers (1952: 59) (diagnosis, distribution), Schouteden (1963: 397) (listed), Gillon (1972: 286, 296, 304) (in key, redescription and figures of adult and larva), Gillon (1974: 218) (ecology), Linnavuori (1982: 15) (records, distribution, bionomics), Czaja (2007: 308) (figure, wing venation), Czaja (2013b: 102) (figure, fore wing), Czaja (2016a: 366) (listed, habitus). Alphocoris lixoidea [inadvertent error]: Herrich-Schäffer (1853: 8) (listed, distribution). Type material examined. Syntype #1: ♂, ‘Senegal \ Buq’ [blue, hw], ‘Cat. No \ 121’ [hw], ‘Alphocoris \ lixoides 310 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. Germ’ [hw] , ‘Alphocoris \ lixoides \ Germ.*’ [hw], ‘Type’ [red]; pinned, tarsus of left hind leg lacking (ZMHB) (Figs. 65–68). Syntype #2: ♀, ‘Seneg. Dup.’ [green, hw], ‘121’, ‘Zoolog. Museum \ Berlin’, ‘Alphocoris [hw] \ lixoides [hw] \ * Germ. [hw] \ F. Schumacher det. [pr] rev. [hw]’ [‘det.’ crossed out by hw], ‘Alphocoris \ lixoides Germ. \ C.H.C. Lyal 1973 \ LECTOTYPE’ [red, hw], ‘Typus’ [red]; pinned, tarsus of left fore leg lacking (ZMHB) (Figs. 69–72). Remarks. Alphocoris lixoides is the type species of the genus Alphocoris. It was described based on an unspecified number of but more than one specimen from “Senegallia” [= Senegal] (Germar 1839), and subsequently recorded from India (Dallas 1851, Walker 1867, Atkinson 1884, Atkinson 1887, Distant 1902, Maxwell-Lefroy 1909, Parveen et al. 2014) and Pakistan (Ahmad & Mushtaq 1977, Ahmad et al. 1979, Ahmad 1980, Ahmad & Moizuddin 1980, Afzal et al. 1982) by a number of authors. FIGURES 34–43. Alphocoris spp., external male genitalia. 34–36, A. caudatus sp. nov.; 37–40, A. naso sp. nov.; 41–43, A. asper sp. nov. Genital capsule, posterodorsal view (Figs. 34, 37); same, posteroventral view (Figs. 38, 41); left paramere, two different aspects (Figs. 35–36, 39–40, 42–43). Scales in mm. Two specimens, a male and a female, both in excellent condition, potentially representing syntypes of A. lixoides were found in ZMHB (Figs. 65–72). The female (Figs. 69–71) is more similar to the illustration provided by Germar (1839: plate I fig. III), but there is no ground to exclude the male from the syntype series either. The female bears a handwritten lectotype label of C.H.C. Lyal, but the lectotype designation has apparently never been published. None of the Alphocoris species occurring in Southern Asia examined by us is conspecific with the two syntypes, therefore we consider all earlier records of A. lixoides from India and Pakistan as misidentifications. We mention that based on these two syntypes the Afrotropical A. lobulatus Stål, 1873, appears to be conspecific with A. lixoides, but as a revision of the Afrotropical fauna of Alphocoris is beyond the scope of the present paper, no further analysis of the problem is offered here, and also a lectotype designation for A. lixoides is left for a future reviser. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 311 FIGURES 44–46. Alphocoris spp., phalli in lateral view. 44, A. caudatus sp. nov.; 45, A. naso sp. nov.; 46, A. asper sp. nov. Lettering: aed = aedagus; cp-II1, cp-II2, cp-II3 = branches of conjunctival process; cpd = conjunctival pad. Scale in mm. Distribution. SENEGAL (Germar 1839).—IVORY COAST. Lamto (Schouteden 1963, Gillon 1972, 1974), Odienne–Madinani (Linnavuori 1982).—NIGERIA (Czaja 2016a).—ERITREA. Halibaret [= Elabered] (Schouteden 1905).—“Tagana” [unlocated] (Schouteden 1903) Discussion Alphocoris species in the neighbouring Palaearctic areas. Described species of Alphocoris are listed in Table 1. Species occurring in the neighbouring areas of the Middle East and Central Asia differ from the three new species described in the present paper in the following characters: Alphocoris curculionoides (Jakovlev, 1877) (Iran, Afghanistan, Turkmenistan): differs from all new species in its shorter and broader body, its clypeus only slightly protruding anteriad of mandibular plates, and the apically broadly rounded scutellum. Diagnostic characters of this species were discussed and figured by Linnavuori (1984) and Hoberlandt (1997); a non-type from Afghanistan (NMPC) was examined during the present study. Alphocoris halophilus Linnavuori, 1984 (Iraq, Iran): differs from all new species in the shorter and broader body, and its clypeus only slightly protruding anteriad of mandibular plates; differs from A. caudatus sp. nov. by its broadly angularly excised apex of scutellum (Linnavuori 1984). This species is known to us only from the original description. 312 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. FIGURES 47–50. Alphocoris spp., phalli. 47–48, A. naso sp. nov.; 49–50, A. asper sp. nov. Conjunctiva and aedeagus, ventral view (47, 49); same, posterior view (Figs. 48, 50). Lettering: cp-II1, cp-II2, cp-II3 = branches of conjunctival process; cpd = conjunctival pad. Scale in mm. FIGURES 51–58. Alphocoris spp., genitalia of both sexes. 51–54, A. caudatus sp. nov.; 55–56, A. naso sp. nov.; 57–58, A. asper sp. nov. Conjunctiva and aedeagus, ventral view (51); terminalia of female, dorsal view (52, 55, 57); same, posteroventral view (53, 56, 58); intermediate part and apical receptacle of spermatheca (54). Lettering: lt9 = laterotergite IX; t8 = tergite VIII; vf8 = valvifer VIII. Scale in mm. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 313 FIGURES 59–64. Alphocoris spp., female genitalia. 59–62, A. caudatus sp. nov.; 63–64, A. naso sp. nov. Detached genital and postgenital segments in posterior view, left valvifer VIII opened, most of the gynatrium cut off (59, 63); same, dorsal view (60, 64); spermatheca, dorsal view (61); same, right lateral view, spermathecal duct shown in cross section (62). Lettering: algp = anterolateral gynatrial pouch; dd = distal duct of spermatheca; dil = dilation of spermatheca; fec = fecundation sclerite; lt8, lt9 = laterotergites VIII and IX; pd = proximal duct of spermatheca; ra8 = sclerotized rami of valvula VIII; t8, t10 = tergites VIII and X; va8, va9 = valvulae VIII and IX; vf8, vf9 = valvifers VIII and IX. Scales in mm. Alphocoris pfeifferi Hoberlandt, 1997 (Iran): differs from all new species in the shorter and broader body and the posteriorly broadly, obtusely truncate scutellum; differs from A. naso sp. nov. and A. asper sp. nov. in the anteriorly less produced clypeus. Photos of the holotype (♂) and paratype (♀) were seen during the present study. Alphocoris species in Pakistan. A species of Alphocoris was recorded from Pakistan (Sind: Mianiforest, Tandojam) by I. Ahmad and coauthors as A. lixoides (Ahmad & Mushtaq 1977: 25, 26, 31–37; Ahmad et al. 1979: 314 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. 29, 32; Ahmad 1980: 134; Ahmad & Moizuddin 1980: 195; Afzal et al. 1982: 378). The inadequate description and drawings do not allow an identification of the taxon examined by these authors, but it is certain that it was not A. lixoides, and very likely it was none of the three species described in the present paper. The short and broad body of the species (Ahmad & Mushtaq 1977: 26, fig. 55) shows a certain resemblance to some Central Asian members of this genus, e.g. A. curculionoides or A. pfeifferi. FIGURES 65–72. Type material of Alphocoris lixoides Germar, 1839. 65–68, male syntype and its labels; 69–72, female syntype and its labels. Dorsal (65, 69), lateral (66, 70) and ventral (67, 71) views; labels (68, 72). Scales in mm. © ZMHB. THE GENUS ALPHOCORIS IN THE INDOMALAYA Zootaxa 4382 (2) © 2018 Magnolia Press · 315 TABLE 1. Described species of Alphocoris and their distribution (AFR = Afrotropical Region, IND = Indomalayan Region, PAL = Palaearctic Region). Species Distribution A. asper sp. nov. IND: western India A. caudatus sp. nov. IND: western India A. crassipes Linnavuori, 1975 AFR: Sahel (Sudan) A. curculionoides (Jakovlev, 1877) PAL: Middle East, Central Asia (Iran, Turkmenistan, Afghanistan) A. halophilus Linnavuori, 1984 PAL: Middle East (Iraq, Iran) A. indutus Stål, 1865 AFR: Sahel, southern Africa A. larinoides Germar, 1839 PAL: northern Africa, AFR: Sahel, Arabian Peninsula A. lixoides Germar, 1839 AFR: western Africa A. lobulatus Stål, 1873 AFR: Sahel A. naso sp. nov. IND: western India A. pachycephalus Bergevin, 1931 PAL: northern Africa (Algeria, Tunisia) A. pfeifferi Hoberlandt, 1997 PAL: Middle East (Iran) Remarks on comparative morphology. The aedeagus (Fig. 45: aed) bearing an invaginated apical portion, documented for the three new species described in this paper, is a peculiar character within the family Scutelleridae. Although they are rather schematic, judging from the illustrations provided by Linnavuori (1984: 4, fig. 6) at least Alphocoris curculionoides and A. halophilus apparently have a similar aedeagus. The illustrations of an Alphocoris species misidentified as A. lixoides by Ahmad & Mushtaq (1977: 26, figs. 59–60) are apparently inaccurate. In several Heteroptera the support bridge complex gives off a pair of projections, the so-called support bridge prolongations (Yang 2007, Tsai et al. 2011), into the inner lumen of the phallotheca. They can be of various shape, frequently fused along the meson; distally they join to the bases of the conjunctival processes and the aedeagus, and they assist the withdrawal of the aedeagus into the phallotheca in rest. They probably belong to the groundplan of Pentatomoidea, and they are present in the vast majority of its subgroups; their lack in some taxa is probably an apomorphy. Support bridge prolongations occur in most of the scutellerids in which this character has ever been explored; its lack has been documented only in Hotea Amyot & Serville, 1843 (Hoteinae) (Tsai et al. 2011) and it is now explicitly documented in all three species of Alphocoris described in the present paper. In all of the three species a specialized structure, termed as conjunctival pad (Figs. 44, 47, 49: cpd) (new term), is present. It is an inward proliferation of the conjunctiva at the base of the conjunctival processes, closely surrounding the endophallic reservoir. It apparently aids the withdrawal of the aedeagus into the phallotheca in lack of support bridge prolongations. A similar but clearly convergent structure occurs in Dismegistus spp. (Parastrachiidae?) (Tsai 2011). The spermathecae of A. caudatus sp. nov., A. naso sp. nov., and A. asper sp. nov. are also unusual. The basal, thin-walled, saccular portion (Figs. 61–62: pd) is recognized as a modified proximal duct based on its association with the fecundation sclerite (Fig. 60: fec) in A. caudatus sp. nov. The structure of the cross-section of its wall suggests that the thick-walled portion (Figs. 61–62: dil) is presumably homologous with the dilation of the spermathecal duct found in several Scutellerinae (cf. Tsai et al. 2011). In one carefully dissected specimen of A. caudatus sp. nov., A. naso sp. nov. and A. asper sp. nov. each, the dilation was withdrawn into the proximal duct (as shown in Fig. 64), but it was easily everted (as in Fig. 60) using a fine forceps. The dilated proximal duct perhaps serves for reception of the terminal portion of the aedeagus during copulation; an analogous dilation restricted to the extreme base of the proximal duct was documented in Coleotichus borealis Distant, 1899 (Elvisurinae) by Tsai & Rédei (2010). Similar spermathecae, nevertheless without clear indication of the distinction between proximal duct and dilation, were figured by Czaja (2016b) for two Afrotropical species of Alphocoris; this author claimed that the whole dilated portion of the spermathecal duct is saccular. The reduction of certain portions of the ovipositor (gonangulum, sclerotized rami) is apparently a trend in certain subgroups of Scutelleridae, particularly in the subfamilies Odontoscelinae, Odontotarsinae, Eurygastrinae and Tectocorinae (together presumably forming a single monophyletic assemblage, cf. Wu et al. 2017). Although 316 · Zootaxa 4382 (2) © 2018 Magnolia Press RÉDEI ET AL. some authors (e.g. McDonald & Cassis 1984, Cassis & Vanags 2006) attributed subfamily importance to the lack of sclerotized gonangulum and/or rami, Alphocoris is a striking example that species with distinctly developed (A. caudatus sp. nov., Fig. 59) or strongly reduced to completely missing (A. naso sp. nov., Fig. 64; A. asper sp. nov.) rami might occur even within the same genus. Acknowledgements We are grateful to Michael D. Webb, Maxwell V. L. Barclay (BMNH), Petr Kment (NMPC), and Jürgen Deckert (ZMHB) for allowing access to the collections under their care. Attilio Carapezza (University of Palermo) sent us photos of the holotype and paratype of A. pfeifferi, he and Petr Kment provided helpful comments to the manuscript, their cooperation is much appreciated. This study received financial support from the National Natural Science Foundation of China (grant no. 31472024) and the One Hundred Young Academic Leaders Program of Nankai University to D. Rédei. References Afzal, M., Shaukat, S.S. & Ahmad, I. (1982) Taxometric study of the family Scutelleridae Leach (Hemiptera: Pentatomoidea) from Pakistan. Oriental Insects, 16 (3), 373–383. https://doi.org/10.1080/00305316.1982.10434321 Agassiz, L. (1843) Nomina systematica generum hemipterorum […]. In: Agassiz, L., Nomenclator zoologicus, continens nomina systematica generum animalium tam viventium quam fossilium, secundum ordinem alphabeticum disposita, adjectis auctoribus, libris, in quibus reperiuntur, anno editionis, etymologia et familiis, ad quas pertinent, in singulis classibus. Fasciculus III et IV. Jent et Gassmann, Soloduri, V + 20 pp. Ahmad, I. (1980) Insect fauna of Pakistan and Azad Kashmir—some groups within the order Hemiptera. 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