Frullania subarctica Vilnet, Borovich. & Bakalin, 2014

Vilnet, Anna A., Borovichev, Eugeny A. & Bakalin, Vadim A., 2014, Frullania subarctica-a new species of the Frullania tamarisci complex (Frullaniaceae, Marchantiophyta), Phytotaxa 173 (1), pp. 61-72 : 67-70

Frullania subarctica Vilnet, Borovich. & Bakalin sp. nov.

The species is recognized by moderate size stylus of 5–8 cells long and 1–2 cells wide; disc strongly reduced, up to 3(–5) cells long; ocelli lacking or forming a short line of 2–5(–7) cells long to 1/3 of leaf lobe length; plants red-brown to blackish brown and almost black.

Type: — Russia. Magadan Province: Ol`skoe Basalt Plateau in upper course of Maltan River , 60°38’46” N, 151°27’25” E), 1166 m elev., tundra belt, wet moss- Sphagnum tundra on slope, in hollows, 9 August 2011, V . Bakalin Mag 54-19-11 (holotype VBGI, isotypes KPABG, MHA). GenBank KC918805 (nrITS1-5.8S-ITS2), KC918789 (trn L – F) .

Plants red-brown to blackish brown and almost black. Leafy shoots (1.1–) 1.2–1.6 mm wide, 1.0– 3.5 cm long, pinnately to bipinnately branched; branches with leaves becoming gradually smaller downward. Leaves conduplicate-trilobed. Dorsal lobes imbricate, apices often incurved, acute or acuminate-apiculate, uniseriate apex of 2–4 cells; cells of leaf apex ± triangular, 8–10(–12) µm, median cells of leaf lobe usually isodiametric or slightly elongated (9–)12–15 × (12–)18-20 µm, usually with thickened walls and coarse trigones; basal leaf cells paler in color (10–)14–16 × (12–)20(–25) µm, often with nodular thickenings; ocelli larger than other cells, ± rounded, (18–)23–28(30) µm in diameter, absent or forming a short line varying from 2–5(–7) cells long to 1/3 of leaf length). Lobule cylindrical to clavate-cylindrical, saccate, with rounded apex, with axis parallel to the stem; stylus small, subulate to filiform, 5–8 cells long and 1(–2) cells wide; disc virtually absent to rarely 3(–5) cells long, very rare with a single tooth of 1-2 cells. Cuticle smooth. Underleaves oblong-quadrate to quadrate-ovate, (0.4–) 0.5–0.6 mm width, with strongly recurved margins, bilobed for 1/3–1/4 (or less) of underleaf length, lobes obtuse to acute, not or slightly auriculate at base. Leaf lobes of lateral branches smaller, commonly with ± well-developed median line of ocelli. Gemmae lacking. [No gametangia and sporophytes seen].

Diagnostic characters: —The species is recognized by moderate in size stylus of 5–8 cells long and 1(–2) cells wide, virtually absent or rarely 3(–5) cells long disk, lacking or a short line of 2–5(–7) cells to ⅓ of leaf length ocelli, red-brown to blackish brown and almost black coloration.

Ecology: —The species mostly occurs in tundra communities on flat surfaces or slopes (from swampy Sphagnum tundra to wet dwarf shrub-moss tundra). In the latter habitat the species is characterized by growing as an epibryophyte over wet moss patches. It also grows on north-facing wet cliffs and their crevices in the tundra belt of mountains, where it mostly forms pure mats.

Distribution: —According to the data at hand, F. subarctica is an arcto-montane, presumable Asian species with some localities in Europe and North America.

Selected specimens examined: — Paratypes: Frullania subarctica Vilnet, Borov. & Bakalin – RUSSIA. Murmansk Province: Lovozero District, on the bank of Purnach River , 66°57’45” N - 39°50’36” E, 160 m., near the bottom of the rock, 5 July 2009, Melekhin 20639 ( KPABG) ; Krasnoyarsk Territory : Dikson District , Ledyanaya Bay , 73°31’ N - 80°20’ E, 1 July 2004, Fedosov 107971 ( KPABG) ; West Sayan , upper course of the Malyi River , 5 July 1968, Bardunov ( KPABG) ; Republic of Yakutiya: Tomponskiy District , upper course of the Chukomnan River , 62°42’ N - 134°42’ E, 25 August 1956, Ivanova s.n. ( KPABG) ; Magadan Province: Ol`skoe Basalt Plateau in upper course of Maltan River , 60°38’46” N - 151°27’25” E, 1166 m, tundra belt, swampy Sphagnum tundra on slope, in hollows, 9 August 2011, Bakalin Mag 54-17-11, Mag-55-11-11 ( KPABG, VBGI) ; ibidem, wet dwarf shrub-moss tundra on slope, 14 August 2012, Bakalin Mag-28-23-12 ( VBGI) ; Sakhalin Province: Sakhalin Island , middle part of Nabilskij Range , more or less wet crevice in north-facing cliff, 50°45’18,4” N - 143°17’55,0” E, 1300 m alt., 11 September 2009, Bakalin , S-61- 14 - 09 ( VBGI) , Vostochno-Sakhalinskiye Mts. , moss tundra on percolate water, 49º52’27,2” N - 143º28’59,1” E, 900 m alt., 20 August 2006, Bakalin , S-32-29 - 06 ( VBGI) . U.S.A. Alaska: Deer Island , 54°55’ N - 162°18’ W, depression in Empetrum tundra, 12 August 1997, Schofield 109173 ( UBC) .

Frullania nisquallensis Sull. U.S.A. California: Humboldt Co., rock outcrops, 9 September 1989, Ignatov s.n., (VBGI); on Acer , 7 August 1989, Ignatov s.n. ( VBGI) ; Oregon: Lane County, willow Creek Natural Area , trunk of Pyris tree, 2 May 2000, Schofield 114240 ( UBC) ; Multnomah County , Oneonta Gorge, Columbia River Gorge, open cliff legde, 4 May 2000, Schofield 114410 ( UBC) ; Multnomah County , J. B. Yeon State Park , Columbia River Gorge, Elwah Falls trail, tree trunk, 4 May 2000, Schofield 114415 ( UBC) ; Washington: Snohomish County , Wiley Creek Campground on Stillaguamish River , alder trunk by river, 25 October 2002, Schofield 120750 ( UBC) ; Cowlitz County , ca 2 mi N of Kalama, 0.5 mi N along Kalama River Road at junction of old 99 Highway , trunk of Fraxinus tree, 1 May 2002, Schofield & Harpel 114163 ( UBC) ; Clallam County , Ray`s Bog , on small shrubs and trees in bog, 30 May 1996, Schofield & Harpel 105328 ( UBC) ; Lewis County , Big Creek Campground, tree trunk, 3 May 2000, Schofield 114319 ( UBC) ; Alaska: Aleutians West County , Analga Bight, Adak Island , cliff face near stream, 26 August 2006, Schofield & Talbot 125344 ( UBC) ; Alaska Peninsula Area , Nakchamik Island , tundra slope near Pyramid Mt. , 16 July 1999, Schofield & Talbot 112914 ( UBC) ; CANADA. British Columbia: Howe Sound , on Abies trunk, 22 May 2001, Schofield, Ellis, Olivia Lee & Gina Choe 117409 ( UBC) .

Frullania tamarisci (L.) Dumort. RUSSIA. Murmansk Province: Pechenga District,vicinity of Nickel`, Orschoayjvi Mt. , on dry rock, 21 June 2013, Borovichev 2-4-2013 ( KPABG) ; Lapland State Biosphere Reserve , Monchegorsk District , Pel`-tundra, on dry rock, 28 June 2008, Melekhin 29-3-08 ( KPABG) ; Krasnodar Territory: Sochi National Park , Schahe River Basin, on Abies trees, 3 October 2008, Konstantinova K 432-1- 08 ( KPABG) ; ibidem, Bzhizhu River , aul Tchagapsch Vicinty , on base of sand rock, 22 November 2011, Konstantinova & Savchenko K 322-2- 11 ( KPABG) ; Karachaevo-Cherkessian Republic: Teberda River Basin, left bank of Azgek River , on stony slope of stream, on rock, 19 September 2005, Konstantinova K 539-5- 05 ( KPABG) ; Dagestan Republic: Charodinskijy District , Nuckatal` Range, valley of confluent of Kara-or River , on rock under mosses, 7 March 2011, Konstantinova K 35b- 11 ( KPABG) ; ibidem, Uruh-Sota Settlement Vicinity , on rock, 7 March 2011, Konstantinova K 44-2- 11 ( KPABG) . GEORGIA. Adjaria: Mtirala National Park , bark of tree, 11 May 2013, Bakalin G-11-55-13 ( VBGI) , cliffs in open place, 12 May 2013, Bakalin, G-12-32-13 ( VBGI) .

Frullania appendiculata Steph. RUSSIA. Sakhalin Province: Kunashir Island , the bark of Abies on the height 1- 2 m from the ground, 7 September 2006, Bakalin K-47-16-06 (VBGI); ibidem, decaying wood, 09 September 2006, Bakalin K-50-10a-06 ( VBGI) ; ibidem, bark of Taxus , 10 September 2006, Bakalin K-53-12-06 ( VBGI) ; ibidem, bark of Abies , 17 September 2006, Bakalin K-63-24-06 ( VBGI) ; Sakhalin Island, Korsakovsky District , crevice filled by fine-grained soil in vertical N-facing dry cliffs, 16 September 2009, Bakalin S-66-12 - 09 ( VBGI) ; Primorsky Territory: Lazovsky District , more or less dry boulders on slope in part shade, 08 September 2010, Bakalin P-42-12-10 ( VBGI) ; Partizansky District , crevices among stones in gravelly barrens field, 10 September 2010, Bakalin P-44-11-10 ( VBGI) ; Snezhnaya Mt. , crevice between rocky blocks, 17 September 2007, Bakalin P-72-7-07 ( VBGI) ; Malaya Margaritovka River , cliff ledge on slope to river, 19 September 2007, Bakalin P-80-9-07 ( VBGI) ; Shkotovsky District, Livadijsky Range , wet boulders along stream, 25 July 2012, Bakalin P-15-65-12 ( VBGI) ; Khabarovsk Territory: Nanajsky District, Tardoki-Yani Range , cliff ledge, 29 August 2012, Barkalov, s.n. ( VBGI) . REPUBLIC OF KOREA. Deokgyusan : shaded stone in broadleaved forest, 24 June 2008, Bakalin Kor-7-10-08 ( VBGI) ; ibidem, rocks in full sun, 25 June 2008, Bakalin Kor-9-11-08 ( VBGI) ; bark of Quercus mongolica , 25 June 2008, Bakalin Kor-10-1-08 ( VBGI) ; KyongNam Province: dry stones along stream, 13 June 2009, Bakalin Kor-2-23-09 ( VBGI) ; decaying wood Abies koreana lighted forest, 15 June 2009, Bakalin Kor-11-14b-09 ( VBGI) ; ChonNam Province: wetted stone near stream, 20 June 2009, Bakalin Kor-22-17-09 ( VBGI) ; Jeonnam Province: more or less dry cliffs in part shade, 19 May 2011, Bakalin Kor-17-40-11 ( VBGI) ; Oinarodo Island : more or less dry stone in part shade, 20 May 2011, Bakalin Kor-18- 16-11 ( VBGI) ; Kangwon Province: dry cliffs in full sun, 11 May 2011, Bakalin Kor-5-32-11 ( VBGI) . JAPAN. Honshu: Ouchi-dani Park , tree trunk, 10 March 2013, Bakalin J-1-22-13 ( VBGI) .