Sphagnum talbotianum R.E. Andrus, Sida

Shkurko, A. V., Kuznetsova, O. I. & Fedosov, V. E., 2023, Sphagnum Warnstorfii complex in Northern Asia, Arctoa 32 (2), pp. 176-188 : 183-186

Sphagnum talbotianum R.E. Andrus, Sida 22: 970, f. 35–40. 2006. ( Fig. 4, 5)

Plants small to medium sized, variegated yellow-pink, sometimes to purplish red, capitula flat to slightly round- ed, not stellate, apex bud typically not visible. Stems pale to pink, sclerodermis well delimited, superficial cortical cells eporose. Stem leaves triangular-lingulate, 0.9–1.1

mm long and 0.45–0.55 mm wide, apex rounded, narrowly erose, hyaline cells rhomboid, 0-1(-2) septate, typically with weak fibrils at apical part of the leaves, sometimes efibrillose, aporose, border strong, widening at the base to more than 0.25 of the width. Hemiisophyllous plants have more triangular and longer stem leaves with strong fibrils and pores. Branch fascicles with 2 spreading and 1(-2) pendent branches, spreading branches 5- ranked, typically curved or straighten up in plans from compact cushions. Branch leaves straight to slightly subsecund at middle part of branches, ovate to ovate-lanceolate, apex involute, 0.9–1.1 mm long and 0.35–0.45 mm wide, hyaline cells on convex surface with rounded to elliptical pores along commissures, grading from small (4–6 µm) pores near the apex to larger ones at the base, concave surface with large round pores in proximal margins of the leaf. Chlorophyllous cells triangular to trapezoidal in transverse section, broadly exposed on concave surface. Sexuality and sporophytes unknown.

Ecology. The plants assigned to this species were collected from arctic fens, wet moss dominated tundra and eutrophic bogs, in localities dispersed from low maritime areas to high mountain elevations up to 1500 m a.s.l.

Distribution in Russia ( Fig. 6): S. talbotianum is widely distributed in Asiatic part of Russia from the western foothills of Ural Mountains in Nenets Autonomous District to the Russian Far East. We also expect it’s presence in western and middle Siberia, and in inland Chukotka. For now, we proved the west boundary of S. talbotianum distribution in inland Eurasia to be situated in Nenets AD (however, its occurrence along northern shore of Kola Peninsula is possible) and the southern one in Amur Province. Along the boundaries of its distribution, S. talbotianum occurs sympatrically with S. warnstorfii , which, however, does not occur in Asian cryolithozone.

Differentiation. S. talbotianum vs S. capillifolium . Although the distributions of the two species overlap, even when growing mixed they readily differ morphologically. In contrast to S. capillifolium , S. talbotianum mostly has 5-ranked branches, smaller and more round- ed pores at the apical part of the branch leaves ( Fig. 7), and weaker fibrils (or its total absence) in the hyaline cells of the stem leaves. Hemyisophyllous plants of S. talbotianum which are very similar to S. capillifolium in morphology of stem and branch leaves and often lack typical small pores, still have more or less prominent 5- ranked branch foliage and slightly subsecund leaves, lacking in S. capillifolium .

S. talbotianum vs S. warnstorfii

In most cases, S. warnstorfii readily differs from S. talbotianum . Macroscopically plants of S. warnstorfii often has a flat star-shaped capitulum with a slightly visible apical bud, straight 5-ranked divergent branches and straight branch leaves, while plants of S. talbotianum are more compact with less star-like aspect of capitulum and hidden apical bud, divergent branches slightly curved or straighten up, often with somehow subsecund branch leaves. Microscopically the two species also differ well, since the tiny (3–4 µm) ringed pores often runs lower towards median part of the branch leaves in S. warnstorfii becoming mixed with bigger ones proximally ( Fig. 7A). Sphagnum talbotianum commonly has slightly bigger pores (5–6 µm) at apices of the branch leaves, which become bigger towards median part of the leaf rather gradually ( Fig. 7B). Sometimes, S. talbotianum also has few tiny pores, however they are more oval, less ringed and often placed closer to commissures than the tiny pores of S. warnstorfii . Similar morphological differentiation was mentioned for the “arctic” vs “boreal” plants of S. warnstorfii in the study of Yousefi et al., 2019, which contributed a lot to our feeling of S. talbotianum .

S. talbotianum vs S. rubellum . Despite the clear phylogenetic differentiation, these two are very similar morphologically ( Fig. 3), which has resulted in numerous reports of S. rubellum from the continental areas of North Asia ( Ignatov et al., 2006; Fedosov et al., 2011). Macroscopically both have pink to sometimes red coloration, flattering capitula, and mostly 5-ranked branches with slightly subsecund branch leaves; however the shoots of S. rubellum typically more weak and slender in contrast to commonly compact plants of S. talbotianum . The shape of the stem leaves is rather triangular in S. talbotianum with weak fibrils at the apical part or without them similarly to S. warnstorfii , while S. rubellum has more lingulate stem leaves, the marginal parts of which runs rather parallel to each other basally and commonly has strong fibrils in upper hyaline cells or throughout the stem leaves. Also, according to our measurements ( Table 1), S. rubellum mostly has higher stem/branch leaf length ratio than S. talbotianum has, which can be used as an additional trait for separating the two species. The apical pores at the convex side of branch leaves also can help to differentiate these species, since S. talbotianum typically has smaller pores, rather similar to tiny pores of S. warnstorfii ( Fig. 7). Ecological preferences of the two species also differ, S. rubellum commonly grows in ombrotrophic bogs, S. talbotianum prefer richer habitats as arctic bogs and rich fens.

Specimens used for molecular and morphological study:

Sphagnum capillifolium : Arkhangelsk Province, Verkhneoemky Disrict, vicinity of Tymoshino Village , oligotrophic bog, 61.94°N, 45.28°E, 19.VII.1999, Churakova 497 (OK3657, MHA9112619 ) ; Buryatia Republic, Kabansk Disrict, Khamar- Daban Range , valley of Bolshoy Mamai Creek , minerotrophic bog, 51.37°N, 104.83°E, 1100 m a.s.l., 6.IX.2018, Ignatov, Ignatova, & Kolenikova 18-4238 (OK3656, MHA9027234 ) ; Yakutia, Khangalasskiy District, “Lenskye stolby” National Park , hanging sphagnum bog, 61.15°N, 127.60°E, 05.VIII.2016, Ignatov & Ignatova 16-488 (OK3521, MW9113201 ) ; Sakhalin Province, Sakhalin Island, Nogliksky Disrict , oligotrophic bog, 51.43°N, 143.43°E, 28.IX.2016, Dudov & Kozhin Br _0113 (OK3522, MW9079504 ) ; Kunashir Island, Doktorskiy Stream valley, fir-spruce-bamboo forest near the thermal Lake Faust , on rock outcrops near the lake water, 43.99°N, 145.77°E, 20.IX.2020, Mamontov 833-1-s34 (OK2964, MHA9121196 ) ; the same place, 833-1-s40 (OK2965, MHA9121197 ) ; Kamchatka Province, Milkovsky Disrict, Kostin Mountain , rocky slope, on soil, 04.IX.2010, Fedosov, 10-4-58 (OK3525, MW9065671 ) ;

Sphagnum rubellum : France, Sainte-Croix-Hague, 14. VI .1888, Cobiere, 3399 ( MW9014338 ) ; Germany, Berlin, Sumpfe zwischen Hundekehle und Grunewald , IX.1902, Prager 40 ( MW9014329 ) ; Lithuania, Kaunas, 12.IV.1953, Tyuremnov ( MW9014344 ) ; Russia, Murmansk Province, Apatity City outskirts, north shore of Imandra Lake , aapa-bog, 67.60°N, 30.00°E, 27. VI .2012, Ignatov & Ignatova 12-14, (OK3526, MW9014350 ) ; Leningradskaya Province, Laryanskoe bog, 06.X.1935, Tyuremnov ( MHA9014357 ) ; Pskov Province, Polistovsky National Reserve , Bezhanitskiy District, southern part of Polistovskiy peatland, oligotrophic bog, 57.23°N, 30.43°E, 18. VI .2021, Shkurko & Fedosov T5 -s1 (OK3284, MHA); Novgorod Province, Rdeyskiy National Reserve, Kholmskiy Disrict, eastern part of Polistovskiy peatland, oligotrophic bog, 57.09°N, 30.73°E, 18. VI .2022, Shkurko & Korolkova, 169, (OK3730, MHA9121195 ) ; Moscow Province, Klinskiy District , “Zavidovo”, minerotrophic bog, 56.33°N, 36.15°E, 28. VI .2008, Ignatov & Notov 08-21 (OK3729, MHA9022810 ) ; Arkhangelsk Province, Primorskiy District, Yagry Lake , oligotrophic bog, 10.IX.2001, Churakova 2037 (OK3727, MW9014412 ) ;

Sphagnum talbotianum : Russia, Nenets Autonomous District, Middle course of Adzva River , moss tundra, 67.09°N, 60.59°E, 04.VIII.2009, Ivanov & Donskov 09-344 (OK3701, MHA9102281 ) ; Khanty-Mansiysk Autonomous District, Berezovsky District, eastern slope of Ner-Oyka Mountain , sedge-sphagnum bog with willows, 64.57°N, 59.59°E, 914 m a.s.l., 06.VIII.2013, Lapshina 13-525 (OK3390, MHA9015818 ) ; the same area, sedge-sphagnum bog, 64.58°N, 59.59°E, 960 m a.s.l., 01.VIII.2013, Lapshina 13-507 (OK3698, MHA9015873 ) ; Yamalo-Nenets Autonomous District, Baydara River basin, sedge-sphagnum tundra, 67.53°N, 67.93°E, 31.VII.2018, Popov 307 (OK3692, MHA9102182 ) ; the same area, sedge mire, 70.85°N, 73.93°E, 27.VIII.2022, Dudov, Poloshevetz & Kopeina 15-2 (OK3531, MHA) ; Tazovsky District , tundra, 70.93°N, 74.13°E, 21.VIII.2022, Dudov, Poloshevetz & Kopeina 19-8 (OK3530, MHA) ; Krasnoyarsk Territory, Taimyrsky District, Dikson village area, Kara Sea shore near Velbotov Bay , rich fen 73.56°N, 80.62°E, 07.VIII.2019 Fedosov & Koltysheva 291-3 (OK3278, MHA9102973 ) ; the same area, vicinity of Dikson settlement, wet tundra at base of ridge slope, rich fen, 73.48°N, 80.57°E, 08.VII.2017, Fedosov 17-2-19-6 (OK3279, MHA9102970 ) ; the same area, Bol’shoy Arctichesky State Reserve, vicinity of Meduza Bay , poor fen, 73.39°N, 80.59°E, 24.VII.2019, Fedosov & Koltysheva 274-8 (OK3281, MHA9102982 ) ; Yakutia, Momsky District, Ulakhan-Chistay mountain ridge, forestless bog, on side of a hummock, 64.9°N, 146.417°E, 840 m a.s.l., 19.VII.2018, Ignatov & Ignatova 18- 1915 (OK3275, MHA9028536 ) ; Tyrekhtyakh River downstream, wet larch forest near a lake, 64.17°N, 146.69°E, 25.VII.2018, Ignatov & Ignatova 18-2476 (OK3691, MHA9028800 ) ; Tomponsky district, Kyurbelyakh Creek , wet tundra, 63.13°N, 139.07°E, 950 m a.s.l., 8.VII.2011, Ignatov & Ignatova 11-2170 (OK3393, MHA9102061 ) ; Amur Province, Zeyskiy National Reserve, Tukuringra Mountain Ridge , the sphagnum bog with sparse trees, 54.21°N, 126.98°E, 28.VIII.2016, Dudov & Kozhin Br _0112 (OK3523, MW9079503 ) ; Norsky Nature Reserve , mire with Larix , 52.97°N, 130.17°E, 6.VII.2010, Bezgodov 220 (OK3694, MHA9105692 ) ; Zeya District, Stanovoy Ridge, Tokinsky-Stanovoy National Park , tundra peat bog, 55.74°N, 130.78°E, 1529 m a.s.l., 16.VII.2021, Dudov TSMF0247-355 (OK3341, MHA) ; Sakhalin Province, Nogliksky District, Bolshoy-Gamoray Riv- er range, sphagnum bog, 52.52°N, 143.12°E, 24.VIII.2009, Pisarenko op03309 (OK3697, MHA9102843 ) ; Smirnykh District, Nature Reserve “Vaida Mountain”, northern slope of mountain, 49.88°N, 143.47°E, 750 m a.s.l., 21.VIII.2006, Ignatov & Teleganova 06-3607 (OK3395, MHA9101781 ) ; Chukotka, Vrangel Island , tundra, 2020, D 20m 2 (OK3283, MHA) .

Sphagnum warnstorfii : Russia, Murmansk Province, Apatity City outskirts, north shore of Imandra Lake , aapa-bog, 67.60°N, 30.00°E, 27. VI .2012, Ignatov & Ignatova 12-16 (OK3301, MHA9105709 ); Pechengskiy District , sedge mire with willows, 69.64°N, 32.43°E, 13.VII.2009, Popova 118-m- 27 ( MW9014349 ) ; Moscow Province, Klinskiy District , “Zavidovo”, minerotrophic bog, 56.33°N, 36.15°E, 28. VI .2008, Ignatov & Notov 08-23 (OK3302, MHA9015728 ) ; Nenets Autonomous District, Middle course of Adzva River , tundra with willows, 67.09°N, 60.60°E, 04.VIII.2009, Ivanov & Donskov 09-687 (OK3399, MHA9102309 ) ; Irkutsk Province, Khamar- Daban Mountains , Vicinity of Chersky Peak, minerotrophic bog near lake, 51.49°N, 103.60°E, 1700 m a.s.l., 05.IX.2022, Fedosov & Shkurko 274-1 (OK3706, MHA) ; Amur Province, Zeya District, Stanovoy Ridge, Tokinsky-Stanovoy National Park , tundra peat bog, 55.74°N, 130.79°E, 1548 m a.s.l., 26.VII.2021, Dudov TSMF0265-393 (OK3342, MHA) ; Khabarovk Territory, Badzhal Mountains, Yarap River tributary upper course, lover subalpine belt, minerotrophic bog, 50.35°N, 134.63°E, 1400 m a.s.l., 08.VIII.2016, Fedosov 16-40-s6 (OK3280, MHA9120573 ); Botchinsky National Reserve , pine forest, along stream, 48.30°N, 139.57°E, 12.VIII.2013, Ignatov & Ignatova 13-365 (OK3655, MHA9101775 ) .

Sphagnum warnstorfii 3: Russsia, Yakutia, Oymyakomskiy Disrict , Ust’-Nera Settlement outskirts, grass-sphagnum mire, 64.5°N, 143.72°E, 500 m a.s.l., 5.VIII.2015, Ignatov & Ignatova 15-1564 (OK3401, MHA9102053 ) ; Amur Province, Zeyskiy National Reserve, Zeya District, Stanovoy Ridge , Tokinsky-Stanovoy National Park , tundra peat bog, 55.74°N, 130.78°E, 1529 m a.s.l., 16.VII.2021, Dudov 355-1 (OK3340, MHA) ; Chukotka, Wrangel Island , tundra, 2020, D 25m 4n1 (OK3283, MHA) .