S.K. Ryndevich, Baranovichi State University, 21, Voykova str., Baranovichi , Belarus.

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ZOOSYSTEMATICA ROSSICA, 26(2): 315 324 25 DECEMBER 2017 New data on Sphaerites perforatus and S. nitidus (Coleoptera: Sphaeritidae): additional diagnostic characters and records from China Новые данные о Sphaerites perforatus и S. nitidus (Coleoptera: Sphaeritidae): дополнительные диагностические признаки и указания из Китая S.K. RYNDEVICH* & D.S. LUNDYSHEV С.К. РЫНДЕВИЧ, Д.С. ЛУНДЫШЕВ S.K. Ryndevich, Baranovichi State University, 21, Voykova str., Baranovichi 225404, Belarus. E-mail: ryndevichsk@mail.ru D. S. Lundyshev Baranovichi State University, 21, Voykova str., Baranovichi 225404, Belarus. E-mail: lundyshevdenis@yandex.by The article deals with diagnostic characters, distribution including additional faunal records, and ecology of Sphaerites perforatus Gusakov, 2017 and S. nitidus Löbl, 1996. A redescription of S. perforatus with indication of additional morphological characters is given. Wings venation of S. perforatus, S. nitidus, and S. glabratus (Fabricius, 1792) is analysed. A key to all species of Sphaerites Duftschmid, 1805 is given. В статье рассмотрены диагностические признаки, распространение, включая дополнительные фаунистические находки, и экология Sphaerites perforatus Gusakov, 2017 и S. nitidus Löbl, 1996. Приводится переописание S. perforatus c указанием дополнительных морфологических признаков. Проанализировано жилкование крыльев S. perforatus, S. nitidus и S. glabratus (Fabricius, 1792). Дана определительная таблица всех видов рода Sphaerites Duftschmid, 1805. Key words: false clown beetles, taxonomy, China, Coleoptera, Sphaeritidae, Sphaerites, new records Ключевые слова: жуки-таежники, таксономия, Китай, Coleoptera, Sphaeritidae, Sphaerites, новые фаунистические указания INTRODUCTION Sphaeritidae, commonly called false clown beetles, include a single genus, Sphaerites Duftschmid, 1805, with seven currently known species. The majority (viz. six) of species occur in the Palaearctic Region: S. dimidiatus Jureček, 1934, S. glabratus (Fabricius, 1792), S. involatilis Gusakov, 2004, S. nitidus Löbl, 1996, S. opacus Löbl et Háva, 2002, and S. perforatus Gusakov, 2017. These species are known only from China (Jureček, 1934; Löbl, 1996; Löbl et Háva, 2002; Gusakov, 2004, 2017) except for S. glabratus widely distributed over the forest zones. A single species, S. politus Mannerheim, 1846, is distributed in the Nearctic Region (Mannerheim, 1846; Kryzhanovskij & Reichardt, 1976). The taxonomic review of the family was published by Gusakov (2004), and more recently the distribution of Sphaerites in the Palaearctic Region was considered by Löbl (2015). The present study is based on the recently collected material of the two rare species, S. perforatus and S. nitidus, from Yunnan and Sichuan (China). The former species was described recently on the basis of seven specimens from five localities in Yunnan; the latter one was known only by the holotype and paratype collected in one locality in Sichuan. 2017 Zoological Institute, Russian Academy of Scienсes

316 The careful study of the morphology of these species, including the hind wings, made it possible to find some additional diagnostic characters and modify the key to the species of the genus proposed by Gusakov (2004). MATERIAL AND METHODS Examined specimens are deposited in the following collections: CDL, D.S. Lundyshev s private collection, Baranovichi, Belarus; CSR, S.K. Ryndevich s collection, Baranovichi, Belarus; ZIN, the Zoological Institute of the Russian Academy of Sciences, St Petersburg, Russia. The males of Sphaerites species were dissected; their genitalia were placed in watersoluble glue on plastic plates pinned below the respective specimens. Beetles were examined using a Nikon SMZ-745T and Nikon SMZ-800 stereomicroscopes. Measurements were taken using an ocular micrometer. Total body length is measured between the anterior angles of the pronotum and the apices of the elytra, width is taken as maximum linear distance between the outer margins of the elytra (Kryzhanovskij & Reichardt, 1976; Lackner, 2016). The following two parameters were also considered: the elytral length (EL) measured along the suture from the basal margin of the elytra to their apices; and the length of the metathoracic wings (WL) measured from the outer (basal) margin of the jugal area to wing apex. Habitus photographs were taken using a Nikon D5100 digital camera with attached Nikon 60 mm 1:2.8G macro lens and Meike Macro Extension Tube Set, and subsequently edited in Adobe Photoshop CS5. Genitalia images were taken with a Nikon Eclipse 50i stereomicroscope with mounted camera from glass slides in glycerol. Figures were prepared using the Photoshop CS5 software. Maps were constructed in Google Maps and subsequently edited in Adobe Photoshop CS5. Terminology for the metathoracic wing venation largely follows Kukalová-Peck & Lawrence (1993) and Newton (2016). TAXONOMY Order COLEOPTERA Family SPHAERITIDAE Genus Sphaerites Duftschmid, 1805 Sphaerites perforatus Gusakov, 2017 (Figs 1 4, 7 9, 10, 11, 14 17, 24) Material examined. China, Yunnan: Laojunshan Mts., NE Liming, 4.2 km S Muzhengdu, 27 6 34 N / 99 45 3 E, 03.VI.2016, H = 3695 m, I. Kabak, G. Davidian leg., 1 male (CDL); Laojunshan Mts., 6.26 km SSW Segengsheng, 27 0 20 N / 99 28 33 E, 06.VI.2016, H = 3575 m, I. Kabak, G. Davidian leg., 1 male (CSR); NNE Wexi City, river tributary of Lapugou Riv., 4.7 km ENE Jizong, 27 27 41 N / 99 23 53 E, 05.VI.2015, H = 3355 m, I. Belousov, I. Kabak, G. Davidian leg., 3 females (CSR); NW of Jianchuan, 6.2 km SW Baiyan Vill., 26 38 31 N / 99 44 36 E, 19.V.2016, H = 3615 m, I. Kabak, G. Davidian leg., 1 male, 1 female (ZIN, CDL). Redescription. Shape and colour. Body broadly oval, moderately convex (Fig.1), length 5.1 5.6 mm, width 2.9 3.7 mm. Dorsum black, with lateral margins of pronotum and elytra dark brown. Antennae reddish, except for greyish-brown antennal club and dark brown first or first and second antennomeres. Maxillary palpomeres dark reddish, terminal palpomere dark brown. Submentum reddish-brown. Pronotum and elytra shiny, with or without very weak bronze lustre. Ventral surface largely dark brown; epipleurae reddishbrown, abdomen black, apex of pygidium reddish. Legs dark brown, with tarsi paler (Figs 1, 2). Tuft of setae on pro- and mesotrochanters yellowish. Head. Dorsal surface covered with dense moderately coarse punctures, without microsculpture. Anterior margin of clypeus narrowly bordered. Eyes small, interocular distance about 9.1 9.3 times as wide as eye width from dorsal view (Fig. 1). Mentum glabrous, with shallow and sparse punctation, about 1.5 1.6 times as wide as long. Submentum semi-circular, about 2.5 times as wide as long; anterior margin strongly arcuate and irregularly elevated. Anten-

317 Figs 1, 2. Sphaerites perforatus, habitus. Dorsal view (1), ventral view (2). Scale bar: 5 mm. Figs 3, 4. Sphaerites perforatus. 3, lateral margin of elytron with denticles (yellow arrow), ventral view; 4, base of elytron with punctate row 6 (yellow arrow), dorsal view. Scale bar: 1 mm.

318 Figs 5 7. Sphaerites spp., metathoracic wing. 5, S. glabratus; 6, S. nitidus; 7, S. perforatus. RP, RP2, RP3+4 radius posterior veins; RP2a, RP2p anterior and posterior branches of radius posterior vein; MP, MP3+4 media posterior veins; CuA2+3+4 cubitus anterior vein; AA3+4 anal anterior vein; J jugal area. Scale bar: 5 mm. nal club compact. Terminal maxillary palpomere almost symmetrical. Thorax. Pronotum about 2.2 times as wide as long. Pronotal punctation (compared to that of head) sparse and shallow on disc, becoming coarser and denser laterally. Lateral margins of pronotum narrowly bordered, border not reaching anterior angle; anterior and posterior margins not bordered. Prosternum, mesoventrite and lateral parts of metaventrite with coarse and shallow punctation. Median area of metaventrite with sparse and very shallow punctation. Prosternum with very long and narrow intercoxal process. Plate of mesoventrite trapezoidal, almost hexagonal, about 1.4 times as long as wide, widest in anterior part near apex, very coarsely and densely punctate (Fig. 2). Elytra asetose, with ten longitudinal rows of punctures; rows 1 to 6 reaching or almost reaching elytral base, 7 to 10 not reaching elytral base; rows 1 to 9 consisting of very large and rather coarse punctures, row 10 consisting of smaller punctures and located almost at lateral edge of elytra. Elytral intervals flat, covered with punctation very similar to that of pronotum (Fig. 1). Humeral and subapical bulges low, but distinct. Elytral epipleuron widest in its anterior part; lateral margin of elytra with very small but distinct teeth in middle and

319 Figs 8, 9. Sphaerites perforatus, aedeagus. Ventral view (8), lateral view (9). Scale bar: 0.5 mm. anterior parts, and with indistinct teeth in posterior part (Fig. 3). Metathoracic wings (Fig. 7) greatly shortened (EL/WL = 1.0 1.03). Legs. Femora with sparse and shallow punctures, without tibial grooves. Pro- and mesotrochanters with dense subapical tuft of erect setae (Fig. 2). Apex of metatrochanters adjoins to femora (Fig. 24). Tarsi with sparse pale setae ventrally; tarsomere 1 as long as tarsomeres 2 and 3 combined; tarsomere 5 about as long as tarsomeres 2 4 combined. Abdomen. Ventrite 1 very short, ventrite 2 distinctly longer than the others, about 1.4 times as long as ventrite 3 (Fig. 2). Pygidium with dense and shallow punctation. Male genitalia (Figs 8, 9). Phallobase asymmetrical. Parameres distinctly longer than phallobase (ratio of their length and phallobase length 2.0 2.2), slightly curved inward subapically, each with membranous apex. Median lobe parallel-sided, slightly narrowed distally, with slightly rounded apex; gonopore situated sub-apically (Fig. 8). Variation. Several specimens have their elytral punctate row 6 not reaching the elytral base and bifurcate near the humeral bulge (Fig. 4). Although all the examined specimens are without metallic shine, the holotype, according to the original description (Gusakov, 2017), is with a very weak bronze lustre on the dorsum. Comparison. Within the genus, S. perforatus is most similar to its Palaearctic congeners S. nitidus, S. glabratus, S. involatilis as well as the Nearctic S. politus in having the elytra black or brownish-black, without reddish-yellow or brownish-red colouration in anterior half. Sphaerites perforatus can be easily distinguished from them by the absence of a distinct metallic shine on the dorsum (at most with a very faint bronze lustre) and by having the larger and much coarser punctures in the elytral rows. Sphaerites dimidiatus and S. opacus differ from all other species, including S. perforatus, by having the anterior half of the elytra reddish-yellow or brownish-red. The important character for identification of the Sphaerites species is the ratio of their elytral length to metathoracic wing length (EL/WL). Sphaerites nitidus and S. glabratus have well developed wings (Figs 5, 6), their EL/WL = 0.5 and 0.6, respectively. Sphaerites perforatus is most similar in this character to S. involatilis and has almost the same ratio (1.0 1.03). Sphaerites involatilis has greatly shortened metathoracic wings, EL/WL = 1.3 (see Gusakov, 2004, fig. 7). The wing venations of Sphaerites are different among species with well-developed wings and species with shortened wings (Figs 5 7). Sphaerites glabratus and S. nitidus have the distinct radius posterior veins (RP2 and RP3+4) reaching the edge of the wing. In S. perforatus, these veins are markedly shortened due to the reduction

320 Figs 10 13. Sphaerites spp., distribution. 10, 11, S. perforatus; 12, 13, S. nitidus. of the wing area. The radius posterior vein (RP3+4) is represented by the short vestige, and the radius posterior vein (RP2) at the edge of the wing forks into two branches, anterior and posterior (RP2a and RP2p), which diverge in almost opposite directions. All the three species have the radius posterior vein (RP), however in S. glabratus and S. nitidus, it is shortened, as about half the length of the media posterior vein and does not reach the base of the wing. Furthermore, in S. perforates, this vein is thinned but reaches the base of the wing. In S. glabratus and S. nitidus, the media posterior vein (MP3+4), cubitus anterior vein (CuA2+3+4) and anal anterior vein (AA3+4) are well pronounced. Two veins, the media posterior vein (MP3+4) and cubitus anterior vein (CuA2+3+4), reach the edge of the wing; the anal anterior vein (AA3+4) is slightly shortened and does not reach the edge of the wing. Sphaerites nitidus has the media posterior vein (MP3+4) with the distinct branch that extends at an acute angle in the direction of the anterior edge of the wing. In S. perforatus, the cubitus anterior vein (CuA2+3+4) and anal anterior vein (AA3+4) are shortened. The anal anterior vein (AA3+4) slightly does not reach the edge of the wing. The media posterior vein (MP3+4) is very thin and in some areas almost indistinguishable. The anterior part of the cubitus anterior vein (CuA2+3+4) is also very thin, the posterior part is almost

321 Figs 14 17. Habitats of Sphaerites perforatus in Yunnan. 14, high mountain forest, 4.7 km ENE of Jizong; 15, high mountain forest, 6.2 km SW of Baiyan Vill.; 16, high mountain forest, 4.2 km S of Muzhengdu; 17, high mountain forest, 6.26 km SSW of Segengsheng. indistinguishable. The jugal area (J) is separated from the rest of the wing by a clear cut in S. glabratus and S. nitidus. Sphaerites perforatus has the almost reduced jugal area. Sphaerites glabratus, S. nitidus, and S. politus have the elytra with the distinct subapical bulges. In S. perforatus, the subapical bulges are distinct, but lower. Based on this character, S. perforatus can be grouped together with S. involatilis which likewise has greatly shortened wings. Sphaerites politus differs from S. perforatus in the apex of the metatrochanter which is spike-shaped and does not adjoin the femora (Fig. 24). On the other hand, the former species has the lateral elytral margin with very distinct denticles in the anterior and middle parts similar to those of S. perforatus (Fig. 3). In other species, the denticles either are absent on the elytra or they are located in different positions. Distribution. Hengduan Mountains (Xue Shan, Dao Chun Shan, Laogu Shan, Yun Ling Shan, and Qingshuila Shan), northern Yunnan, China (Figs 10, 11). Type locality: 4.1 km WNW Yanguangcan (27 15 32 N / 99 24 28 E, 3595 m), NE Weixi City, Yun Ling Shan. Ecology. A terrestrial species inhabiting high-mountain forests at an altitude of 3355 4035 m (Figs 14 17). Beetles were collected on fallen tree trunks at night. Sphaerites nitidus Löbl, 1996 (Figs 12, 13, 18, 19, 20, 21, 23) Material examined. China, Sichuan: WSW Lixian, 5.9 km SW of Shangzhai, 31 20 54 N /

322 Figs 18, 19. Sphaerites nitidus, habitus, dorsal view. Scale bar: 5 mm. Figs 20, 21. Habitats of Sphaerites nitidus in Sichuan. 20, high mountain forest, 5.9 km SW of Shangzhai; 20, high mountain forest, 13.4 km SW of Shangzhai. 102 50 26 E, 15.VI.2016, H = 3025 m, I. Kabak, G. Davidian leg., 2 specimens (ZIN, CSR); WSW Lixian, 13.4 km SW of Shangzhai, 31 17 45 N / 102 47 18 E, 17.VI.2016, H = 3595 m, I. Kabak, G. Davidian leg., 1 specimen (CDL). Diagnosis. Body broadly oval, moderately convex (Figs 18, 19). Length 5.5 5.7 mm, width 3.7 4.0 mm. Black; pronotum and elytra with blue-green, violet-green or greenish metallic shine. Dorsal surface without microsculpture. Elytra with nine longitudinal rows of punctures: rows 1 to 3 shortened anteriorly, 4 and 5 originating near basal edge, 6 to 9 originating behind humeral bulges; an additional, very short

323 Figs 22 24. Sphaerites spp., metatrochanter and base of metafemur. 22, S. politus; 23, S. nitidus; 24, S. perforatus. Scale bar: 0.4 mm. row of punctures lying laterad of humeral bulges. Punctures in rows small and shallow, in rows 1 9 becoming indistinct near elytral apex. Elytral intervals flat, with extremely fine punctation. Humeral and subapical bulges low, but distinct. Apex of metatrochanter adjoining to femur (Fig. 23). Metathoracic wings long (EL/WL = 0.5). Remark. Löbl (1996) provided a very detailed description of the species and illustrated it with photographs of high quality but without the habitus photograph. Thus, the reader is referred to Löbl s publication (1996) for the details. Distribution. Described from the Gongga Mountain, Daxue Shan, Sichuan. According to the examined material, the species occurs in the Hengduan Mountains (Mazhi Shan and Daxue Shan), central Sichuan, China (Figs 12, 13). Ecology. A terrestrial species inhabiting high mountain forests at an altitude of 2800 3600 m (Figs 20, 21). Beetles were collected on fallen tree trunks at night (the similar collecting circumstances were observed for S. perforatus). Key to the species of Sphaerites 1. Elytra black, their anterior half reddish-yellow or brownish-red..................... 2 Elytra black or brownish-black........... 3 2. Elytra without distinct microsculpture, their anterior part from base to middle reddishyellow; border between light and dark parts distinct....................... S. dimidiatus Elytra with distinct microsculpture, their anterior part (about one-seventh of elytron length) brownish-red; border between light and dark parts indistinct, blurred.............................................. S. opacus

324 3. Pronotum and elytra with blue-green, violetgreen or greenish metallic lustre (Figs 18, 19)............................... S. nitidus Pronotum and elytra shiny, either without metallic lustre or with more or less distinct bronze lustre............................ 4 4. Apex of metatrochanter adjoining to femora, not spike-shaped (Figs 23, 24)............ 5 Apex of metatrochanter spike-shaped, not adjoining to femora (Fig. 22)...... S. politus 5. Punctures in elytral rows very large and rather coarse (Fig. 1). Metathoracic wings greatly shortened, about as long as elytra (Fig. 7)....................... S. perforatus Punctures in elytral rows shallow and sparse (Figs 18, 19). Metathoracic wings either markedly longer or markedly shorter than elytra.................................... 6 6. Metafemora not curved. Pygidium and abdominal ventrites with dense and fairly coarse punctation. Metathoracic wings well developed, markedly longer than elytra, EL/ WL = about 0.6 (Fig. 5)........ S. glabratus Metafemora curved. Pygidium and abdominal ventrites with sparse and shallow punctation. Metathoracic wings markedly shorter than elytra, EL/WL = about 1.3............................................. S. involatilis ACKNOWLEDGEMENTS We are very grateful to I.A. Belousov, G.E. Davidian, I.I. Kabak, and A.G. Koval (the All-Russian Institute of Plant Protection, St Petersburg, Russia) for the material treated in the paper (I.I. Kabak also for the photographs of the habitats of Sphaerites perforatus and S. nitidus), V.A. Safonov (the Ecological Laboratory, Baranovichi State University, Baranovichi, Belarus) for preparation of the habitus photographs, T. Lackner (Zoologische Staatssammlung München, Munich, Germany), A.A. Gusakov (the Zoological Museum of Moscow State University, Moscow, Russia) for valuable comments on the manuscript, and to E.G. Karapetova (Belarus State Linguistic University, Minsk, Belarus) for language correction. REFERENCES Gusakov A.A. 2004. A review of species of the family Sphaeritidae (Coleoptera). Euroasian Entomological Journal, 3(3): 179 183. (In Russian). Gusakov A.A. 2017. New species of false clown beetles Sphaerites perforatus (Coleoptera: Sphaeritidae) from the highlands of Yunnan province, China. Gumanitarnoe prostranstvo [Humanitarian space], 6(1): 6 11. (In Russian). Jureček S. 1934. Dvě nové formy palaearktických brouků. Zwei neue palaearktische Käferarten. Časopis Československé Společnosti Entomologické, 31: 45. Kryzhanovskij O.L. & Reichardt A.N. 1976. Beetles of the superfamily Histeroidea (families Sphaeritidae. Histeridae. Synteliidae). Fauna SSSR. Zhestkokrylye [Fauna of the USSR. Coleoptera], 5(4). Leningrad: Nauka. 434 p. (In Russian). Kukalová-Peck J. & Lawrence J.F. 1993. Evolution of the hind wing in Coleoptera. The Canadian Entomologist, 125(2): 181 258. Lackner T. 2016. Re-description of Margarinotus (Paralister) longus (Bickhardt, 1910) with a key to the species of the subgenus Paralister of the genus Margarinotus from the Balkans (Coleoptera, Histeridae, Histerinae). Zootaxa, 4079(1): 103 118. Löbl I. 1996. A new species of Sphaerites (Coleoptera: Sphaeritidae) from China. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 69(2): 195 200. Löbl I. 2015. Family Sphaeritidae. In: Löbl I. & Löbl D. (Eds.). Catalogue of Palaearctic Coleoptera. Hydrophiloidea Staphylinoidea. Revised and updated edition, 2(1): 76. Leiden, Boston: Brill. Löbl I. & Háva J. 2002. A new species of Sphaerites (Coleoptera: Sphaeritidae) from China. Entomological Problems, 32(2): 179 181. Mannerheim C.G. 1846. Nachtrag zur Käfer- Fauna der Aleutischen Inseln und der Insel Sitkha. Bulletin de la Société Impériale des Naturalistes de Moscou, 19(2): 501 516. Newton A.F. 2016. 13.1 Sphaeritidae Shuckard, 1839. In: Beutel R.G. & Leschen R.A.B. (Eds.). Handbook of Zoology, Arthropoda: Insecta. Coleoptera, Beetles. Vol. 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2nd Edition: 274 277. Berlin, Boston: Walter de Gruyter. Received 30 Oct. 2017 / Accepted 28 Nov. 2017 Editorial responsibility: B.M. Kataev