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15The lichens and allied fungi of the Zapadny Kotlin Protected Area and its vicinities (Saint Petersburg)
I. S. Stepanchikova1, 2, D. E. Himelbrant1, 2, A. V. Dyomina1, G. M. Tagirdzhanova1
'St. Petersburg State University, Universitetskaya emb., 7-9, St. Petersburg, 199034, Russia; [email protected], [email protected], [email protected], [email protected]
2Komarov Botanical Institute, Professora Popova Str., 2, St. Petersburg, 197376,
Russia
Abstract. The lichens of the Zapadny Kotlin Protected Area and neighbouring area of the fort «Rif» (Saint Petersburg, Russia) have been investigated for the first time. Altogether 181 species of lichen-forming, lichenicolous and non-lichenized fungi are reported from the protected area. Bacidina sulphurella (Samp.) M. Hauck et V. Wirth is new for Russia, Arthonia phaeophysciae Grube et Matzer is published for the first time for North-Western European Russia, and 10 species are new to St. Petersburg. Three species included in the Red Data Book of St. Petersburg (Prilozhe-nie..., 2014) are known from the Zapadny Kotlin Protected Area: Arthonia helvola (Nyl.) Nyl., Pleurosticta acetabulum (Neck.) Elix et Lumbsch and Ramalina dilacera-ta (Hoffm.) Hoffm. In spite of maritime position, there are no specific maritime elements in the lichen flora of this area.
Keywords: lichens, new records, red-listed species, Arthonia phaeophysciae, Bacidina sulphurella, Saint Petersburg, Baltic Sea, Gulf of Finland, Kotlin Island, the Zapadny Kotlin Protected Area.
Лихенофлора заказника «Западный Котлин» и его окрестностей (Санкт-Петербург)
И. С. Степанчикова1, 2, Д. Е. Гимельбрант1, 2, А. В. Дёмина1, Г. М. Тагирджанова1
1Санкт-Петербургский государственный университет, Университетская наб., д. 7-9, Санкт-Петербург, 199034, Россия; [email protected], [email protected], [email protected], [email protected]
^Ботанический институт им. В. Л. Комарова РАН, ул. Профессора Попова, д. 2, Санкт-Петербург, 197376, Россия
Резюме. Впервые исследована лихенофлора государственного природного заказника «Западный Котлин» и прилежащей территории форта «Риф» (Санкт-Петербург, Россия). Выявлен 181 вид лишайников, лихенофильных и родственных лишайникам нелихенизированных грибов. Bacidina sulphurella (Samp.) M. Hauck et V. Wirth указана для России впервые, Arthonia phaeophysciae Grube et Matzer — новый вид для Северо-Запада Европейской России, еще 10 видов являются новыми для Санкт-Петербурга. На территории заказника найдены три
вида, занесенных в Красную книгу Санкт-Петербурга (Prilozhenie..., 2014) — Arthonia helvola (Nyl.) Nyl., Pleurosticta acetabulum (Neck.) Elix et Lumbsch и Ramalina dilacerata (Hoffm.) Hoffm. Несмотря на островное положение территории, лихенофлора западной части о. Котлин не обладает какой-либо приморской спецификой.
Ключевые слова: лишайники, новые находки, охраняемые виды, Arthonia phaeophysciae, Bacidina sulphurella, Санкт-Петербург, Балтийское море, Финский залив, остров Котлин, заказник Западный Котлин.
Introduction
The Zapadny Kotlin State Natural Protected Area (zakaznik in Russian) (ZK) is situated in the western part of Kotlin Island (the Gulf of Finland, Baltic Sea). The Island administratively belongs to Kronshtadt District of St. Petersburg. ZK was organized in 2012 to protect ecosystems and landscapes of Kotlin Island and occupies 1.02 km2. We also investigated small (ca. 0.13 km2) westernmost part of Kotlin Island west of ZK (the fort «Rif» and its vicinities), which was used for military aims up to the beginning of 2014. The total investigated area occupies about 1.15 km2 only, including the territories of the old marine forts «Rif» and «Shanets» (Fig.). It is a low and narrow peninsula (ca. 2.2 km x 150— 300 m) with swampy inner part and sandy seashores. Most part of the area is covered by considerably disturbed middle-aged or young black alder [Alnus glutinosa (L.) Gaertn.], aspen (Populus tremula L.) and willow (Salix spp.) stands, coming to the seashores; along the shoreline old
Fig. Western part of the Kotlin Island (with sample plots and additional collecting points).
willow trees are also rather common. Significant part of the territory is represented by different anthropogenic open places, especially in the territories of the forts. Present in some places are short lines of Populus bal-samifera L., Quercus robur L. and Tilia sp. as well as separate trees or groups of Acer platanoides L., Fraxinus excelsior L., Larix sp., Malus sp. and Ulmus sp. (Atlas..., 2013).
The investigated territory is of special interest for lichen researches in St. Petersburg. The lichens of Kotlin Island as well as those of Baltic forts and fort islands were not investigated before, at the same time the lichen flora of island areas could be particularly specific due to marine influence. This paper presents all the obtained data on the lichen diversity of the western part of Kotlin Island and adds some new species to the lichen flora of St. Petersburg. In course of our investigations the records of 21 species only were published recently from the area (Stepanchikova et al., 2010; Vondrak et al., 2010; Pykala et al., 2012; Atlas..., 2013).
Materials and Methods
Field investigations were undertaken in 2007, 2012 and 2014. Lichens have been collected on 29 sample plots (mainly 20 x 20 m or within natural limits of plant communities) in typical habitats, and 13 additional samples were taken in different biotopes (additional collecting points). Representative specimens are mainly deposited in the lichen herbarium of the Department of Botany of the Saint-Petersburg State University (LECB), otherwise designated specially in the text (herbarium of the University of South Bohemia, Czech Republic — CBFS, herbarium of the Botanical Museum of University of Helsinki, Finland — H, lichen herbarium of Komarov Botanical Institute RAS, St. Petersburg, Russia — LE). Lichen substances were analyzed by standard technique of thin-layer chromatography (TLC) using the solvent system C (Orange et al., 2001). WGS 84 system was used for geographical (GPS) coordinates. The lichen nomenclature generally follows Nordin et al. (2011).
Abbreviations and symbols: AD — Alexandra V. Dyomina, DH — Dmitry E. Himelbrant, LG — Ludmila V. Gagarina, LK — Ludmila A. Konoreva, MK — Martin Kukwa, EK — Ekaterina S. Kuznetsova, JP — Juha Pykala, IS — Irina S. Stepanchikova, GT — Gulnara M. Ta-girdzhanova, JV — Jan Vondrak, LY — Lidia S. Yakovchenko, ZK — the Zapadny Kotlin Protected Area, fn — field number(s), * — new for St. Petersburg, # — lichenicolous fungus, + — non-lichenized saprobic fungus.
Localities. Sample plots: 1 — open place with separate old trees on ruins of the concrete construction on the shore, S shore of the island, W part of ZK,
60°01'45" N, 29°38'55" E, 01.07.2007, LG and IS; 2 — old aspens and willows on the dunes, S shore of the island, W part of ZK, 60°01'42" N, 29°39'00" E, 01.07.2007, LG and IS; 3 — black alder forest on dunes, S shore of the island, central part of ZK, 60°01' 33" N, 29°39 '43" E, 01.07.2007, LG and IS; 4 — concrete walls around S arch of the fort «Shanets», E part of ZK, 60°01'31" N, 29°40 '24" E, 03.06.2007, DH and IS; 01.07.2007, LG and IS; 5 — old aspens and willows with separate elderberry bushes, N shore of the island W of the fort «Shanets», central part of ZK, 60°01' 39" N, 29°40 '15" E, 05.07.2007, EK and IS; 6 — young aspen grove on dunes, N shore of island, central part of ZK, 60°01'47" N, 29°39'25" E, 05.07.2007, EK and IS; 7 — group of old elms on dunes, N shore of the island in the central part of ZK, 60°01'47" N, 29°39'22" E, 05.07.2007, EK and IS; 8 — separate rowans on dunes, N shore of the island in the central part of ZK, 60°01 47" N, 29°39 22" E, 05.07.2007, EK and IS; 9 — aspen grove, W part of ZK, 60°01'49" N, 29°38 '59" E, 05.07.2007, EK and IS; 10 — black alder forest, W part of ZK, 60°01'47" N, 29°39 ' 01" E, 05.07.2007, EK and IS; 11 — elm line along the road, central part of ZK, 60°01'41" N, 29°39'51" E, 05.07.2007, EK and IS; 12 — black alder forest with separate old rowans, central part of ZK, 60°01'42" N, 29°39 '35" E, 08.07.2007, LK and IS; 13 — Rosa sp. shrubs, S shore of the island, central part of ZK, 60°01'39" N, 29°39 33" E, 08.07.2007, LK and IS; 14 — remnants of basement on herb rich meadow, fort «Shanets», E part of ZK, 60°01'33" N, 29°40 '33" E, 12.07.2007, LK and IS; 15 — trees on the glade near the road, E of the fort «Shanets», E part of ZK, 60°01'31" N, 29°40 '34" E, 12.07.2007, LK and IS; 16 — old willows with black alders on the shore, S shore of the island SE of the fort «Shanets», E part of ZK, 60°01'20" N, 29°40'37" E, 12.07.2007, LK and IS; 17 — separate trees along the E side of the fort «Shanets» wall, E part of ZK, 60°01 33" N, 29°40 27" E, 12.07.2007, LK and IS; 18 — ashes near the fort «Shanets» wall, E part of ZK, 60°01' 31" N, 29°40'24" E, 21.07.2007, LK and IS; 19 — Salixphylicifolia L. shrubs and young aspen grove on the sandy shore, N shore of the island in vicinity of the fort «Shanets», E part of ZK, 60°01'39" N, 29°40 '13" E, 20.09.2012, DH and GT; 10.05.2014, DH, IS and GT; 20 — black alder forest with willows, aspens and rowans on the sandy shore, S shore of the island, central part of ZK, 60°01 34" N, 29°39'43" E, 20.09.2012, DH and GT; 10.05.2014, DH, IS and GT; 21 — concrete constructions of the fort «Rif», westernmost part of the island, 60°01'54" N, 29°38'06" E, 28.09.2014, DH, IS and AD; 22 — open place inside the fort «Rif», westernmost part of the island, 60°01' 55.4" N, 29°38 '11.5" E, 28.09.2014, DH, IS and AD; 23 — poplar alley and concrete constructions along the N shore E of the fort «Rif», 60°01'54" N, 29°38'32" E, 28.09.2014, DH, IS and AD; 24 — black alder forest with rowans and birdcherry shrubs, E of the fort «Rif», 60°01'51" N, 29°38 '44" E, 12.10.2014, DH, IS and AD; 25 — black alders and willows on the sandy seashore, S shore E of the fort «Rif», 60°01'53" N, 29°38 '20" E, 12.10.2014, DH, IS and AD; 26 — small maple forest E of the fort «Rif», 60°01'55" N, 29°38'26" E, 12.10.2014, DH, IS and AD; 27 — old linden alley E of the fort «Rif», 60°01'55" N, 29°38'27" E, 12.10.2014, DH, IS and AD; 28 — young oaks and caragana shrubs, N shore E of the fort «Rif», 60°01' 54" N, 29°38 '40" E, 12.10.2014, DH, IS and AD; 29 — old willow forest, N shore E of the fort «Rif», 60°01'51" N, 29°38'50" E, 12.10.2014, DH, IS and AD. Additional collection points: a1 —
concrete constructions and separate trees, fort «Shanets», E part of ZK, 60°01'31" N, 29°40'24" E, 21.07.2007, LK and IS; a2 — black alder trees on the sandy shore, N shore of the island, W part of ZK, 60°01'53" N, 29°38 '59" E, 05.07.2007, EK and IS; a3 — granite boulders on the open place, fort «Shanets», E part of ZK, 60°01' 31" N, 29°40 '31" E, 21.07.2007, LK and IS; a4 — black alder forest E of the fort «Shanets», E part of ZK, 60°01'36" N, 29°40' 37" E, 21.07.2007, LK and IS; a5 — apple-tree near the sand road, central part of ZK, 60°01'45" N, 29°39 '30" E, 05.07.2007, EK and IS; a6 — old willows on the sandy shore, N shore of the island, E part of ZK close to the fort «Shanets», 60°01'41" N, 29°40'10" E, 03.06.2007, DH and IS; a7 — group of aspens on the N shore of the island, central part of ZK, 60°01'46" N, 29°39'27" E, 03.06.2007, DH and IS; a8 — group of aspens, W part of ZK, 60°01'48" N, 29°39'03" E, 03.06.2007, DH and IS; a9 — group of black alders near the path, S shore of the island E of the fort «Rif», 60°01'49" N, 29°38 '31" E, 03.06.2007, DH and IS; a10 — lichen-moss communities on the concrete walls of the fort «Rif» (S part), 60°01'51" N, 29°38'04" E, 10.05.2014, DH, IS and GT; a11 — lichen-moss communities on the concrete wall and soil, N part of the fort «Rif», 60°01' 57" N, 29°38 '13" E, 10.05.2014, DH, IS and GT; a12 — lichen-moss communities on the concrete walls and soil, W part of the fort «Rif», 60°01'55" N, 29°38 ' 04" E, 10.05.2014, DH, IS and GT; a13 — asphalted pathway and separate pine tree, W of the fort «Rif», 60°01'53" N, 29°38 '23.8" E, 28.09.2014, DH, IS and AD.
The species
Absconditella trivialis (Willey ex Tuck.) Vezda — 22: soil (LECB; fn Rif-2).
Acarospora fuscata (Schrad.) Th. Fr. — 1, 17, 18: granite (Atlas..., 2013; H; fn Kr-20).
A. glaucocarpa (Ach.) Körb. — 18: concrete.
A. moenium (Vain.) Räsänen — 14, 23, a1, a10: concrete.
*A. veronensis A. Massal. — 17, a1, a11, a12: granite, concrete, iron (H; fn Kr-20ad).
Agonimia gelatinosa (Ach.) Brand et Diederich — 21: mosses (LECB; fn Rif-1).
Alyxoria varia (Pers.) Ertz et Tehler — 23: bark of old Populus balsamifera.
Amandinea punctata (Hoffm.) Coppins et Scheid. — 7, 8, 12, 13, 15-17, 20, 23-28, a4, a9, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Larix sp., Pinus sylvestris L., Populus balsamifera, Rosa sp., Salix spp., Sorbus aucuparia L., Tilia sp. and Ulmus sp., lignum, asphalt.
Anisomeridium polypori (Ellis et Everh.) M. E. Barr — 23, 24, 29: bark of Populus balsamifera and Salix spp.
Arthonia apatetica (A. Massal.) Th. Fr. — 21: lignum (LECB; fn Rif-1).
A. dispuncta Nyl. — 26: bark of branches of Acer platanoides (LECB; fn Rif-6).
A. fusca (A. Massal.) Hepp — 14: granite (LECB; fn Kr-15).
A. helvola (Nyl.) Nyl. — 12: bark of Alnus glutinosa. Protected in Saint-Petersburg (Prilozhenie..., 2014; LECB; fn Kr-13).
A. mediella Nyl. — 12, 15, 29: bark of Alnus glutinosa, Salix sp. and Sorbus au-cuparia.
A. patellulata Nyl. — 2: bark of Populus tremula.
*#A. phaeophysciae Grube et Matzer — a10: thalli of Phaeophyscia sciastra on concrete (LECB; fn 1062). — New for North-Western European Russia. In Russia also known from Republic of Bashkortostan (Urbanavichus, Urbanavichene, 2011) and Republic of Dagestan (Urbanavichus, Ismailov, 2013).
The species is characterized by black apothecia breaking through the cortex of the host thallus. Our specimen has dark (slightly olivaceous-)brown epihymenium, pale brown subhymenium, colourless hymenium with hymenial gel I+ red, spores hyaline, 2-celled, obovate, ca. 12-15 x 4-5 mkm — i. e. generally corresponds the protologue (Grube, Matzer, 1997).
+A. punctiformis Ach. — 1, 3, 7, 17, 20, 28, a5: bark of Alnus glutinosa, Caragana sp., Malus sp., Sambucus sp., Sorbus aucuparia and Ulmus sp. (LECB; fn ZK-2).
A. ruana A. Massal. — 26: bark of Acer platanoides.
Arthrosporum populorum A. Massal. — 6, 18, 19: bark of Populus trémula and Salix sp.
Aspicilia cinerea (L.) Körb. — 17: granite (Atlas..., 2013).
Athallia holocarpa (Hoffm.) Arup, Fr0dén et Sechting — 1, 14, 17, 21, a1, a11, a12: concrete, granite incorporated in concrete and brick.
A. pyracea (Ach.) Arup, Fr0dén et Sechting — 1, 2, 5, 6, 9, 16-20, 23, 26, 29: bark of Acer platanoides, Populus balsamifera, P. trémula, Salix spp., Sambucus sp., Sorbus aucuparia and lignum.
#Athelia arachnoidea (Berk.) Jülich — 11, 15, 25, 28, 29, a13: crustose lichens and algae growing on bark of Acer platanoides, Padus avium Mill., Pinus sylvestris, Salix spp., Sorbus aucuparia and Ulmus sp.
Bacidia subincompta (Nyl.) Arnold — 5, 9, 11, 23, 29, a5: bark of Malus sp., Populus balsamifera, P. tremula, Salix spp. and Ulmus sp.
Bacidina arnoldiana (Körb.) V. Wirth et Vézda — 4, 14, 18, 21, a11: concrete, granite, iron, mosses.
*B. sulphurella (Samp.) M. Hauck et V. Wirth — 9, 10: bark of Populus tremula and Salix spp. (LECB; fn Kr-9). — New for Russia. The species is very close to B. arnoldiana, differing by regular presence of pycnidia, hook-shaped conidia, and substrate (Brand et al., 2009).
B. caligans (Nyl.) Llop et Hladún — 21, 23, 28, a11: concrete and lignum (LECB; fn 1060).
B. chloroticula (Nyl.) Vézda et Poelt — 12, 20, 21: bark of Fraxinus excelsior and Salix spp., lignum.
B. delicata (Larbal. ex Leight.) V. Wirth et Vézda — 29: bark of Salix spp. (LECB; fn Rif-9).
Biatora efflorescens (Hedl.) Räsänen — 29, a2: bark of Alnus glutinosa and Salix spp.; det. MK et IS, 2009 (LECB). Thalli contain argopsin and norargopsin.
B. ocelliformis (Nyl.) Arnold — 29: bark of Salix spp.
Bryoria capillaris (Ach.) Brodo et D. Hawksw. — 3: bark of Alnus glutinosa.
B. subcana (Nyl. ex Stizenb.) Brodo et D. Hawksw. — 10, 20: bark of Alnus glutinosa. Protected in the Leningrad Region (Krasnaya..., 2000).
Buellia disciformis (Fr.) Mudd — 1: bark of Sorbus aucuparia.
B. griseovirens (Turner et Borrer ex Sm.) Almb. — 12, 15, 24: bark of Acer platanoides, Alnus glutinosa and Salix spp.
Calogaya decipiens (Arnold) Arup, Fröden et Sechting — 1, 4, 14, 17, 18, 21, a1: concrete and granite (Atlas..., 2013).
C. pusilla (A. Massal.) Arup, Fröden et Sechting — a10: concrete (Himelbrant et al., 2015).
Caloplaca cerina (Hedw.) Th. Fr. — 6, 23, a1: bark of Populus tremula and lignum.
C. chlorina (Flot.) H. Olivier — 17, 21, 23, 28, a1, a10-13: concrete and granite; confirm. JV, 2010 (LE L7649).
C. obscurella (J. Lahm. ex Körb.) Th. Fr. — 5, 29: bark of Salix spp.; confirm. JV, 2010 (Vondrak et al., 2010; LE L7399).
C. saxicola (Hoffm.) Nordin — 4, 18, 21, a10, a11: concrete.
Candelariella aurella (Hoffm.) Zahlbr. — 1, 14, 23, 25, 28, a1, a11, a12: concrete and granite.
C. efflorescens R. C. Harris et W. R. Buck — 2, 5, 7-9, 12, 15, 16, 18, 24, 25, 29: bark of Acer platanoides, Fraxinus excelsior, Padus avium, Populus tremula, Salix spp., Sorbus aucuparia, Ulmus sp. and lignum.
C. vitellina (Hoffm.) Müll. Arg. — 1, 2, 4, 5, 9, 16-18, 29, a1, a13: bark of Populus tremula and Salix spp., concrete, granite and iron; confirm. LY, 2012 (LECB; fn Kr-2 and Kr-18).
C. xanthostigma (Ach.) Lettau — 23, 24, 26, 28, 29: bark of Acer platanoides, Caragana sp., Populus balsamifera, Salix sp. and lignum.
Cetraria sepincola (Ehrh.) Ach. — 3, 10, 13, 20: bark of Alnus glutinosa and Rosa sp.
Chaenotheca trichialis (Ach.) Th. Fr. — 15, 27: bark of Larix sp. and Tilia sp.
+Chaenothecopsis pusilla (Ach.) A. F. W. Schmidt — 15: bark of Larix sp.
Circinaria contorta (Hoffm.) A. Nordin, S. Savic et Tibell — 1, 17: concrete (LECB; fn Kr-18).
Cladonia chlorophaea (Flörke ex Sommerf.) Spreng. s. l. — 1, 17, 21, 22, 29, a3, a12: bark of Salix spp. and soil.
C. coniocraea (Flörke) Spreng. — 14, 21, 22: soil.
C. pyxidata (L.) Hoffm. — a3: soil.
C. rei Schaer. — 21, a3, a9: soil.
Coenogonium pineti (Ach.) Lücking et Lumbsch — 12, 24: bark of Alnus glutinosa.
Enchylium limosum (Ach.) Otalora et al. — a10: soil (LECB; fn 1062).
Evernia prunastri (L.) Ach. — 5, 7-10, 12, 15, 18, 20, 28, a1, a8: bark of Acer platanoides, Alnus glutinosa, Fraxinus excelsior, Padus avium, Populus tremula, Quercus robur, Salix spp., Sorbus aucuparia and Ulmus sp.
Flavoplaca citrina (Hoffm.) Arup, Fröden et Sechting — 17, 21, 23: concrete (H; fn Kr-18, in the specimen of Verrucaria christiansenii).
F. dichroa (Arup) Arup, Fröden et Sechting — a1, a10, a11: concrete; det. JV, 2010 (Vondrak et al., 2010; LE L7398).
F. flavocitrina (Nyl.) Arup, Fröden et Sechting — 21, 23, 28: concrete and granite (LECB; fn Rif-1).
Fuscidea arboricola Coppins et Tensberg — a2: bark of Alnus glutinosa; det. MK, 2009.
F. pusilla Tensberg — 3, 10, 12, 16, 20: bark of Alnus glutinosa and Fraxinus excelsior.
Graphis scripta (L.) Ach. — 29: bark of Salix spp.
Gyalecta fagicola (Hepp) Kremp. [= Pachyphiale fagicola (Hepp) Zwackh] — 1, 5, 11, 12, 15, 18, 23, 24, 29: bark of Acer platanoides, Fraxinus excelsior, Populus balsamifera, Salix spp., Sorbus aucuparia and Ulmus sp.
Gyalolechia flavorubescens (Huds.) Sechting, Frôdén et Arup — 23: lignum.
Hypocenomyce scalaris (Ach.) M. Choisy — 12, 15, a2: bark of Alnus glutinosa and Larix sp.
Hypogymnia farinacea Zopf — 1, 3, 11, 16, 20: bark of Alnus glutinosa, Betula sp. and Ulmus sp. (Atlas..., 2013; LECB; fn Kr-1).
H. physodes (L.) Nyl. — 1-3, 5-18, 20, 24-26, 28, 29, a1-3, a5, a9, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Fraxinus excelsior, Larix sp., Malus sp., Padus avium, Pinus sylvestris, Populus tremula, Quercus robur, Rosa sp., Salix spp., Sorbus aucuparia, Ulmus sp., lignum and granite.
H. tubulosa (Schaer.) Hav. — 1, 2, 5, 6, 8, 10, 14-17, 20: bark of Acer platanoides, Alnus glutinosa, Populus tremula, Salix spp., Sorbus aucuparia and granite.
#Illosporiopsis christiansenii (B. L. Brady et D. Hawksw.) D. Hawksw. — 4, 14, 25, 28, 29: thalli of Phaeophyscia orbicularis and Physcia spp. growing on bark of trees and concrete (Atlas..., 2013).
#Illosporium carneum Fr. — 22: thalli of Peltigera didactyla and P. extenuata growing on soil.
*Lathagrium undulatum (Flot.) Otalora et al. (= Collema undulatum Laurer ex Flot. var. granulosum Degel.) — 4, a1: concrete and mosses along the margin of wall (Atlas..., 2013; LECB, Kr-20ad).
Lecania cyrtella (Ach.) Th. Fr. — 1, 2, 5, 7, 9, 15, 16, 19, 20, 23, 26, 29, a6: bark of Acer platanoides, Populus balsamifera, P. tremula, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp. and lignum.
L. cyrtellina (Nyl.) Sandst. — 9: bark of Populus tremula.
L. dubitans (Nyl.) A. L. Sm. — 6: bark of Populus tremula.
L. naegelii (Hepp) Diederich et van den Boom — 2, 5-7, 9, 12, 19, 20, 23, 25, 29: bark of Populus balsamifera, P. tremula, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp. and lignum.
L. sylvestris (Arnold) Arnold — 4, 14, 21, 23: concrete.
Lecanora albellula (Nyl.) Th. Fr. — 1-3, 5, 11, 15, 16, 24, 29, a5, a13: bark of Acer platanoides, Alnus glutinosa, Larix sp., Malus sp., Pinus sylvestris, Salix spp., Ulmus sp. and lignum.
L. albescens (Hoffm.) Branth et Rostr. — 4, 14, a10, a11: concrete and brick (Atlas..., 2013).
L. allophana Nyl. — 2, 18, 23, 25, 29: bark of Populus balsamifera, P. tremula and Salix spp.
L. argentata (Ach.) Malme — 9, 10, 16: bark of Alnus glutinosa, Populus tremula and Salix spp.
L. carpinea (L.) Vain. — 1, 2, 5-12, 15-20, 23-26, 28, 29, a1, a5, a6: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Malus sp., Padus avium, Populus balsamifera, P. tremula, Quercus robur, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp. and lignum.
L. chlarotera Nyl. — 1-3, 5, 7, 8, 10, 12, 15, 16, 20, 23, 25, al, a6, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Fraxinus excelsior, Padus avium, Pinus sylvestris, Populus tremula, Salix spp., Sorbus aucuparia, Ulmus sp. and lignum.
L. crenulata Hook. — 4: concrete.
L. dispersa (Pers.) Sommerf. — 1, 4, 14, 17, 18, 21, 23, 25, 28, a1, a11, a12: concrete, granite incorporated in concrete and iron.
L. hagenii (Ach.) Ach. — 1, 2, 5-20, 23-26, 28, 29, a5, a6: bark of Acer platanoides, Betula sp., Fraxinus excelsior, Malus sp., Populus balsamifera, P. tremula, Rosa sp., Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp., concrete, lignum and granite (H; fn Kr-15).
L. intricata (Ach.) Ach. — 17, 21: granite and iron.
L. polytropa (Ehrh. ex Hoffm.) Rabenh. — 1, 17, 21, a1, a13: granite and rubber.
L. populicola (DC.) Duby — 6: smooth bark of young Populus tremula.
L. pulicaris (Pers.) Ach. — 1-3, 7, 12, 15-17, 20, 24, 25, a1: bark of Acer platanoides, Alnus glutinosa, Betula sp., Fraxinus excelsior, Padus avium, Salix spp., Sorbus aucuparia and Ulmus sp.
*L. saligna (Schrad.) Zahlbr. — 20: lignum.
L. sambuci (Pers.) Nyl. — 6: bark of Populus tremula (H; fn Kr-6).
L. semipallida H. Magn. — a1, a11: concrete (LECB; fn 1060).
*L. stenotropa Nyl. — 1, 14, 17: granite (H; fn Kr-18).
L. subintricata (Nyl.) Th. Fr. — 25: bark of Salix sp.
L. symmicta (Ach.) Ach. — 1-3, 5, 7-13, 15-20, 24, 25, 28, a1, a5: bark of Acer platanoides, Alnus glutinosa, Betula sp., Malus sp., Padus avium, Populus tremula, Quercus robur, Rosa sp., Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp. and lignum.
L. varia (Hoffm.) Ach. — 3, 7, 20, a1, a13: bark of Acer platanoides, Alnus glutinosa, Pinus sylvestris and Ulmus sp.
Lecidea nylanderi (Anzi) Th. Fr. — a2: bark of Alnus glutinosa; det. MK, 2009. Thallus contains divaricatic acid.
*L. praenubila Nyl. — a1: granite (H; fn Kr-20ad).
Lecidella elaeochroma (Ach.) M. Choisy — 2, 5, 6, 16, 18-20, 23, 25, 29: bark of Populus tremula, Salix spp. and lignum.
L. euphorea (Flörke) Hertel — 6: bark of Populus tremula.
L. stigmatea (Ach.) Hertel et Leuckert — 1, 4, 14, a1: granite and concrete.
Lepraria elobata Tensberg — 20, 24: bark of Alnus glutinosa.
L. incana (L.) Ach. — 12, 15, 25, a4, a9: bark of Alnus glutinosa and Larix sp.
L. lobificans Nyl. — 24: bark of Alnus glutinosa.
Leproplaca chrysodeta (Räsänen) J. R. Laundon — 1: concrete.
Leptorhaphis atomaria (Ach.) Szatala — 23: bark of Populus balsamifera.
Melanelixia glabratula (Lamy) Sandler et Arup — 1, 9, 15, 17, 24: bark of Alnus glutinosa, Populus tremula, Salix spp. and Sorbus aucuparia.
M. subaurifera (Nyl.) O. Blanco et al. — 1, 12, 18, 20, 24, a1: bark of Acer platanoides, Alnus glutinosa, Fraxinus excelsior, Populus tremula and Salix spp.
Melanohalea exasperata (De Not.) O. Blanco et al. — 1, 2, 5, 7-9, 14, 16-18, 20, 25, 28, a1: bark of Acer platanoides, Alnus glutinosa, Populus tremula, Quercus robur, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp. and granite.
M. exasperatula (Nyl.) O. Blanco et al. — 1-3, 5-18, 20, 24, 25, 28, 29, a1, a3, a5, a6: bark of Acer platanoides, Alnus glutinosa, Betula sp., Fraxinus excelsior, Malus sp., Padus avium, Populus tremula, Quercus robur, Rosa sp., Salix spp., Sorbus aucuparia, Ulmus sp., granite and lignum.
M. olivacea (L.) O. Blanco et al. — 1-3, 7, 10, 14, 16, 18, 20: bark of Alnus glutinosa, Salix spp., Ulmus sp. and granite.
Micarea erratica (Körb.) Hertel, Rambold et Pietschm. — a1: granite.
M. prasina Fr. s. l. — 24: bark of Alnus glutinosa.
Mycobilimbia epixanthoides (Nyl.) Vitik. et al. — 21: mosses.
Naetrocymbe punctiformis (Pers.) R. C. Harris — 1, 2, 5-8, 11, 12, 15, 17, 18, 20, 28, a1, a5: bark of Acer platanoides, Alnus glutinosa, Betula sp., Fraxinus excelsior, Malus sp., Populus tremula, Sambucus sp., Sorbus aucuparia and Ulmus sp.
Parmelia sulcata Taylor — 1-3, 5-20, 23-25, 28, 29, a1, a5, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Larix sp., Malus sp., Padus avium, Pinus sylvestris, Populus balsamifera, P. tremula, Quercus robur, Rosa sp., Salix spp., Sorbus aucuparia, Ulmus sp., lignum, granite and iron.
Parmeliopsis ambigua (Wulfen) Nyl. — 10, 20, a9: bark of Alnus glutinosa.
Peltigera canina (L.) Willd. — 1, a12: mosses and soil (LECB; fn 1061).
P. didactyla (With.) J. R. Laundon — 1, 17, 18, 21, 22, a3, a12: mosses and soil.
P. extenuata (Nyl. ex Vain.) Lojka — 14, 17, 21, 22, a12: soil.
P. rufescens (Weiss) Humb. — 1, 4, 17, 18, 21, 22, a10-12: mosses and soil.
Phaeophyscia ciliata (Hoffm.) Moberg — 2: bark of Populus tremula.
P. endophoenicea (Harm.) Moberg — 5, 12, 24: bark of Sambucus sp. and Sorbus aucuparia.
P. nigricans (Flörke) Moberg — 1, 5, 9, 11, 14, 16-18, 21, 23, a1, a11: bark of Betula sp., Fraxinus excelsior, Populus balsamifera, P. tremula, Salix spp., Sorbus aucuparia, Ulmus sp., concrete, lignum and iron.
P. orbicularis (Neck.) Moberg — 1, 2, 4-7, 9, 11, 12, 14-21, 23-26, 28, 29, a1, a5, a6, a11: bark of Acer platanoides, Alnus glutinosa, Caragana sp., Fraxinus excelsior, Malus sp., Padus avium, Populus balsamifera, P. tremula, Quercus robur, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp., concrete, granite, lignum and iron.
P. sciastra (Ach.) Moberg — 1, 17, 29, a1, a10-12: bark of Salix spp., concrete, granite, mosses, iron.
Phlyctis argena (Spreng.) Flot. — 9, 16, 25, 29: bark of Populus tremula and Salix spp.
Physcia adscendens H. Olivier — 1, 2, 5-12, 14-16, 18-20, 23-26, 28, 29, a5, a6: bark of Acer platanoides, Alnus glutinosa, Caragana sp., Fraxinus excelsior, Malus sp., Populus balsamifera, P. tremula, Quercus robur, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp., granite and lignum.
P. aipolia (Ehrh. ex Humb.) Fürnr. — 1, 2, 5-12, 15-20, 23, 25-29, a1, a5, a6: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Malus sp., Populus balsamifera, P. tremula, Quercus robur, Salix spp., Sambucus sp., Sorbus aucuparia, Tilia sp., Ulmus sp. and lignum.
P. alnophila (Vain.) Lohtander et al. — 1, 2, 8, 9, 16, 17, 29: bark of Populus tremula, Salix spp. and Sorbus aucuparia.
P. caesia (Hoffm.) Fürnr. — 1, 4, 14, 17, 18, 21, 25, 28, 29, a1, a3, a11, a13: bark of Caragana sp., Fraxinus excelsior, Quercus robur, Salix spp., Sorbus aucuparia, concrete, granite, iron, lignum, mosses and rubber (H; fn Kr-1).
P. dubia (Hoffm.) Lettau — 1, 4, 12, 14-18, 20, 21, 24, 25, 28, 29, a1, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Padus avium, Pinus sylvestris, Populus tremula, Quercus robur, Salix spp., Sorbus au-cuparia, concrete, granite, lignum and iron.
P. stellaris (L.) Nyl. — 1, 5, 8, 10, 11, 17, 18, 20, 25: bark of Acer platanoides, Alnus glutinosa, Populus tremula, Salix spp., Sambucus sp., Sorbus aucuparia and Ulmus sp.
P. tenella (Scop.) DC. — 1, 2, 5, 6-18, 20, 23-26, 28, 29, a1, a5, a6: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Larix sp., Malus sp., Populus balsamifera, P. tremula, Quercus robur, Rosa sp., Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp., concrete, granite and lignum.
Physconia distorta (With.) J. R. Laundon — 1, 2, 5, 7, 16, 23, 29: bark of Populus balsamifera, Salix spp., Ulmus sp. and lignum.
P. enteroxantha (Nyl.) Poelt — 5, 15, 18, 23, 24, 29: bark of Acer platanoides, Fraxinus excelsior, Populus balsamifera and Salix spp.
Piccolia ochrophora (Nyl.) Hafellner — 5: bark of Salix spp. and Sambucus sp. (Stepanchikova et al., 2010).
Placynthiella icmalea (Ach.) Coppins et P. James — 19, 25, 29: bark of Salix spp. and lignum.
Placynthium nigrum (Huds.) Gray — 4, 14, 17, a1: concrete (Atlas..., 2013).
Platismatia glauca (L.) W. L. Culb. et C. F. Culb. — 9, 10: bark of Alnus glutinosa and Populus tremula.
Pleurosticta acetabulum (Neck.) Elix et Lumbsch — 2: bark of Populus tremula (Atlas..., 2013; LECB; fn Kr-2). Protected in the Leningrad Region and Saint-Petersburg (Krasnaya..., 2000; Prilozhenie..., 2014).
*Polyblastia agraria Th. Fr. — 1: dead thallus of Peltigera rufescens on mosses on soil (H; fn Kr-1).
Polycauliona candelaria (L.) Frôdén, Arup et Sechting — 25: bark of Alnus glutinosa.
P. phlogina (Ach.) Arup, Frôdén et Sechting — 14: concrete (Vondrak et al., 2010; LE L7400).
P. polycarpa (Hoffm.) Frôdén, Arup et Sechting — 1-3, 5-20, 23-26, 28, 29, a1, a5, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Larix sp., Malus sp., Padus avium, Pinus sylvestris, Populus balsamifera, P. tremula, Quercus robur, Rosa sp., Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp., concrete, granite and lignum.
Protoparmeliopsis muralis (Schreb.) M. Choisy — a1, a13: granite and concrete (Atlas..., 2013).
Pseudevernia furfuracea (L.) Zopf — 3, 7, 8, 10, 20: bark of Alnus glutinosa, Sorbus aucuparia and Ulmus sp.
Pseudoschismatomma rufescens (Pers.) Ertz et Tehler — 23, 26: bark of Acer platanoides and Populus balsamifera.
*Psoroglaena abscondita (Coppins et Vézda) Hafellner et Türk — 5: bark of Sambucus sp. (H; fn Kr-5).
Psorotichia schaereri (A. Massal.) Arnold — 18: soil (H; fn Kr-20).
Ramalina dilacerata (Hoffm.) Hoffm. — a8: bark of Populus tremula (Atlas..., 2013; LECB; fn 321). Protected in the Leningrad Region and Saint-Petersburg (Kras-naya..., 2000; Prilozhenie..., 2014).
R. farinacea (L.) Ach. — 2, 7, 9, 20, 24, a8: bark of Alnus glutinosa, Populus tremula, Salix spp. and Ulmus sp.
R. fraxinea (L.) Ach. — 5, 6, 9, 16, 25, a6-8: bark of Populus tremula and Salix spp. Protected in the Leningrad Region (Krasnaya..., 2000).
Rhizocarpon lavatum (Fr.) Hazsl. — 1: granite.
Rinodina pyrina (Ach.) Arnold — 1, 2, 5, 6, 15-17, 20, 23, a1: bark of Acer platanoides, Alnus glutinosa, Populus tremula, Salix spp., Sambucus sp., Sorbus au-cuparia and lignum.
R. septentrionalis Malme — 1, 2, 7-13, 17-19, a5: bark of Alnus glutinosa, Fraxinus excelsior, Malus sp., Populus tremula, Rosa sp., Salix spp., Sambucus sp., Sorbus aucuparia and Ulmus sp.
Ropalospora viridis (Tensberg) Tensberg — 1, 16: bark of Alnus glutinosa and Sorbus aucuparia.
*Rufoplaca arenaria (Pers.) Arup, Sechting et Frôdén — 17: granite; confirm. JV, 2009 (LECB; fn Kr-18).
Rusavskia elegans (Link) S. Y. Kondr. et Kärnefelt — 14, 17, 21, a1: concrete.
Scoliciosporum chlorococcum (Graewe ex Stenh.) Vézda — 1, 3, 5, 7, 8, 12, 13, 16-18, 29, a5, a6: bark of Acer platanoides, Alnus glutinosa, Betula sp., Malus sp., Rosa sp., Salix spp., Sorbus aucuparia and Ulmus sp.
S. sarothamni (Vain.) Vézda — 1, 2, 5-13, 15-20, 24-26, 28, 29, a1, a2, a5, a6, a13: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Larix sp., Malus sp., Padus avium, Pinus sylvestris, Populus tremula, Rosa sp., Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp. and lignum.
S. umbrinum (Ach.) Arnold — 17, 21, a12: granite and iron (Atlas..., 2013).
Steinia geophana (Nyl.) Stein — a3: mosses and soil.
*Strigula jamesii (Swinscow) R. C. Harris — 19: lignum of burnt mossy tree.
Thelenella pertusariella (Nyl.) Vain. — 19, 23, 24, 26, 29: bark of Acer platanoides, Alnus glutinosa, Padus avium, Populus balsamifera and Salix sp. (Atlas..., 2013).
Thelidium minutulum Körb. — 14: granite; confirm. JP, 2011 (Pykälä et al., 2012; H; fn Kr-15).
Thelocarpon cf. epibolum Nyl. — 15: soil.
Trapeliopsis flexuosa (Fr.) Coppins et P. James — 20, 23: bark of Salix sp. and lignum.
Tuckermannopsis chlorophylla (Willd.) Hale — 1, 8, 20, 24: bark of Alnus glutinosa, Salix spp. and Sorbus aucuparia.
Usnea hirta (L.) Weber ex F. H. Wigg. — 10, 18, 20: bark of Alnus glutinosa and Salix spp.
U. subfloridana Stirt. — 10, 12: bark of Alnus glutinosa, Fraxinus excelsior and Salix spp.
Verrucaria boblensis Servit — 14, brick; det. JP, 2011 (Pykälä et al., 2012; H; fn Kr-15).
V. bryoctona (Th. Fr.) Orange — a3, a10: soil (LECB; fn 1062).
V. christiansenii Servit — 17: concrete; det. JP, 2011 (Pykälä et al., 2012; H; fn Kr-18).
V. elevata (Nyl.) Zschacke — a1: concrete; det. JP, 2011 (Pykälä et al., 2012; H; fn Kr-20ad).
V. tectorum (A. Massal.) Körb. — 17, 21, 23, 28, a1: concrete and granite; det. JP, 2011 (Pykälä et al, 2012; H; fn Kr-18 and Kr-20ad).
V. xyloxena Norman — 15: soil.
Vulpicida pinastri (Scop.) J.-E. Mattsson et M. J. Lai — 9, 10, 12, 13, 15, 20, a2, a9: bark of Alnus glutinosa, Padus avium, Populus tremula, Rosa sp. and Sorbus aucuparia.
Xanthocarpia crenulatella (Nyl.) Frôdén, Arup et Sechting — 21, a11: concrete; det. JV, 2015 (CBFS; fn Rif-1).
Xanthoria parietina (L.) Th. Fr. — 1-3, 5-12, 14-21, 23-26, 28, 29, a1, a3, a5, a6: bark of Acer platanoides, Alnus glutinosa, Betula sp., Caragana sp., Fraxinus excelsior, Malus sp., Populus balsamifera, P. tremula, Quercus robur, Salix spp., Sambucus sp., Sorbus aucuparia, Ulmus sp., concrete, granite, lignum and mosses.
Results and Discussion
The revealed lichen flora of ZK and the area of the fort «Rif» altogether comprises 181 species: 175 lichens, four lichenicolous and two saprobic fungi related to lichens. The lichen Bacidina sulphurella is new for Russia, the lichenicolous fungus Arthoniaphaeophysciae is published for the first time for North-Western European Russia, 10 other species are new to St. Petersburg.
The lichen diversity of the western part of Kotlin Island is mostly concentrated in corticolous and saxicolous lichen communities: 112 species (62 % of the lichen flora) were recorded on bark of trees and shrubs, 57 (31 %) on stones, 36 (20 %) on lignum, 15 (8 %) on soil, 12 (7 %) on iron, 10 (6 %) on mosses, 5 (3 %) were lichenicolous and 2 species (1 %) were found on old rubber. The richest phorophytes were Salix spp. (68 species, 38 % of the lichen flora), Alnus glutinosa (59 species, 33 %) and Populus tremula (52 species, 29 %). It is noteworthy that willow is the species-richest tree in the study area. Salix spp. in ZK are represented by very old single trees growing mostly along the seashore; these trees are large, branched and have coarse texture of bark, so far they are able to give shelter to many lichen species. In spite of relatively big amount of aspens, epiphytic lichens on Populus tremula are not extremely diverse. The reason is that the most of aspens in ZK are rather young; the older ones on the seashore are partly covered with sand, so only young aspen bark is available for lichens. Other trees and shrubs are not so common within the study area (and usually some of them are rather young as well), therefore the diversity of their epiphytes is lower: Sorbus aucu-
paria — 42 species (23 %), Acer platanoides — 38 (21 %), Ulmus sp. — 36 (20 %), Fraxinus excelsior — 28 (15 %), Populus balsamifera — 25 (14 %), Sambucus spp. — 22 (12 %). It is noteworthy that such regionally common phorophytes as birch and pine are presented by several trees only in the western part of Kotlin Island. Their lichen cover is very poor: on Betula sp. 19 species (10 %) and on Pinus sylvestris only 10 species (6 %) were recorded, mostly Lecanora spp. and Physcia spp. All of them are widespread lichens, no one can be considered specific to phorophyte or preferring acidic bark. Saxicolous lichen communities are most diverse on concrete — 41 species (23 % of the lichen flora), 36 species (20 %) were found on granite and 6 (3 %) on brick. Granite and concrete are usually characterized by dissimilar lichen biota, but in the western part of Kotlin Island rather many (17) species present on both substrates. The main reason is that the granite within the study area is mostly incorporated in concrete constructions and evidently contaminated by concrete. Granite boulders along the seashore in ZK do not bear any lichens, probable reasons are water pollution and frequent changes of water level. Ter-ricolous lichen cover is poor and represented mostly by pioneer species of disturbed soil: Absconditella trivialis, Enchylium limosum, Psoroti-chia schaereri, Steinia geophana, Thelocarpon cf. epibolum, Verrucaria bryoctona, V. xyloxena, several species of Cladonia and Peltigera.
The most common lichen species (found at least on 20 sample plots) in the study area are: Polycauliona polycarpa, Xanthoria parietina, Lecanora hagenii, Parmelia sulcata, Hypogymnia physodes, Phaeophy-scia orbicularis, Physcia tenella, Lecanora carpinea, Melanohalea ex-asperatula, Physcia adscendens, Physcia aipolia, Scoliciosporum saro-thamni, Lecanora symmicta. Three species found in ZK are protected in St. Petersburg: Arthonia helvola, Pleurosticta acetabulum and Ramalina dilacerata (Prilozhenie..., 2014).
In general, the lichen flora of the western part of Kotlin Island can be characterized as not very rich (in comparison to other protected areas of St. Petersburg) and significantly transformed by human activity. In spite of geographic position, there are no specific marine elements in the lichen flora of this area. Sandy soil in ZK is very poor substrate with almost no lichens: sandy ground is moving because of winds. Terricolous lichens present on primary soil on fort walls, but such places are heavily trampled and therefore the lichens there are not common as well.
In comparison to other investigated areas of St. Petersburg, ZK is characterized by extremely low amount and abundance of common forest lichens growing on acidic bark (e. g. Parmeliopsis spp., Vulpicidapinas-tri) and lignicolous species, on one hand, and high abundance of some
species growing on subneutral bark of deciduous trees (e. g. Physcia spp., Ramalina fraxinea) on the other hand. The lichenological peculiarities of the territory are also rather high diversity of saxicolous (especially on concrete) and pioneer terricolous species.
Acknowledgements. We thank M. Kukwa (Gdansk, Poland), J. Pykälä (Helsinki, Finland), J. Vondrak (Ceske Budejovice, Czech Republic) and L. Yakovchenko (Vladivostok, Russia) for help in identification or revision of the selected specimens. Our special thanks are due to L. V. Gagarina, L. A. Konoreva and E. S. Kuznetsova (St. Petersburg, Russia) who participated in the field investigations, and to anonymous reviewer for valuable comments and corrections. The work was carried out within the framework of the institutional research project (no. 01201255601) of the Komarov Botanical Institute of the Russian Academy of Sciences. The study was also supported by Russian Foundation for Basic Research (grant 14-04-01411) (TLC investigations), by Saint-Petersburg State University (research grant 1.37.151.2014) and by the Program of Basic Research of the Presidium of RAS «Wildlife: Current Status and Problems of Development» and also supported by the Directorate of Nature Protected Areas of St. Petersburg.
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Литература
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