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Revision of the Shield-Bug Genera <Emphasis Type= ISSN 0013-8738, Entomological Review, 2007, Vol. 87, No. 6, pp. 701–739. © Pleiades Publishing, Inc., 2007. Original Russian Text © E.N. Belousova, 2007, published in Entomologicheskoe Obozrenie, 2007, Vol. 86, No. 3, pp. 610–654. Revision of the Shield-bug Genera Holcostethus Fieber and Peribalus Mulsant et Rey (Heteroptera, Pentatomidae) of the Palaearctic Region E. N. Belousova Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia Received December 11, 2005 Abstract—A complex of the heteropteran genera centering around Peribalus Mulsant et Rey and Holcostethus Fieber is considered. The genus Dryadocoris Kirkaldy reveals no relationship with the above genera and is be- lieved to represent a separate clade of the family Pentatomidae. The genera Peribalus and Holcostethus are revised. The former includes three subgenera: Peribalus s. str. with two species, Asioperibalus subgen. n. (type species Ci- mex inclusus Dohrn) with six species, and Tianocoris subgen. n. (type species Holcostethus manifestus Kiri- tshenko) with two species. Holcostethus embraces two subgenera: Holcostethus s. str. and the monotypic Enigmo- coris subgen. n. (type species H. fissiceps Horváth). Two new species are described: Peribalus tianshanicus sp. n. from the Tien Shan Mts. and P. przewalskii sp. n. from the northern part of China (Huan He River). P. capitatus Jakovlev and P. vernalis (Wolff) are downgraded to subspecies of P. strictus (F.). P. ovatus Jakovlev is syn- onymized with P. inclusus (Dohrn). Two new monotypic genera related to the revised complex of genera are estab- lished, Paraholcostethus gen. n. (type species Peribalus breviceps Horváth) and Himalayastethus gen. n. (type species H. pilosus sp. n. from Kashmir). A key to, and morphometric characters for all the taxa considered are pro- vided. The key characters, including both male and female genitalia, are illustrated, and distributional maps are given. DOI: 10.1134/S0013873807060085 Species of the genera Holcostethus and Peribalus The author of the present study agrees to this division were repeatedly transferred from one to the other ge- and reconsiders the species composition of these gen- nus. Fieber (1861) described the genus Holcostethus era in the Palaearctic fauna. In the present study, two for species with the open clypeus, and placed there new subgenera are described in the genus Peribalus, Cimex sphacelatus Fabricius (type species by subse- Asioperibalus subgen. n. with the new species quent designation), H. congener Fieber (= Cimex albi- P. (A.) przewalskii sp. n., and Tianocoris subgen. n. pes F.), and H. jani Fieber (= Cimex apicalis Herrich- with the new species P. (T.) tianshanicus sp. n. Schaeffer). Mulsant and Rey (1866) included species In the genus Holcostethus, the subgenus Enigmoco- with closed clypeus in the genus Peribalus, and pro- ris subgen. n. is described. Peribalus vernalis and posed an unnecessary new name Dryocoris, instead of P. capitatus are downgraded to subspecies of P. stric- Holcostethus, for species with the open clypeus. Later, tus, P. ovatus is synonymized with P. (A.) inclusus. Stål (1872) and some subsequent authors left in the Peribalus breviceps, exhibiting a set of unique charac- genus Holcostethus only Pentatoma analis Costa ters, is separated in a genus of its own, Parahol- (a junior synonym of Dryadocoris apicalis), all the costethus gen. n. The new genus Himalayacoris gen. other species were transferred to the genus Peribalus. n. is described from the southern spurs of the Himala- However, Kirkaldy (1909) designated Cimex spha- yas. Examination of representatives of the genus Dry- celatus as the type species of Holcostethus, in which adocoris has revealed no relationship to the genera he included most species of the group discussed. For considered. The members of this generic complex, Pentatoma analis and closely related African species with rare exceptions, are very similar in external char- he erected the genus Dryadocoris. Ribes and Schmitz acters. Therefore, features of the genital structure, (1992) divided Holcostethus into the two genera: differing between the genera and subgenera, gane in Peribalus (erroneously referred to as Dryocoris) with importance. In addition, the number, shape, and degree closed clypeus and Holcostethus with open clypeus. of sclerotization of conjunctive processes of the 701 702 BELOUSOVA aedeagus and also the type of sclerotized median 2(1). Head moderately wide, gradually narrowed ante- plates and parameres of males correlate with charac- riorly, weakly, if at all, emarginate before eyes ters of the female genitalia: the size and shape of the (Figs. 95, 99). Pronotum with costa or carina bulb and its processes, the dilation at the base of the along margin (e.g., Figs. 84–89). Scutellum sclerotized duct of the spermatheca, and the shape of without pits in inner corners. Aedeagus with 1–3 the genital plates. Diversity of shapes and sizes of the pairs of membranous conjunctive processes; conjunctive processes is illustrated by the example of sclerotized median plates not clavate, frequently the structure of the aedeagus of Holcostethus abbre- absent (Paraholcostethus breviceps). Parameres viatus Uhler, a representative of the New World fauna much more slender; hypophysis narrower, with (Figs. 47–49). smaller wrinkled area; sensory lobe poorly de- veloped (Figs. 61–62) or absent (Figs. 50–52). The study is based on examination of the collection of the Zoological Institute, Russian Academy of Sci- 3(8). Clypeus open. ence (St. Petersburg) [ZIN] and also the collections of 4(5). Aedeagus without sclerotized median plates; Institute of Zoology, National Academy of Sciences of apices of conjunctive processes bifurcate and the Ukraine (Kiev) [IZASU] and National Museum in sclerotized (Fig. 38). Paramere (Fig. 68) with Prague. The type material is deposited in ZIN. For rounded lobe at base of hypophysis. Anterior an- more common species, only a part of the material from gle of bucculae rounded ....................................... ZIN collections is listed. ..................................... Paraholcostethus gen. n. METHODS 5(4). Aedeagus with sclerotized median plates (e.g., Figs. 29–34); when apices of conjunctive proc- The methods of measurements and genital prepara- esses sclerotized, they do not bifurcate. Paramere tion technics were described earlier (Belousova, without lobe at base of hypophysis. Anterior an- 1996). About 600 specimens were examined, over 300 gle of bucculae pointed. preparations of genitalia were made, and 14 parame- 6(7). Head and pronotum distinctly pubescent dorsally. ters were measured in 230 specimens. Morphometric Paramere pointed apically in lateral view data were processed statistically. The following abbre- (Fig. 69). Sclerotized median plates of aedeagus viations are used in the paper: b. lw, body length to wide, connected at base (Figs. 35, 36) ................. width ratio; h. wl, head width to length ratio; l. bh, ......................................... Himalayacoris gen. n. body to head length ratio; ds, diatone to synthlipsis ratio (i.e., ratio of the width of the head, including the 7(6). Body without distinct pubescence. Paramere eyes, to the distance between the inner margins of the rounded apically in lateral view. Sclerotized me- eyes; the greater this value, the greater the relative size dian plates of aedeagus narrow, separate of the eye); l. ba, body to antenna length ratio; p. wl, (Figs. 39, 40) ............ Holcostethus Fieber, 1861. pronotum width to length ratio; s. lw, scutellum length 8(3). Clypeus covered by jugae in front view to width ratio; l. bb, ratio of the length of the body to (Figs. 84–93). Aedeagus with 1–3 pairs of con- the circumference of spermathecal bulb. junctive processes; sclerotized median plates wide, connected at base (Figs. 18–34) ................. A KEY TO THE GENERA OF THE ......................... Peribalus Mulsant et Rey, 1866. HOLCOSTETHUS GROUP Genus PERIBALUS Mulsant et Rey, 1866 1(2). Head rather wide, with distinct emargination Peribalus Mulsant et Rey, 1866 : 262. Type species before eyes (Fig. 109). Pronotum without costa Cimex vernalis Wolff, 1804 (Kirkaldy, 1909: XXIX). or carina along margin. Scutellum with pits in inner corners (Fig. 108). Aedeagus with 3 Diagnosis. The genus is most closely related to membranous conjunctive processes; sclerotized Holcostethus but differs in the following characters: median plates clavate (Figs. 113–117). Pa- clypeus closed; sclerotized median plates of aedeagus wide, connected at base; outgrowths of spermathecal rameres large, flattened; hypophysis wide, bulb short, forming no process. with large wrinkled area; sensory lobe well de- veloped (Figs. 110–112) ..................................... Composition. The species of the genus are classi- ............................... Dryadocoris Kirkaldy, 1909. fied into three subgenera, two of which are described ENTOMOLOGICAL REVIEW Vol. 87 No. 6 2007 REVISION OF THE SHIELD-BUG GENERA 703 as new based on the number of conjunctive processes duct not widened at base (Figs. 6–9); occasion- of the aedeagus, shape of the inner surface of the pa- ally with bulb spherical and sclerotized duct ramere, and structure of the spermatheca. widened at base to varying extent (Figs. 10, 11, 15, 17). The range of the genus covers nearly entire Palae- arctic Region from Mediterranean to the coast of the 5(16). Aedeagus with 2 pairs of conjunctive processes; Pacific Ocean. The ten species occur in the plains and vesica strongly
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