Bryoxiphium norvegicum (Brid.) Mitt., J. Linn. Soc., Bot.

Bryoxiphium norvegicum (Brid.) Mitt., J. Linn. Soc., Bot. View in CoL 12: 580. 1869.

— Phyllogonium norvegicum Brid., Bryologia Universa 2: 674. 1827. — Eustychia norvegica (Brid.) Brid., Bryologia Universa 2: 789. 1827. — Eustychia norvegica (Brid.) Bruch et al., Bryologia Europaea 2: 161. 195. 1849. Figs.6 View Fig , 8 View Fig .

Type: Krýsuvík , Iceland, A. Mörch, 1820 (C, not seen) .

Stems 1–15 mm long. Leaves in the middle part of stem 1.2–2.1×0.5–0.8 mm, oblong-lanceolate, obtuse and occasionally notched at apex, blunt or shortly apiculate, rarely with stout awn to 0. 2 mm long; costa percurrent or shortly excurrent, with dorsal lamella 1–2(–3) cells high, usually hardly visible in profile, reaching leaf tip, but not descending to the leaf base; margin entire, crenulate, rarely minutely serrulate at leaf apex; cells in upper and middle part of leaf 9–16 µm wide and 12–20 long, short rectangular, triangular, rhomboidal or hexagonal, firm-walled, along margins in 10-15 rows narrower, elongate, 4–6 µm wide and 28–50 µm long, thin-walled, gradually transiting to mid-leaf cells; basal laminal cells larger, 11–18 µm wide and 16–40 µm long, with length/width ratio (1.2–)1.6–3(–4), short rectangular, weakly differentiated from the rest of lamina. Perichaetial and perigonial leaves from ovate-lanceolate base gradually narrowed into a flexuose and twisted, yellowish, brownish or hyaline filiform awn 1–3(–5) mm long, with entire to crenulate margins. Seta 1.5–2.5 mm. Urn ca. 1 mm long. Spores 14–19(–21) µm.

Distribution. Bryoxiphium norvegicum s. str. is distributed mainly in humid areas of Northern Hemisphere. It is known from Europe ( Iceland), North America ( Greenland, U.S.A.) and Asia ( Russia and China). The identity of Macaronesian, Mexican, Dominican and Indonesian populations needs further study. Distribution area of B. norvegicum in Russia ( Fig. 3 View Fig ), includes northern Far East: Chukotka Autonomous District, Magadan Province, Kamchatka Peninsula and Kommander Islands (Afonina, 2004; Czernyadjeva, 2012, as Bryoxiphium norvegicum subsp. japonicum ; Fedosov et al., 2012; Pisarenko et al., 2015). Scattered records were also reported from continental regions of Siberia: Yakutia and Zabaikalsky Territory (Ivanova et al., 2005; Afonina et al., 2015). All specimens from Kamchatka Peninsula previously referred to B. japonicum , in fact, represent B. norvegicum , and molecular data confirm this identification.

Ecology. The species occurs on bare cliffs and boulders composed of acidic to basic volcanic rocks, and on sandstone; it grows on cliff surfaces, in crevices and niches, and on mineral soil and humus between rocks. In most cases it occurs near lakes, creeks or along sea coasts. In Chukotka Peninsula it also grows in rocky tundra. The species often forms pure carpets, especially extensive in shaded niches, or grows intermixed with Distichium capillaceum (Hedw.) Bruch et al. , Ditrichum flexicaule (Schwägr.) Hampe , Eurhynchiastrum pulchellum (Hedw.) Ignatov & Huttunen , Amphidium lapponicum (Hedw.) Schimp. , Hypnum cupressiforme Hedw. , Pohlia cruda (Hedw.) Lindb. , Polytrichastrum alpinum (Hedw.) G.L.Sm. , Mnium spp. , etc. Four specimens from Kamchatka Peninsula and one from Onekotan Island (North Kurils) contained plants with sporophytes.

Specimens examined: ICELAND: Gullbringusysla : Kleifarvatn, Innro Stapi (63°55´N, 22°00´W), 150 m alt., 3 IX.1950 Askel & Doris Löve 2-805 ( ICEL, dupl. MHA) GoogleMaps ; Southwest Iceland , Sveifluhals, 26.VI.1964, Steere s.n. ( LE) ; Kleifarvatn auf Reykjanes, Sudvesturland, 26.VI.1973, Frahm s.n. ( LE) . DENMARK: Northeast Greenland , Wollaston Foreland, 14.VIII.1949, Holmen s.n. ( LE) .

RUSSIA: SIBERIA: Zabaikalsky Territory, Kalar Distr., Stanovoye Upland (56°14´36.4´´N, 117°26´23.8´´E), 1516 m alt., 26.VIII.2014, Mamontov 506/1 ( LE) GoogleMaps . Yakutia: Olekminskij Distr., Udokan Range , 1200 m. alt., 29.VII.2001, Kuznetsova s.n. ( SASY) ; Ust-Majskij Distr., Aldan River valley , 18.VI.2001, Krivoshapkin s.n. ( SASY, MW) . FAR EAST: Magadan Province, vicinityof Magadan City (59°34´N, 150°39´E), 330 m alt., 10.VIII.2014, Pisarenko 05358 ( VBGI, LE, MW) GoogleMaps . Chukotka Autonomous District: Iultinskij Distr., vicinity of Egvekinot Settl. (66°20´N, 179°07´W), 21.VI.1970, Afonina s.n. ( LE) GoogleMaps ; Providenskij Distr., vicinity of Yanrakynnot Settl. (64°53´N, 172°30´W), 27.VII.1976, Afonina s.n. ( LE) GoogleMaps ; Anadyrskij Distr., vicinity of Pekulnejskoe Lake, Kakanaut Gulf (62°43´N, 176°58´E), 15.VIII.1984, Afonina s.n. ( LE) GoogleMaps ; the same area, Byanranaj Mt. (62°43´N, 176°58´E), 10.VIII.1984, Afonina s.n. ( LE) GoogleMaps . Kamchatsky Territory: Kamchatka: Ust’- Kamchatskij Distr., Klyuchevskaya group of volcanoes: Bilchenok glacier (56°11´N, 160°21´E), 1000 m alt., 16.VII.2003, Czernyadjeva 18 ( LE) GoogleMaps ; the same area 950 m alt., 19.VII.2003, Czernyadjeva 37, 64 ( LE) ; the same area, 750 m alt., 22.VII.2003, Czernyadjeva 53 ( LE) ; Bilchenok River (56°11´N, 160°21´E), 550 m alt., 22.VII.2003, Czernyadjeva 18 ( LE) GoogleMaps ; Ushkovsky volcano (55°58´N, 160°15´E), 1000 m alt., 19.VIII.2005, Czernyadjeva 40 ( LE) GoogleMaps ; the same place, 15.VIII.2004, Czernyadjeva 78 ( LE) ; vicinity of Kozyrevsk Settl. , valley of Kamchatka River, (56°00´N, 160°02´E), 220 m alt., 12.VIII.2004, Czernyadjeva 63 ( LE) GoogleMaps ; Bystrinskij Distr., vicinity of Esso Settl. (55°55´N, 158°42´E), 700 m alt., 30.VII.2001, Czernyadjeva 47 ( LE) GoogleMaps ; Middle Kamchatskij Range, Icha volcano (55°32´N, 157°37´E), 800 m alt., 4 & 5.VIII.2007, Czernyadjeva 6, 10 ( LE) GoogleMaps ; Elizovo Distr., Kronotskij State Reserve, Death Valley in upper course of Geyzernaja River (54°28´N, 160°16´E), ca. 840 m alt., 7.IX.2012, Fedosov 12-132 ( MW) GoogleMaps ; Ust’-Bol’sheretskij Distr., middle course of Bannaya River (52°54´N, 157°30´E), 250 m alt., 12.VIII.2002, Czernyadjeva 67 ( LE, MHA, MW) GoogleMaps ; Aleutskij Distr., Kommander Islands, Bering Island , Nakoval’nya Mt. near Nikol’skoe settl. (55°13´N, 166°01´E), ca. 20 m alt., 5.VIII.2010, Fedosov 10-3- 494, 10-3-593, 10-3-1104 ( MW) GoogleMaps . Sakhalinskaya Province: Northern Kuril Islands: Paramushir Island, vicinity Severo-Kurilsk Settlement (~ 50°23´N, 155°41´E), 4.VIII.1978, Barkalov s.n. ( VLA) GoogleMaps ; Onekotan Island, Nemo Bay (~ 49°27´N, 154°46´E), 12.VII.1981, Barkalov s.n. ( VLA, LE) GoogleMaps ; the same place, 29.VII.2008, Koroteeva 0-32.2-08 ( MHA) ; Shumshu Island, Bolshoe Lake (~ 50°45´N, 156°21´E), 22.VII.1978, Zavorotniy s.n. ( VLA) GoogleMaps .

U.S.A.: Washington, Glacier Peak Wilderness Area, Kennedy Ridge above Pacific Crest Trail , VIII.1977, B.R. Jenkinson s.n. ( MHA) ; Arkansas, Newton County, 24.IV.1993, P.L. Redfearn, Jr. & A. Redfearn (Mosses of the Interior Highlands Exsiccatae 36) ( LE) ; Wisconsin Dells, Potsdam Sandstone , 19.VIII.1893, Britton s.n. ( LE) ; Missouri, St. Clair County, 30.V.1973, Redfearn 28522 ( LE) ; Missouri, St. Clair County, 28.VII.1969, Ireland 10092 ( LE) ; Ohio, Hocking County, VI.1937, Welch 208 ( LE) .