Composition of the Genera Gnaptorina Reitter and Pseudognaptorina Kaszab of the Tribe Blaptini (Coleoptera, Tenebrionidae)

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1 ISSN , Entomological Review, 2009, Vol. 89, No. 4, pp Pleiades Publishing, Inc., Original Russian Text G.S. Medvedev, 2009, published in Entomologicheskoe Obozrenie, 2009, Vol. 88, No. 2, pp Composition of the Genera Gnaptorina Reitter and Pseudognaptorina Kaszab of the Tribe Blaptini (Coleoptera, Tenebrionidae) G. S. Medvedev Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia Received November 18, 2008 Abstract Based on the structure of the segments of the male fore and middle tarsi, three subgenera are distinguished in the genus Gnaptorina Reitter: Gnaptorina s. str. (brushes or tufts of pale setae are present on the sole surface of the 1st 3rd segments of the fore tarsus and also on the 1st and 2nd segments of the middle tarsus, the type species Gnaptorina felicitana Reitter, 1887), Boreoptorina subgen. n. (a brush is present on the sole surface of the 1st segment of the fore tarsus, but is absent on the sole surface of the segments of middle tarsus, the inner margin of the male fore tibia with a hairy brush, the type species Gnaptorina cordicollis G. Medvedev, 1998), and Hesperoptorina subgen. n. (a flat hairy brush is present only on the sole surface of the 1st segment of the fore tarsus, type species Gnaptorina brucei Blair, 1923). The genus Pseudognaptorina Kaszab, 1977 is considered monotypical. Lectotypes of four species of Gnaptorina are designated. DOI: /S The number of species described in the subtribe Gnaptorinina considerably have increased during the last decade, which contributed to elaboration of the taxonomic structure of the genera belonging to the tribe and to specification of their diagnostic characters. In the present study, the approaches to perfection of the infrageneric classification, based on analysis of the diagnostic significance of the characters, are discussed by the example of the genera Gnaptorina Reitter and Pseudognaptorina Kaszab. GENUS GNAPTORINA REITTER Reitter, 1887 : ; 1889 : ; Koch, 1965 : ; Medvedev, 1998 : ; 2008 : ; Li et Ren, 2004 : ; Shi et al., 2007 : Type species Gnaptorina felicitana Reitter, 1887, by monotypy. Gnaptorina is among the largest genera in the subtribe Gnaptorinina. It comprises 29 species, but some of them have been erroneously included in the genus. It was mentioned earlier (Medvedev, 2008 : 814) that the species described as Gnaptorina quadrata Li et Ren, 2004 should be placed in the genus Asidoblaps Fairmaire. Gnaptorina rufipes Li et Ren, 2004 possesses tibiae strongly widened in the distal part (Li et Ren, 2004: Plate I, 7), spurs at the apex of the fore tibia short, and parameres weakly arcuately converging toward the apex, these characters (first of all, the structure of the legs) indicate that this species belongs to the genus Agnaptoria. The species described as Gnaptorina tenuicorpra Li et Ren, 2004 belongs to the same genus, since the spurs at the apex of the female fore tibia are poorly developed in this species, all the tibiae are considerably widened, and the body is elongate (Li et Ren, 2004: figs ; Plate I, 6). I do not include G. longicornis Li et Ren, 2004 in the genus Gnaptorina, since at least the hind tibiae of this species are S-curved (Li et Ren, 2004: Plate II, 15), which is not characteristic of representatives of the genus Gnaptorina, but is observed in species of the genus Tagonoides Fairmaire, in particular, in T. zamotailovi G. Medvedev (Medvedev, 1998: figs. 18, 19). However, judging by the structure of the parameres distinctly elongate and straightly narrowed toward the apex, G. longicornis does not belong to the genus Tagonoides. Based on the structure of the sole surface of the fore and middle tarsi and the inner surface of the fore tibia of the male, I subdivide the genus Gnaptorina into three groups of species treated here as the subgenera because of morphological isolation of these taxa and compactness of the distribution of their species. Subgenus GNAPTORINA s. str. Description. Sole surfaces of 1st 3rd segments of fore tarsus, and also 1st and 2nd segments of middle 451

2 452 MEDVEDEV Fig. 1. Distribution of species of the genus Gnaptorina Reitter: (1) G. lii Li et Ren, (2) G. platytarsia Li et Ren, (3) G. montana G. Medvedev, (4) G. media G. Medvedev, (5) G. kozlovi G. Medvedev, (6) G. miroshnikovi G. Medvedev, (7) G. kashkarovi G. Medvedev, (8) G. rugosipennis G. Medvedev, (9) G. potanini potanini Reitter, (10) G. potanini minxiana G. Medvedev, (11) G. australis G. Medvedev, (12) G. proxima Reitter, (13) G. felicitana Reitter, (14) G. cylindricollis Reitter, (15) G. reitteri Koch, (16) G. cordicollis G. Medvedev, (17) G. sikkimensis Kaszab, (18) G. brucei Blair, (19) G. pilifera Shi et al., (20) G. compressa Shi et al., (21) G. kangmar Shi et al., (22) G. himalayana Shi et al., (23) G. globithoracalis Shi et al., (24) G. nigra Shi et al. A, collecting sites of species of nominotypical subgenus; B, the same of the subgenus Boreoptorina subgen. n.; C, the same of the subgenus Hesperoptorina subgen. n. tarsus of male with hairy brush or hairy tuft. Inner surface of fore tibia rather sparsely covered with semierect pale setae. Parameres strongly narrowed in apical part. Males frequently with rudiment of lower spur at apex of fore tibia (G. australis G. Medvedev, G. cyllndricollis Reitter, G. potanini Reitter, G. rugosipennis G. Medvedev, etc.). The range of the subgenus covers the territory lying within the limits of the upper Huang He, Mekong, and Yangtze rivers (Fig. 1). The subgenus includes 15 species (Fig. 1, 1 15): G. australis G. Medvedev (11), G. cylindricollis Reitter (14), G. felicitana Reitter (13), G. fairmairei Koch (Sichuan, Sunpan, 32 42'N / 'E), G. kashkarovi G. Medvedev (7), G. kozlovi G. Medvedev (5), G. lii Li et Ren (1), G. media G. Medvedev (4), G. miroshnikovi G. Medvedev (6), G. montana G. Medvedev (3), G. platytarsia Li et Ren (2), G. potanini Reitter (9), G. potanini potanini Reitter, (10) G. potanini minxiana G. Medvedev), G. proxima Reitter (12), G. reitteri Koch (15), G. rugosipennis G. Medvedev (8). Subgenus BOREOPTORINA G. Medvedev, subgen. n. Type species Gnaptorina cordicollis G. Medvedev, Description. Only 1st segment of fore tarsus of male with flat hairy brush at apical margin of sole surface. Sole surface of segments of middle tarsus of male without hairy brushes or tufts. Inner surface of male fore tibia with dense pale hairy brush in apical part. Parameres elongate, cuneiform, strongly narrowed from base to apex, with outer margins arcuately emarginate along entire length and with non-separated widened basal part (Medvedev, 1998: fig. 6). Surface of prosternum vertical before fore coxae. Pronotum distinctly cordate. The range of the subgenus occupies the northwestern part of the range of the genus Gnaptorina, extending from the upper Huang He River to the upper Mekong River and partly coinciding with the range of the nominotypical subgenus. The subgenus includes only the type species G. cordicollis (Fig. 1, 16).

3 COMPOSITION OF THE GENERA 453 Subgenus HESPEROPTORINA G. Medvedev, subgen. n. Type species Gnaptorina brucei Blair, Description. Only 1st segment of male fore tarsus with flat hairy brush at apical margin of sole surface, which is also typical of G. cordicollis belonging to subgenus Boreoptorina subgen. n. However, in species of Hesperoptorina subgen. n., inner surface of male fore tibia with dense brush of pale setae in apical half, in contrast to sparse setae of G. cordicollis. In addition, parameres of species of Hesperoptorina either similar to those of species of Gnaptorina s. str. or differing from them in outer margins rectilinearly converging toward apex (G. pilifera Shi et al., G. globithoracalis Shi et al.), whereas in G. cordicollis they are strongly elongate and sharply narrowed in apical part, with outer margins arcuately emarginate along entire length. The range of the subgenus Hesperoptorina subgen. n. occupies the southern part of Tibet and the northern Himalayas and extends from the northeastern edge of Nepal and Sikkim (India) in the west to the source of the Salween River in the northeast, being rather distinctly isolated from the ranges of the other subgenera of the genus Gnaptorina. The subgenus includes 9 species (Fig. 1, 17 24): G. brucei Blair (18), G. compressa Shi et al. (20), G. globithoracalis Shi et al. (23), G. himalayana Shi et al. (22), G. kangmar Shi et al. (21), G. nigra Shi et al. (24), G. pilifera Shi et al. (19), G. sikkimensis Kaszab (17), G. tishkovi G. Medvedev ( Thibet, N. Himalaya, Šnišnašma ). The naturalness of the subgenus Hesperoptorina subgen. n. as a group of the species descending from a common ancestor is confirmed not only by the morphology of the species but also by geographical criteria. At the same time, some representatives of this subgenus exhibit a considerable peculiarity of their characters, which testifies to a long existence of the southwestern center of the evolution of the genus Gnaptorina. In particular, G. pilifera bears longitudinal granulate prominences on the elytra (Shi et al., 2007: figs 73, 74), which are absent in the other species of the genus Gnaptorina. Among the species of the subgenus Hesperoptorina subgen. n., the elongately angular shape of the parameres (not occurring in Gnaptorina) is inherent in G. pilifera and G. globjfithoracalis (Shi et al., 2007: figs 6, 54). The following descriptions of G. brucei and G. sikkimensis supplement the original descriptions with measurements and contain figures of details of the structure of these species. Gnaptorina (Hesperoptorina) brucei Blair (Figs. 1, 18; 2 20) Blair, 1923 : 282 (type locality: Tengkye-La, Thibet, ft. ); Koch, 1965 : 129; Kaszab, 1965 : 110. Description. Body black; antennae (especially apical segments), labrum, and segments of maxillary and labial palpi reddish; legs brownish. Male. Temples almost rectilinearly narrowed toward neck constriction (Fig. 2), their outer margins smoothly passing into outer margins of eyes. Outer margins of genae parallel immediately before eyes, then more sharply converging toward base of clypeus, or regularly arcuately converging anteriorly from very base. Outer margin of head above base of antennae with widely obtuse-angled emargination. Anterior margin of clypeus shallowly arcuately emarginate, its anterior angles rounded. Frontoclypeal suture rather distinct. Surface of head with relatively coarse, dense, simple punctation. Antennae (Fig. 3) with apices almost 1/4 of length of pronotum not reaching base of pronotum. Length(width) ratio of 2nd 11th antennal segments 10(7) : 22(7) : 10(7) : 10(7) : 10(7) : 10(10) : 10(10) : 10(10) : 10(10) : 12(10). Pronotum (Fig. 5) transverse ( times as wide as long), widest in anterior 1/3, times as wide there as head. Anterior margin of pronotum straight or only shallowly arcuately emarginate; base straight or slightly arcuately convex in medial 1/3; sides weakly arcuately convex, very finely edged; anterior angles widely obtuse, rounded; posterior angles obtuse, narrowly rounded apically. Surface of pronotum regularly convex between outer margins, covered with moderately coarse, dense, distinct simple punctures. Propleura not depressed along outer margin, with fine longitudinal rugosity. Prosternum vertical before fore coxae. Prosternal process steeply sloping behind coxae, with small projection at end of sloping surface. Elytra oblong-oval ( times as long as wide), times as wide as pronotum. Surface of elytra regularly convex, with rather coarse, rugosepunctate sculpture. Outer margin of epipleura visible in dorsal view in anterior 2/3 and at apex, posteriorly reaching sutural elytral angle. Surface of epipleura finely rugose. Visible abdominal sternites lustrous,

4 454 MEDVEDEV Figs Gnaptorina brucei Blair: (2) head of male; (3) antenna of male; (4) antenna of female; (5) pronotum of male; (6) pronotum of female; (7) fore tibia of female; (8) fore tibia of male; (9) aedeagus, lateral view; (10) parameres, dorsal view. A, scale to Figs. 2, 7, 8; B, to Figs. 3, 4; C, to Figs. 5, 6; D, to Figs. 9, 10. covered with fine punctures and fine, rather sparse, pale setae. Legs relatively short. Fore tibia (Fig. 8) narrow, slightly S-curved; its outer margin with shallow emargination at apex. Apical spur on fore tibia of uniform width along entire length, rounded at apex; rudiment of lower spur absent. Middle tibia slightly arcuately curved, hind tibia straight. All tarsi narrow, only 1st segment of fore tarsus with flat hairy brush on sole surface at apical margin. Aedeagus (Fig. 9) in lateral view distinctly curved ventrally in basal 1/3. Length of aedeagus 1.75 mm, width mm (body length 10 mm). Parameres (Figs. 9, 10) in dorsal view with wide base and strongly narrowed apical part, their outer margins deeply arcuately emarginate. In lateral view, parameres slightly S-curved. Length of parameres 0.5 mm, width 0.4 mm. Female. Body wider than that of male, distinctly convex, lustrous. Pronotum times as wide as head, times as wide as long. Elytra wideoval, times as long as wide, times as wide as pronotum. Head and pronotum with rather coarse, dense punctation. Surface of pronotum with 2 symmetrically lying mirror spots at center of disc (obsolete in male). Sculpture of elytra coarsely rugosepunctate. Antennae (Fig. 4) with apices extending slightly beyond midlength of pronotum. Shape of 7 10th antennal segments more regular than that in male, transversely oval. Fore tibia slightly curved; its outer margin emarginate in apical 1/3 and then, toward base, with obsolete sinuosity bearing small spines. Apical spur on fore tibia very wide, rounded at apex; its apex reaching apical margin of 3rd tarsal segment. Body length of male mm, width 4.9 mm; body length of female , width 6.2 mm.

5 COMPOSITION OF THE GENERA 455 Material examined (Hungarian Natural History Museum, Budapest). Nepal, Mustangbhot, 29 11' Br., 83 58' L., 3700 m, 12.VIII.1955, 1, 1, leg. F. Lobbichler; same locality, 3800 m, 14.VIII.1955, 1, leg. F. Lobbichler; same locality, 3800 m, 16.VIII.1955, 1, leg. F. Lobbichler. All specimens with label: Brucei Blair, Dr. Z. Kaszab det. Gnaptorina (Hesperoptorina) sikkimensis Kaszab (Figs. 1, 17; 11 18) Kaszab, 1965 : 110. G. sikkimensis was described from three males collected in Sikkim (Donkung, ft., 22.VI.1959), with indication only of its differences from G. brucei Blair, In the only figure accompanying the list of diagnostic characters, the pronotum is shown to be more elongate than that which is typical of this species. In the present description of G. sikkimensis, the male antenna, the male and female pronotum, parameres, and details of the structure of the legs are illustrated. The female of G. sikkimensis is described for the first time. In male, body black; in female, elytra brownish red. Ventral side of body and legs of male also black, apical antennal segments slightly brownish. Male. Outer margins of temples slightly arcuately converging to neck constriction. Genae parallel-sided immediately before eyes. Outer margins of head almost rectilinearly converging from apex of genal angle to anterior margin of clypeus. Anterior margin of clypeus shallowly arcuately emarginate. Frontoclypeal suture distinct. Surface of head with dense fine punctation denser on clypeus. Antennae (Fig. 11) fine, their apices not reaching base of pronotum. Length (width) ratio of 2nd 11th antennal segments 9(8) : 20(9) : 9(9) : 9(9) : 9(9) : 9(9) : 9(11) : 9(11) : 9(11) : 12(11). Pronotum (Fig. 12) moderately transverse (1.23 times as wide as long), widest before middle, 1.67 times as wide there as head. Base of pronotum straight; anterior margin with very shallow arcuate emargination; sides arcuately convex in anterior half, shallowly arcuately emarginate in basal half, finely edged, distinctly S-curved in lateral view; posterior angles weakly obtuse, rounded apically; anterior angles widely obtuse, more narrowly rounded apically than anterior angles. Surface of pronotum regularly convex between outer margins, densely and finely punctate, with 2 small spots at center of disc. Figs Gnaptorina sikkimensis Kaszab: (11) antenna of male; (12) pronotum of male; (13) pronotum of female; (14) fore tibia of male; (15) fore tibia of female; (16) apical spurs of hind tibia of female; (17) parameres, dorsal view; (18) parameres, lateral view. A, scale to Figs. 11, 16; B, to Figs. 12, 13; C, to Figs. 14, 15; D, to Figs. 17, 18. Propleura with fine irregular rugosity, without depression along outer margin. Prosternum vertical before fore coxae. Prosternal process steeply sloping behind fore coxae, with flat dentiform process at end of sloping surface. Elytra oblong-oval (length 1.45 times width), widest in middle, 1.43 times as wide there as pronotum. Surface of elytra regularly convex, densely covered with fine punctures and rugae. Outer margin of epipleura visible in dorsal view in anterior 1/3. Epipleura strongly narrowed at apex, reaching sutural angle. 1st 3rd visible abdominal sternites with very weak wrinkles and hardly visible minute setae at anterior margin. 4th sternite without wrinkles and setae, 5th sternite with minute recumbent setae in apical part. Legs rather strong. Ratio of length(width) of fore, middle, and hind femora 66(15) : 66(15) : 70(16), that

6 456 MEDVEDEV of corresponding tibiae 46(9) : 50(10) : 67(12). Outer margin of fore tibia (Fig. 11) emarginate at apex, slightly undulate in basal 2/3; inner surface with rather sparse setae. Apical spur of fore tibia parallel-sided, rounded at apex; rudiment of lower spur absent. Middle tibia slightly arcuately curved, distinctly (as well as hind one) widened at apex. Sole surface of only 1st segment of fore tarsus with flat hairy brush on anterior margin. Aedeagus moderately strongly curved in basal half in lateral view. Parameres (Figs. 17, 18) sharply narrowed in apical half in dorsal view, slightly S-curved in lateral view. Length of aedeagus 2 mm, width 0.5 mm; length of parameres 0.7 mm, width 0.4 mm (body length 9.5 mm). Female. Elytra wider than those in male, 1.34 times as long as wide. Pronotum 1.71 times as wide as head, 1.26 times as wide as long. Elytra 1.4 times as wide as pronotum. Surface of pronotum with 2 symmetrically situated mirror spots at center of disc. Antennae similar to those of male, rather short. Ratio of length(width) of 2nd 11th antennal segments 8(7) : 18(7) : 8(7) : 8(7) : 8(8) : 8(8) : 8(10) : 9(11) : 10(12) : 11(11). Fore tibia (Fig. 15) wider than that of male, with large apical spur rounded apically, ventrally with granules bearing spiniform setae. Inner apical spur of hind tibia (Fig. 16) rather smoothly widened toward apex, slightly longer than outer spur. 1st 3rd visible abdominal sternites covered with smooth longitudinal rugae and minute pale setae more strongly developed on last visible sternite. Structure of spermatheca characterized by common base of sphincter and 2nd reservoir separated from base of 1st reservoir by short duct. Body length of male 9.5 mm, width 4.8 mm; body length of female 10.7 mm, width 5.6 mm. Material examined. N. Sikkim, Gayozamy, 4000 m, 25.VII 8.X.1938, 1 ; Oberhalb Gayamtsona Lake, c m, VIII.1938, 1. Both specimens are also provided with the labels: Schäfer-Exped. and Gnaptorina brucei Blair, A. Bogačev det. LECTOTYPES OF THE SPECIES OF THE GENUS GNAPTORINA, DESCRIBED BY REITTER On the basis of the data included by Reitter (1887, 1889) in descriptions of species of the genus Gnaptorina (G. felicitana Reitter, G. cylindricollis Reitter), it was established that each of the species had been described from no less than two specimens, which requires designation of lectotypes. The body size was reported in the descriptions for G. felicitana and G. cylindricollis ( mm and mm, respectively), the characters of the male and the female, for G. cylindricollis, and the number of individuals of the type series ( 2/Ех. ), for G. proxima. The collection of the Zoological Institute, Russian Academy of Sciences (St. Petersburg) [ZIN] includes one syntype of each species with the data listed below. I designate here these specimens as lectotypes, and specimens from E. Reitter s collection in the Hungarian Natural History Museum (Budapest), as paralectotypes. For each species, I indicate the number of the specimens which are part of the material of expeditions of N.M. Przhevalsky and N.G. Potanin but have not been examined by E. Reitter and, consequently, cannot be considered as paralectotypes. Gnaptorina felicitana Reitter Reitter, 1887 : 365. G. felicitana was described from the material of N.M. Przhevalsky s expedition to the basins of the Yangtze (Blue in English) and Huang He (Yellow River in English) rivers in June and July of The ZIN collection includes one specimen (female) labeled: Amdo, 1884, Przevalsky (with the mark Bytschu on the underside) and Gnaptorina m. n. g. felicitana m. n. sp. (E. Reitter s handwriting). This specimen is designated here as lectotype. The specimens examined by E. Reitter and deposited in the Hungarian Natural History Museum (Budapest) are paralectotypes. The ZIN collection includes a long series of the specimens (19 males, 4 females) collected together with the lectotype but, probably, not considered by E. Reitter during preparation of the species description. Gnaptorina proxima Reitter. Reitter, 1889 : 694. G. proxima was described from two specimens collected in Gansu ( Kan-ssu, 12/V Ex. ). The ZIN collection includes a specimen (female) labeled Kan-ssu, 1885, G. Patanin [Potanin] ( 12/V on the underside) and Gnaptorina proxima m. n. sp. (E. Reitter s handwriting). This specimen is designated here as lectotype, and the specimen deposited in the collection of the Hungarian Natural History Museum (Budapest), as paralectotype.

7 COMPOSITION OF THE GENERA 457 One specimen (male) in the ZIN collection was collected, as well as the lectotype, on 12.V.1885; the four others, on 11.V Gnaptorina potanini Reitter Reitter, 1889 : 694. Gnaptorina potanini was described from Gansu and Sichuan ( Kan-ssu, April Mai; Sze-tschuan, August, 1885 ). The ZIN collection includes one specimen (female) labeled: Kan-ssu, 1885, G. Patanin [Potanin] ( 22/V on the underside) and Gnaptorina Potanini m. n. sp. (E. Reitter s handwriting). This specimen is designated here as lectotype, the other specimens examined by E. Reitter and deposited in the Hungarian Natural History Museum (Budapest), as paralectotypes. 11 males and 8 females in the ZIN collection were found by the expedition of G.N. Potanin on 22.V.1885, 1 female 11.V and 1 female on 14.V, 1 male and 3 females on 15.V, 8 males and 2 females on 26.V, 2 males on 27.V, 2 males and 1 female on 28.V, 1 male on 29.V, and 1 male and 2 females on 17.VI. Gnaptorina cylindricollis Reitter Reitter, 1889 : 693. G. cylindricollis was described from Gansu ( Kanssu, April Mai, 1885 ). The ZIN collection includes one specimen (male) labeled: Kan-ssu, 1885, G. Patanin [Potanin] ( 14 V on the on underside) and Gnaptorina cylindricollis n. sp. (E. Reitter s handwriting). This specimen is designated here as lectotype. The ZIN collection includes 3 males and 1 female collected on 14.V.1885 and also 7 males and 2 females collected on 15.V GENUS PSEUDOGNAPTORINA KASZAB Kaszab, 1977 : 250; Shi et al., 2005 : 163. Type species Pseudognaptorina nepalica Kaszab, 1977, by original designation. Pseudognaptorina was described as a monotypical genus from the central part of Nepal (Kaszab, 1977). Recently P. exsertogena Shi et al., 2005 and P. obtusa Shi et al., 2005 from the Tibet Autonomous Region (China) were described in this genus. Considerable differences of these species from the type species of the genus Pseudognaptorina, represented in the fauna of the Himalayas, have necessitated finding additional characters of the genus. In Z. Kaszab s opinion, the genus Pseudognaptorina is most closely related to the genus Agnaptoria. However, it should be noted that Kaszab s paper with description of the genus Pseudognaptorina was published when only two species, A. rubripes Reitter, 1887 (type species of the genus) and A. seidlitzi Reitter, 1893, were known in the genus Agnaptoria. At present, this genus comprises 28 species. Analysis of the characters of the known species of Agnaptoria has shown their wide variability. In particular, the type species of the genus is characterized by the tibiae strongly widened in the apical half and the vertical position of the prosternum before the fore coxae. Being clearly pronounced in the type species, these characters are obsolete in others, and, thus, a complex of characters (including those of the internal structure) should be used for generic identification of the individuals. Inclusion of P. exsertogena and P. obtusa in Pseudognaptorina has made this genus collective. This conclusion is based on the fact that the dorsal margin of the inner surface of the fore femur in P. obtusa forms an angular projection, similarly to that in the species of the genus Asidoblaps Fairmaire. The angular or arcuate projection on the inner surface of the fore femur in species of the genus Asidoblaps is always developed, but very widely varies in shape from the apically rounded to acute-angled (Medvedev, 2002: figs ). This character rather clearly distinguishes the species of the genus Asidoblaps from those of the genera Agnaptoria and Gnaptorina. The presence of a projection on the inner surface of the fore femur in P. obtusa was mentioned in the original description and shown in the figure of the general view of the male (Shi et al., 2005: fig. 19). The structure of the parameres in P. obtusa (Shi et al., figs. 18, 19) sharply differs from that in P. nepalica. In P. obtusa, the parameres are strongly elongate (length is 2.79 times width), and their outer margins form a very shallow arcuate emargination; in P. nepalica, the parameres are 2.27 times as long as wide, and their outer margins are deeply arcuately emarginate, which makes the parameres similar to those of the species of the genus Gnaptorina. The elongate parameres almost cuneiformly narrowed toward the apex and similar to those in P. obtusa are characteristic of a number of species of the genus Asidoblaps (A. modica G. Medvedev, A. munda G. Medvedev, A. bifida G. Medve-

8 458 MEDVEDEV Figs Pseudognaptorina nepalica Kaszab: (19) head of male; (20) head of female; (21) antenna of male; (22) pronotum of male; (23) pronotum of female; (24) fore tarsus of male; dorsal view; (25, 27, 28) fore, middle, and hind tibiae of male; (26) fore tibia of female; (29) fore tarsus of male; lateral view; (30) apical spurs on hind tibiae of male; (31) apical spurs on fore tibia; (32) apical spurs on fore tibia of female; (33) apical spurs on hind tibiae; (34, 35) parameres, dorsal view; (36) parameres, lateral view; (37) apical lobes of ovipositor. A, scale to Figs. 19, 20, 25 28; B, to Figs. 21, 29, 37; C, to Figs. 22, 23; D, to Fig. 24; E, to Figs dev, etc.), and also of Sintagona miranda G. Medvedev which is also similar to P. obtusa in the shape of the parameres in lateral view. The sides of the pronotum in P. nepalica are strongly roundly convex, and the propleura are widely and strongly flattened in the outer part, appearing nearly lamelliform. Such a structure of the pronotum has not been observed in P. exsertogena and P. obtusa. According to the original description, the prosternum of P. obtusa is vertically positioned before the fore coxae, which is not typical

9 COMPOSITION OF THE GENERA 459 of species of the genus Asidoblaps. Therefore, the status P. obtusa can be reliably determined only with the use of additional data on the structure of the prothorax, propleura, epipleura, and female genital tubes, though I can assert for certain that P. nepalica and P. obtusa belong to different genera. In my opinion, the inclusion of the species described as P. exsertogena in the genus Pseudognaptorina is questionable. This species is similar to P. obtusa in the structure of the parameres: its parameres cuneiformly narrow toward the apex, are curved ventrally in lateral view only at the apex, but are not S-shaped, in contrast to those of P. nepalica. An important difference of P. exsertogena from P. obtusa is the position of the prosternum before the fore coxae: in P. obtusa, the prosternum is gently sloping, and not vertical, and the anterior margin of the pronotum is arcuately emarginate, and not straight. Estimating the degree of affinity between P. nepalica, P. obtusa, and P. exsertogena, the presence of setae or tufts of setae on the sole surface of each segment of the fore and middle tarsi of males should be taken into consideration, since these formations are usually similar in all the species of the genus. Fig. 38 shows a significant morphological heterogeneity of the species of the genus Pseudognaptorina in this character. Characteristic of the genus Pseudognaptorina is the combination in its only representative, P. nepalica, of characters important in diagnostics of such genera as Gnaptorina, Agnaptoria, and Asidoblaps. Pseudognaptorina is similar to these genera in the structure of the ovipositor: the lobes of the ovipositor are short and wide, with the oblique apical margin. Pseudognaptorina is similar to Gnaptorina in the structure of the parameres, the presence of widened, apically rounded spurs at the apices of the female fore and hind tibiae, and the scantiness of pale setae in the apical part of the inner surface of the male fore tibia; to the genus Asidoblaps, in the prosternum gently sloping before the fore coxae, and to the genus Agnaptoria, in the simple structure of the fore femur which does not form an angular projection at the dorsal margin of the inner surface. It should be also noted that the distribution ranges of P. nepalica, P. exsertogena, and P. obtusa are very widely separated. P. nepalica is represented in the fauna of the Himalayas, P. exsertogena, in the fauna of Tibet, and P. obtusa, in the fauna of the eastern part of the Tibet Plateau. The characters of the orography of southwestern China essentially influence the fauna and determine the situation of the faunal boundaries. The genera plentifully represented in the fauna of the eastern part of the Tibet Plateau (Gnaptorina, Agnaptoria, Asidoblaps) yield to a complex of the Himalayan genera (Montagona G. Medvedev, Nepalindia G. Medvedev, Blaptogonia G. Medvedev) on the border of this system with the Himalayas. Morphologically peculiar but not species-rich genera (Pseudognaptorina, Sintagona, Belousovia) occur in the transitional zone between the faunas of the Central Asian and Himalayan types. In this zone, the taxa show instability of some characters of diagnostic significance. In particular, the southwestern part of China is inhabited by species of the genus Agnaptoria, which are distinguished from the other species in the structure of the tibiae (the tibiae are weakly widened in the apical half, which is not typical of this genus), and the female genital ducts (a short duct is present between the anterior end of the vagina and the base of the 1st reservoir). In representatives of the genus Belousovia G. Medvedev described from there and rather closely related to the genus Colasia Koch, the 1st 4th segments of the male fore and middle tarsi are provided on the sole surface with dense hairy brushes which are absent in Colasia akisoides Koch known from the northern part of Yunnan and Sichuan. It should be also noted that the genus Gnaptorina is represented in southwestern China by the subgenus Hesperoptorina subgen. n. Thus, based on analysis of the morphological characters of the species included in the genus Pseudognaptorina and on the considerable isolation of their ranges, I can suppose that this genus includes only the type species. Pseudognaptorina nepalica Kaszab (Figs ) Kaszab, 1977 : 251; Shi et al., 2005 : 164. Description. Body black or brownish dorsally, brownish ventrally; mandibles black; maxilla, labium, and their palpi pale brown. Male. Temples almost rectilinearly narrowed backwards (Fig. 19). Eyes weakly convex, outer margins of genae arcuately converging from anterior margin of eyes toward base of clypeus. Outer margins of head above base of antennae with rather deep obtuse-angled emargination. Anterior margin of clypeus straight. Frontoclypeal suture fine. Surface of head with moderately small, dense punctation.

10 460 MEDVEDEV Fig. 38. Scheme of distribution of hairy brushes (A) and hairy tufts (B) on sole surfaces of segments of fore and middle tarsi of male among species described in the genus Pseudognaptorina Kaszab. 1 5, 1st 5th segments of fore and middle tarsi. Antennae (Fig. 21) with apices reaching base of pronotum. Ratio of length(width) of 2nd 11th antennal segments 10(8) : 21(8) : 11(8) : 11(8) : 10(8) : 12(8) : 10(12) : 10(12) : 10(12) : 17(12). Pronotum (Fig. 22) transverse ( times as wide as long, n = 3), widest in middle, times as wide there as head. Ratio of width of pronotum at anterior margin to that in middle, and to that at base 0.58 : 1.00 : Sides of pronotum roundly convex, with distinct sharply raised edging along entire length; base straight; anterior margin with shallow arcuate emargination; posterior angles widely obtuse-angled, relatively sharp apically; anterior angles widely rounded. Surface of pronotum very narrowly flattened along outer margins from base nearly to anterior angles, covered with fine dense punctation. Propleura sharply flattened along outer margin; outer margin sharp, nearly lamelliform. Inner part of propleura covered with longitudinal rugae. Prosternum before fore coxae gently sloping. Prosternal process gently sloping behind fore coxae, forming there obtuse projection. Elytra oblong-oval (length times width), widest in apical 1/3, times as wide there as pronotum. Dorsal surface of elytra passing into outer (deflexed) surface without traces of humeral carina. Outer margin of epipleura visible in dorsal view in anterior 1/3 and at apex. Surface of elytra with minute, dense, rather smooth punctation almost vanishing on apical declivity. Visible abdominal sternites rather sparsely covered with minute, pale, recumbent setae becoming longer on last sternite. Femora and tibiae moderately thickened. Ratio of length(width) of fore, middle, and hind femora 60(15) : 64(15) : 74(15), that of corresponding tibiae 51(9) : 52(12) : 70(12). Fore tibia (Fig. 25) narrow, straight, with upper spur (Fig. 32) slightly longer than lower one; middle tibia (Fig. 27) slightly arcuately curved; hind tibia (Fig. 28) S-curved, narrow. Tarsi (Fig. 24) narrow. Sole surfaces of 1st and 2nd segments of fore tarsus with hairy brush, 3rd segment with small hairy tuft at apical margin (Fig. 38). Sole surface of 1st and 2nd segments of middle tarsus with small hairy tuft at apical margin. Length ratio of 1st 4th segments of hind tarsus 10 : 5 : 5 : 9. Aedeagus small, moderately strongly arcuately bent in lateral view. Parameres (Figs. 34, 35) more sharply narrowed in apical part, their outer margins distinctly arcuately emarginate. In lateral view (Fig. 36), parameres slightly S-curved. Length of aedeagus 2.6 mm, width 0.3 mm; length of parameres 0.63 mm, width 0.27 mm (body length 10 mm). Female. Body larger than that in male. Anterior margin of clypeus straight (Fig. 20). Outer margin of head above base of antennae with widely obtuseangled emargination less sharp than that in male. Antennae with apices reaching base of pronotum. Pronotum (Fig. 23) much wider than long (width 1.38 times length), 1.84 times as wide as head. Ratio of width of pronotum at anterior margin to its maximum width and to width at base 0.58 : 1.00 : Sides of pronotum regularly arcuately convex. Surface rather widely flattened along outer margins and slightly depressed in medial part before base. Edging of outer margin fine and sharp, keel-like elevated in basal 1/3. Punctation of pronotum very dense, rather coarse. Elytra oval,

11 COMPOSITION OF THE GENERA 461 relatively short (length 1.49 times width), widest in apical 1/3, 1.34 times as wide there as pronotum. Outer margin of epipleura visible in anterior 1/3 and at apex. Punctures on elytra slightly smaller and slightly sparser than those on pronotum. Visible abdominal sternites covered with inconspicuous pale recumbent setae. Outer margin of fore tibia (Fig. 26) serrate, in contrast to that in male. Middle tibia slightly arcuately curved, hind tibia S-curved. Spurs on fore (Fig. 32) and hind (Fig. 33) tibiae with rounded apical margin, somewhat differing in length from each other in all tibiae. Apical lobes of ovipositor short and wide (Fig. 37). In 3 males examined, body length mm, width mm; in female, body length 12 mm, width 5.9 mm. Material examined (3, 1, paratypes). Nepal. Expeditionen Jochen Martens, Gompabei Tarakot, m, V.1970, Tagonoides (Pseudognaptorina) nepalica Kasz., det. N. Skopin, 1976, 1 ; Dolpo, Tal der oberen Barbung Khola, zwischen Terang und Tukot, 4000 m, 19.VI.1970, 1 ; Dolpo, Weg von Kangar nach Shimen, m, 18.VI.1973, 1 ; Dolpo, Tal der oberen Barbung Khola, Charka, m, VI.1973, 1. ACKNOWLEDGMENTS The author is grateful to Dr. O. Merkl (Hungarian Natural History Museum, Budapest) for supplying four paratypes of Pseudognaptorina nepalica Kaszab and four specimens of Gnaptorina brucei Blair, identified by Dr. Z. Kaszab. The study was supported by the Russian Foundation for Basic Research (grant no ) and was performed with the use of the collection of the Zoological Institute, the Russian Academy of Sciences (St. Petersburg). REFERENCES 1. Blair, K.G., Coleoptera of the Second Mt. Everest Expedition, 1922: II. Heteromera, Ann. Mag. Nat. Hist. 9 (11), (1923). 2. Kaszab, Z., Wissenschaftliche Ergebnisse der von Dr. F. Schmidt in Indien gesammelten Tenebrioniden (Coleoptera), Miscell. Zool. 2 (1), (1965). 3. Kaszab, Z., Tenebrionidae der Nepal-Expeditionen von Dr. J. Martens ( ), Senckenbergiana Biol. 57 (416), (1977). 4. Koch, C., Sur les types de Fairmaire des tribus Blaptini et Platyscelini conserves au Museum de Paris (Col. Tenebrionidae), Ann. Soc. Entomol. France 1 (1), (1965). 5. Li, She and Ren, Guo-dong, A Systematic Study of Gnaptorina Reitter (Coleoptera, Tenebrionidae) from China, Oriental Insects 38, (2004). 6. Medvedev, G.S., New Species of Tenebrionid Beetles of the Tribe Blaptini (Coleoptera, Tenebrionidae) from Hissaro-Darvaz Mountains and the Plateau of Tibet, Entomol. Obozr. 77 (3), (1998) [Entomol. Rev. 78 (5), (1998)]. 7. Medvedev, G.S., New Species of the Tenebrionid- Beetle Genera Tagonoides Fairm., Gnaptorina Rtt. and Agnaptoria Rtt. (Coleoptera, Tenebrionidae) from China, Entomol. Obozr. 87 (4), (2008) [Entomol. Rev. 88 (9), (2008)]. 8. Reitter, E., Insecta in itinere cl. N. Przewalskii in Asia centrali novissime lecta IX. Tenebrionidae, Horae Soc. Entomol. Ross. 21, (1887). 9. Reitter, E., Insecta, a cl. G. N. Potanin in China et in Mongolia novissime lecta, Horae Soc. Entomol. Ross. 23, (1889). 10. Shi, A.M., Ren, G.D., and Merkl, O., Six New Species of Gnaptorina Reitter, 1887 (Coleoptera, Tenebrionidae: Blaptini) from the Tibet Plateau, Acta Zool. Acad. Sci. Hung. 53 (3), (2007).

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