Memoranda Soc. Fauna Flora Fennica 95: 1–35. 2019 ISSN 0373-6873 (print) Helsinki 24 January 2019 ISSN 1796-9816 (online)

The flora and vegetation of Sosnovets Island, the

Mikhail N. Kozhin*, Ekaterina O. Golovina, Ekaterina I. Kopeina, Stanislav A. Kutenkov & Alexander N. Sennikov

Kozhin, M. N., Department of Geobotany, Faculty of Biology, Moscow State University, Leninskye Gory 1–12, GSP–1, 119234 Moscow, ; & Avrorin Polar-Alpine Botanical Garden and Institute of Kola Scientific Centre of Russian Academy of Sciences, 184250 Kirovsk, Region, Russia. E-mail: [email protected] (*Author for correspondence) Golovina, E. O., Laboratory of Vegetation Geography and Mapping, Komarov Botanical Institute of Russian Academy of Sciences, Prof. Popov str. 2, 197376 St. Petersburg, Russia. E-mail: [email protected] Kopeina, E. I., Avrorin Polar-Alpine Botanical Garden and Institute of Kola Scientific Centre of Russian Academy of Sciences, 184250 Kirovsk, Murmansk Region, Russia. E-mail: kopeina-e@ yandex.ru Kutenkov, S. A., Institute of Biology of Karelian Research Centre of the Russian Academy of Sciences, Pushkinskaya str. 11, 185910 Petrozavodsk, Karelia, Russia. E-mail: effort@krc. karelia.ru Sennikov, A.N., Botanical Museum, Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland; & Herbarium, Komarov Botanical Institute of Russian Academy of Sciences, Prof. Popov str. 2, 197376 St. Petersburg, Russia. E-mail: alexander.sennikov@ helsinki.fi

The flora and vegetation of Sosnovets Island (White Sea Throat, Murmansk Region, Russia) has been studied and described in detail. This is a small island situated within the tundra zone, largely covered by a permafrost peatland with the presence of flarks, a palsa mire, and rock outcrops. Vascular of Sosnovets Island include 167 species and subspecies, of which 134 species and subspecies are considered native and 33 species are alien. The number of tundra species is higher and that of boreal species is lower than on the other White Sea islands; a few species with east- ern distributions in East Europe are present; 6 protected species are recorded. Alien species were mostly transported from Region but partly from Central Russia; main pathways were forage, construction and gardens; one species (Alchemilla cymatophylla) was likely introduced as a polemochore. The vegetation of Sosnovets Island is represented by a complex of lichen, dwarf- shrub, cottongrass-sphagnous and sedge-sphagnous communities of the peatland, which covers the major part of the island, as well as dwarf-shrub and lichen tundras, coastal vegetation, willow thickets, dwarf cornel (Cornus suecica) and secondary anthropogenic meadows and grasslands. A palsa mire, marshes with Calamagrostis deschampsioides, highly dissected peatlands with cloud- berry-crowberry-lichen communities on elevated sites and cottongrass-sphagnous communities in depressions are the unique features of the island’s vegetation. 2 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

Introduction Vascular plants of the White Sea islands were included into larger studies in Murmansk Region Insular ecosystems have a certain level of peculi- (Gorodkov 1953–1954; Pojarkova 1956–1966), arity because of their restricted size and isolation, Karelia (Kravchenko 2007) and Arkhangelsk Re- which makes them suitable for testing ideas and gion (Schmidt 2005), although their particular de- principles of ecology and evolution but also leads scriptions and analyses are far from being com- to their vulnerability under climatic changes and prehensive. Nevertheless, there is a long tradition anthropogenic pressure (Drake et al. 2002). The of biodiversity studies of the White Sea, includ- biogeographic context of such studies concerns ing numerous islands along its coasts. species richness and distribution, and also region- Ivan I. Lepekhin was the first naturalist to vis- alization patterns of target study groups. Island it the White Sea islands. In June 1772, upon com- biogeography (MacArthur & Wilson 1967) has pleting his four-year journey across various prov- been developed when the biodiversity of insular inces of the Russian Empire, he visited the So- ecosystems is studied (Haila et al. 1982; Patiño et lovetsky Islands and provided a brief descrip- al. 2017); its results are of special interest when tion of them (Lepekhin 1805). Almost a centu- larger archipelagoes, harbouring many endemic ry thereafter, in 1861, the islands were visited species, are examined (e.g. Bramwell 1976), but by Karl Emil Inberg, an entomologist, and Gus- may also reveal important dependencies when taf Selin, a botanist, as a group of the Finnish ex- smaller and minor insular groups are considered pedition to Russian Lapland supported by Socie- (e.g. Rebassoo 1987). tas pro Fauna et Flora Fennica and the Consisto-

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38° 40° Figure 1. Sosnovets Island as a study area, and the route of our botanical expeditions to the south-eastern coast of the , July–August 2015–2016. Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 3 ry of the University of Helsinki. They stayed on published based on an expedition of the North- the islands for two weeks, 11–25 July, and pre- ern (Arctic) Federal University in 2012 (Churak- pared the first scientific collections (insects and ova et al. 2016). plants) from the White Sea islands (Sennikov & As a result of these activities, numerous an- Kozhin 2018). For a certain period, further floris- notated check-lists, vegetation descriptions and tic studies on the White Sea islands were sporad- analyses of human and natural dynamics of the is- ic and non-organized (Lindberg 1914; Pobedimo- land floras were published. However, all the pre- va et al. 1960; Uotila 2013). vious studies were performed in the taiga zone When the Kandalaksha Strict Nature Reserve and mainly in the western part of the White Sea. was founded in 1932, regular studies of the flo- To extend the studies into the tundra zone, we ra and vegetation had been established on the is- have started a new initiative that aimed at reveal- lands. A number of insular systems in the Kan- ing the flora and describing the vegetation of the dalaksha Bay were explored, first floristic check- islands situated along the north-eastern coast of lists (e.g. Bogdanova & Vekhov 1969a, 1969b; the White Sea. Vorob’eva 1986, 1989, 1996b; Zherikhina & On this way, Sosnovets Island was selected as Moskvicheva 2006), descriptions and classifica- a case study. We described its complete flora and tions of their flora and vegetation (Breslina 1987; communities in order to provide insights Vorob’eva 1996a; Kozhin 2011) were compiled. into the features of the northern islands’ species The flora of the , the largest ar- composition, the vegetation of the tundra zone chipelago situated in the outer part of the and its anthropogenic transformation. Bay, was regularly studied by the Moscow State University in 1980–2000 (e.g. Kiseleva et al. 1997), with later additions from other research- Study area ers (e.g. Savinov & Semashko 2014). Minor is- lands at the border of Murmansk Region and Ka- Sosnovets Island (Figs. 1, 2) is located 8 km south relia were explored by student expeditions from of the Arctic Circle near the south-eastern part of Moscow (Shipunov & Abramova 2006). Since the Tersky Coast in the White Sea Throat, at 66° 2000, the floras of the other islands 29′ N, 40° 41′ E. The island has an area of 40.6 have been studied by Karelian Research Centre ha, a length of 1385 m, a width of 445 m, and a of the Russian Academy of Sciences (Kravchen- perimeter of ca 3.5 km. The summit plain is about ko et al. 2002, 2008, 2015). Recently, the infor- 10–15 m above sea level and the maximum ele- mation on the island floras of the was vation situated in the southern part of the island

Figure 2. General view of Sosnovets Island from a powered paraglider. Photo: Boris B. Vakhmistrov. 4 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 is 16.2 m above sea level. The Sosnovets light- part of the Tersky Coast, the climate is most se- house is located in the central part of the island. vere among all the other regions of the Sea (Yak- The southern shore is lower and flatter than the ovlev 1961). As observed at the Sosnovets weath- northern and western ones. The tideland width er station (ID 22355), the average annual temper- comprises from 10 to 90 m. Sosnovetskaya Sal- ature during a long-term observation period ac- ma Strait with the depth of 6–11 m separates the counted for –0.66°C. However, due to the trend island from the mainland. The minimal distance towards temperature rise in the last two dec- to the mainland is 3.3 km from the cape on the left ades the average annual temperature has reached side of the Sosnovka River mouth. +0.24°C (www.rp5.ru; Novakovsky & Elsakov The geological structure of the Tersky Coast 2014). The average long-term annual frostless is formed by the eastern margin of the Baltic period comprises 150 days. The annual precipi- Shield. The island is formed by Neoarchean gran- tation equals ca. 400 mm mostly occurring in the ite-gneisses, gneisses, migmatites and amphib- warm season with the recent tendency to increase. olites, which belong to the Purnach Block near The snow layer with a depth of 10–20 cm, seldom its border with the Tersky Block (Pozhilenko et 30 cm, usually does not cover the entire land sur- al. 2002). The island’s basement rock is overlaid face. The air humidity is constantly high through- with soft, mostly sandy deposits of the early Hol- out the year, varying from 85.9 (May) to 91.8 ocene marine terrace. Along with the eastern part (August), with the average value of 88.2%. High of the Kola Peninsula, Sosnovets Island is under- tides are typical of the study area, comprising 3.5 going tectonic subsidence (Koshechkin 1979). m on average and reaching 4.3 m in syzygy. Tidal The sea-level rise leads to continuous erosion currents are widespread in the White Sea Throat; of loose deposits on the island’s periphery. The in Sosnovetskaya Salma Strait, the high tide cur- abrasion of the Holocene deposits results in steep rent is directed southwards, whereas the low tide shore slopes as well as basement rock exposure. current is directed northwards with a stream ve- The major part of the island, namely its sum- locity of 1–2 knots (Anonymous 1954). mit plain and some slopes, is covered by a heavy layer of peatland with a thickness of 1–2 m. Such peatlands were termed ’arctic bog’ by Zinserling (1934). The surface of the peatland is undulate; its elevated and marginal parts are divided by Attributing the study area to the cracks into polygonal blocks with an area of 1–10 existing vegetation divisions m2, whereas flarks are formed in the lowest parts with a relatively small area. According to macro- Sosnovets Island is located on the limit between fossil analysis, the island’s peat deposit belongs the tundra and forest-tundra zones, which ac- to the raised-bog (ombrotrophic) type; its high- counts for the ambiguity in attributing it to a par- er layer consists of loose, dwarf-shrub peat, while ticular unit of vegetation division. According to the lower one is composed of denser, cottongrass the vegetation map of Murmansk Region (Cher- peat (Kutenkov et al. 2018). The flarks are filled nov 1956), a narrow stripe along the coasts of the with fen sedge peat. Permafrost peatlands cover Barents and White seas from the Sem’ Ostrovov the major part of the island with a seasonal thaw to the Ponoi River mouth belongs to the ’typical depth of about 50–60 cm. In the central part of the tundra’. Besides, the islands situated along this island, north of the weather station, the peatland stripe and further south (including Sosnovets Is- topography is more prominent and approaching land) were attributed to the ’typical tundra’ as that of palsa bogs. The southern part of the island well. In later vegetation studies, all the tundra ar- is characterized by an undulating relief, stone and eas of Murmansk Region were referred to as the sand deposits with rock outcrops, the absence of southern tundra subzone, which was reflected on permafrost and a considerably thinner peat layer. vegetation maps of Murmansk Region and the The study area has a subarctic climate with USSR. In particular, the vegetation of the White long mild winters and cool summers, and a con- Sea coast in the vicinity of Sosnovka Village and siderable impact of the White Sea. In the eastern the adjacent islands was classified as dwarf shrub Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 5 tundra (Chernov 1971; Isachenko & Lavrenko ago and Rusinga Bay (Fig. 1). The field work on 1975). Sosnovets Island was performed in two days, 7–8 According to the scheme of vegetation zones August 2016. of the Nonblack-Earth Region of Russia, which The island flora was recorded and described includes the boreal and tundra regions of the Eu- during field excursions. Firstly, we walked ropean part of the USSR (Alexandrova & Yurk- around the island twice to study the shore habi- ovskaya 1989), Sosnovets Island occupies an in- tats in detail. Secondly, we studied the whole is- termediate position between the Iokanga-Ponoi land by a network of parallel routes paying par- vegetation district, Kola subprovince, European ticular attention to specific natural and human-in- and West Siberian tundra province, and the Po- fluenced habitats. As a result of visiting all the noi-Tersky vegetation district, Kola-Karelia sub- habitats throughout the island, the entire checklist province, North European taiga province. Ac- was compiled and the occurrence of species was cording to the vegetation zones of north-western assessed using a five-grade scale: Un (unique) – Europe (Ahti et al. 1968), the coastal area and the one or two occurrences of a species on the island; islands in the White Sea Throat south of the Po- Rar (rare) – rare plants (less than 10 individuals) noi River mouth do not belong to the tundra zone. observed in several locations; Sp (sparse) – plants This delimitation of tundra is accepted in Alex- are quite abundant but occupy less than 1/20 of androva (1977). In Safronova et al. (1999), the the island; Fr (frequent) – plants cover from 1/20 eastern part of the Kola Peninsula is attributed to 1/2 of the island; Com (common) – plants are to the East Scandinavian (Kola) vegetation dis- widespread and cover more than a half of the is- trict of the southern hypoarctic tundra subzone. land. The checklist included all the natural and In the up-to-date circumpolar map of the Arc- alien species. When detecting alien species, par- tic (CAVM Team 2003; Walker et al. 2005), the ticular emphasis was placed on their habitat type Kola Peninsula has been excluded from the low- and degree of naturalization. The checklist with shrub tundra subzone. However, a recent analysis species attributions to particular habitat types is of the vegetation and climate patterns by Korole- placed in Appendix 1. va (2006b) and Virtanen et al. (2015) suggested Phytosociological data were collected accord- that the scheme of Walker et al. (2005) may need ing to the relevé method of vegetation sampling revision. (Yunatov 1964) at sites with homogeneous mi- Based on our flora and vegetation knowledge, crotopography and moisture. Plots of 25 m2 were we accept the attribution of Sosnovets Island to used for large-scale communities, mostly in tun- the south subarctic tundra of Alexandrova (1977), dra, whereas small-scale communities, most- otherwise called the south tundra in Chernov & ly in flarks and on meadows, were described in Matveeva (1979). A more precise zonation of the their natural limits. When compiling phytosocio- Kola Peninsula is outside the scope of the pre- logical relevés, precise geographical coordinates sent paper. (WGS-84) as well as meso-relief position and mi- cro-relief features were recorded; species abun- dance according to Braun-Blanquet scale or cov- Field work and data analysis er was estimated. Phytosociological data covered all types of the island’s vegetation. In total, 76 rel- The research material was collected during our evés were obtained. botanical expedition to the south-eastern coast of A map of the island’s vegetation was com- the Kola Peninsula from the Varzuga River mouth piled according to the authors’ field observations to the Rusinga River (Lapponia Varsugae and and a set of relevés. The vegetation was visually Lapponia Ponojensis) in July – August 2015 and interpreted using open-access satellite high res- 2016. The expedition proceeded largely along olution images. The plant community classifica- the coastline near Sosnovka Village, Danilov Is- tion was based on the dominant (ecological-phy- land, Danilovskaya Bay, the Ponoi River mouth, tocoenotic) approach (Alexandrova 1969). To abandoned Ponoi Village, Goryainov Island, Al- identify plant communities, dominant species and dobinskaya Guba Bay, the Tri Ostrova Archipel- species-indicators of environmental conditions 6 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 were taken into account. In homogeneous vege- exander von Schrenk, a plant collector of the Bo- tation, the main mapping units were community tanical Garden in Saint-Petersburg, and Wilhelm types, corresponding to association, subassocia- Boehtlingk, a geographer and geologist. On their tion or variant of local hierarchy. For heteroge- route the naturalists travelled near Sosnovets Is- neous vegetation that may be difficult to visual- land (Fellman 1869). In 1840, the expedition of ize as separate contours, with regular alternations Baer with Alexander Theodor von Middendorff in a certain topographic site, complexes (in case as a zoologist and student Pankevich as a collect- of peatland vegetation) and ecological series (in ing assistant explored the Kola Peninsula. Their case of shore meadows) were used as mapping vessel landed near Sosnovka Village on 15 June, units. The legend consists of 24 basic units and and foul weather made them stay there for four 4 extensions marked with an asterisk, which de- days, during which the naturalists explored the note the communities found with the basic ones. neighbouring territories (Raikov 1961). The map and its legend are placed in Appendix 2. In 1841, the route of Franz Josef Ruprecht’s Collected specimens of vascular plants, moss- expedition to Malozemelskaya tundra (’ter- es, liverworts and lichens were deposited in the ra Samojedorum’: Ruprecht 1845) passed near Herbaria of Moscow State University (MW), Sosnovets Island, where he stopped and collect- University of Helsinki (H), Forest Research In- ed several specimens probably on the adjacent stitute of Karelian Research Centre (PTZ), Kan- mainland. dalaksha State Nature Reserve and Polar-Alpine Botanical collections of these academic ex- Botanical Garden-Institute (KPABG). peditions, supplied with extremely brief labels, were deposited in Komarov Botanical Institute (LE). To determine whether the specimens from History of botanical studies ’Sosnovets’ were actually collected on the main- land or on the island poses unavoidable difficul- In the early 19th century, scientists started explor- ties as in the 19th century the village and the island ing the coast and the islands of the White Sea were known under the same name (Baer 1837). Throat. The first expeditions to the area were di- We assume that, most likely, the island was not rected by the Imperial Saint-Petersburg Academy visited by these early expeditions. of Sciences. The first naturalists known to visit Sosnovets In 1837, naturalist Karl von Baer, botanist Island were entomologist Johan [John] Reinhold Alexander Lehmann and draughtsman Karl von Sahlberg and botanist August Johan Malmberg Röder travelled from Arkhangelsk to Novaya (later Aukusti Juhana Mela) supported by Soci- Zemlya. They reached the Kola Peninsula in the etas pro Fauna et Flora Fennica. In 1870, they vicinity of Pyalitsa Village on 2 July and stayed travelled across Russian Lapland and obtained there for a few days. Then they passed by Sos- vast collections of vascular plants (Uotila 2013). novets Island and arrived at the Ponoi River mouth Sahlberg and Malmberg arrived at the island and later at the Tri Ostrova Archipelago and fur- on 26 August and collected a few specimens of ther proceeded to Novaya Zemlya via the Barents Arctanthemum arcticum subsp. polare, Stellaria Sea (Raikov 1961). On the way back the expedi- humifusa and Tripleurospermum maritimum (de- tion vessel anchored near Sosnovets Island at the posited in H). beginning of September and stayed here for sev- In 1889, Alfred Oswald Kihlman (later Kaira- eral days due to foul weather. They explored the mo) visited Sosnovka Village (’Sosnovets’), ex- nearest vicinities of the Sosnovka River mouth, a plored the vicinities and went upstream the Sos- place populated by the Saami. They provided sev- novka River to Babozero (Akjawr) Lake in 12–18 eral naturalistic descriptions and noted that the lo- August. He studied the adjacent tundra, paludi- cal forest almost reached the coastline, which is fied places and marshes (Kihlman 1890). Kihl- untypical of the south-eastern part of the Tersky man never visited the island itself; all his speci- Coast (Baer 1837a, 1837b). mens deposited in H and attributed to ’Sosnovets’ In 1839, the Academy organized an expedi- refer to the village rather than the island. tion to Eastern Lapland with participation of Al- Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 7

In 1913, Constantin Regel travelled across century. Nevertheless, the results of these stud- the Kola Peninsula and stayed at Sosnovka Vil- ies were left unpublished and the available data lage (’Sosnovets’) in 12–18 August. Following remained scarce. The name ’Sosnovets’ was fre- the previous researchers, he studied the tundra, quently mentioned in historical literature and on marshes and riparian vegetation on plains and herbarium labels (Ruprecht 1845; Hjelt 1895, small hills in the vicinities of the village; Regel 1902, 1926), but this information was uncertain also visited Babozero (Akjawr) Lake and the Ba- because the village and the island were known bja (Akjok) River (Regel 1927). He never reached under the same name in early usage. In herbari- the island either. um collections, this name may refer to the main- In 1926, the first phenological observations land part of the locality, i.e. Sosnovka Village (the were conducted on the island by B. Vorontsov, collections of Baer, Kihlman and Schrenk), or to a staff member of the weather station (Smirnov the island (the collections of Malmberg and Sahl- 1927). He was also interested in botany and was berg). the first to detect the larch on the Kola Penin- sula near Sosnovka Village (Tolmatschev 1925; Kozhin & Sennikov 2016). The following year, Human occupation an expedition of the Botanical Garden in Saint- Petersburg directed by Yuri D. Zinserling stud- Sosnovets Island is situated in the White Sea ied the island. Zinserling (1934) described the is- Throat near the sailing lines from the west and land’s peatlands as ’arctic bog’, a bog type that north to Arkhangelsk. Sosnovetskaya Salma seldom occurs in the north-west of the European Strait is the sole safe roadstead for vessels in the part of the USSR. He recorded permafrost peat- White Sea Throat, which attracted seafarers af- lands throughout the island, which were thaw- ter crossing the White Sea from the Summer and ing to depths of 30–40 cm at the end of summer, Winter coasts to the Tersky Coast and helped and Empetrum nigrum subsp. hermaphroditum them to avoid storms and wait out foul weather. and Rubus chamaemorus-dominated oligotroph- On the western shore of the island several wood- ic bogs with patches of crust-lichen vegetation, en crosses were installed to facilitate marine nav- which he found widespread. igation. Because of this feature, Sosnovets Island More than half a century after Zinserling, in commonly passed under the name of ’Cros ilond’, July and August of 1986, the Kola geobotanical or ’Krestovii’ on early foreign maps and in trav- expedition from the Polar-Alpine Botanical Gar- elogues (Reineke 1850; Komaritsin et al. 2001). den-Institute of Russian Academy of Sciences, In 1822, a navigation watchtower was con- directed by Irina P. Breslina, worked in the south- structed on the island at the direction of Arkhan- east of the Kola Peninsula (Breslina 1986). The gelsk Marine Service. The tower was fitted with landscape of Sosnovets Island was described by wooden planks and painted red. Saami, who lived the expedition, several relevés were obtained, a near the Sosnovka River mouth, were in charge preliminary checklist of vascular plants was com- of maintaining the tower, whereas the island re- piled, and a substantial number of herbarium mained uninhabited (Reineke 1850). specimens (deposited in KPABG and MW) were In 1862, a new lighthouse was construct- collected. A quantitative and qualitative analyses ed, with five buildings erected on its southern of the island’s flora was planned but this study side for the lighthouse keeper and seven main- had not been eventually completed, and even a tenance staff, and for food and kerosene supply. draft floristic checklist had not been preserved in Since that year, meteorological data were regu- the archives. In 2000–2002, Natalya R. Kirillo- larly recorded by the lighthouse keeper. In 1896, va (Kaneva), a student of the Kola Branch of the Russian Hydrographic Service organized a sepa- Petrozavodsk State University, visited the island rate weather station on the island to continuous- and collected field material for her diploma the- ly obtain meteorological and hydrological data. sis (Kaneva 2002). In 1909, the lighthouse area was reconstructed, As we see, botanists have repeatedly visit- a new lighthouse was built, with a chapel, two ed Sosnovets Island since the middle of the 19th dwelling houses, an alarm flagpole, two fog guns 8 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 and a weather box. In 1920, the weather station peat mounds with a diameter of 15–40 m and a was handed over to Meteorological Service of the height of up to 2 m are located among the dells Russian Soviet Federative Socialist Republic. At (Fig. 3.2). On its southern margin, near the weath- that time the station already had a separate house er station, the peatland is morphologically simi- and its own staff. In addition to the other activi- lar to palsa mire and is treated as such (Fig. 3.3). ties, rock blasting and drilling were conducted on Small pools occasionally occur in highly dissect- the island to construct a tide gauge in the 1930s ed parts of the peatland. South of the weather sta- (Komaritsin et al. 2001). tion, where the summit plain gradually decreas- Consequently, a permanent population and es towards its southernmost tip, the central part a number of residential and technical buildings of the island is covered with a thin discontinuous have been present on the island for more than layer of dry peat without permafrost; the relief of one and a half centuries. At present, most of the the peatland is smooth, lacking mounds and ex- buildings have not been used for a long time and pressed depressions (dells). are currently abandoned. Food supply and fuel to The vegetation cover of the permafrost peat- maintain the weather station and the lighthouse land is represented by several community types are annually delivered to the island from Arkhan- distributed across different topographic sites. gelsk. Noteworthy is that the weather station and Peat mounds, flat surfaces, round gently elevat- the lighthouse have always belonged to marine ed sites with indistinct wide depressions in be- and weather services of Arkhangelsk and almost tween, characteristic of weakly dissected parts of all the cargoes have been carried from the remote the peatland, are occupied by cloudberry (Rubus in Arkhangelsk Region rather chamaemorus)-crowberry (Empetrum nigrum than from the neighbouring Tersky Coast in Mur- subsp. hermaphroditum)-lichen communities mansk Region. (Fig. 3.4). Species composition of their herb layer (cover of up to 40–50%) is extremely poor; these communities often lack species other than dom- Vegetation inants: Empetrum nigrum subsp. hermaphrodi- tum and Rubus chamaemorus. Sometimes Vac- The vegetation of the island is represented by a cinium vitis-idaea and some other species may complex of lichen, dwarf-shrub, cottongrass- occur in such communities. In the lichen cover, sphagnous and sedge-sphagnous communities of Cladonia arbuscula, C. rangiferina, Flavocetrar- the peatland, which covers the major part of the ia nivalis and Cetraria islandica dominate and island, as well as dwarf-shrub and lichen tundras, some chionophobous species (Bryocaulon diver- coastal vegetation, willow thickets, dwarf cornel gens, Alectoria ochroleuca, Gowardia nigricans, (Cornus suecica) and secondary anthropogenic Sphaerophorus globosus) are constant. The cover meadows and grasslands (Appendix 2). of mosses (Dicranum elongatum and Pohlia nu- The summit plain of the island is largely cov- tans, typical of dry peatlands) and liverworts (Pti- ered with a permafrost peatland in its northern lidium ciliare, Sphenolobus minutus) does not ex- part. It is weakly dissected in the north and north- ceed 1%. The tops of mounds and gently elevat- east: flat surface areas of the summit plain alter- ed sites are cracked into polygonal blocks when nate with round gently elevated sites separated by drying. In their centres, lichen patches of crus- wide and shallow depressions; dells draining the tose Ochrolechia frigida and cup-shaped Clad- peatland, shallow and low-branched, are direct- onia species (C. deformis, C. borealis, C. squa- ed from the centre of the island to its east and mosa, C. fimbriata etc.) sporadically occur (Fig. west shores (Fig. 3.1). The permafrost peatland 3.1); patches of bare peat are uncommon. is more dissected in the western and especially In the north and east of the island, the li- southern parts of the island, near the lighthouse chen abundance markedly decreases in the peat- village; the dells are also directed mainly to the land margin near the shore slopes and cloudber- eastern and western shores of the island, where ry-crowberry communities are formed, similar they become deeper, longer and forming a high- to those described above but with the increased ly branched network. Round and elongated frost participation of Empetrum nigrum subsp. her- Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 9 maphroditum (75–90%) and the reduced cov- Schistochilopsis incisa etc.) appear with a cover er of lichens (5–10%) (Fig. 3.5). We frequently of up to 7%. observed this negative correlation between the In deep and well-expressed depressions be- lichen and crowberry abundances in the shore tween peat mounds, cottongrass- and sedge- parts of islands and on remote islets also in the sphagnous flarks are formed. The former are typ- other regions of the White Sea. Apparently, this ical of relatively narrow depressions. The cotton- trend may result from the greatest values of sur- grass-sphagnous flarks with a small number of face air moisture at such sites, which favours ac- species are most frequent. In such communities, tive crowberry growth (Tybirk et al. 2000; Vasi- Eriophorum scheuchzeri dominates with Carex levich & Bibikova 2011). rariflora as a co-dominant in the herb layer (Fig. Similar cloudberry-crowberry communi- 3.7). Other species of vascular plants are repre- ties occur also in the inner part of Sosnovets Is- sented by single individuals; yet only cloudber- land, around the lighthouse village and the weath- ry is constant while the others (Carex aquatilis, er station, as well as on a thin peat layer with- C. canescens, Cornus suecica, Stellaria borea- out permafrost. These communities are charac- lis) sporadically occur. In the moss layer, Sphag- terized by the presence of herbs including mead- num riparium and S. lindbergii are dominant and ow and weedy species: Chamaenerion angustifo- S. balticum and Warnstorfia fluitans are common lium, Calamagrostis neglecta subsp. groenlandi- with a cover of up to 5% and less than 1%, corre- ca, Poa alpigena, Solidago virgaurea subsp. lap- spondingly; liverworts are not regular. ponica, Avenella flexuosa, Festuca ovina and Eq- Cottongrass-sphagnous flarks with Eriopho- uisetum arvense subsp. alpestre. In the herb layer, rum angustifolium as a dominant and E. scheuchz- Cornus suecica is occasionally co-dominant; the eri as a co-dominant are less frequent. They are cover of Vaccinium vitis-idaea is increased; in the similar to the abovementioned community type moss layer, Polytrichum hyperboreum is present. in accompanying species of the herb layer; how- The low abundance of lichens and the presence of ever, Sphagnum angustifolium dominates in their herbs in these communities is more likely to be moss layer, whereas the cover of S. riparium and connected with constant trampling, not only by S. lindbergii does not exceed 1%. people but also by horses which used to be kept Sedge-cottongrass-sphagnous communities there, as well as with other types of long-term hu- with the dominance of Eriophorum russeolum man disturbance rather than with habitat features. and Carex rariflora in the herb layer occur in Shallow depressions among elevated sites of flarks of the palsa mire near the weather station permafrost peatland are occupied by cloudber- at slightly less moist sites. In these communities, ry (Rubus chamaemorus)-dominated commu- dwarf shrubs Andromeda polifolia, Empetrum ni- nities (Fig. 3.6), which are either monodominant grum subsp. hermaphroditum (cover of up to 7%) or with crowberry and dwarf cornel as co-domi- and Vaccinium vitis-idaea (cover less than 1%) nants (cover of 15–25%). Other species of vascu- are common. Except the dominants, herbs are lar plants (Dryopteris expansa, lapponi- not abundant yet quite numerous; Coptidium pal- cus, Luzula wahlenbergii, Vaccinium vitis-idaea) lasii, Stellaria borealis, Epilobium palustre, Lu- and cryptogams (Dicranum elongatum, D. majus, zula wahlenbergii, Eriophorum angustifolium, E. Lophozia silvicola, Cladonia rangiferina, Cetrar- vaginatum, Rubus chamaemorus etc. being most ia islandica etc.) are represented by single indi- typical. In the moss layer, Sphagnum flexuosum viduals. As the depressions deepened, Carex rar- dominates whereas Aulacomnium palustre and iflora with a cover of 10%, Andromeda polifolia Straminergon stramineum are abundant. Apart and Vaccinium uliginosum appear in the commu- from the common species like Sphagnum ripar- nities. In the moss layer, the abundance of Dicra- ium and S. lindbergii, other mosses (Sphagnum num elongatum increases and sphagnous and oth- russowii, S. fimbriatum, Dicranum laevidens) and er hydrophilous mosses (Sphagnum lindbergii, S. liverworts (Lophozia silvicola, Calypogeia muel- riparium, Polytrichum jensenii, Warnstorfia flu- leriana, Cephalozia spp.) are present. itans) and liverworts (Cephalozia lunulifolia, C. Carex aquatilis-dominated sedge-sphag- bicuspidata, Mylia anomala, Riccardia latifrons, nous communities are characteristic of wide 10 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

Figure 3. Plant communities of Sosnovets Island. 1. Weakly dissected permafrost peatland with cloudberry-crowberry-lichen­ communities in the north-eastern part of the island; in the foreground, on the tops of peatland mounds, patches of crust lichen­ Ochrolechia frigida and cup-shaped Cladonia sp. are visible. 2. Highly dissected peatland with cloudberry-crowberry-li- chen communities on elevated sites and cloudberry, cottongrass-sphagnous communities in depressions. 3. Palsa mire be- tween the weather station and the lighthouse. 4. Cloudberry-crowberry-lichen community. 5. Cloudberry-crowberry commu- nity in the peatland’s margin near the shore slope. 6. Cloudberry community in a shallow depression among elevated sites of the permafrost peatland. 7. Flark with the dominance of Eriophorum sсheuchzeri, Sphagnum lindbergii, S. riparium among ele- vated sites of the permafrost peatland. 8. Sedge-sphagnous community with Carex rariflorain the herb layer and Sphagnum fimbriatum, S. lindbergii, and S. riparium in the moss layer in the lower part of a dell near the coastline. Photos: 1 – E. I. Kopeina; 2, 3, 7 – M. N. Kozhin; 4, 8 – E. O. Golovina; 5, 6 – S. A. Kutenkov. Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 11 flarks; they mainly occur in the palsa mire near nii with a cover of up to 10–15%. The herb layer the weather station. The cover of Carex aquatil- is represented by Carex rariflora as a dominant, is amounts to 40–50%, with other species of the as well as dwarf shrubs Empetrum nigrum subsp. herb layer: Carex rotundata (cover of up to 5%), hermaphroditum and Andromeda polifolia along Epilobium palustre, Ranunculus hyperboreus, with Rubus chamaemorus and Cornus suecica as Rumex lapponicus, Luzula wahlenbergii, Agros- co-dominants with a cover of 5–20%. Due to the tis mertensii, vivipara, Carex canescens, increase in moisture, sedge-sphagnous commu- Rubus chamaemorus, Montia fontana etc. Dwarf nities are formed with the dominance of Carex shrubs, namely Andromeda polifolia, Empetrum aquatilis and C. rariflora in the herb layer and nigrum subsp. hermaphroditum, Vaccinium ox- Sphagnum balticum in the moss layer, with the ycoccos, are sparse. Sphagnum riparium domi- absence of dwarf-shrubs and the increasing abun- nates in the moss layer; Sphagnum squarrosum, dance of S. riparium. Larger waterlogged depres- Straminergon stramineum, Aulacomnium palus- sions on the western shore of the island are occu- tre, Warnstorfia exannulata, Scapania paludicu- pied by sedge-sphagnous communities with the la are common but not abundant; other species, dominance of Carex aquatilis and Sphagnum ri- for example, Sphagnum lindbergii, Polytrichum parium, similar to those formed in flarks of the jensenii, were found seldom with low cover. palsa mire. Nevertheless, these communities dif- At waterlogged sites, which have degraded as fer by a high abundance of Carex rariflora and a result of human transformation of flarks, dense Polytrichum jensenii, as well as by the presence sedge-grass communities are formed. Calama- of coastal species such as Calamagrostis des- grostis neglecta subsp. groenlandica, Phleum al- champsioides, Parnassia palustris, Arctanthe- pinum and Carex aquatilis dominate in the herb mum arcticum subsp. polare. layer, accompanied by Chamaenerion angustifoli- On slopes of the eastern and western shores of um, Rumex lapponicus and Rubus chamaemorus. the island, large areas are covered with peat de- Sphagnum spp. and Warnstorfia fluitans are pre- posits. On the eastern shore, which is flatter, the sent in the moss layer. summit plain slope and the adjacent shore line The dells draining the permafrost peatland 15–30 m wide are occupied by dwarf cornell- gradually descend and widen towards the is- cloudberry-crowberry community. Its dense land’s shoreline. In these depressions, various herb layer (cover of about 95%) is represented by sedge-sphagnous communities with a peculiar Empetrum nigrum subsp. hermaphroditum as a moss layer composition occur. The communities dominant and Cornus suecica (cover of 25–30%) of small depressions are characterized by a poly- with Rubus chamaemorus as co-dominants. Vac- dominant moss-lichen layer. In particular, Sphag- cinium vitis-idaea (cover of about 10%) is abun- num fimbriatum, S. lindbergii, S. riparium, Pol- dant; Vaccinium uliginosum, Luzula wahlenber- ytrichum jensenii and Warnstorfia fluitans are gii, Poa alpigena, Equisetum arvense subsp. al- abundant there, and Lophozia silvicola and Cetra- pestre are typical; low shrubs of Salix glauca are riella delisei are typical (Fig. 3.8). Carex rariflo- sometimes present. The moss-lichen layer repre- ra is the major dominant of the herb layer, some- sented by Cladonia arbuscula, C. stellaris, Fla- times accompanied by Eriophorum scheuchzeri vocetraria nivalis, Dicranum majus and Ptilidi- or Carex recta as a co-dominant. Cornus sueci- um ciliare is fragmentary. The extensive cover of ca, Rubus chamaemorus and Empetrum nigrum Cornus suecica is probably determined by a high- subsp. hermaphroditum are constant; other herbs er snow accumulation. Along the peatland margin and dwarf shrubs (Epilobium palustre, Stellaria on the western shore, which is steeper, cloudber- borealis, Arctanthemum arcticum subsp. polare, ry and crowberry-cloudberry communities sim- Andromeda polifolia, Vaccinium uliginosum) are ilar to those occupying shallow depressions on sparse. Shrub willows, Salix phylicifolia and S. the summit plain are widespread. glauca, sometimes appear in these communities. Carex rariflora appears as a co-dominant In larger depressions, the moss layer mostly con- in paludified crowberry communities on peat- sists of Sphagnum balticum accompanied by S. lands of shore slopes with slightly moist condi- flexuosum, S. lindbergii and Polytrichum jense- tions. In more moist depressions below the sum- 12 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 mit plain slope, where snow is accumulated, Em- A peculiar dwarf shrub community was once petrum nigrum subsp. hermaphroditum, Rubus observed on the SE-faced slope of the summit chamaemorus, Carex rariflora and Cornus sueci- plain in the south of the island. Its species com- ca dominate, and Rumex lapponicus, Stellaria position is influenced not only by a greater snow borealis, Bistorta vivipara, Arctanthemum arcti- accumulation in winter and a sufficient insola- cum subsp. polare are typical, whereas only sin- tion in summer but also by anthropogenic distur- gle thallomes of Cladonia rangiferina and single bance. Its herb layer is polydominant; Empetrum individuals of Dicranum majus are present in the nigrum subsp. hermaphroditum, Vaccinium uligi- moss-lichen layer. nosum, V. vitis-idaea, V. myrtillus, Cornus sueci- Sedge (Carex rariflora)-crowberry commu- ca and Equisetum arvense subsp. alpestre domi- nities are formed in marginal parts of the peat- nate; dwarf shrub Arctous alpina and numerous land, adjacent to the shore, with a thin peat lay- herb species such as Avenella flexuosa, Solidago er and a constant water inflow (Fig. 4.1). They virgaurea subsp. lapponica, Hieracium atratum, have a lesser number of species: H. nigrescens, Euphrasia wettsteinii, Phleum al- Rubus chamaemorus, Cornus suecica, Rhodio- pinum, Festuca ovina, Lysimachia europaea, Lu- la rosea are present with low abundance in ad- zula multiflora subsp. frigida, Trifolium repens dition to Empetrum nigrum subsp. hermaphrodi- are present with low abundance. Dicranum sco- tum and Carex rariflora. The moss-lichen layer is parium and Polytrichum juniperinum dominate; more prominent due to a high abundance of nu- Pleurozium schreberi, Hylocomium splendens, merous lichen species and their greater cover of Sciuro-hypnum curtum, Sanionia uncinata, Bar- up to 3–5%, which may be connected with a less- bilophozia lycopodioides are typical in the sparse er moisture and a lesser level of snow accumula- moss-lichen layer. These species are otherwise tion. Along with Cladonia rangiferina, other li- rare in dwarf shrub and lichen communities of chens such as C. arbuscula, C. uncialis, C. amau- the island. Lichen-crowberry communities with rocraea, Cetraria islandica, Sphaerophorus glo- the presence of dwarf cornel in the herb layer and bosus, Flavocetraria nivalis are present. Sedge- Cladonia arbuscula, Cetraria islandica, Flavoce- crowberry communities are more widespread on traria nivalis in the moss-lichen layer are charac- the western shore and in the southern part of the teristic of the north-western slope of the summit eastern shore, where a peat layer is thinner. plain in the southernmost tip of the island. The Tundra communities are formed on loose de- area of these communities is not large. posits and occupy a limited area in the southern- Willow thickets with forbs of slopes and val- most tip of the island. Crowberry tundras with leys shielded from strong winds are widespread a low species number are typical of the summit in the eastern side of the island’s southern part. plain, whereas crowberry-lichen Flavocetraria Salix phylicifolia, S. glauca, S. lapponum form a nivalis- and Cladonia arbuscula-dominated tun- dense shrub layer with a height of 0.5–1.5 m. The dras with the presence of chionophobous Gowar- herb layer usually consists of Calamagrostis ne- dia nigricans and Alectoria ochroleuca occur at glecta subsp. groenlandica, Rumex lapponicus, elevated sites. Single individuals of cloudber- Vaccinium uliginosum, Empetrum nigrum subsp. ry are seldom observed in these communities. hermaphroditum, Filipendula ulmaria, Bistor- Crowberry and lichen communities are usually ta vivipara, Cornus suecica, Veratrum album, in- disturbed by anthropogenic impact. Meadow and cluding Comarum palustre at waterlogged sites. weedy species such as Chamaenerion angusti- The sparse moss layer is represented by the cover folium, Trifolium repens, Calamagrostis neglec- of Sanionia uncinata often containing hygrophil- ta subsp. groenlandica, Poa alpigena, Equisetum ous mosses such as Calliergon giganteum, Bryum arvense subsp. alpestre also appear in these com- pseudotriquetrum, Aulacomnium palustre. Alien munities. The tundra does not form a continuous species Deschampsia cespitosa, Poa pratensis, cover and alternates with homogeneous groups Vicia cracca, Phleum pratense, Sagina procum- of Chamaenerion angustifolium, Deschamp- bens indicate that these communities were part- sia cespitosa and Calamagrostis neglecta subsp. ly transformed as a result of human activity in the groenlandica. 20th century. Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 13

Dwarf cornel (Cornus suecica) communi- of up to 40%) consists of Sphagnum squarrosum, ties are the characteristic feature of the island’s S. riparium, S. fimbriatum, Polytrichum jensenii, shore slope vegetation and are usually locat- Dicranum majus etc. ed in shallow depressions. In these communities On the other hand, these communities may be dwarf cornel constantly dominates with a high represented by dense stands of Carex recta be- cover, whereas other dominant species are usu- ing the single dominant. All the abovementioned ally absent. Epilobium palustre, Bistorta vivipa- herb species occur there while the dwarf shrubs ra, Rumex lapponicus, Dryopteris expansa, Ru- are untypical and the moss layer is almost ab- bus chamaemorus are typical in the herb layer. sent. Being located above the littoral and supralit- Under wetter conditions, sedges appear in these toral zones, the Carex recta communities are ex- communities: Carex recta on the eastern shore, C. posed to wave run-up during heavy gales. As a aquatilis on the western shore, and C. rarifloraon result, marine water accumulates there, which fa- both shores. Polytrichastrum alpinum dominates vours the growth of Carex recta. These commu- in the sparse moss layer, whereas other moss and nities occasionally alternate with small mono- liverwort species (Dicranum majus, Warnstor- dominant groups of Equisetum sylvaticum, Eri- fia fluitans, Polytrichum jensenii, Cephaloziel- ophorum scheuchzeri and Calamagrostis phrag- la sp.) are rare. Under drier conditions, Calama- mitoides. On the western shore, Carex aquatilis grostis neglecta subsp. groenlandica, Poa alpige- forms communities similar in accompanying spe- na, Festuca rubra subsp. arctica, Cerastium font- cies of the herb and moss layers. Carex recta sel- anum subsp. fontanum, Equisetum arvense subsp. dom occurs in plant communities of the western alpestre, Empetrum nigrum subsp. hermaphrodi- shore, mainly in its southern part. tum are common. In the northern part of the island, on steep Peculiar sedge communities with a sparse or cliffs with drip moisture, which are not- direct even absent moss layer are located on the shore ly exposed to the sea, sparse plant communities slopes. Communities with the presence of halo- of rock vegetation with herbs (Carex lachenalii, philous Carex recta are characteristic of the east- Saxifraga rivularis) and several tufts of mosses ern shore. They are situated in shallow depres- (Bryum spp., Aulacomnium palustre) occur. sions on gentle shore slopes and also occur asso- Coastal vegetation is weakly developed on the ciated with sedge-sphagnous communities in dis- island and is mostly represented by sparse plant tal parts of the dells passing through the slopes communities of coastal rock outcrops. There are from the summit plain with the permafrost peat- rocks, sometimes steep, on almost all the shores land. Sometimes these communities ascend along of the island. Along cracks of coastal rocks in the the cracks reaching the summit plain of the island. wave run-up zone, sparse groups of Puccinellia On the one hand, these may be polydominant capillaris subsp. pulvinata, Cochlearia arctica, communities with the dominance of Carex recta, Ligusticum scothicum, Arctanthemum arcticum C. rariflora, Empetrum nigrum subsp. hermaph- subsp. polare, Tripleurospermum maritimum sub- roditum, Rubus chamaemorus and Cornus sueci- sp. phaeocephalum, Mertensia maritima, Carex ca in the herb layer (Fig. 4.2). While Rumex lap- glareosa, Rhodiola rosea, and separate tufts of ponicus, Comarum palustre, Bistorta vivipara Bryum sp. occur. and Epilobium palustre are constant, dwarf shrubs Above the limit of the stony or rocky littoral Vaccinium uliginosum, V. vitis-idaea, Rhododen- zone, medium-level coastal meadows stretch nar- dron tomentosum and herbs such as Juncus fili- rowly along the shore. They are slightly flooded formis, Equisetum sylvaticum, Dryopteris expan- during ordinary tides, submerged during syzygy sa, Calamagrostis neglecta subsp. groenlandica, and permanently influenced by wave run-up, es- Poa alpigena are also present. Sometimes Erio- pecially during heavy gales. At well-drained sites phorum angustifolium and Carex subspathacea, on flat rocks,marshes with Festuca rubra ­subsp. seldom Carex aquatilis are abundant. Tiny plants arctica and Sanionia uncinata form a dense cover of an annual herb, Koenigia islandica, can be (up to 60–100%). Coastal species such as Carex found in moss tufts. The sparse moss layer (cover glareosa, Arctanthemum arcticum subsp. polare, 14 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

Figure 4. Plant communities of Sosnovets Island. 1. Sedge-crowberry community with Carex rariflora on the peatland margin in the western shore of the island. 2. Polydominant community with the dominance of Carex recta, Empetrum nigrum subsp. her- maphroditum, Rubus chamaemorus and Cornus suecica on the eastern shore of the island. 3. Medium-level marshes with Festu- ca rubra subsp. arctica, Carex glareosa and C. mackenziei in the southern part of the island. 4. Coastal high-level meadow with Sonchus arvensis var. maritimus, Ligusticum scothicum, Festuca rubra subsp. arctica in the southern part of the island. 5. Sep- arate patches of Honckenya peploides subsp. diffusa on a cobble shore in the southern part of the island. 6. Coastal low-level meadow with Plantago maritima subsp. borealis on fine gravel deposits in the southern part of the island. 7. Willowherb (Cha- maenerion angustifolium) meadows alternating with cloudberry-crowberry communities and crowberry tundra in the south- ern part of the island. 8. Mesophytic anthropogenic grass-forb meadows with alien species in the foreground and willowherb community in the background. Photos: 1, 2 – E. O. Golovina; 3, 4, 7, 8 – M. N. Kozhin; 5, 6 – E. I. Kopeina. Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 15

Argentina anserina subsp. groenlandica, Parnas- cetraria nivalis are frequent along with Sanionia sia palustris and Ligusticum scothicum are typi- uncinata. The vegetation cover of these meadows cal of these communities (Fig. 4.3). is characterized by mosaic structure, correlated At moderately drained sites on flat rocks, with microtopography conditions, and by a con- along the margin of crowberry communities, siderable abundance of species from surrounding marshes of Carex subspathacea and Carex glare- communities, e.g. Cornus suecica, Empetrum ni- osa sometimes occur. They are characterized by grum subsp. hermaphroditum, Calamagrostis ne- a relatively small area (from 2 to 10 m2) and a glecta subsp. groenlandica. These meadows are sparse herb layer. In the sodded rock cracks, Ar- remarkable due to their highest species diversity gentina anserina subsp. groenlandica, Festuca among coastal communities. rubra subsp. arctica, Calamagrostis deschamp- Below the level of lime-grass meadows, on sioides and single tufts of Bryum spp. grow. small areas of supralittoral sand and cobble, sepa- On shore slopes with some constant inflow rate patches of Honckenya peploides subsp. diffu- of fresh water, marshes with the dominance of sa (with an area of 1–6 m2) occur as a narrow dis- Calamagrostis deschampsioides and Carex rar- continuous belt (Fig. 4.5). In the northern part of iflora are located. They form a relatively dense the island, along the tideline, among heaps of de- cover (of 50–95%). Along with coastal species caying algae, a single patch of five shoots ofBol - Stellaria humifusa, Arctanthemum arcticum sub- boschoenus maritimus has been observed. sp. polare, their herb layer is composed of typi- In the southern part of the island, on its east- cal species of paludified places such asEpilobium ern side, there is a shallow depression that in- palustre and Triglochin palustris. In the south- cludes a small sea bay, which is occupied by pe- western part of the island, a community with a culiar coastal meadow and marsh communities. considerable abundance of Juncus gerardi subsp. The whole ecological series of low-level coast- atrofuscus apart from the aforementioned species al meadows, which are flooded daily, and of me- was observed. Warnstorfia fluitans and W. exan- dium-level coastal meadows, which are flooded nulata usually dominate in its dense moss layer, only in syzygy, is developed on sandy and silty whereas Sanionia uncinata seldom occurs there. deposits as well as on sandy and gravelly ones. Along elongated rock depressions, on steep In the lower level of silty littoral with more stag- and flat shore slopes, grasslands of Calamagros- nant water, a sparse community of Puccinellia tis neglecta subsp. groenlandica sometimes oc- phryganodes was recorded in the only locality on cur. In the herb layer, narrow small reed domi- the island. At a level above it, on sandy substrate nates and is constantly associated with Parnassia with fine gravel, low-level meadows of Plantago palustris, Festuca rubra subsp. arctica, Rumex maritima subsp. borealis occur (Fig. 4.6). Seaside lapponicus. Sometimes Allium schoenoprasum plantain (Plantago maritima) forms a homogene- appears in these communities, possibly as a co- ous dense cover, which includes single individu- dominant. The moss layer (Sanionia uncinata, als of Triglochin maritima and Stellaria humifu- Sciuro-hypnum curtum and Bryum sp.) is weak- sa. Further above, they are replaced by a dense ly developed. tuft of Carex subspathacea communities with Supralittoral lime-grass (Leymus arenarius) single individuals of Triglochin palustris, Argen- grasslands are present as separate fragments (with tina anserina subsp. groenlandica, Stellaria hu- an area of 1–2 m2) in the northern and southern mifusa, Arctanthemum arcticum subsp. polare, parts of the island. They are usually formed along and sporadic clusters of Bryum sp. the shores on loose drift deposits overlaying the At sites with constant inflow of fresh water, basement rock. Leymus arenarius conspicuously or in flat shallow depressions with stagnant sea dominates in these communities but may be re- water on silty deposits, marshes of Carex mac- placed by Lathyrus maritimus, Ligusticum sco- kenziei are located. The sedge cover reaches 80% thicum, Sonchus arvensis var. maritimus, Festu- and includes single individuals of Stellaria hu- ca rubra subsp. arctica and Conioselinum tatari- mifusa and Atriplex nudicaulis, with intrusion of cum (Fig. 4.4). In the moss-lichen layer, lichens sparse shoots of Festuca rubra subsp. arctica and Cetraria islandica, Cladonia arbuscula, Flavo- Calamagrostis deschampsioides from adjacent 16 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 medium-level coastal communities. Carex rari- tium fontanum subsp. fontanum, Cornus suecica, flora is common at desalinated paludified sites. Empetrum nigrum subsp. hermaphroditum, Fes- Further up, on the supralittoral, medium-lev- tuca ovina, Vaccinium vitis-idaea and V. uligino- el coastal meadows are represented by commu- sum. Alien species, Deschampsia cespitosa, Ra- nities of Festuca rubra subsp. arctica and Sanio- nunculus acris subsp. acris, Stellaria graminea nia uncinata, which are similar to those described and Trifolium repens are present as single individ- above. However, apart from typical coastal spe- uals or in groups. The cover of the moss-lichen cies, Galium palustre and Lathyrus palustris sub- layer varies from 1 to 45%, which is determined sp. pilosus were recorded in their composition. In not only by intensive disturbance and succession the uppermost part of the supralittoral, the sole stage, but also by the density of the willowherb stripe of coastal grassland of Agrostis stolonifera cover. In the moss-lichen layer, Polytrichum juni- subsp. straminea on the island occurs in the in- perinum and Hylocomium splendens are present, ner part of the bay. In this community, Festuca as well as some other species of mosses (Dicra- rubra subsp. arctica, Cerastium fontanum subsp. num majus) and lichens (Cetraria islandica, Cla- fontanum, Stellaria crassifolia are present; at wet donia arbuscula and Flavocetraria nivalis) that sites, Comarum palustre is also observed. A sin- are widespread in crowberry and dwarf cornel gle patch of Alopecurus arundinaceus (with an communities of the island. area of 1 m2) was found among creeping bent- Mesophytic anthropogenic grass-forb mead- grass and arctic fescue meadows, although Alope- ows of small size are located in the southern part curus arundinaceus communities are quite com- of the island in the nearest vicinity of the weather mon on mainland shores and islands of the White station, village buildings and the pier. They are Sea. polydominant with irregular species patterns On the top of a ridge between the weather sta- (Fig. 4.8). Alien meadow species such as Lathy- tion and the southernmost tip of the island, sandy rus pratensis, Trifolium repens, Alchemilla sub- blowouts with sparse plant communities of crenata, Vicia cracca, Ranunculus acris subsp. small size are found. They may have natural (ero- acris play a considerable role in these communi- sion) or anthropogenic (sand extraction) origin. ties. However, the greatest species diversity in The vegetation is represented by highly sparse these meadows comes from the native flora groups (cover of up to 5%) of Rumex acetosella (Calamagrostis neglecta subsp. groenlandica, subsp. arenicola, Luzula spicata, Juncus trifidus, Anthoxanthum alpinum, Cornus suecica, Coni- Omalotheca supina and Poa alpina. Bromopsis oselinum tataricum, Rumex lapponicus, Rubus inermis, an alien species, was discovered here chamaemorus, Empetrum nigrum subsp. her- close to dilapidated buildings. maphroditum, Achillea apiculata, Solidago vir- Intensive anthropogenic activities on the is- gaurea subsp. lapponica, Carex brunnescens and land have already been in progress for one centu- Cerastium fontanum subsp. fontanum). ry and a half. The central and southern parts of the At waterlogged disturbed sites, on the sum- island near the lighthouse village and the weather mit plain and in shallow ravines on slopes in the station have been most transformed, with the de- island centre, Deschampsia cespitosa-dominated velopment of anthropogenic vegetation. anthropogenic grasslands are most widespread. Chamaenerion angustifolium-dominated an- Such grasslands are similar to true meadows but thropogenic meadows are located on the sum- differ from the latter in degraded species composi- mit plain and the shore slope; they have the great- tion and the presence of species otherwise untyp- est extension at moderately moist, disturbed sites ical of meadows. They replace cottongrass- and (Fig. 4.7). These meadows are characterized by sedge-sphagnous flarks as well as adjacent cloud- the absolute dominance of willowherb (Chamae- berry-crowberry communities, which had degrad- nerion angustifolium), which forms continuous ed in the result of permafrost thawing, increased pink-coloured stands with a height of 0.7–1.5 drainage and unstable water regime. These pro- m. The herb layer consists of apophytes: Luzu- cesses destroying natural communities were trig- la multiflorasubsp. frigida, Taraxacum repletum, gered by human activities in the lighthouse area. Avenella flexuosa, Lysimachia europaea, Ceras- In present-day communities, tufted hairgrass (De- Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 17 schampsia cespitosa) dominates with a cover of is represented by small patches of alien Trifolium over 50%. The herb layer is characterized by mo- repens near the weather station at sites exposed saic structure and the presence of apophytes, e.g. to permanent trampling. Otherwise, habitats typi- Rumex lapponicus, Dryopteris expansa, Equise- cal of synanthropic communities are occupied by tum arvense subsp. alpestre, Cornus suecica, Eq- nitrophilous coastal plants. For example, Coch- uisetum sylvaticum, Phleum alpinum, Cerastium learia arctica frequently occurs on waste places fontanum subsp. fontanum and Lysimachia euro- with a humus substrate, whereas Tripleurosper- paea. Alien species such as Achillea ptarmica, mum maritimum subsp. phaeocephalum is found Alchemilla subcrenata, Phleum pratense occur in on disturbed soils along the paths and near the small groups. Sometimes single shrubs of Salix houses. glauca and S. phylicifolia with a height of up to 0.5 m are present. Since litter covers the surface almost completely, the moss-lichen layer is weak- Features of the island’s vegetation ly developed (cover of less than 3%) and repre- sented by Sanionia uncinata, Sciuro-hypnum The vegetation of Sosnovets Island is typical starkei and S. curtum. Some communities resem- of the tundra zone. Dwarf-shrub (Empetrum ni- ble a tussock field, where hairgrass tufts reach a grum subsp. hermaphroditum) and lichen (Cla- height of 10–15 cm and spaces between the tufts donia arbuscula, Flavocetraria nivalis) tundras are filled with litter. are formed on sandy deposits on the summit plain On the steep slopes and at their foot in the of the island. They belong to the class Loiseleu- south-eastern part and near the pier in the western rio procumbentis-Vaccinietea Eggler ex Schubert part of the island, vast grasslands of Calamagros- 1960 and are similar to the tundras widespread in tis phragmitoides are located. They are confined the plains and mountains of the Kola Peninsula to anthropogenically disturbed sites with con- (Koroleva 2006a). Since the major part of the is- stant fresh water inflow. Small-reed forms a dense land is overlaid by peatland, the tundra commu- monodominant (over 70%) cover with a height of nities occupy only a small area, i.e. 0.4 ha (1% of 1–2 m. Comarum palustre, Carex aquatilis, Ra- the island’s area). nunculus repens, Veratrum album are present in Plant communities on drained elevated sites such communities in the western part, whereas of the permafrost peatland (cloudberry-crowber- Dryopteris expansa and Phalaroides arundina- ry-lichen and cloudberry-crowberry) are similar cea are found in the south-eastern part of the is- to those described from vast peatlands of the tun- land. dra area east of the Kola Peninsula in the alliance At the southernmost tip, severely disturbed Rubo chamaemori-Dicranion elongati Lavrinen- vegetation of various origin is situated. Some of ko & Lavrinenko 2015, belonging to the class these plant formations resulted from anthropo- Oxycocco-Sphagnetea Br.-Bl. & Tx. ex West- genic transformation of high-level coastal mead- hoff et al. 1946 (Lavrinenko & Lavrinenko 2015). ows and dwarf cornel communities, the others ap- The absence of characteristic species of the class peared by overgrowing of bare man-made lands. Oxycocco-Sphagnetea, namely Andromeda poli- These plant groups emerged at sites of coal stor- folia, Rhododendron tomentosum s. l., Eriopho- age, technogenic waste, abandoned iron con- rum vaginatum and Sphagnum spp., is peculiar to structions and batteries. In this area Poa alpige- these island’s communities. This absence distin- na, Festuca rubra subsp. rubra, F. rubra sub- guishes these communities from those of the same sp. arctica, Solidago virgaurea subsp. lapponica, alliance that are situated on the mainland of the Chamaenerion angustifolium, Empetrum nigrum Malozemelskaya and Bolshezemelskaya tundras subsp. hermaphroditum, Cornus suecica sporad- and brings them closer to those of the ically occur. Rumex acetosella subsp. tenuifolius islands (Vaygach, Kolguev and Dolgii), where and Poa pratensis rapidly overgrow bare soil and these species are also absent or rare (Lavrinen- coal piles. ko & Lavrinenko 2015). However, the differenc- Synanthropic vegetation is present only in es in the species composition do not allow us to the closest proximity to residential buildings and attribute these communities of Sosnovets to any 18 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 of the associations or subassociations described their vegetation significantly differs from that of from the Barents Sea islands, which lay mainly Sosnovets Island, which is situated in the tundra within the subzone of typical tundra. zone. In the abovementioned bays, a considera- The majority of cottongrass- and sedge- ble number of islands are also almost complete- sphagnous communities of the island belong to ly overlaid with peatland, although the morpholo- the class Scheuchzerio palustris-Caricetea fuscae gy of these peatlands is different: they are formed Tx. 1937. They are formed in flarks and in wid- by a layer of dry peat without permafrost, often er coastal parts of dells draining the peatland, and cracked into polygonal peat blocks when drying; can be attributed to the suballiance Caricenion for this reason, flarks with their sedge-, cotton- rariflorae Lavrinenko et al. 2016 of the alliance grass-sphagnous communities, which occur on Sphagnion baltici Kustova ex Lapshina 2010 due Sosnovets Island, are absent here. The dominants to the presence of the characteristic species of of plant communities of these dry peatlands (Em- these syntaxa (Lavrinenko et al. 2016): Sphag- petrum nigrum subsp. hermaphroditum, Rubus num lindbergii, S. balticum, Polytrichum jensenii, chamaemorus, Cladonia arbuscula, Flavocetrar- Eriophorum scheuchzeri. Several sedge-sphag- ia nivalis) are the same as on Sosnovets Island nous communities of palsa mire flarks, which but the other species are partly different due to lack the abovementioned species but include the presence of several boreal and boreal-nemor- Calliergon stramineum, Warnstorfia exannula- al species (Linnaea borealis, Luzula pilosa, Pla- ta, Epilobium palustre, are analogous to commu- tanthera bifolia) and the absence or rareness of nities of the alliance Drepanocladion exannula- arcto-montane species, mostly in the moss-lichen ti Krajina 1933. The communities of the suballi- layer (Thamnolia vermicularis, Alectoria ochro- ance Caricenion rariflorae are widespread in pal- leuca). On the other hand, the large peatland of sa mire flarks of East European tundras, where- Sosnovets Island (29.6 ha, 72.9% of the island’s as the communities of the northern and high-el- area) is typical of the south tundra and the forest- evation alliance Drepanocladion exannulati oc- tundra but is not found on other small islands of cur, although rarely, in flarks with flowing water the White Sea within the tundra zone. (Lavrinenko et al. 2016). At the same time, the vegetation of Sosnovets The coastal vegetation of the island belongs differs from that of the Barents Sea islands, which to intrazonal classes Juncetea maritimi Br.- are also situated in the tundra zone and over- Bl. in Br.-Bl. et al. 1952 (alliances Puccinellion laid with peat. In the peatland’s vegetation on phryganodis Hadač 1946, Caricion glareosae the summit plain of the island, the communities Nordh. 1954, Armerion maritimae Br.-Bl. & De with the dominance of fruticose lichens occupy Leeuw 1936) and Ammophiletea Br.-Bl. & Tx. almost 12.0 ha or 29.8% of the total area. The ex Westhoff et al. 1946 (alliance Agropyro-Hon- crowberry-dominated communities, natural and ckenyion peploidis Tx. in Br.-Bl. et Tx. 1952) and anthropogenic, are twice smaller (almost 6.5 ha, is similar to the vegetation of the Kola Peninsula or 16.0%). In the vegetation of the summit plain coast in both the tundra and taiga zones (Korole- peatlands of the Barents Sea islands, lichen com- va et al. 2011; Popova et al. 2017). The presence munities are rare or absent, whereas cloudberry- of Calamagrostis deschampsioides, which has a crowberry and cloudberry-fern (Dryopteris ex- more eastern distribution in Fennoscandia and is pansa) communities prevail (Breslina 1969; Par- very rare west of Lapponia Ponojensis, in some fentieva 1969). of these communities brings the island’s coastal The communities with the dominance of vegetation together with that of the eastern part of Carex recta, a protected species in Murmansk Re- the Barents Sea (Matveeva & Lavrinenko 2011). gion, occupy a relatively large area (1 ha, 2.7%) Although the vegetation of Sosnovets is typ- and contribute to the peculiarity of the Sosnovets ical of the tundra zone, it drastically differs from Island’s vegetation. that of the other White Sea islands. The anthropogenic vegetation of Sosnovets On the one hand, the majority of small islands Island is specific as well. As the island has been of the White Sea are located in the Kandalaksha populated for more than 150 years, the latter is and Onega bays within the boreal zone. Hence, quite diverse and extensive (5.0 ha, 12.3%), Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 19 which is not typical of small islands of the White south arctic regions. Species composition, distri- Sea. The anthropogenic vegetation is predomi- bution across habitat types and frequency are pre- nantly composed of alien meadow species, which sented in Table 1. form communities according to their ecological preferences: Deschampsia cespitosa in degraded wet flarks of the peatland, and the other species at Taxonomic diversity of the native flora moderately moist sites. Small islands (9–120 ha) of the Kola Subarctic considerably differ in the native species diversi- Floristic composition ty. On average, the Kandalaksha Bay islands that are similar in size to Sosnovets have floras up to Vascular plants of Sosnovets Island include 167 50 species (or 30%) richer than that of Sosnovets species and subspecies from 53 families, of which Island. This difference may be explained by sev- 134 species and subspecies from 36 families are eral reasons. Firstly, the Kandalaksha Bay islands considered native. In the native flora, Poaceae (21 are located within the boreal zone (north taiga species), Cyperaceae (19 species) and Asteraceae subzone), where the species diversity is higher in (13 species) are characterized by the highest spe- general. According to Morozova (2008), the spe- cies diversity, which is typical of north boreal and cies richness of local floras within the mainland

Table 1. Vascular plant diversity of small (9–120 ha) islands of the White and Barents seas Taxonomic diversity Island Area, ha Sources Total Native Alien Sosnovets 40.6 167 134 33 original data Kandalaksha Bay, White Sea Ovechii (Olenii Archipelago) 34 181 170 11 Vorob’eva 1996b Bolshoi Berezovyi 39 190 174 16 Vuzman & Kozhin (pers. comm.) Bolshoi Kurtyazhnyi 36 160 153 7 Vuzman & Kozhin (pers. comm.) Tarasikha 50 163 155 8 Vorob’eva 1986 Medvezhii (Por’ya Guba) 57 240 230 10 Kozhin 2016 Bolshoi Asaf’ev 70 201 200 1 Bogdanova & Vekhov 1969a Berezhnoi Vlasov 106 207 193 14 Vuzman & Kozhin (pers. comm.) Vachev 107 207 204 3 Vorob’eva 1989 Telyachii (Olenii Archipelago) 120 181 173 8 Vorob’eva 1996b Onega Bay, White Sea Luda Gorelka 9 142 132 10 Kravchenko et al. 2015 Sharapikha 11 120 120 0 Kravchenko & Timofeeva 2008 Lambasluda 15 129 127 2 Kravchenko et al. 2015 Rishluda 26 131 129 2 Kravchenko et al. 2015 Ropaki 36 151 148 3 Kravchenko et al. 2015 Kurich’ya Niloksa 43 106 105 1 Kravchenko & Timofeeva 2002 Oleshin 50 132 131 1 Kravchenko & Timofeeva 2002 Malyi Zayatskii 102 166 160 6 Kravchenko & Timofeeva 2002 Barents Sea Bolshoi Gavrilovskii 43 74 73 1 Paneva et al. 2006 Bolshoi Gusenets 11 55 53 2 Paneva et al. 2006 Bolshie Voronukhi 15.5 35 32 3 Paneva et al. 2006 Malyi Ainov 59 74 70 4 Parfentieva & Breslina 1969 20 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 part of the boreal zone in Murmansk Region is novets than in the White Sea and richer than in estimated to be slightly more than 300 species, the Barents Sea. whereas that of the tundra zone is less than 300 species. We have noted similar patterns in island ecosystems. Secondly, the Kandalaksha Bay is Phytogeoraphic features of the native most diverse among the White Sea bays in terms flora of natural conditions (Bianki 1996), which deter- mines the species richness of its islands. Third- Sosnovets Island and the adjacent territories are ly, the majority of the Kandalaksha Bay islands referred to the tundra zone in this work, but their have been thoroughly studied by the staff of the zonal classification has been debated for a long Kandalaksha State Nature Reserve, and their flo- time. Its flora contains a number of species that ras are consequently better known and more com- are typical of the Hypoarctic phytogeographic pletely revealed. belt (Yurtzev 1966), which is an extrazonal ter- Unexpectedly, the floristic diversity of Sos- ritory embracing the southern part of the tundra novets Island is comparable to that of the Onega zone, forest tundra and the north taiga. Accord- Bay islands of similar size, although the latter are ing to Yurtzev (1966), the Hypoarctic phytogeo- situated in the southern part of the north taiga sub- graphic belt is separated from the Arctic by the zone (Table 1). On the one hand, this low diversi- presence of a certain floristic complex and ex- ty results from severe climatic conditions, mostly cludes the ’true’ cryophilous tundra, whereas strong cold winds, which account for the natural the limit between this area and the Boreal belt is absence of forests on the majority of islands and formed by the northern limit of taiga on water- the peculiarities of their vegetation cover (Zinser- sheds. ling 1934). These were the reasons for Zinserling Irrespective of the classification applied, a (1934) to separate this territory as a geobotanical specific trait of the vegetation in the north Kola district, the White Sea islands. On the other hand, section of the Hypoarctic (Yurtzev 1966), or in Zinserling’s conclusions were based on relative- the Kola tundras (Safronova et al. 1999), or in the ly short-term expeditions rather than comprehen- coastal parts of the entire Kola Subarctic (Bres- sive studies. lina 1987) is a high abundance of Empetrum ni- On the contrary, all the Barents Sea islands of grum subsp. hermaphroditum. This is confirmed similar size have a twice less diverse flora than by our observations on Sosnovets Island, whose that of Sosnovets Island. Despite all these islands area is covered by crowberry for more than three (Sosnovets and the Barents Sea islands) are locat- fourth of its total surface. ed in the same (tundra) zone, a considerable dif- In comparison with the other White Sea is- ference in their floristic diversity may stem from lands studied, the flora of Sosnovets stands out several reasons. First and foremost, the Barents by its high specificity connected with the island’s Sea islands stretch along the coastline exposed to position within the tundra zone and at the eastern- heavy gales, whereas Sosnovets Island is situated most edge of Fennoscandia. In particular, 14 spe- in the White Sea Throat and is therefore shielded cies of its flora are absent from the other White against the storms. Notably, the eastern part of the Sea islands. These are predominantly the species Tersky Coast features the most severe climate of of plain and mountain tundra, common to Fen- the White Sea, yet the number of days with heavy noscandia, namely Comastoma tenellum, Hiera- gales there is 2.5 times lower than on the East cium alpinum, H. atratum, H. nigrescens, Tarax- Murman Coast (Yakovlev & Kozlova 1971). Sec- acum hjeltii, Koenigia islandica, Luzula confusa, ondly, due to proximity to the taiga zone, a few L. spicata, L. wahlenbergii, Ranunculus subbore- boreal species appear also on Sosnovets Island. alis subsp. pumilus and Omalotheca supina. An- Thus, the native species richness of Sosnovets other set of specific taxa are typical arctic, such Island occupies an intermediate position among as Arctanthemum arcticum subsp. polare, Cop- the Kola Subarctic islands. Comparing the spe- tidium pallasii, Calamagrostis deschampsioides; cies diversity of Sosnovets Island with that of the and one specific boreal taxon is present,Lathyrus other islands of similar size, it is poorer on Sos- palustris subsp. pilosus. All these species pene- Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 21 trate into Fennoscandia only at its eastern margin, Protected species Lapponia Ponojensis and Lapponia Murmanica. Another characteristic feature of the island’s There are six vascular plant species included in flora is a low number of typical boreal species. the Red Data Book of Murmansk Region (Kon- For example, it completely lacks native trees. stantinova et al. 2014), of which one species Apart from widespread tundra shrub willows, (Rhodiola rosea) is included in the Red Data woody plants are represented by prostrate shrubs Book of the Russian Federation (Bardunov et al. of Betula pubescens subsp. czerepanovii with a 2008). The majority of protected species occur in height of up to 0.5 m. Bilberry (Vaccinium myr- coastal habitats. tillus) has a particularly low abundance on the is- Arctanthemum arcticum subsp. polare spo- land; its distribution is limited to the places with a radically occurs on sea shores within the tundra higher snow accumulation in winters and a great- zone of the Kola Subarctic; in particular, the re- er insolation in summers. The majority of com- cord from Sosnovets is one of the southernmost. mon boreal species, e.g. Linnaea borealis L., Bolboschoenus maritimus is rare on the White Luzula pilosa (L.) Willd., Maianthemum bifoli- Sea coast; its locality on the island is the eastern- um (L.) F.W.Schmidt, were not detected on Sos- most in Murmansk Region and the only one in novets Island although they are widespread on the the tundra zone of the Kola Peninsula. Only four other White Sea islands studied. shoots, two of which being reproductive (with The difference between the flora of Sosnovets spikes), were discovered. Island and floras of the Barents Sea islands is less Carex recta sometimes occurs along the coasts significant. Firstly, scattered individuals of sev- of the White and Barents seas. Its population on eral boreal species (Carex paupercula, Carex Sosnovets, estimated at several thousands of re- chordorrhiza, Vaccinium oxycoccos) still occur productive shoots, is possibly the largest in Mur- on Sosnovets Island, whereas they are complete- mansk Region. This species occupies a vast part ly absent from the northern Barents Sea islands. of the eastern shore of Sosnovets and penetrates Secondly, species with eastern distributions in into crowberry communities in the inner part of Fennoscandia (e.g. Calamagrostis deschamp- the island along dells and waterlogged sites. sioides) seldom reach the Barents Sea islands or Comastoma tenellum was recorded in a small are completely absent from the latter. Thirdly, the number of individuals on forb-grass meadows flora of the Barents Sea islands is considerably with minor anthropogenic impact, which are its poorer due to the decrease in the number of coast- typical habitats in the east of the Kola Peninsula. al species, which is connected with a harsh cli- Coptidium pallasii was once recorded in a mate and frequent heavy gales in the Barents Sea. small number of individuals in flarks of the pal- It is noteworthy that one common arctic species, sa mire. Koenigia islandica, was discovered only on Sos- Rhodiola rosea sporadically occurs on coast- novets Island and was not recorded on the other al rocks throughout the island. It is frequent in the Kola Subarctic islands studied; however, this spe- east of the Tersky Coast. cies is not rare in the adjacent mainland tundra. A considerable number of coastal species as well as a high activity and abundance of crowber- Alien species and their origin ry and cloudberry are common features of Sos- novets and the other islands of the White and Bar- The flora of Sosnovets Island includes 33 alien ents seas. The absence of Betula nana in crowber- species of vascular plants, which constite 20% of ry communities on dry peatlands on all the oth- its total species number. This is the highest num- er islands of the White and Barents seas is also ber of alien species among the islands of simi- remarkable. Dwarf birch was observed on mires lar size in the White and Barents seas (from 9 to only on a few islands in the boreal zone, where- 120 ha) (Table 1). This high diversity and wide as it is widespread and plays a crucial role in the distribution of alien species is connected with the vegetation on the mainland in the tundra zone. island being explored and populated for a long time. In particular, a permanent human popula- 22 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 tion has been living on the island for more than ca, V. sepium. These species appeared on the is- 150 years, making it the only small island with land as a result of long-term hay supply to feed permanent population in the Kola Subarctic. In horses, which were kept there in the 20th century. the 19th and the 20th centuries, various econom- Plant diaspores dropped from the hay gave rise to ic activities took place also on the other islands of populations of alien plants that have established small size in the White and Barents seas but these and still remain in disturbed communities. No ev- islands remained unpopulated. The pomors used idence of these species having been cultivated on different parts of the White Sea for trap fishing the island was observed. and built small houses to store their fishing equip- The majority of these species are widespread ment, e.g. on Malyi Berezovyi and Luda Gorel- in the boreal zone and are common on fields and ka (Nikolsky 1927, Kravchenko et al. 2015). On semi-natural meadows in the middle taiga (Pa- Medvezhii Island in the Por’ya Guba Bay, silver rinova 2010), whereas in the tundra zone of the was mined and barracks for workers were built Kola Subarctic they are strictly alien. In the north in the 18th and the 19th centuries (Efimov 1970). taiga of Russian Lapland, only a few of these spe- In various parts of the Kandalaksha Bay (Bolshoi cies, namely Agrostis capillaris, A. gigantea, De- Kurtyazhnyi and Berezhnoi Vlasov islands) and schampsia cespitosa, Hieracium umbellatum, on the Ainov Islands in the Barents Sea, hayfields Scorzoneroides autumnalis, Phalaroides arundi- were situated (Matsak 2005). On average, 10 al- nacea, Sagina procumbens, Trifolium pratense ien species were discovered on each of these is- var. pratense and Veronica longifolia, may be re- lands; many of these species began to natural- garded as native. Apart from the meadow spe- ize also on the other islands. The threefold great- cies, a single shrub of Salix caprea subsp. caprea er number of alien plants detected on Sosnovets was recorded, whose diaspores may also have Island can be explained by its long history of di- emerged on the island with forage. This mass oc- verse economic activities, which drastically dif- currence of plants typical of middle taiga mead- fers from that of the other small islands studied in ows suggests that the hay was delivered to Sos- the Kola Subarctic. novets from Arkhangelsk Region. This conclu- Since the village of the lighthouse keepers sion is further supported by the presence of Achil- and the weather station staff had been established lea ptarmica, an alien species that is absent from on Sosnovets, a regular supply of food, construc- the Kola Peninsula, whereas it widely occurs on tion materials, and fuel on the basis of the supply floodplain forb-grass meadows, shrub thickets system for Arctic polar stations was organized, and riverbanks in the middle taiga of Arkhangel- which had a strong impact on the island’s flora. sk Region (Schmidt 2005). Five grass species directly used as forage, Among alien plants typical of meadows, Al- namely Phleum pratense, Alopecurus pratensis, chemilla cymatophylla is an extraordinary find- Bromopsis inermis, Festuca rubra subsp. rubra ing as this species was unknown from the east- and Elytrigia repens, were discovered on the is- ern part of Murmansk Region despite its presence land. These species are usually sown on agricul- in the western part (Kurtto et al. 2007). This spe- tural fields in the boreal zone. Apart from them, cies cannot have been transported with hay from a considerable number of plants characteristic of Arkhangelsk Region because it was only once managed or disturbed meadows and abandoned collected there in a remote area of the Mezen Riv- fields were found as accompanying species, such er basin (Schmidt 2005), where it is treated as al- as Agrostis capillaris, A. gigantea, Alchemil- ien (Kurtto et al. 2007). In Finland, A. cymato- la subcrenata, Deschampsia cespitosa, Galium phylla is considered of polemochore origin, i.e. mollugo, Geranium pratense, Hieracium umbel- a war-time immigrant (Heikkinen 1969); it oc- latum, Lathyrus pratensis, Leucanthemum vul- curs mostly along the present-day Russian bor- gare, Phalaroides arundinacea, Sagina procum- der (Piirainen 2019). Its secondary distribution bens, Ranunculus acris subsp. acris, R. aggr. au- in East Fennoscandia runs largely along the Rus- ricomus, R. repens, Scorzoneroides autumna- sian-Finnish border and continues from the north- lis, Stellaria media, Trifolium pratense var. prat- ernmost part of Karelian Republic to the western ense, T. repens, Veronica longifolia, Vicia crac- part of Murmansk Region, as do the distributions Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 23 of other polemochore species of Alchemilla that in the White Sea and richer than on islands in the occur in northern Finland; native distributions of Barents Sea. Its peculiar features are explained by such species are centered in the hemiboreal zone the eastern localtion of Sosnovets within the Kola from the Baltic countries to Central Russia (Pi- Peninsula. There are six protected species in the irainen & Chkalov 2018). Considering this pat- flora of Sosnovets, of which one species, Carex tern, we think it possible that A. cymatophylla had recta, plays a considerable role in the vegetation. been brought to Sosnovets as a result of long-dis- As the island has been populated for more tance transportation of hay supply, most likely than 150 years, its plant cover was partly trans- in the war time when the Russian army required formed. The alien flora of Sosnovets island is large quantities of hay from Central Russia. most diverse compared with similar-size marine A small population of Euphrasia stricta var. islands of the Kola Subarctic and contains typi- brevipila was observed on a single spot near the cal meadow species of the middle taiga subzone, lighthouse tower, on a small meadow with a dense which were transported with regular forage sup- moss cover and other apophytes. This communi- ply (hay transportation) from Arkhangelsk Re- ty represents a stage of plant cover demutation, gion. The introduction of a few other alien spe- which has been ongoing for decades. The species cies can be linked with construction activities and may have appeared on the island when the light- war-time long-distance forage supply. house tower was reconstructed in 1960. In Mur- The vegetation of the island is remarkable due mansk Region, E. stricta var. brevipila occurs at to a significant diversity of plant communities, sites with strong anthropogenic disturbance. We which were developed because of natural condi- think its presence on the island is linked with car- tions (extensive permafrost peatlands and marine go transportation from Arkhangelsk, since it is severe climate) and anthropogenic impact. The widespread on meadows, along roads, on fields palsa mire, the marshes with Calamagrostis des- and fallow lands in Arkhangelsk Region up to the champsioides, the highly dissected peatland with limit of the north taiga (Schmidt 2005). cloudberry-crowberry-lichen communities on el- Rumex acetosella subsp. tenuifolius, an alien evated sites and the cottongrass-sphagnous com- subspecies of anthropogenically disturbed sites, munities in depressions are the unique features of is widespread in the southern part of the island. the island’s vegetation; among small islands of A few juvenile plants of Melilotus sp., which be- the White Sea, they were recorded only on Sos- long to a contemporary ephemeral introduction, novets. were observed on a recent coal pile in the eastern part of the island. Due to severe climatic conditions, kitch- en gardens were little used in the island’s vil- Acknowledgements. We are obliged to Ludmila A. Konor- lage. Only a single depauperate shrub of Ribes ni- eva and Yurii S. Mamontov (Avrorin Polar-Alpine Botan- grum (with a height of up to 30 cm) was observed ical Garden-Institute, Kola Research Centre of Russian among crowberry communities near the weather Academy of Sciences, Kirovsk) for identification of li- station as a relic of cultivation. chens and liverworts, correspondingly. We are grateful to Elena A. Ignatova (Moscow State University, Moscow) for verification of some moss specimens. Henry Väre (Helsin- ki) and Pertti Uotila (Helsinki) provided detailed advice Conclusions and clues to the historical literature. Boris B. Vakhmistrov (Kirovsk) shared his photograph of Sosnovets Island and Sosnovets Island significantly differs from the allowed its publication here. We also thank local residents other islands of the White and Barents seas in its Nikolai A. Kozhin, Pavel A. Kozhin, Andrey P. Vaganov flora and vegetation. and Pavel V. Vaganov for transport support. The work of The native species richness of Sosnovets Is- E. Golovina and S. Kutenkov was supported by institu- tional research projects of the Komarov Botanical Insti- land occupies an intermediate position among the tute RAS (no. AAAA-A18-118031690042-9) and of the Kola Subarctic islands. Comparing the species di- Institute of Biology of Karelian Research Centre RAS (no. versity of Sosnovets Island with that of the other AAAA-A17-117031710038-6), correspondingly. The her- islands of similar size, it is poorer than on islands barium work of M. Kozhin at the Botanical Museum, Uni- 24 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019 versity of Helsinki, was generously supported by the So- ogy in the Canary Islands. Monographiae Biologicae cietas pro Fauna et Flora Fennica, and our expedition to 30: 207–240. 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Sennikov, A.N. & Kozhin, M.N. 2018: The history of the Walker, D.A., Raynolds, M.K., Daniëls, F.J.A., Einarsson, Finnish botanical exploration of Russian Lapland in E., Elvebakk, A., Gould, W.A., Katenin, A.E., Kholod, 1861 and 1863. — Memoranda Societatis pro Fauna S.S., Markon, C.J., Melnikov, E.S., Moskalenko, et Flora Fennica 94: 1–35. N.G., Talbot, S.S., Yurtsev, B.A. & The other mem- Shipunov, A.B. & Abramova, L.A. 2006: (Floristic chang- bers of the CAVM Team 2005: The Circumpolar Arc- es on the Kem-Ludy Islands (Russian White Sea, tic vegetation map. — Journal of Vegetetion Science 1962–2004).) — Bulleten’ Moskovskogo Obshchest- 16: 267–282. va Ispytatelei Prirody. Otdel Biologicheskii 111(1): Westhoff, V. & Maarel, E. van der 1973: The Braun-Blan- 45–56. (In Russian with English summary.) quet approach. — In: Whittaker, R.H. (ed.) Handbook Smirnov, N. 1927: (Phenological observations in Karelia.) of vegetation science, Part V. Ordination and Classifi- — Karelo-Murmanskii krai 3: 35–36. (In Russian.) cation of vegetation: 617–726. Junk, The Hague. Tolmatschev, A.I. 1925: Larix sibirica Led. auf der Kola- Yakovlev, B.A. 1961: (The climate of Murmansk Region.) Halbinsel. — Svensk Botanisk Tidskrift 19: 523. — 200 pp. Murmansk, Murmansk publishing house. Tybirk, K., Nilsson, M.-C., Michelsen, A., Kristensen, (In Russian.) H.L., Shevtsova, A., Strandberg, M.T., Johansson, Yakovlev, B.A. & Kozlova, L.G. 1971: (Precipitation, air M., Nielsen, K.E., Riis-Nielsen, T., Strandberg, B. & temperature, direction and velocity of wind. Insola- Johnsen, I. 2000: Nordic Empetrum dominated eco- tion.) — In: Durov, A.G. (ed.), (Atlas of Murmansk systems: Function and susceptibility to environmen- Region): 9. Main Department of Geodesy and Cartog- tal changes. — AMBIO: A Journal of the Human En- raphy, Research Geographical and Economic Institute vironment 29(2): 90–97. of Leningrad State University, Moscow. (In Russian.) Uotila, P. 2013: Finnish botanists on the Kola Peninsu- Yunatov, A.A. 1964: (Types and details of geobotanical la (Russia) up to 1918. — Memoranda Societatis pro study. Choosing sample plots and ecological profiles.) Fauna et Flora Fennica 89: 75–104. — In: Korchagin, A.A. & Lavrenko, E.M. (eds.), Vasilevich, V.I. & Bibikova, T.V. 2011: (Cowberry pine (Field geobotany) 3: 9–36. Science Publishers, Len- forests (Empetri nigri–Pinetum) in North-Western ingrad & Moscow. (In Russian.) European Russia.) — Botanicheskii Zhurnal 96(9): Yurtzev, B.A. 1966: (Hypoarctic botanical-geographical 1153–1161. (In Russian with English summary.) belt and origin of its flora.) — 95 pp. Science Publish- Virtanen, R., Oksanen, L., Oksanen, T., Cohen, J., Forbes, ers, Leningrad & Moscow. (In Russian.) B.C., Johansen, B., Käyhkö, J., Olofsson, J., Pulliai- Zherikhina, V.N. & Moskvicheva, L.A. 2006: (Checklist nen, J. & Tømmervik, H. 2015: Where do the tree- of vascular plants of the Severny Archipelago (Kan- less tundra areas of northern highlands fit in the glob- dalaksha Bay, White Sea).) — In: VIII–IX Interna- al biome system: Toward an ecologically natural sub- tional workshop ”Rational exploration of the coastal division of the tundra biome. — Ecology and Evolu- zone of the northern seas”, 17 July, 2004, Kandalak- tion 6: 143–158. sha. Reports: 64–77. Russian State Hydrometeorolog- Vorob’eva, E.G. 1986: (Flora of Tarasikha and the Ro- ical University, Saint-Petersburg. (In Russian.) govye Ludy islands in the Kandalaksha Bay.) — Pri- Zinserling, Yu.D. 1934: (Plant cover geography of the roda i Khoziaistvo Severa 14: 47–60. (In Russian.) north-west of the European part of the USSR). — 377 Vorob’eva, E.G. 1989: (Flora and vegetation of the Vachev pp. Academy of Sciences of the USSR, Leningrad. (In Archipelago in the middle part of the Kandalaksha Russian with German summary.) Bay.) — In: Pulyaev, A.I. (ed.), (Flora and fauna of protected islands: 5–33.) The Head Committee on Hunting at the Council of Ministers of the RSFSR & Central Research Laboratory of Hunting and Nature Reserves, Moscow. (In Russian.) Vorob’eva, E.G. 1996a: (Analysis of the island flora in the Kandalaksha Bay.) — In: Filippova, L.N. (ed.), (Flo- ra and vegetation of islands of the White and Barents seas): 89–100. Murmarnsk Provincial-level Institute of Regional Education and Development of Teachers & Kandalaksha State Nature Reserve, Murmansk. (In Russian.) 28 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

Appendix 1. Vascular plant flora of Sosnovets Island, the White Sea.

Status 6. Paludified crowberry communities with Carex rariflora and ● – native; ○ – alien. small fens with C. rariflora. Main habitats 7. Paludified coastal community with Carex recta. 1. Crowberry tundra on mineral deposits and crowberry- and 8. Sedge- and cottongrass-sphagnous flarks. cloudberry-dominated communities on the peatland. 9. Coastal meadows, grasslands, marshes and rocks. 2. Lichen tundra on mineral deposits and cloudberry-crow- 10. Sandy blowouts. berry-lichen communities on the peatland. 11. Anthropogenic and synanthropic vegetation and dis- 3. Dwarf shrub (Empetrum nigrum subsp. hermaphroditum, turbed sites. Vaccinium myrtillus, V. uliginosum, Cornus suecica) tundra on Regional protected status is given in accordance with Red SE-facing slopes. Data Book of Murmansk Region (Konstantinova et al. 2014). 4. Cornus suecica-dominated communities. Plant names are provided according to FinBIF (2019). 5. Willow thickets.

Main habitats

Species 1 2 3 4 5 6 7 8 9 Status 10 11 Regional Regional Frequency protected status protected ● Achillea apiculata N.I.Orlova + + + + Rar ○ Achillea ptarmica L. + Rar ○ Agrostis capillaris L. + + Rar ○ Agrostis gigantea Roth + Rar ● Agrostis mertensii Trin. + + Un ● Agrostis stolonifera L. subsp. stolonifera + + Rar ● Agrostis stolonifera subsp. straminea (Hartm.) Tzvelev + Rar ○ Alchemilla cymatophylla Juz. + Rar ○ Alchemilla subcrenata Buser + Rar ● Allium schoenoprasum L. + + Rar ● Alopecurus arundinaceus Poir. + Un ○ Alopecurus pratensis L. + Rar ● Andromeda polifolia L. + + + Rar ● Angelica archangelica L. + Un ● Anthoxanthum alpinum Á.Löve & D.Löve + + Rar ● Arctanthemum arcticum subsp. polare (Hultén) Tzvelev + + + + + Rar 3 ● Arctous alpina (L.) Nied. + + + Rar ● Argentina anserina subsp. groenlandica (Tratt.) Á.Löve + Rar ● Atriplex nudicaulis Boguslaw + Rar ● Avenella flexuosa(L.) Drejer + + + Rar ● Betula pubescens subsp. czerepanovii (N.I.Orlova) Hämet-Ahti + + Un ● Bistorta vivipara (L.) Delarbre + + + + + + + + Rar ● Bolboschoenus maritimus (L.) Palla + Un 3 ○ Bromopsis inermis (Leyss.) Holub + Rar ● Calamagrostis deschampsioides Trin. + + + Rar ● Calamagrostis neglecta subsp. groenlandica (Schrank) Matuszk. + + + + + + + Sol Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 29

App. 1. cont. Main habitats

Species 1 2 3 4 5 6 7 8 9 Status 10 11 Regional Regional Frequency protected status protected ● Calamagrostis phragmitoides Hartm. + + + + + + Sp ● Carex aquatilis Wahlenb. + + + + Sol ● Carex bigelowii Torr. ex Schwein. + + Rar ● Carex brunnescens (Pers.) Poir. + + + + Rar ● Carex canescens L. + + + Rar ● Carex cespitosa L. + Un ● Carex chordorrhiza Ehrh. + Un ● Carex glareosa Wahlenb. + + Rar ● Carex lachenalii Schkuhr + Un ● Carex mackenziei V.I.Krecz. + Rar ● Carex paupercula Michx. + Un ● Carex rariflora (Wahlenb.) Sm. + + + + Sol ● Carex recta Boott + + + + + Rar 3 ● Carex rotundata Wahlenb. + Un ● Carex subspathacea Wormsk. ex Hornem. + + + + Rar ● Cerastium alpinum L. subsp. alpinum + + + Rar ● Cerastium fontanum Baumg. subsp. fontanum + + + + + + + + Rar ● Chamaenerion angustifolium (L.) Scop. + + + + + + + Sol ● Cochlearia arctica Schltdl. ex DC. + + Rar ● Comarum palustre L. + + + Rar ● Comastoma tenellum (Rottb.) Toyok. + Rar 2 ● Conioselinum tataricum Hoffm. + + + Rar ● Coptidium pallasii (Schltdl.) Á.Löve & D.Löve + + Un 2 ● Cornus suecica L. + + + + + + + + + + Sol ○ Deschampsia cespitosa (L.) P.Beauv. + + Sol ● Dianthus superbus L. + + + Rar ● Dryopteris expansa (C.Presl) Fraser-Jenk. & Jermy + + + + + Rar ○ Elytrigia repens (L.) Nevski + Rar ● Empetrum nigrum subsp. hermaphroditum (Hagerup) Böcher + + + + + + + + + Com ● Epilobium palustre L. + + + + + + Rar

Equisetum arvense subsp. alpestre (Wahlenb.) Schönswetter & + + + + + + + + Rar ● Elven ● Equisetum pratense Ehrh. + + Rar ● Equisetum sylvaticum L. + + + + Rar ● Eriophorum angustifolium Honck. + + + + + Rar ● Eriophorum russeolum Fr. + + Un 30 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

App. 1. cont. Main habitats

Species 1 2 3 4 5 6 7 8 9 Status 10 11 Regional Regional Frequency protected status protected ● Eriophorum scheuchzeri Hoppe + + + Rar ● Eriophorum vaginatum L. + + Rar ○ Euphrasia stricta var. brevipila (Burnat & Gremli) Hartl + Un ● Euphrasia wettsteinii G.L.Gusarova + + + Rar ● Festuca ovina L. + + + + + Rar ○ Festuca rubra L. subsp. rubra + + Rar ● Festuca rubra subsp. arctica (Hack.) Govor. + + + + + + + + + Rar ● Filipendula ulmaria (L.) Maxim. + Un ○ Galium mollugo L. + Rar ● Galium palustre L. + + Rar ● Galium trifidum L. + + Un ○ Geranium pratense L. + Un ● Geranium sylvaticum L. + Un ● Hieracium alpinum L. + + + Rar ● Hieracium atratum Fr. s.l. + + + + Rar ● Hieracium coniops Norrl. + + + Rar ● Hieracium lapponicum Fr. + + Un ● Hieracium nigrescens Willd. s.l. + Rar ○ Hieracium umbellatum L. + Un ● Hierochloë hirta subsp. arctica (J.Presl) G.Weim. + Rar ● Honckenya peploides subsp. diffusa (Hornem.) Hultén + Rar

Huperzia selago subsp. appressa (Bach. Pyl. ex Desv.) D.Löve + Un ● ex Tzvel. ● Juncus filiformis L. + + + + Rar ● Juncus gerardi subsp. atrofuscus (Rupr.) Printz + Rar ● Juncus trifidus L. + + Un ● Koenigia islandica L. + + + Un ● Lathyrus maritimus (L.) Bigelow + Rar ● Lathyrus palustris subsp. pilosus (Cham.) Hultén + Un ○ Lathyrus pratensis L. + + Rar ○ Leucanthemum vulgare Lam. + Un ● Leymus arenarius (L.) Hochst. + + Rar ● Ligusticum scothicum L. + + + Rar ● Luzula confusa Lindeb. + Rar ● Luzula multiflora subsp. frigida (Buchenau) V.I.Krecz. + + + + + + Rar ● Luzula spicata (L.) DC. + + Rar Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 31

App. 1. cont. Main habitats

Species 1 2 3 4 5 6 7 8 9 Status 10 11 Regional Regional Frequency protected status protected ● Luzula wahlenbergii Rupr. + + + + + Rar

Lycopodium clavatum subsp. monostachyon (Grev. & Hook.) + + Rar ● Selander ● Lysimachia europaea (L.) U.Manns & Anderb. + + + + + + + + Rar ○ Melilotus officinalis L. + Un ● Mertensia maritima (L.) Gray + Un ● Montia fontana L. + + + Rar ● Nardus stricta L. + Un ● Omalotheca supina (L.) DC. + Rar ● Parnassia palustris L. + + + Rar ○ Phalaroides arundinacea (L.) Rauschert + Un ● Phleum alpinum L. + + + + + Rar ○ Phleum pratense L. + Rar ● Pinguicula vulgaris L. + Un ● Plantago maritima L. subsp. maritima + Rar ● Plantago maritima subsp. borealis (Lange) Blytt & Dahl + Rar ● Poa alpigena (Blytt) Lindm. + + + + + + + + + + Sol ● Poa alpina L. + Rar ● Poa humilis Ehrh. ex Hoffm. + Rar ● Poa pratensis L. + + + + Rar ● Puccinellia capillaris subsp. pulvinata (Fr.) Tzvelev + Rar ● Puccinellia phryganodes (Trin.) Scribn. & Merr. + Un bio ● Pyrola minor L. + + Un ○ Ranunculus acris L. subsp. acris + Rar ○ Ranunculus aggr. auricomus L. + Rar ● Ranunculus hyperboreus Rottb. + Un ○ Ranunculus repens L. + + Rar ● Ranunculus subborealis subsp. pumilus (Wahlenb.) Elven + + Rar ● Rhinanthus minor subsp. groenlandicus (Chabert) Neuman + Un ● Rhodiola rosea L. + + Rar 3 ● Rhododendron tomentosum Harmaja + + + Rar ○ Ribes nigrum L. + Un ● Rubus chamaemorus L. + + + + + + + + Fr ● Rumex acetosella subsp. arenicola Y.Mäkinen ex Elven + + Rar ○ Rumex acetosella subsp. tenuifolius (Wallr.) O.Schwarz + Rar ● Rumex lapponicus (Hiitonen) Czernov + + + + + + + Rar 32 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

App. 1. cont. Main habitats

Species 1 2 3 4 5 6 7 8 9 Status 10 11 Regional Regional Frequency protected status protected ○ Sagina procumbens L. + + + Rar ○ Salix caprea L. subsp. caprea + Un ● Salix glauca L. subsp. glauca + + + + + + + + Rar ● Salix hastata L. + Rar ● Salix lapponum L. + + Un ● Salix myrsinifolia subsp. borealis (Fr.) Hyl. + + Un ● Salix myrsinites L. + Un ● Salix phylicifolia L. + + + + + + Rar ● Saxifraga rivularis L. + Un ○ Scorzoneroides autumnalis (L.) Moench + Rar ● Solidago virgaurea subsp. lapponica (With.) Tzvelev + + + + + Rar ● Sonchus arvensis var. maritimus Wahlenb. + Rar bio ● Spinulum annotinum subsp. alpestre (Hartm.) Uotila + + Rar ● Stellaria borealis Bigelow + + + + + + Rar ● Stellaria crassifolia Ehrh. + + + + + Rar ● Stellaria fennica (Murb.) Perfil. + Rar ● Stellaria graminea L. + Rar ● Stellaria hebecalyx Fenzl Rar ● Stellaria humifusa Rottb. + + Rar ○ Stellaria media (L.) Vill. + Rar ● Taraxacum hjeltii (Dahlst.) Dahlst. + Rar ● Taraxacum repletum (Dahlst.) Dahlst. + Un ○ Trifolium pratense L. var. pratense + Un ○ Trifolium repens L. + + + Rar ● Triglochin maritima L. + Un ● Triglochin palustris L. + + Un

Tripleurospermum maritimum subsp. phaeocephalum (Rupr.) + + Rar ● Hämet-Ahti ● Trollius europaeus L. + + Rar ● Vaccinium myrtillus L. + + + + + Rar ● Vaccinium oxycoccos L. + + Un ● Vaccinium uliginosum L. + + + + + + + Sol ● Vaccinium vitis-idaea L. + + + + + + + + Sol ● Veratrum album L. + + + Rar ○ Veronica longifolia L. + Rar ○ Vicia cracca L. + Rar ○ Vicia sepium L. + Rar Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 33

Appendix 2. Vegetation map of Sosnovets Island, the White Sea. Scale 1:6500.

5 ³ 8* 20 17

8 12 11 7*

15 20 5 8

20 5 14 10 7*

14 12

14 14 5 14 22 14 9 17* 5 7 12 8 16 9

20 6 13 12 20 2 23 3 7 24 1 7 7 7 21 4 18 6* 7

19 20 0 100 Metres 34 Kozhin et al. • Memoranda Soc. Fauna Flora Fennica 95, 2019

Appendix 2. Legend Area PLANT COMMUNITY ha %

TUNDRA

Crowberry (Empetrum nigrum subsp. hermaphroditum) communities in combination with 1 0.25 0.6 crowberry-lichen (Flavocetraria nivalis, Cladonia arbuscula) communities at elevated sites

2 Lichen (Cladonia arbuscula)-crowberry communities 0.12 0.3

3 Bilberry (Vaccinium myrtillus)-dwarf cornel (Cornus suecica)-crowberry communities 0.01 0.04

VEGETATION OF PEATLANDS

Cloudberry-crowberry-lichen (Cladonia arbuscula, Flavocetraria nivalis, Cladonia rangiferina) 4 0.04 0.1 communities

5 Cloudberry-crowberry communities 1.56 3.8

Cloudberry-crowberry communities including meadow and weed species: Chamaenerion an- 6 gustifolium, Calamagrostis neglecta subsp. groenlandica, Poa alpigena, Equisetum arvense 0.47 1.2 ­subsp. alpestre

6* in combination with crowberry tundra on mineral deposits 0.59 1.5

Dwarf cornel-cloudberry-crowberry communities in combination with sedge (Carex rariflora)- crowberry and sedge (Carex recta) communities, at some sites with willow (Salix phylicifolia, S. 7 2.00 4.9 glauca, S. lapponum) thickets and tufted hairgrass (Deschampsia cespitosa) communities with Dryopteris expansa and Veratrum album in elongated hollows on slopes

in combination with sedge (Carex rariflora)-sphagnous (Sphagnum fimbriatum, S. balticum) 7* 1.28 3.2 communities with Polytrichum jensenii and Andromeda polifolia

Cloudberry communities in combination with sedge (Carex rariflora)-crowberry communities, 8 at some sites with sedge (Carex aquatilis) and small-reed (Calamagrostis phragmitoides) com- 0.93 2.3 munities in elongated hollows on slopes

in combination with sedge (Carex rariflora)-sphagnous (Sphagnum fimbriatum, S. lindber- 8* gii, S. riparium) communities with Polytrichum jensenii, Andromeda polifolia and Eriopho- 1.02 2.5 rum scheuchzeri

9 Estuary sedge (Carex recta) communities 0.31 0.8

Sedge (Carex aquatilis) and small-reed (Calamagrostis phragmitoides) communities in combi- nation with sedge (Carex aquatilis, C. rariflora)-sphagnous (Sphagnum riparium) communities 10 0.90 2.2 with Calamagrostis deschampsioides in depressions, and communities of Calamagrostis neglec- ta subsp. groenlandica and Allium schoenoprasum on dry steep shore slopes

Community complex of weakly dissected peatland Cloudberry-crowberry-lichen (Cladonia arbuscula, Flavocetraria nivalis, Cladonia rangiferina) 11 6.56 16.2 communities at elevated sites and cloudberry communities in depressions Community complexes of peatland highly dissected by dells

Cloudberry-crowberry-lichen (Cladonia arbuscula, Flavocetraria nivalis, Cladonia rangiferina) communities at elevated sites, cloudberry communities and cottongrass-sphagnous (with the 12 7.87 19.4 dominance of Sphagnum riparium, S. lindbergii, Eriophorum scheuchzeri or S. angustifolium and E. angustifolium) communities with Carex rariflora in depressions

Cloudberry-crowberry communities at elevated sites; cloudberry communities, cottongrass- sphagnous (with the dominance of Sphagnum riparium, S. lindbergii, Eriophorum scheuchzeri 13 1.41 3.5 or S. angustifolium and E. angustifolium) communities with Carex rariflora in depressions; small reed (Calamagrostis phragmitoides) communities in degraded flarks Memoranda Soc. Fauna Flora Fennica 95, 2019 • Kozhin et al. 35

App. 2. Legend cont. Area PLANT COMMUNITY ha % Cloudberry-crowberry communities including meadow and weedy species: Chamaenerion angustifolium, Calamagrostis neglecta subsp. groenlandica, Poa alpigena, Equisetum arven- 14 se subsp. alpestre at elevated sites; tufted hairgrass (Deschampsia cespitosa) and sedge (Carex 3.62 8.9 aquatilis)-small reed (Calamagrostis neglecta subsp. groenlandica) communities in degraded flarks

Cloudberry communities in shallow depressions and cottongrass-sphagnous (with the dom- inance of Sphagnum riparium, S. lindbergii, Eriophorum scheuchzeri or S. angustifolium and E. 15 angustifolium) communities with Carex rariflora in deep depressions; sedge (Carex aquatilis, C. 0.34 0.8 rariflora)-sphagnous (Sphagnum balticum, S. riparium) communities in the lower part of dells near the shoreline

Community complex of palsa mires

Cloudberry-crowberry-lichen (Cladonia arbuscula, Flavocetraria nivalis, Cladonia rangiferina) and cloudberry-crowberry communities on peat mounds; sedge (Carex aquatilis)-sphagnous 16 0.73 1.8 (Sphagnum riparium) and sedge (Carex rariflora)-cottongrass (Eriophorum russeolum)-sphag- nous (Sphagnum flexuosum) flarks

DWARF CORNEL (Cornus suecica) COMMUNITIES

Forb (Rumex lapponicus, Equisetum arvense subsp. alpestre)-grass (Calamagrostis neglecta 17 0.07 0.2 subsp. groenlandica, Poa alpigena)-dwarf cornel communities

in combination with forb (Rumex lapponicus, Conioselinum tataricum)-grass (Calamagros- 17* tis neglecta subsp. groenlandica, Poa alpigena, Festuca rubra subsp. arctica) anthropogen- 0.06 0.2 ic meadows

COASTAL VEGETATION

Fescue (Festuca rubra subsp. arctica) coastal meadows with Ligusticum scothicum and Argenti- 18 0.03 0.1 na anserina subsp. groenlandica

Coastal meadows with Festuca rubra subsp. arctica, Calamagrostis neglecta subsp. groenland- ica, Cornus suecica, Ligusticum scothicum in combination with dwarf cornel communities, and 19 sparse plant communities with Poa alpigena, Festuca rubra subsp. rubra, Solidago virgaurea 0.34 0.8 subsp. lapponica, Chamaenerion angustifolium and Rumex acetosella subsp. tenuifolius in se- verely disturbed habitats and on bare man-made lands

Sparse plants communities of Puccinellia capillaris subsp. pulvinata, Cochlearia arctica, Arctan- themum arcticum subsp. polare, Tripleurospermum maritimum subsp. phaeocephalum and Rho- diola rosea in combination with medium-level coastal marshes with the dominance of Festu- 20 4.89 12.0 ca rubra subsp. arctica and Sanionia uncinata or Carex subspathacea and C. glareosa or Calama- grostis deschampsioides and Carex rariflora and high level coastal meadows with Leymus are- narius and Sonchus arvensis var. maritimus, Ligusticum scothicum, Festuca rubra subsp. arctica

Ecological series of low- and medium-level coastal meadows, grassland and marshes with the dominance of Puccinellia phryganodes Plantago maritima subsp. borealis Carex sub- 21 → → 0.18 0.4 spathacea → Carex mackenziei → Agrostis stolonifera subsp. straminea → Festuca rubra subsp. arctica, Sanionia uncinata

ANTHROPOGENIC VEGETATION

Community complex: tufted hairgrass (Deschampsia cespitosa), sedge (Carex aquatilis)-small reed (Calamagrostis neglecta subsp. groenlandica), willowherb (Chamaenerion angustifolium) 22 communities and cloudberry-crowberry communities including meadow and weedy species: 3.32 8.2 Chamaenerion angustifolium, Calamagrostis neglecta subsp. groenlandica, Poa alpigena, Equi- setum arvense subsp. alpestre

Willowherb (Chamaenerion angustifolium) and grass (Calamagrostis neglecta subsp. groenland- ica, Anthoxanthum alpinum)-forb (Lathyrus pratensis, Trifolium repens, Alchemilla subcrenata, 23 1.54 3.8 Solidago virgaurea subsp. lapponica) meadows in combination with crowberry tundra on min- eral deposits and cloudberry-crowberry communities on a thin peat layer

24 Willowherb meadows and small reed (Calamagrostis phragmitoides) grasslands 0.18 0.4