Mannerheimia arctica ( Erichson, 1840 )

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 : 308-315

Mannerheimia arctica ( Erichson, 1840)

( Figs. 2 , 12–17, 20–22 , 26–47 )

Deliphrum arcticum Erichson, 1840: 873

Deliphrum arcticum: Thomson 1861: 198 , J. Sahlberg 1871: 424, 1876: 214, Eppelsheim 1875: 409, Seidlitz 1875: 236

Deliphrum (Mannerheimia) arcticum: Sparre-Schneider 1888: 172

Mannerheimia arctica: J. Sahlberg 1880: 108 , Ganglbauer 1895: 725, Sparre-Schneider 1910: 102, Dodero 1908: 95, Luze 1905: 253, Poppius 1905: 153, 1907: 314, 1909: 15, 1910: 362, Jakobson 1908: 460, Ammann & Knabl 1912: 72, Gridelli, 1924: 135, 136, Jansson 1926: 914, Porta 1926: 24, Lindberg 1927: 31, Strand & Hanssen 1932: 20, Brundin 1934: 276, Lindroth 1935: 31, Stenius 1936: 9, Renkonen 1938: 73, Holdhaus & Lindroth 1939: 150, Strand 1946: 216, Palm 1948: 96, 1949: 67, Wörndle 1950: 126, Linder 1953: 65, Krogerus 1960: 30, Horion 1963: 100, Handschin 1963: 54, Lohse 1964: 50, Coope 1969: 100, Holmström 1972: 168, Focarile 1978: 34, Comellini 1974: 515, Lundberg 1977: 92, Peez & Kahlen 1977: 129, Muona & Viramo 1986: 15, Kahlen 1987: 218, Zanetti 1987: 274, 2012: 73, Schatz 1988: 271, Koch 1989: 228, Siitonen 1993: 230, Lagunov & Novozhenov 1996: 27, Olsvik et al. 2001: 21, Franzen & Molander 2011: 111, Schneider et al. 2012: 43

<?!> Mannerheimia arctica: Kastcheev 1999: 143 , Kastcheev & Ishkov 2001: 101, Ryabukhin 2002: 198, Gabdullina 2016: 58 Homalium affine Mäklin, 1878: 27

Mannerheimia affinis: Mäklin, 1880b: 40

Homalium confusum Mäklin, 1878: 27

Mannerheimia confusa: Mäklin, 1880b: 40

Homalium saginatum Mäklin, 1878: 28

Mannerheimia saginata: Mäklin, 1880b: 41

Type material examined: Type of Homalium affine Mäklin, 1878 , ♂: ‘ Fl. Jenisej’ <printed>, ‘ Lt 68°45’’ <printed>, ‘ Ndsk & Stbg’ <printed>, ‘affine Maklin’ <printed>, ‘7455 E91’ <printed> ( SMNH) .

Type of Homalium confusum Mäklin, 1878 , ♀: ‘Fl. Jenissej’ <printed>, ‘Lt. 68°3’’ <handwritten>, ‘Ndsk & Stbg’ <printed>, ‘confusum’ <printed>, ‘Mäklin’ <handwritten>, ‘7457 E91+’ <printed>, ‘ arctica Er. ’ <handwritten> ( SMNH).

Type of Homalium sagunatum Mäklin, 1878 , ♀: ‘ Kr [a]sn[o]j[arsk]-T[o]msk’ <printed>, ‘ Sibiria’ <printed>, ‘ Ndsk & Stbg. ’ <printed>, ‘saginatum Mäklin’ <printed>, ‘7456 E61’ <printed> ( SMNH) .

Material examined: ITALY: 1 ♀: Stelvio. 13.07.1905 ( NMPC); 4 ♀♀: Stilfserj. L. Ganglbauer leg. ( MZHF, NMPC); NORWAY: 1 ♂, 1 ♀: Syd Varanger. T.G. Münster leg. ( NMPC); 1 ♀: same locality, B. Poppius leg. ( MZHF); 4 ♂♂: Kalvhau [Kalvåg]. A. Strand leg. ( NMPC, MZHF); 1 ♂: ‘Norvegia bor. Lysholm’ ( MZHF); SWE- DEN: 1 ♂: Abisko. E. Klefbeck leg. ( ZML); 1 ♀: same locality. 17.07.1949. O. Lundblad leg. ( SMNH); 1 ♀: same locality and collector. 20.07.1949. ( SMNH); 3 exs.: same locality. 08.1948. Pfeiffer leg. ( SNMB); 1 ♂: same data, Torne trask. E. Selman leg. ( MZHF); 6 ♂♂, 2 ♀♀: same locality. L. Brundin leg. ( MZHF, NMPC); 1 ♀: Karesu- ando. 08- 26.06.1935. N. Bruce leg. ( SMNH); 1 ♀: Luopakte 21.07.1949. O. Lundblad leg. ( SMNH); FINLAND: 4 ♂♂, 1 ♀: Kuhmo, “Friendship park”. Pitfall traps. 1998-1999. L.B. Rybalov leg. ( ZMM, cSh); 1 ♀: Suomussalmi ( NMPC); 1 ♂, 2 exs.: same locality. O. Sorsakovski leg. ( SNMB); 1 ♀: Ivalojoki ( NMPC); 2 ♂♂: Ivalo. E. Thune- berg ( NMPC); 1 ex.: Petsamo. 04.07.1929. H. Lindberg leg. ( ZIN); 1 ♂, 1 ♀: Saariselkä. B. Poppius leg. ( NMPC); 1 ♂: Ruhtinassalmi. O. Sorsakovski leg. ( NMPC); SCANDINAVIAN REGIONS NOT LOCATED: 1 ♂: ‘Lapponia Reitter’ ( SNMB); 3 exs.: ‘Lapponia’ ( ZIN); 1 ♀: ‘Lapponi J. Sahlb.’ ( IRSNB); 1 ♂: ‘Lapponia’, ‘J. Sahlb.’, ‘Rambousek Collection’ ( NMPC); 1 ♂: ‘Lp. b. Bhn.’ ( SMNH); RUSSIA: MURMANSK AREA: 185 ♂♂, 65 ♀♀: Khibiny, near Kirovsk, Botanical Garden. 13- 23.09.1986. T. Rossolimo leg. (cR, ZMM, cSh); 1 ♂, 1 ♀: Porjeguba. J. Sahlberg leg. ( IRSNB); ARKHANGELSK AREA: 1 ex.: Velsk. 16.05.1902. D.V. Pomerantsev leg. ( ZIN); 1 ♂: ‘Fl. Mesen’, ‘B. Poppius’ ( MZHF); 1 ♀: ‘Solovetsk’, ‘Edgren’ ( MZHF); NENETS AUTONOMOUS REGION: 44 ♂♂, 12 ♀♀: Kanin. B. Poppius leg. ( MZHF); YAMALO- NENETS AUTONOMOUS REGION: 1 ♂: “ОбдорскаЯ тундра, Тобольск. губ., Ф. Зайцев [Obdor tundra, Tobolskaya guberniya, F. Zaitsev leg.] 18 vi [19]09’ ( MZHF); 1 ♂: same data and collector, but 05.07.1909 ( MZHF); 1 ♂: Priuralskiy District, near Shchuchye trading station. Picea abies, Vaccinum vittisidaea, Pleurozium schreberi . 26.07.1980. E.M. Veselova leg. ( ZMM); 2 ♂♂, 1 ♀: same data and collector. 08.08.1980. ( ZMM); 1 ♂: same data and collector. Larix forest, mosses. 12- 14.08.1980. (cR); 2 ♂♂, 2 ♀♀: same data and collector. Alder forest on the border with Carex swamp. 14.08- 21.09.1980. ( ZMM, cSh); 38 ♂♂, 13 ♀♀: same data and collector. Larix , Pyrola . 14.08- 21.09.1980. (cR, ZMM, cSh); 1 ♀: same locality, floodplain of Shchuchya River. 23.07- 07.08.1980. P.K. Eryomin leg. ( ZMM); 1 ♀: same data. Myosotis . 12- 24.08.1980. T.R. Andreeva leg. ( ZMM); 1 ♂: same data, Shchuchya River, 15 km downstream of Shchuchye trading station. Larix . 07- 25.08.1980. T.R. Andreeva leg. ( ZMM); 2 ♂♂, 1 ♀: Labytnangi. 07- 08.09.1987. T. Rossolimo leg. (cSh, ZMM); 34 ♂♂, 23 ♀♀: same locality and collector. Forest tundra. 25.08- 04.09.1987. (cSh, ZMM); 21 ♂, 11 ♀♀: Sob` St., valley of Sob` River. Tundra. 02- 05.09.1987. L.B. Rybalov leg. (cR, ZMM, cSh); 3 ♂♂: same data, traps. 08- 18.07.1987. ( ZMM); 34 ♂♂, 24 ♀♀: same data, southern slope, taiga, traps. 26.08- 05.09.1987. ( ZMM, cSh); KOMI: 1 ex.: Ust`-Tsilma, Pechora. 16.04.1907. A.V. Zhuravskiy leg. ( ZIN); 1 ♀: ‘Коми ивнЯк [Komi, willows]’ (cR); CHELYABINSK AREA: 1 ♀: Krasnyi Kamen`. 16.07.1987 (cSh); KHANTY- MANSI AUTONOMOUS REGION: 1 ♀: Sur- gutskiy District, Yuganskiy Nature Reserve, Basin of Ay-Magromsy River, “Medvezhiy Ugol” cordon. Mosses and litter near stream with Calamagrostis sp. , Ribes spp. , Rosa sp. , Filipendula ulmaria , Matteuccia struthiopteris , Equisetum sylvati cum, Sphagnum squarrosum, Sph. spp., Plagiomnium sp. etc. +15C°. 20.09.2002. A.B. Ryvkin leg. (cSh); TYUMEN AREA: 3 ♂♂, 1 ♀: st. Sob`, taiga, traps. 26.08- 05.09.1987. T. Rossolimo leg. (cSh, ZMM); 1 ♂: Uvatskiy District, 9 km S Gornoslinkino, near Tobolsk Field Research Station of Severtsov Institute. Marshy flood plain near left side of Irtysh River: Carex spp. , Comarum palustre , Salix sp. , Climacium dendroidea , small green mosses, duckweeds etc. 15.09.2003. A.B. Ryvkin leg. ( ZMM); TAYMYR AUTONOMOUS REGION: 1 ♂: Khatangskiy District, Taimyrskiy Biosphere Reserve, Ary-Mas: right bank of Novaya River, near mouth of Bogatyr`-Yuryakh River. River bank: sand, silt, shingle beds. 21.07.1992. A.B. Ryvkin leg. (cR); 16 ♂♂, 19 ♀♀: environs of Potapovo. 1995. L.B. Rybalov leg. ( ZMM, cSh); 2 ♂♂, 2 ♀♀: same data and collector. 07- 12.08.1994. ( ZMM); KRASNOYARSK TERRITORY: 2 ♂♂: Turukhanskiy District, Bakhta River basin, near Keteollo Lake. 450 m a.s.l., summit of bald mountain (1 st transect): mosses and litter under Pinus sibirica , Picea obovata , Betula sp. , Vaccinium uliginosum , V. myrtillus , Ledum sp. etc. 03.08.1992. A.B. Ryvkin leg. (cSh); 5 ♂♂, 1 ♀: same locality and collector. 370 m a.s.l., mosses and litter under Betula , Picea , Larix , Pinus sibirica , Abies with Pleurozium , Hylocomium , Vaccinum myrtillus , V. vitis-idaea , Lycopodium , Dicranum etc. 11.08.1992. (cSh, ZMM); 2 ♂♂, 2 ♀♀: same District, Yeloguy Na- ture Refuge, Yeloguy River mouth. In litter on swamped bank of forest lake with Filipendula ulmaria , Ribes nigrum , Alnus sp. , Betula sp. 27.07.1992. V.B. Semenov leg. (cR, cSh); 3 ♂♂, 1 ♀: same data and collector, Yeloguy River, 6 km below mouth of Tyna River. Clay-sandy bank of stream with Carex , Sphagnum sp. : in litter, dried sedges and Sphagnum sp. 21.07.1992. (cSh, ZMM); 1 ♀: same District, Yeloguy River. 07- 17.07.1989. L.B. Rybalov leg. ( ZMM); 1 ♂: same District, Yeloguy River, 125 km below Kellog, 15 km downstream of mouth of Altus River. Abies , Pleurozium , Hylocomium and Betula . 23.09.1993. V.B. Semenov leg. ( ZMM); 3 ♂♂: same data and collector, 40 km above of Kellog, 15 km upstream of Kepches River. Litter in Abies forest with Betula , Abies , Pinus sibirica , Equisetum , Hylocomium . 01.09.1993. ( ZMM); 2 ♂♂: same data and collector. Mosses and litter in swampy Betula forest with Alnus , Carex , Mnium . 01.09.1993. (cSh, ZMM); 2 ♂♂, 1 ♀: same District, Bolshaya Varlamovka River, 25 km up-stream of Malaya Varlamovka River near Bolshoy Stream. Swampy mouths of streams. 21.09.1984. A.B. Ryvkin leg. ( ZMM); 5 ♂♂, 5 ♀♀: same District and collector, Vorogovka River (I) near mouth of Bolotnaya River. Litter and mosses under Picea , Betula , Larix with Pleurozium , Hylocomium , Trientalis etc. among swampy Betula forest with Carex spp. , grasses, Allium ursinum etc. 26.06.1991. (cSh, ZMM); 4 ♂♂, 3 ♀♀: same District and col- lector, Podkamennaya Tunguska River basin, Rybnaya River (I), near seismological profile (about 40 km up-stream of river mouth). 190 m a.s.l., slope with Sphagnum spp. , Ledum , Oxycoccus sp. , Eriophorum sp. , Vaccinium vitisidaea , V. myrtillus , Betula , Pinus sylvestris , Pinus sibirica , Abies sibirica , Picea obovata etc. 13.09.1990. (cSh, ZMM); 2 ♂♂: same locality and collector. 170 m a.s.l., swampy and stony banks of stream: mosses, grasses, Carex spp. etc. 14.09.1990. ( ZMM); 5 ♂♂, 1 ♀: same District and collector, Podkamennaya Tunguska River basin, Ryb- naya River ( II), about 15 km up-stream of mouth. 100 m a.s.l., Betula forest with Picea , Abies , Pinus sibirica , grasses, Linnaea , Vaccinum vitis-idaea , V. myrtillus etc. (old fire-site). 17.09.1990. ( ZMM, cSh); 1 ♂, 2 ♀♀: same District and collector, Podkamennaya Tunguska River, Sukhaya Lebyazhya River mouth. 60 m a.s.l., afforested swamp with Picea , Pinus sibirica , Larix , Sphagnum spp. , Eriophorum , Vaccinum vitis-idaea , Hylocomium , Pleurozium etc. 18.09.1990. (cR, cSh); 1 ♂, 1 ♀: same District, Kamenniy Dubches River, 60 km below mouth of Teulch- es River, near hut. 150 m a.s.l., lake connected by stream with river. Carex , Filipendula , Sphagnum , Mnium and in Salix old leaves. 22.09.1992. V.B. Semenov leg. (cSh); 2 ♂♂: same locality and collector. Picea forest, with Pinus sibirica , Betula , Polytrichum commune , Pleurozium schreberi , Hylocomium . 24.09.1992. ( ZMM); 132 ♂♂, 41 ♀♀: same District, environs of Goroshikha. 31.08.1994. L.B. Rybalov leg. ( ZMM, cSh); 21 ♂♂, 65 ♀♀: same data. 24- 30.08.1994. ( ZMM, cSh); 1 ♀: same data. 29.07.1994. (cSh); 1 ♂: same District, Central Siberian Biosphere Re- serve, near Lebed`. 40 m a.s.l., litter and mosses in Picea - Pinus forest with Pleurozium schreberi . 29.08.1988. A.B. Ryvkin leg. (cSh); 3 ♂♂, 1 ♀: same District and Reserve, middle flow of Bolshaya Rakosaya River. In mosses and litter at firebreak with undergrowth ( Picea , Betula , Populus tremula ), Pleurozium , Polytrichum commune, Spahgnum spp. etc. 07.06.1992. V.B. Semenov leg. (cSh, ZMM); 2 ♂♂: same District, right side of Yenisei River 15 km S Komsa, near cordon at S border of Central Siberian Biosphere Reserve, 40 m a.s.l., mosses and litter at swampy bald mountain slope with Carex spp. , Poaceae gen. spp., ferns etc. 09.08.1988. A.B. Ryvkin leg. (cR); EVENKIA: 3 ♀♀: Baykitskiy District, Central Siberian Biosphere Reserve, Stolbovaya River basin, Kulingna River near mouth of Usas River. Sweeping on Carex sp. and grasses at riverbank. 09.09.1989. A.B. Ryvkin leg. ( ZMM); 3 ♂♂: same data and collector, Stolbovaya River basin: Kulingna River near mouth of Pravyi Usas River. Flight in forest near campfire. 04.09.1989. ( ZMM); 1 ♂: same data and collector, basin of Stolbovaya River: Dulkuma River ( II), 7-8 km up-stream mouth of Topkaya River. In Sphagnum sp. in flood plain forest. 28.06.1990. ( ZMM); 1 ♀: same data and collector, basin of Stolbovaya River, upper flowing of Raskol Stream (confluent of Dulkuma River). Bank of stream (left confluent of Raskol Stream), litter under Alnus , Abies , Picea and mosses near water. 16.07.1991. (cSh); 1 ♂: same data and collector, Stolbovaya River basin: middle flow of Raskol River (right confluent of Dulkuma River). 200 m a.s.l., litter and mosses under Picea near stream. 11.07.1991. ( ZMM); 1 ♂: same data and collector, Stolbo- vaya River basin: Kulingna River near mouth of Bolotnaya River. 95 m a.s.l., flood plain forest with Picea , Alnus , Betula , Salix , Ribes , Lonicera etc. 08.09.1989. (cSh); 3 ♂♂, 1 ♀: same District and collector, Podkamennaya Tun- guska River, 1 km above Sulomay. 110 m a.s.l., litter and mosses on slope near small streams: Alnus sp. , Salix sp. , Betula sp. etc. 04.10.1989. ( ZMM, cSh); 1 ♂, 1 ♀: same District and collector, basin of Bakhta River: Mochektakon River, 2 km up-stream of mouth. 220 m a.s.l., bank with Carex grasses, Salix . 01.09.1991. (cSh); TUVA: 1 ♀: Sange- len Mt., Dzen-Aryk, upper flow, 50°29‘N 95°25‘E. 1750 m a.s.l. Y.M. Marusik leg. ( FMNH); IRKUTSK AREA: 4 ♂♂, 1 ♀: Bodaibinskiy District, Vitim Nature Reserve, first confluent of Pravaya Polovinka River, 57°07’25,6’’N 116°35’58,3’’E. 829 m a.s.l., wet Sphagnum sp. among stones near river bank. 13.07.2016. I.V. Enushchenko leg. (cSh); 1 ♀: same data, watershed of Vitim and Olekma rivers, Aunakit Lake, 58°19’01,3’’N 114°51’33,1’’E. 1019 m a.s.l. I.V. Enushchenko leg. (cSh); BURYATIA: 2 ♂♂, 1 ♀: Baikalskiy Mts., upper of Kurkula River, 2 km lower from Gitara Lake. Near streams. 22- 24.07.2010. A.V. Shavrin & I.V. Enushcheko leg. (cSh); 1 ♂, 3 ♀♀: same data and collectors, upper reaches of Kurkula River, Gitara Lake. 20- 26.07.2010. (cSh); REGIONS NOT LOCATED: 1 ♂: ‘Lapponie russe’ ( IRSNB); 1 ♂: ‘Lapp. Ross.’ ( IRSNB); 1 ♂: ‘Sibir. Arct. J. Sahlb.’ ( IRSNB).

Redescription. Measurements (n=200): HW: 0.44–0.65; HL: 0.24–0.43; OL: 0.09–0.18; AL: 0.88–1.19; PL: 0.35–0.58; PW: 0.71–1.20; EL: 0.63–1.05; EW: 0.88–1.45; AW: 1.07–1.45; MTbL (averaged): 0.65; MTrL (averaged): 0.40 (MTrL 1–4: 0.25; MTrL 5: 0.15); AedL: 0.40–0.62; TL: 1.90–4.24.

Body small or medium-sized, convex, wide and moderately short or elongate, with extended abdomen ( Fig. 2 ). Coloration yellow-brown to reddish-brown, usually with darker head and abdomen and paler lateral portions of pronotum and paratergites; mouthparts, antennae and legs yellow to reddish-brown; antennomeres 1–2 or 1–4 and tarsi yellow. Body shiny, without microsculpture except for latero-apical portion of head between antennal bases and anterior margins of eyes with indistinct and fine diagonal meshes, neck and sometimes scutellum with very fine, transverse sculpture and abdomen with fine, isodiametric microreticulation. Head with irregular and sparse, moderately large and deep punctation, without visible punctures on clypeus, punctation sparser in middle and denser and sometimes coarser in laterobasal and/or basal portions; neck with moderately dense and deep punctation in middle; punctation of pronotum denser, larger and deeper than that on head, sparser and finer in lateral and basal portions, sometimes with very fine punctation or without punctures in median third; scutellum with or without several very fine punctures in middle; punctation of elytra similar to that of pronotum, each elytron with or without distinct five to six longitudinal rows of punctures, these rows confused in basal and apical portions; abdominal tergites without visible punctation, rarely with very sparse and indistinct punctures on abdominal tergites IV–V. Latero-apical portions of head with very sparse, short and erect setae; abdomen with or without very sparse short setae.

Head 1.5–1.8 times as broad as long, with slightly elevated median portion and moderately deep, short or slightly elongate anteocellar foveae reaching about level of middle length of eyes; anterior portion between antennal insertion and anterior margins of eyes slightly concave. Eyes very large, moderately convex. Ocelli small, slightly convex, and sometimes indistinct, situated at about level of posterior margins of eyes; distance between ocelli similar to distance between ocellus and posterior margin of eyes, or slightly shorter. Labrum as in Fig. 12 . Mandibles as in Fig. 13 . Labium and submentum as in Fig. 15 . Ventral aspect of head as in Fig. 26 . Apical segment of maxillary palpi distinctly more than twice as long as preceding segment, sometimes slightly broader than preapical segment, from middle gradually narrowing apically ( Fig. 14 ). Antennae ( Figs. 16–17 ) reaching basal portion of elytra when reclined, with (sometimes significantly) elongate antennomeres 4–10; basal antennomere ovoid, moderately wide, antennomere 2 slightly narrower and shorter than basal antennomere, 3 narrower and about as long as or longer than 2, 4 slightly longer than wide, 5–6 ovoid, longer than 4, 7–9 usually longer and broader than 6, 10 slightly shorter than 9, apical antennomere about as broad and twice as long as preapical segment, from middle sharply narrowing toward acute apex.

Pronotum twice as broad as long, 1.6–1.8 times as broad as head, widest in or slightly in front of middle, markedly more narrowed posteriad than anteriad; apical margin widely rounded, about as long as, or slightly narrower than somewhat straight posterior margin; anterior angles widely rounded, slightly protruded anteriad; posterior angles subacute or obtuse, with indistinctly ( Figs. 20–21 ) or distinctly ( Fig. 22 ) sinuate laterobasal margins; lateral portions with shallow laterobasal impressions. Ventral aspect of prothorax as in Fig. 27 . Mesothorax as in Fig. 28 . Scutellum as in Fig. 31 . Metathorax as in Fig. 29 . Metendosternite as in Fig. 30 .

Elytra 1.3 times as broad as long, slightly to strongly widened apicad, shorter than pronotum ( Fig. 32 ); apical margins straight or widely rounded. Hind wings fully developed.

Metatarsus 1.8 times as long as metatibia ( Fig. 33 ).

Abdomen about as wide as elytra or slightly broader, with two small transverse tomentose wing-folding spots in middle of abdominal tergite V; apical margin of abdominal tergite VII with narrow palisade fringe.

Male. Antenna as in Fig. 16 . Apical margins of abdominal tergite VIII rather deeply ( Fig. 34 ) and abdominal sternite VIII slightly ( Fig. 35 ) concave. Aedeagus with wide basal portion, significantly narrowed apically toward subacute or rounded apex; parameres moderately wide, exceeding apex of median lobe, with widened apical portions; internal sac with two wide and long fields of elongate thorns ( Figs. 42–43, 45–47 ). Lateral aspect of aedeagus as in Fig. 44 .

Female. Antenna as in Fig. 17 . Apical margin of abdominal tergite VIII straight ( Fig. 36 ) or rounded. Apical margin of abdominal sternite VIII widely rounded or truncate ( Fig. 37 ). Female genital segment as in Fig. 38 .

Comparative notes. Regarding the general shapes of the body and the aedeagus, M. arctica is similar to M. brevipennis and M. pleshanovi sp.n. It differs from M. brevipennis by the shape of laterobasal portion 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 elongate antennomeres 4–10 and the somewhat different shape of the apical part of the median lobe.

Distribution. Mannerheimia arcica is distributed in the boreal zone of the Palaearctic Region from Scandinavia across Northwest Russia, Ural and Siberia to the Baikal Lake and north-eastern part of Irkutsk Area; several isolated populations are located in the Alps ( Fig. 39 ).

Bionomics. The species inhabits mixed forests with Larix , Abies , Pinus sibirica , Picea , Betula , Alnus and sometimes flood plain deciduous forests with Alnus , Salix and Populus . It was also recorded from lichen-mosses tundra in the Arctic zone and in high mountain habitats ( Fig. 41 ). It most often inhabits mosses and litter in vegetation with Sphagnum , Vaccinium , Carex , Hylocomium , Pleurozium , etc. The species is recorded here from low to moderately high elevations between 40 m a.s.l. to 2700 m a.s.l. The beetles were observed during the period from April to October, with the greatest numbers of specimens captured in August-September.

Remarks. Records of M. arctica from Kazakhstan ( Kastcheev 1999; Kastcheev & Ishkov 2001; Gabdullina 2016) and from Magadan Area ( Ryabukhin 2002) require confirmation. A late-glacial Pleistocene subfossil record is known from Great Britain ( Coope 1969).

Figures

FIGURE 41. Habitat of Mannerheimia arctica in Buryatia, Baikalskiy Mts., Gitara Lake (photograph: K. Vershinin).

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 12–15. Mouthparts of Mannerheimia arctica: 12—labrum (dorsal view), 13—mandibles (left and right, dorsal view), 14—maxilla (left, dorsal view), 15—labium and mentum (ventral view). Scale bar: 0.1 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 26–38. Body parts of Mannerheimia arctica: 26—ventral aspect of head, 27—ventral aspect of prothorax, 28—ventral aspect of mesoventrite, 29—ventral aspect of metaventrite (without metanepisternum), 30—metendosternite (ventral view), 31—scutellum (without lateral structures, dorsal view), 32—ventral aspect of elytron, 33—hind tibia and tarsus (anterior view), 34—male tergite VIII (dorsal view), 35—male sternite VIII (ventral view), 36—female tergite VIII (dorsal view), 37—female sternite VIII (ventral view), 38—female genital segment (dorsal view). 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.