Mannerheimia brevipennis ( Motschulsky, 1860 )

Shavrin, Alexey V., 2021, On the genus Mannerheimia Mäklin, 1880 (Coleoptera: Staphylinidae: Omaliinae: Anthophagini), with taxonomic and faunistic notes on some species, Zootaxa 5040 (3), pp. 301-333 : 319-323

Mannerheimia brevipennis ( Motschulsky, 1860)

( Figs. 4 , 18–19, 23–24 , 40 , 42–56 )

Omalium brevipenne Motschulsky, 1860: 545 Omalium (Omalium) brevipenne: Bernhauer and Schubert 1910: 51 Mannerheimia brevipennis: Hammond 1970: 67 , 70, Muona & Viramo 1986: 15, Herman 2001: 37, Assing 2004: 675, 2007: 5,

Zanetti 2008: 982, 2012: 73, Schatz 2009: 324, Kahlen 2011: 180, Hlavač et al. 2016: 3, Assing & Schülke 2019: 99 Homalium divergens Mäklin, 1878: 26 Omalium (Omalium) divergens: Bernhauer and Schubert 1910: 52 Mannerheimia divergens: J. Sahlberg 1880: 108 , Mäklin 1881: 39, Luze 1905: 254, Jakobson 1908: 460, Bernhauer & Schubert

1910: 63, Frisendahl 1916: 31, Gridelli 1924: 135, 137, Palm 1948: 96, Hammond 1970: 67, 70, Lundberg 1974: 87, Leiler

& Prütz 1977: 132, Kastcheev, 1999: 143, Sokolov 2003: 1272 Phyllodrepoidea divergens: Poppius 1910: 363 Mannerheimia doderoi Gridelli, 1924: 135 , 137, Horion 1963: 101, Lohse 1964: 50, Peez & Kahlen 1977: 129, Hammond 1970:

68, Focarile 1978: 34, 1981: 35, Zanetti, 1987: 277, Koch 1989: 228, Kastcheev 1999: 143 Mannerheimia kirschenblatti Iablokoff-Khnzorian, 1956: 138 ; Kastcheev 1999: 143, Assing 2005: 60 Deliphrum (Mannerheimia) micros Fauvel, 1900: 222 syn. n. Mannerheimia micros: Luze 1905: 255 , Gridelli 1924: 138, Hammond 1970: 68, Kastcheev 1999: 143 Mannerheimia arctica: Shavrin et al. 2008: 82

Type material examined: Cotype of Omalium divergens , ♀: ‘Baklanowskij’ <printed>, ‘Stuxberg’ <printed>, ‘Luze det.’ <printed>, ‘Spec. typ.’ <printed>, ‘ Deliphrum ? | divergens | Mäkl.’ <handwritten>, ‘ Mannerheimia | brevipennis (Motsch.) | Shavrin A.V. det. 2015’ <printed> ( MZHF).

Syntypes of Mannerheimia micros : 1 ♂ [antennomeres 3–11 missing]: ‘Mts. Altai’ <handwritten>, ‘Coll. et det. A. Fauvel’ <printed>, ‘Mannerheimia’ <handwritten>, ‘ micros Fauv. ’ <handwritten>, ‘R.I.Sc.N.B. 17.479’ <print- ed>, ‘Syntype’ <red label with black frame> ( IRSNB); 1 ♀: ‘Mts. Altai Turkestan’ <handwritten>, ‘ Deliphrum mi- cros Fvl.’ <handwritten>, ‘Coll. et det. A. Fauvel’ <printed>, ‘ Mannerheimia micros Fauv. ’ <handwritten>, ‘R.I.Sc. N.B. 17.479’ <printed> ( IRSNB).

Material examined: ITALY: 1 ♀: St. Pierre, M. Fallere. 2300 m a.s.l. 04.08.1978. Rosa leg. < Mannerheimia doderoi Grid. det. Focarile> ( ZFMK); SWEDEN: 1 ♀: Nuolja. 20.07.1927. L. Brundin leg. ( SMNH); 1 ♀: same data, 22.6.1927. ( SMNH); 1 ♂: Abisko. 18.07.1927. L. Brundin leg. ( SMNH); 1 ♀: same data, 14.09.1927 ( SMNH); 1 ♂: Hamra. O. Sjöberg leg. ( MZHF); 1 ♂: Sappisatsi. 03.06.1968. T.E. Leiler leg. ( SMNH); 1 ♂: Lögdö. 03.07.1977. T.E. Leiler leg. ( SMNH); 3 ♂♂: Revsund. 02.04.1967. R. Bergvell leg. ( SMNH); 1 ♂: Bruksvallar. 01.09.1989. T.E. Leiler leg. ( SMNH); 1 ♂: ‘Jamtl.’, ‘J. Sahlb.’ ( MZHF); RUSSIA: YAMALO- NENETS AUTONOMOUS REGION: 1 ♂: Polar Ural, “Kharp” research station, Calamagrostis meadow, herpetobium. 19.07.1971. V. Olshvang leg. ( ZMM, collection of A.L. Tichomirova); 1 ♂: same data amd collector. Salix . 29- 30.06.1971. ( ZMM, collection of A.L. Tichomirova); 2 ♂♂: Priuralskiy District, near Shchuchye trading station, Shchuchye River bank. 11.08- 21.09.1980. E.M. Veselova leg. (cSh, ZMM); 1 ♂: same data and collector. Larix , Pyrola . 12- 14.08.1980. (cSh); 50 ♂♂, 55 ♀♀: same data and collector. 14.08- 21.09.1980. (cR, ZMM, cSh); 4 ♂♂, 4 ♀♀: South Yamal, Sopkey. Larix . 07- 25.08.1980. E.M. Veselova leg. (cSh, ZMM); KHANTY- MANSI AUTONOMOUS REGION: 1 ♀: Sovetskiy District, Malaya Sosva Nature Reserve, near Belaya Gora cordon, 61°.78953’N E 64°52015’E. 69 m a.s.l., mosses, and litter in forest of Pinus sylvestris with Vaccinium vitis-ideae , Ledum palustre , Cladonia spp. , Hylocomium splendens , Pleurozium schreberi etc., with undergrowth of Pinus sibirica & Picea obovata . 24.06.2017. A.B. Ryvkin leg. (cR); 5 ♂♂, 2 ♀♀: same data and collector, left side of Malaya Sosva River up-stream of Shukhtungort cordon, 62°37053’N E 64°08785’. 27 m a.s.l., mosses and litter in spruce forest with Abies sibirica , Pinus sibirica , Betula sp. , Oxalis acetosella , Maianthemum bifolium , Hylocomium splendens , Pleurozium schreberi etc. 21.08.2018. ( ZMM, cSh); 1 ♂, 1 ♀: same data and collector, Malaya Sosva Nature Reserve (buffer zone), right side of Malaya Sosva River below Zapadnyi (=Western) cordon, N 61°90686’E 63°.69856’. 83 m a.s.l., mosses and litter in flood-plain forest with Picea obovata , Larix sibirica , Pinus sylvestris , P. sibirica , Betula sp. , Vaccinium vitis-idaea , Equisetum pratense , Pyrola spp. , Poaceae gen. spp., Carex sp. , Maianthemum bifolium , Trientalis europaea , Gymnocarpium sp. , Lycopodium sp. , Lonicera sp. , Hylocomium splendens , Pleurozium schreberi etc. 14.07.2017. (cR); 1 ♀: Sur- gutskiy District, Yuganskiy Nature Reserve, Nyogus`yakh River, Bisarkina cordon. Mosses and litter on slope (from stream to peak) near cordon: Abies , Picea , Pinus sibirica , Betula , aspen, Sorbus , Vaccinium vitis -idaea, V. myrtillus , Equisetum sylvaticum , Pleurozium schreberi , Hylocomium splendens , Maianthemum , Trientalis , Carex ? globularis, grasses etc. 22.07.1998. A.B. Ryvkin leg. ( ZMM); 1 ♀: ‘Samarovo’, ‘Sundman’, ‘ Mannerheimia n.sp. Unicum’, ‘18’’ ( MZHF); 1 ♀: ‘Surgutskoi’, ‘Ndsk. & Stbg’ ( MZHF); TAYMYR AUTONOMOUS REGION: 2 ♂♂: ‘Dudinka’, ‘Wuorentaus’ ( MZHF); 5 ♀♀: ‘Tolstoinos’, ‘J. Sahlb.’ ( IRSNB, MZHF); 1 ♀: ‘Tolstoinos 70°10’’ ( SMNH); 1 ♀: ‘Tolstoinos J. Sahlb.’, ‘Sibérie Jénisséi’, ‘ divergens Mäkl. (vicarium J. Sahlb. in litt.)’ ( IRSNB); KEMEROVO AREA: 17 ♂♂, 8 ♀♀: Kuznetskiy Alatau Reserve, Mt. Bolshaya Tserkovnaya, SSW Belogorsk. 1300-1400 m a.s.l., upper timberline. 26.08.1993. S. Golovatch & V. Gratchev leg. (cSh, ZMM); KRASNOYARSK TERRITORY: 1 ♂: 18 km E Norilsk, Talnakh. 150 m a.s.l. 25.06.2003. S.V. Saluk leg. (cSh); 2 ♂♂: Turukhanskiy District, Yeloguy Nature Ref- uge, Yeloguy River, 6 km below Tyna River mouth. Dried channel of a rill with Alnus sp. and Betula sp. , in leaf litter. 27.07.1992. V.B. Semenov leg. ( ZMM); 3 ♂♂, 3 ♀♀: same District and collector, Yeloguy River, 125 km bel- low Kellog, 15 km below mouth of Altus River. 35 m a.s.l. moss and litter in fir-spruce forest with Betula , sparse Pinus sibirica , Pleurozium schreberi , Hylocomium splendens . 17.10.1993. (cSh, ZMM); 1 ♂, 1 ♀: same District, Mirnoye. 15.08.1989. L.B. Rybalov leg. (cSh); 11 ♂♂, 8 ♀♀: same District, Podkamennaya Tunguska River, Sukhaya River, mouth of Lebyazhya River. 60 m a.s.l., mosses and litter under Picea , Abies , Betula with Vaccinium myrtillus , V. vitis -idaea, Pleurozium , Hylocomium , Polytrichum commune , Sphagnum ? girgensohni etc. 29.09.1990. A.B. Ryvrin leg. (cSh, ZMM); 2 ♂♂, 1 ♀: same District, Kamennyi Dubches River, 3 km below mouth of Teulches River. Mosses in forest with Pinus sibirica , Larix , Abies , Betula , Vaccinium vitis -idaea, V. myrtillus , Hylocomium , Pleurozium , Cladonia spp. 18.09.1992. V.B. Semenov leg. (cSh); 4 ♂♂, 2 ♀♀: same data and collector, 60 km be- low Teulches River, near hut. 150 m a.s.l., Pinus sibirica - Abies forest, with Betula , Polytrichum commune , Pleurozium and Hylocomium . 22.09.1992. (cSh, ZMM); 2 ♀♀: same locality, date and collector. 150 m a.s.l., lake, con- nected by stream with river, Carex on tussocks, Filipendula , Sphagnum , Mnium , and in Salix litter ( ZMM); 1 ♂: ‘Baklanowskij’, ‘Sibir Jeniseie’, ‘Stuxb.’, ‘divergens’ ( SMNH); 1 ♀: ‘Seliwaninskoj’, ‘Sibir Jeniseie’, ‘Stuxb.’ ( SMNH); 2 ♂♂: Turukhanskiy District, Central Siberian State Reserve, Bol`shaya Varlamovka River basin, middle reaches of Bolshaya Raskosaya River. Pleurozium schreberi , Hylocomium splendens etc. in forest with Picea obovata , Betula sp. , Pinus sibirica , Abies sibirica , Juniperus sp. , Larix sibirica , Vaccinium vitis-idaea and V. myrtillus . 24.05.1992. V.B. Semenov leg. (cSh); 1 ♂: same locality and collector. 23.05.1992. (cSh); 2 ♂♂, 1 ♀: same local- ity and collector. Under bark of Picea obovata . 27.05.1992. ( ZMM); 1 ♂: same locality and collector. In Polytrichum commune , Hylocomium splendens , Pleurozium schreberi , and in birch leaf litter in forest with Betula sp. , Abies sibirica , old trees of Pinus sibirica , undergrowth of Picea obovata , Vaccinium myrtillus , Empetrum sp. 27.05.1992. (cSh); EVENKIA: 4 ♂♂: Baykitskiy District, Central Siberian Biosphere Reserve, Stolbovaya River basin: Birapchana River below Khorlokta River mouth. 360 m a.s.l., litter and mosses ( Pleurozium , Hylocomium , Dicranum etc.) on slope with Picea , Abies , Betula etc. 07.07.1989. A.B. Ryvkin leg. (cSh, ZMM); 3 ♂♂, 1 ♀: same data and collector, Stolbovaya River basin: Kulingna River mouth of right confluent, 6 km below of Usas River mouth. 90 m a.s.l., slope forest with Betula , Picea , Pinus sibirica , Ledum , Lonicera , Spiraea , Vaccinium vitis -idaea, Pleurozium , Hylocomium , Polytrichum commune , Rubus saxatilis etc. 11.09.1989. (cSh, ZMM); 1 ♂, 3 ♀♀: same data and collector, Stolbovaya River basin: lower reaches of Birapchana River near mouth of Krutenkiy Stream. 130 m a.s.l., mosses and litter on slope with Picea obovata , Abies sibirica , Ledum , Hylocomium , Pleurozium , Pyrola , Linnaea etc. 23.07.1991. (cSh, ZMM); 3 ♀♀: same district and collector, basin of Velmo River, Svetlana River, 2-4 km up-stream of river mouth. Mosses and litter, slope forest with Picea , Alnus , Pinus sibirica , Betula , Abies , Pyrola , Trientalis , Majanthemum , Vaccinium vitis -idaea, V. myrtillus etc. 14.08.1990. ( ZMM); 1 ♀: same District and collector, Podkamennaya Tunguska River, 1 km up-stream Sulomay. 70 m a.s.l., litter and mosses on slope near stream, Alnus , Betula , Picea , etc. 02.10.1989. ( ZMM); 5 ♂♂, 3 ♀♀: same District and collector, Bakhta River near mouth of Severnaya Kachikangna River. 470 m a.s.l., mosses and litter under Picea , Betula , Pinus sibirica with Vaccinium uliginosum , V. vitis -idaea, Linnaea , Pleurozium , Hylocomium , etc. 17.08.1991. ( ZMM, cSh); ALTAI: 1 ♂: Kosh-Ashachskiy District, 7 km NE Chagan-Uzun. Bank of mountain stream, traps. 13.06.1989. A. Pisanenko leg. ( ZMM); 1 ♀: near Onguday. Forest litter. 10.06.1989. S. Saluk leg. (cSh); 3 ♂♂, 1 ♀: Korgon Mts., upper reaches of Mokhnatyi Stream (confluent of Kumir River). 1750 m a.s.l. 10- 24.07.1984. V.G. Shilenkov leg. (cSh); 2 ♀♀: ‘ Altai Central’ ( IRSNB); KHAKASSIA: 1 ♂, 1 ♀: Tashtypskiy District, near Zapadnosayanskiy Pass, sources of Bolshoy On River. 2000 m a.s.l., upper forest border, litter and moss under Pinus sibirica with Linnaea , Vaccinium vitis -idaea etc. 15.06.1990. A.B. Ryvkin leg. (cR); TUVA: 1 ♀: near Zhoigan Pass, 52°35’10’’N 98°47’57’’. 1929 m a.s.l. 28.06.2016. I.V. Enushchenko leg. ( CS); 3 ♂♂: left side of Arzhan-Khem River, 52°33’48’’N 98°45’47’’. 1341 m a.s.l. 30.06.2016. I.V. Enushchenko leg. (cSh); BURYATIA: 10 ♂♂, 5 ♀♀: Okinskiy District, Pik Topografov, valley of Khaelgin River, 52°31’05’’N 98°50’02’’E. 2135 m a.s.l. 19.07.2015. I.V. Enushchenko leg. (cSh); 3 ♂♂, 3 ♀♀: same data and collector, 52°31’13’’N 98°49’44’’E. 2172 m a.s.l., near glacier. 24.06.2017. (cSh); 18 ♂♂, 3 ♀♀: same data and collector, right side of Khaelgin River. Thickets of Salix , in litter. 20.07.2015. (cSh); 1 ♂: same data and collector, 52°30’08’’N 98°49’26’’E. 2117 m a.s.l. 21.08.2017. (cSh); 3 ♂♂: same data and collector, south slope of Pyatiozyornyi Pass, 52°31’49’’N 98°49’02’’E. 20.07.2015. (cSh); 5 ♂♂, 1 ♀: same data and collector, left side of Daergyl River, 52°34’18’’N 98°56’18’’E. 24.07.2015. (cSh); 7 ♂♂, 3 ♀♀: same data and collector, Sherpov Pass, 52°31’44’’N 98°52’93’’E. 18.07.2015. (cSh); 5 ♂♂: same data and collector, left side of Dunda-Gol River, 52°35’6’’N 98°50’5’’E. 27.06.2017. (cSh); 1 ♂: same District, near Kropotkin Volcano. 2070 m a.s.l. 14- 17.07.1998. A.V. Shavrin leg. (cSh); YAKUTIA: 2 ♂: Central Yakutia, Spasskaya Pad` Forest Station. Larix , soil sample: layer 0-5 cm. 26.07.2002. IBPC SB RAS (cSh); SIBERIAN REGIONS NOT LOCATED: 2 ♀♀: ‘Tun- guska med.’, ‘Sibir Jeniseie’ ( SMNH); 4 ♂♂, 5 ♀♀: ‘Sibir Jeniseie’ ( SMNH); TURKEY: 1 ♂: Elazið Province, Merkez/Harput, environs of Obuz. 23.05.2013. E.A. Khachikov leg. (cKh); IRAN: 1 ♂, 1 ♀: Esfahan Province, 15 km NNO Semirom, Mt. Aljud, 31°32’09’’N 051°37’23’’E. 2650 m a.s.l. 12.05.2007. J. Frisch & S. Serri leg. ( MNB); KAZAKHSTAN: 1 ♂, 1 ♀: south of Alma-Ata, Zailiyskiy Ala-Tau, Bol`shoye Alma-Atinskoye Lake. 2600 m a.s.l., Picea schrenkiana forest. 29.08- 02.09.1993. K. Es`kov leg. ( ZMM); 1 ♂: south of Alma-Ata, upper reaches of Bol`shaya Almatinka River. 2300-2500 m a.s.l., Picea schrenkiana forest. 08.06.1993. S. Golovatch leg. ( ZMM); 1 ♂, 3 ♀♀: Altai, confluence of Karakoba and Tautekeli rivers. 06.08.1989. V. Kastcheev leg. ( ZIN); 3 ♂♂: Rakhmanovskiye Klyuchi. 14- 16.06.1980. V. Kastcheev leg. ( ZIN); 1 ♂: West Tian Shan, Chimkent Region, Aksu- Zhabagly Nature Reserve, Ulken-Kaindi, 70°35’N 42°23’E. 2100 m a.s.l., grubbing at snow melt. 11.06.1999. J. Cooter leg. ( OUMNH); 1 ♂, 1 ♀: same data and collector. 2200 m a.s.l., in bear dung ( OUMNH, cSh); 9 ♂♂, 5 ♀♀: same data and collector, Kishi-Kaindi, 70°35’N 42°24’E. 2000 m a.s.l, Betula , Juniperus litter. 18.06.1999. ( OUMNH); 1 ♂, 1 ♀: same data and collector, Kalbaidjailau, 70°42’N 42°25’E. 2500 m a.s.l., grubbing near snow melt. 16.06.1999. (cSh, OUMNH); 1 ♀: same Nature Reserve. 1900 m a.s.l., Juniperus seravschanica . 09.05.1966. R.N. Fisechko leg. ( ZIN); 1 ♂, 1 ♀: Ketmen Mts., Bolshoy Aksu Gorge. 2300 m a.s.l. 22- 23.07.1991. S. Saluk leg. (cSh); 1 ♂, 1 ♀: Kungey Alatau, Kulbastau Gorge. 01- 06.06.1988. V. Kastcheev leg. ( ZIN); 6 ♂♂, 7 ♀♀: same data and collector. 20- 27.7.1988 ( ZIN); 1 ♀: same data and collector, Kulbastau River, pass. 22.07.1987. ( ZIN); 1 ♂: Karatau Mt., Zhamantas River, 43°52’05’’ 68°11’55’’. 731 m a.s.l. 10.05.2010. A.V. Ivanov leg. (cSh); 2 ♂♂, 2 ♀♀: Karatau Mts. (plateau), 36 km W Karatau Mt., 43°10’09’’N 69°56’32’’E. 1130 m a.s.l. 06.05.2015. A.S. Prosvirov leg. (cSh); 1 ♀: NW slope of Karatau Mt. range, 30 km WNW Karatau Mt. 43°12’40’’N 70°03’43’’E. 815 m a.s.l. 04.05.2015. A.S. Prosvirov leg. (cSh); 1 ♂: Karatau, Mikhailovka, bank of Aktau River. 16.05.1974. S.Yu. Gryuntal leg. ( ZMM); TAJIKISTAN: 1 ♀: Kondar Gorge, tract Kvak. 2000 m a.s.l., near glacier. 20.05.1988. V.G. Shilenkov leg. (cSh); KYRGYZSTAN: 2 ♂♂, 1 ♀: Terskey Ala-Too, north part of Non-Kilznyagu Gorge, Ailan. 2950 m a.s.l., Picea , Caragana . 29.06.1988. V. Yanushev leg. ( ZMM); TURKMENISTAN: 1 ♂: Kopet-Dagh Mts., Dushak, 57°54‘N 37°57‘E. 2200 m a.s.l. 01- 02.10.1991. A. Podlussány, L. Ronkay & Z. Varga leg. ( HNHM); CHINA: 1 ♂: Xinjiang, W Erenhaberga Shan, road Narat-Kuytun Pass, 90 km SWS Kuytun (tunnel env.). 4000 m a.s.l. 20- 21.7.1993 J. Kaláb leg. (cZan).

Redescription. Measurements (n=200): HW: 0.44–0.57; HL: 0.29–0.43; OL: 0.12–0.17; AL: 0.82–1.15; PL: 0.39–0.52; PW: 0.54–0.86; EL: 0.60–0.92; EW: 0.70–1.37; AW: 1.06–1.33; MTbL (averaged): 0.50; MTrL (averaged): 0.27 (MTrL 1–4: 0.17; MTrL 5: 0.10); AedL: 0.42–0.60; TL: 2.05–3.70.

Body moderately small, slightly or distinctly convex, wide and short or elongate ( Fig. 4 ). Coloration yellowbrown to reddish-brown, usually with darker head, disc of pronotum and abdomen (specimens from Iran with yellow-brown elytra, specimen (teneral) from Turkey with yellow body except brown head); mouthparts, antennae and legs yellow to yellow-brown (sometimes antennomeres 1–2 or 1–4 yellow). Forebody without microsculpture except for traces of very fine diagonal meshes in latero-apical portion of head between bases of antennae and anterior margins of eyes and/or fine transverse microreticulation on clypeus in some specimens, and scutellum with fine transverse meshes (some specimens from Kazakhstan without punctation on head or with fine transverse meshes in middle and dense isodiametric microreticulation in basal portion of head and neck); abdomen with distinct isodiametric microsculpture. Head with irregular, fine or moderately large punctation, denser in middle and/or laterobasal portions (sometimes middle portion with very sparse or denser and coarser (some specimens from Kyrgyzstan) punctation); punctation of neck with sparse or dense; punctation of pronotum about as that on head, but slightly larger and deeper, sparser in middle, sometimes with impuncated area in mediobasal third (specimens from Iran and some specimens from Kazakhstan with finer and sparser punctation; some specimens from Kyrgyzstan with denser and coarser punctation); scutellum with or without several fine punctures; punctation of elytra as that on pronotum, or larger and deeper, middle portions of each elytron with tangled punctures or with indistinct four to six longitudinal rows of punctures; abdominal tergites without visible punctation or with fine punctures.

Head 1.3–1.5 times as broad as long, slightly or distinctly elevated in median portion, with or without short and moderately deep anteocellar foveae. Ocelli small or moderately large, located about level of posterior margins of eyes; distance between ocelli variable: about as distance between ocellus and posterior margin of eyes, slightly shorter or larger. Apical segment of maxillary palpi twice as long as preceding segment or slightly longer. Antenna moderately short ( Figs. 18–19 ) reaching apical margin of basal portion of elytra when reclined, with transverse (sometimes significantly) antennomeres 4–10 or 5–10; antennomere 2 narrower and shorter than basal antennomere, 3 about as long as and as broad as 2 or slightly narrower, 4 about twice shorter than 3, transverse or subovoid, 5 about as long as or slightly shorter and broader than 4, 6 slightly broader than 5, 7–8 slightly broader than 6, 9–10 broader than 8.

Pronotum 1.3–1.6 times as broad as long, 1.2–1.5 times as broad as head, widest in middle, usually evenly narrowed both posteriad and anteriad ( Fig. 23 ), but sometimes slightly more narrowed posteriad ( Fig. 24 ); apical margin somewhat straight or rounded; anterior margins widely rounded, usually not protruded anteriad; posterior angles widely rounded; lateral portion sometimes with indistinct laterobasal impressions.

Elytra distinctly broader than long, slightly or significantly widened apicad, 1.5–1.7 times as long as pronotum; apical margins straight or widely rounded.

Male. Antenna as in Fig. 18 . Apical margin of abdominal tergite VIII somewhat straight or slightly concave. Apical margin of abdominal sternite VIII usually deeply concave. Aedeagus with wide basal portion, gradually narrowed toward truncate ( Fig. 48 ), rounded ( Figs. 50–51, 53 ) or moderately narrow, subacute apex ( Figs. 52, 54–56 ), latero-apical portions sometimes forming wide projections ( Figs. 55–56 ); parameres widened in apical portion, exceeding apex of median lobe ( Fig. 48 ). Lateral aspect of aedeagus as in Fig. 49 .

Female. Antenna as in Fig. 19 . Apical margin of abdominal tergite VIII straight or rounded. Apical margin of abdominal sternite VIII rounded.

Comparative notes. Regarding the general shapes of the body and the aedeagus, M. brevipennis is similar to M. arctica and M. pleshanovi sp.n. From M. arctica it differs by widely rounded posterior angles of the pronotum and from M. pleshanovi sp.n. by narrower and longer elytra, the presence of hind wings and tomentose wing-folding patches in the middle of the abdominal tergite V. From both species it can be distinguished by transverse antennomeres 4–10 and the wider apical part of the median lobe. Besides that, the general shape of the body and aedeagus of M. brevipennis is similar to M. aprutiana from Italy, from which it can be distinguished only by the shape of the broader preapical antennomere ( Zanetti 1987).

Distribution. Mannerheimia brevipennis is distributed in the Palaearctic Region from Scandinavia to Eastern Siberia (with two wide disjunctions between North Europe and north-western Siberia, and basin of Yenisey River and central Yakutia) and from Turkey and Iran to Middle Asia, north-western China and the mountains of southern Siberia; several populations are located in the Alps ( Fig. 40 ). Apparently, the record from “Himalaya” (e.g. Tikhomirova 1973, Schülke & Smetana 2015) is based on misidentification.

Bionomics. Mannerheimia brevipennis inhabits mixed forests with Pinus , Abies , Picea , Larix , Betula , Alnus ( Juniperus , Caragana in Middle Asia) and sometimes flood-plain forests with Salix , Betula , etc. It is known also from the alpine zone. It most often inhabits mosses and litter in vegetation with Vaccinium , Equisetum , Hylocomium , Pleurozium , Carex , etc. ( Cedrus , Juniperus , Picea in Turkey ( Assing 2004, 2007)). Occasionally, it was collected under bark, near glaciers under stones, and in bear dung. The species is recorded from elevations between 27 m a.s.l. to 4000 m a.s.l. Based on published and new data, the beetles were observed during the period from April to October, with the greatest number of specimens captured in July-August. Specimens from Shchutchye Lake (Yamal) were collected together with M. arctica , and specimens from Kulbastau ( Kazakhstan) together with M. asiatica .

Remarks. Omalium brevipenne was originally described without citing of exact locality. Hammond (1970) studied the type specimen from ZMM, designated the lectotype, transferred the species to Mannerheimia and noted that “...the specimen is clearly labelled as originating from Mt. Ula-Tau, in the Kazakh Uplands and not far to the west of the Altais”. Sahlberg (1880) transferred Homalium divergens to Mannerheimia . Mannerheimia doderoi was originally described from “Valsavaranche (Valle d`Aosta), Alpi Graie”. Zanetti (2008) synonymized it with M. brevipennis . Mannerheimia kirschenblatti was described from Armenia. Assing (2005) synonymized it with M. brevipennis . Deliphrum (Mannerheimia) micros was originally described based on two specimens from “Monts Alai”. I studied both types, which regarding external morphological features are similar to other specimens of M. brevipennis and can be distinguished from them only by denser and coarser punctation of the forebody. The aedeagus of the specimen from “Ala-Too” ( Fig. 53 ) is also similar to that of some specimens of M. brevipennis . Consequently, M. micros was synonymized with the latter name.

Based on Central European keys (e.g. Lohse 1964, Zanetti 2012), M. brevipennis differs from the related M. arctica by the coloration, a smaller body, the shape of transverse preapical antennomere ( Figs. 17–18 ), rounded laterobasal margins of the pronotum (as in figure 3 in Zanetti (2012) or Fig. 23 ), and the shape of the aedeagus. The study of huge material of M. brevipennis revealed that body size, coloration, and the shape of laterobasal margin of the pronotum are variable. The coloration of the body varies from reddish-brown to pale in specimens from southern localities of the distribution. The punctation of head and pronotum may be fine and sparse or large, deep and dense. Assing (2004) noted for Turkish populations of M. brevipennis that the punctation of head, as well as the punctation and the shape of the pronotum were variable, and that these specimens have a “...more distinctly bicoloured forebody and antennae, as well as the lower average size” in comparison with European specimens. The shape of the pronotum of some Siberian specimens is somewhat similar to that of some specimens of M. arctica , which have sinuate laterobasal margins in front of posterior angles ( Fig. 24 and Figs. 20–22 ). The internal structure of the aedeagus is very similar to that of M. arctica and other related species ( Figs. 42 , 48 , 57 ). Sometimes, the external morphology of the aedeagus is similar to that of M. arctica ( Fig. 46 and Fig. 52 ), but a study of series of aedeagi showed more diverse shapes of the apical portion of the median lobe in M. brevipennis . The apex of the aedeagus in M. brevipennis is widely rounded ( Figs. 50, 51, 53 ), truncated ( Fig. 48 ), or somewhat narrow ( Figs. 54–56 and figure 178 in Assing (2004)). The latero-apical portions of the median lobe sometimes form wide latero-apical projections ( Figs. 55–56 ). These modifications can be observed also in M. arctica . In conclusion, the shapes of the antennomeres are the most reliable feature for the separation of these species: antennomeres 4–10 are transverse in M. brevipennis ( Figs. 18–19 ) and elongate in M. arctica ( Figs. 16–17 ). The unusual distribution of M. brevipennis ( Fig. 40 ) somewhat coincides with that of M. arctica ( Fig. 39 ), with wide disjunctions in some regions, suggesting that these species are potential objects for the forthcoming modern studies using molecular methods which can lead to interesting results, including possible taxonomic changes, and for the clarification of the taxonomic status of related species (e.g. M. aprutiana ).

Figures

FIGURES 1–9. Habitus of Mannerheimia: 1—M. afghanica (paratype), 2—M. arctica, 3—M. asiatica, 4—M. brevipennis, 5—M. japonica, 6—M. obscura, 7—M. pleshanovi (paratype), 8—M. yasudai, 9—M. zanettii (paratype). Scale bar: 1.0 mm.

FIGURES 16–25. Antenna (Figs. 16–19) and pronotum (Figs. 20–25) of Mannerheimia arctica (Figs. 16–17, 20–22), M. brevipennis (Figs. 18–19, 23–24) and M. pleshanovi (Fig. 25): 16, 18—right antenna of male, 17, 19—right antenna of female, 20—Irkutsk Area: Vitimskiy Nature Reserve, 21—Krasnoyarsk Territory: Potapovo, 22—Murmansk Area: Khibiny, 23—Krasnoyarsk Territory: Lebyazhya River, 24—China (Xinjiang), 25—Irkutsk Area: Khamar-Daban (paratype). Scale bar: 0.1 mm.

FIGURES 39–40. Distribution of Mannerheimia arctica (Fig. 39) and M. brevipennis (Fig. 40). Black circles: material studied by authors, white circles: previously published material not seen by author.

FIGURES 42–47. Aedeagus of Mannerheimia arctica (42: Buryatia: Gitara Lake, 43–44: Murmansk Area: Khibiny, 45: Tymen Area: Sob` St., 46: Yamal: Shchuchye, 47: Kemerovo Area: Belogorsk): 42–43, 45–47—parameral view, 44—lateral view. Scale bar: 0.1 mm.

FIGURES 48–56.Aedeagus of Mannerheimia brevipennis (48–49: Kazakhstan:Aksu, 50: Krasnoyarsk Territory:Turukhanskiy District, 51: Buryatia: Okinskiy District, 52: Yamal: Sopkey; 53: Kyrgyzstan:Ala Too; 54: Yakutia: Spasskiy; 55: Iran: Esfahan; 56: China: Xinjiang): 48, 50–56—parameral view, 49—lateral view. Scale bar: 0.1 mm.

FIGURES 57–62. Aedeagus of Mannerheimia asiatica (Figs. 57–58), M. japonica (Figs. 59–60), M. obscura (Figs. 61–62): 57, 59, 61—parameral view, 58, 60, 62—lateral view. Scale bar: 0.1 mm.