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23LICHENS - ЛИШАИНИКИ
Are lichens coming back? Strelninsky Bereg Protected Area (St. Petersburg, Russia)
D. E. Himelbrant1, 2, I. S. Stepanchikova1, 2, E. S. Kuznetsova1, 2, V. V. Pankova2
'St. Petersburg State University, St. Petersburg, Russia 2Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia Corresponding author. D. E. Himelbrant, d_brant@mail.ru
Abstract. The total revealed lichen diversity of Strelninsky Bereg Protected Area includes 130 species, 123 of them are lichenized, four are lichenicolous, and three are non-lichenized saprobic fungi. The lichen biota of Strelninsky Bereg is quite rich, despite extremely small size of the area. The lichen Lecania olivacella is new to Russia. Bacidina pycnidiata, Fellhanera bouteillei, and Lecidella subviridis are new to North-Western European Russia. Nine lichen species reported from Strelninsky Bereg are red-listed in St. Petersburg. The reappearance of Leptogium saturninum, which was not recorded in St. Petersburg more than last 90 years, could reflect a decline of atmospheric pollution in the city suburbs. Successful occasional reintroduction of foliose lichens Flavoparmelia caperata, Hypotrachyna revoluta, and Punctelia jeckeri could be possible as a result of modern climate change in the region of St. Petersburg and in the World.
Keywords: lichen biota, new records, protected species, modern climate change, North-West European Russia.
Лишайники возвращаются? Памятник природы Стрельнинский берег
(Санкт-Петербург, Россия)
Д. Е. Гимельбрант1, 2, И. С. Степанчикова1, 2, Е. С. Кузнецова1, 2,
В. В. Панькова2
1Санкт-Петербургский государственный университет, Санкт-Петербург, Россия
2Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия Автор для переписки: Д. Е. Гимельбрант, d_brant@mail.ru
Резюме. Выявленное разнообразие лихенобиоты памятника природы «Стрельнинский берег» насчитывает 130 видов, в том числе 123 вида лишайников, четыре вида лихено-фильных и три вида нелихенизированных сапротрофных грибов. Лихенобиота памятника природы достаточно богата, несмотря на незначительную его площадь. Лишайник Lecania olivacella обнаружен впервые в России, а Bacidina pycnidiata, Fellhanera bouteillei и Lecidella subviridis — на Северо-Западе европейской части России. Девять видов лишайников, обнаруженных на территории Стрельнинского берега, занесены в Красную книгу Санкт-Петербурга. Находка лишайника Leptogium saturninum, который не регистрировали в черте Санкт-Петербурга более чем 90 лет, может свидетельствовать о снижении уровня атмосферного загрязнения в пригородах. Успешная непреднамеренная реинтродукция комплекса листоватых лишайников Flavoparmelia caperata, Hypotrachyna revoluta и Punctelia jeckeri
https://doi.org/10.31111/nsnr/2023.57.1.55 Accepted. 14 March 2023
Published: 2 April 2023
стала возможна в результате современного изменения климата в Санкт-Петербурге и в мире в целом.
Ключевые слова: лихенобиота, новые находки, охраняемые виды, современное изменение климата, Северо-Запад Европейской России.
The network of the protected areas of St. Petersburg nowadays counts 16 territories with a total area of ca. 65 km2 (Osobo..., 2012-2022). The lichen diversity in most of them has been investigated properly (see, e. g., Atlas., 2016; Stepanchikova et al., 2020). Noteworthy that a huge part of the contemporary lichen biota of St. Petersburg, ca. 90%, is known from the protected areas (Himelbrant et al., 2017). Although St. Petersburg is a city with the most studied lichen diversity in Russia (Krasnaya..., 2018; Stepanchikova et al., 2020), the investigations on the protected areas still can bring new discoveries.
Nowadays Strelninsky Bereg is a one of the smallest (0.4 km2) protected areas in St. Petersburg. It is situated in Strelna in the easternmost part of the Petrodvorets District just on the shore of the Nevskaya Bay, Gulf of Finland (Fig. 1). Strelninsky Bereg is declared to protect original landscape, plant communities, and biodiversity of the southern shore of the Nevskaya Bay. In addition, small shoreline territory East of Strelninsky Bereg was investigated as proposed for inclusion in the protected area (Fig. 2).
Relief of Strelninsky Bereg is generally uniform and is represented by low and flat accumulative terrace of ancient Littorina Sea with low and flattened coastal bars, with elevations up to 1.5 m a. s. l. Most part of the territory is covered with modern turf and in some places with sandy sediments. The vegetation is represented mainly by black alder forests [Alnusglutinosa (L.) Gaertn.], willow stands (Salix fragilis L.), and willow thickets (S. cinerea L., S. myrsinifolia Salisb., S. pentandra L.). All these types of vegetation are swampy with tall grasses [Filipendula ulmaria (L.) Maxim., Urtica dioica L., Geum rivale L., Dactylisglomerata L., Athyriumfilix-femina (L.) Roth, etc.]. Reed beds [Phragmites australis (Cav.) Trin. ex Steud.] are another type of dominant plant communities situated on the seashore. Most drained parts earlier were occupied by gardens and now are overgrown by young deciduous stands with birch (Betula pendula Roth and B. pubescens Ehrh.), aspen (Populus tremula L.), and gray alder [Alnus incana (L.) Moench], sometimes with undergrowth (Prunuspadus L., rarely Acerpla-tanoides L.). Of special interest are small planted groups of spruces [Picea abies (L.) H. Karst. and P. glauca (Moench) Voss.] on bulk areas. Almost the entire study area was used as agricultural in different periods of XIX and XX centuries, but at least over the last four decades typical shoreline tree and shrub vegetation is gradually recovering (Strelninsky., 2005).
Strelninsky Bereg Protected Area at first sight is not extremely interesting for li-chenological investigations due to its small square, uniform lowland landscape, and trivial vegetation. However, it deserves special attention due to the presence of wet black alder forests which are typical natural communities on the shores of the Gulf of Finland (Strelninsky., 2005).
Fig. 1. The location of the study area in the limits of St. Petersburg.
The first data on lichens of Strelna were published at the end of the 18th century by Grigory F. Sobolewski, who reported five taxa under the generic names "Lepra" and "Lichen" from the localities "Strelinae" and "circa Strelinae" (Sobolewski, 1799, 1802). According to the modern taxonomy, these taxa belong to four species: Anapty-chia ciliaris, Placynthium nigrum (Huds.) Gray, Stereocaulon paschale (L.) Hoffm., and Vulpicida juniperinus (L.) J.-E. Mattsson et M. J. Lai. The fate and place of storage of the Sobolewski's collections is still unclear, probably all specimens were lost (Kolchin-sky et al., 2004). However, later some of these species were found in the same area by other specialists. A specimen of Anaptychia ciliaris collected in Strelna in the beginning of 20th century is stored in the lichen herbarium of the Komarov Botanical Institute of the Russian Academy of Science (LE L-10608, 30 V 1926, N. M. Kartashova), and a specimen of Ramalina baltica Lettau — in the lichen herbarium of the Museum
Fig. 2. Map of the study area, with the location of sampling areas (indicated with numbers 1-11) and additional collection points (indicated with a1-a6).
of Natural History of the Uppsala University (UPS s. n., 9 VII 1931, K. A. Rassa-dina). Unfortunately, the exact localities for all cited specimens were not indicated, so it is impossible to estimate whether these materials originate from the modern territory of the protected area. Remarkable record of Flavoparmelia caperata was published for Strelna in the first edition of the Red Data Book of nature of St. Petersburg (Krasnaya..., 2004), but there are no data on the source of this record and place of storage of the specimen. In addition, Malysheva (1994) reported 21 and 19 common and widespread species for Konstantinovsky and Orlovsky parks in Strelna, respectively. First special investigation of Strelninsky Bereg Protected Area was conducted in June and September 2004 by Olga A. Kataeva, who reported 41 species (Kataeva, 2005). Some specimens in the frame of this work were collected by Lyubov E. Kurbatova. Our current paper is devoted to unexpected results of our recent study of lichens and allied fungi of Strelninsky Bereg Protected Area and its vicinities proposed for inclusion in the protected area.
Material and Methods
The specimens were collected by D. E. Himelbrant (DH) and I. S. Stepanchikova (IS) in May 2022. Altogether 17 localities were investigated (Fig. 2): 11 standard sample areas (SA) (20 x 20 m, otherwise in natural boundaries of the community), where the lichen diversity on each substrate was described as detailed as possible, and six additional plots, where only most interesting substrates and species were recorded. Geographical coordinates are given in the coordinate system WGS 84. Photos were made by IS using digital camera Olympus Tough 5.
Chromatography was performed by IS according to the standard techniques of high performance thin-layer chromatography using solvent systems A, B, and C (Orange
et al, 2001). The specimen of Bacidina pycnidiata was DNA-analyzed. DNA was extracted by V. V. Pankova with magnetic beads-based protocol using "Fitosorb" kit by Syntol. Internal transcribed spacer region (ITS) of the nuclear ribosomal gene was amplified and sequenced using the primer ITS1F (Gardes, Bruns, 1993). The NCBI (GenBank) accession number is provided in the species list.
The nomenclature of taxa generally follows recently published checklists for Scandinavia (Westberg et al., 2021) and Germany (Printzen et al., 2022); the genera of the family Gyalectaceae are understood according to the Lichen Flora of Russia (Gagarina, 2017). The recently collected lichen specimens, as well as the specimens deposited in LE, were identified by DH, IS, and E. S. Kuznetsova (EK). Most important records are confirmed by specimens kept in lichen herbarium of Komarov Botanical Institute (LE, the numbers are provided in the text), other material is deposited in the herbarium of the Department of Botany, St. Petersburg State University (LECB, s. n.).
Sampling locations (Fig. 2)
Standard sample areas. Saint Petersburg, Petrodvorets District, Strelna, NE to Konstanti-novsky Palace, Strelninsky Bereg Protected Area: 1 — 59°51'43.5"N, 30°04'51.0"E, old-growth willow stand (Salix fragilis) with tall grasses, 5 V 2022, Himelbrant, Stepanchikova Streln-1; 2 — 59°51'40.2"N, 30°04'58.8"E, black alder forest with tall grasses, with bird cherry, 5 V 2022, Himelbrant, Stepanchikova Streln-2; 3 — 59°51'37.9"N, 30°04'55.7"E, white spruce (Picea glauca) planted stand ca. 35 years old on a meadow, 5 V 2022, Himelbrant, Stepanchikova Streln-3; 4 — 59°51'33.5"N, 30°04'48.7"E, black alder forest with tall grasses, with rowan and birch, 5 V 2022, Himelbrant, Stepanchikova Streln-4; 5 — 59°51'39.4"N, 30°05'28.0"E, willow thickets with tall grasses, 5 V 2022, Himelbrant, Stepanchikova Streln-5; 6 — 59°51'33.7"N, 30°05'27.9"E, young birch and aspen stand on the place of gardens, 5 V 2022, Himelbrant, Stepanchikova Streln-6; 7 — 59°51'31.4"N, 30°05'15.2"E, spruce alley along the road, 5 V 2022, Himelbrant, Stepanchikova Streln-7; 10 — 59°51'35.9"N, 30°05'01.8"E, old-growth swampy willow stand (Salix fragilis) with tall grasses, 25 V 2022, Himelbrant, Stepanchikova Streln-10; 11 — 59°51'45.4"N, 30°05'12.3"E, old-growth willows (Salix fragilis) on seashore, 25 V 2022, Himelbrant, Stepanchikova Streln-11; Krasnoselsky District, E to Strelna, NE to Konstantinovsky Palace, territory proposed for the inclusion in Strelninsky Bereg Protected Area: 8 — 59°51 '41.7"N, 30°06'24.7"E, sea buckthorn and willow thickets along the track, 5 V 2022, Himelbrant, Stepanchikova Streln-8; 9 — 59°51'37.2"N, 30°06'17.5"E, willow thickets on the seashore, 5 V 2022, Himelbrant, Stepanchikova Streln-9.
Additional collection points. Saint Petersburg, Petrodvorets District, Strelna, NE to Konstantinovsky Palace, Protected Area Strelninsky Bereg: a1 — 59°51'42.2"N, 30°05'31.3"E, granite boulder on seashore, 5 V 2022, Himelbrant, Stepanchikova Streln-ad1; a2 — 59°51'36.8"N, 30°04'55.8"E, black alder in swampy black alder stand with tall grasses, 25 V 2022, Himelbrant, Stepanchikova Streln-ad2; a3 — 59°51'38.0"N, 30°05'01.7"E, black alder in gray alder stand with tall grasses, 25 V 2022, Himelbrant, Stepanchikova Streln-ad3; a4 — 59°51'41.4"N, 30°05'15.9"E, black alder stand with tall grasses, with willows (Salix fragilis), 25 V 2022, Himelbrant, Stepanchikova Streln-ad4; a5 — 59°51'33.0"N, 30°04'56.9"E, swampy black alder stand with tall grasses, 25 V 2022, Himelbrant, Stepanchikova Streln-ad5; a6 — 59°51'49.8"N, 30°05'26.2"E, concrete constructions on the seashore, 25 V 2022, Himelbrant, Stepanchikova Streln-ad6.
Protected Area Strelninsky Bereg (Kataeva, 2005): shore of Finnish Gulf in Strelna, [59°51'N, 30°05'E], 24 VI and 17 IX 2004, Kataeva or Kurbatova.
Results
In the annotated species list presented below the following symbols and abbreviations are used: # — lichenicolous and algicolous fungi, + — non-lichenized saprobic fungi, * — species new to St. Petersburg, R — species protected by the Red Data Book of St. Petersburg (Krasnaya..., 2018), s — habitat specialist, and i — indicator species (Vyyavlenie..., 2009), LR — Leningrad Region, SPb — St. Petersburg. Lichen substances are given for HPTLC-analyzed specimens. For the species already reported from Strelninsky Bereg (Kataeva, 2005), the references are given. For the species not known before from SPb and LR, information on their presence in Baltic and Scandinavian countries, as well as nearest localities in European Russia, is provided.
Acarospora moenium (Vain.) Räsänen — on concrete; a6.
Amandinea punctata (Hoffm.) Coppins et Scheid. — on bark of Betula sp., Piceaglauca, and Salix spp.; 3, 4, 6, 9-11.
RAnaptychia ciliaris (L.) Körb. — on bark of Picea glauca; 3.
Anisomeridium polypori (Ellis et Everh.) M. E. Barr — on bark of Alnusglutinosa and Salix fragilis; 10, a4.
R Arthonia helvola (Nyl.) Nyl. — on bark of Alnus glutinosa; 2, a2, a3, a5.
A. ruana A. Massal. — on bark of Acer platanoides, Alnus glutinosa, and Prunus padus; 2, 4.
Athallia cerinelloides (Erichsen) Arup et al. — on bark of Ulmus sp. in the garden, 17 IX 2004, Kurbatova, LE L-3800 (sub Phaeophyscia orbicularis).
A. holocarpa (Hoffm.) Arup et al. — on concrete; a6.
A. pyracea (Ach.) Arup et al. — on bark of Populus tremula and Salix fragilis; 1, 6, 10. On bark of Betula sp. and Salix mirsinifolia Salisb. in willow thickets [Kataeva, 2005, as Caloplaca holocarpa (Hoffm. ex. Ach.) A. E. Wade, LE L-3804].
#Athelia arachnoidea (Berk.) Jülich — on thalli of Scoliciosporum spp. and green algae growing on bark of Salix sp. and Picea abies; 2, 5, 7.
Bacidia arceutina (Ach.) Arnold — on bark of Salix fragilis; 10.
Bacidina arnoldiana (Körb.) V. Wirth et Vezda — on concrete; 6.
B. inundata (Fr.) Vezda — on granite boulder and on concrete; 6, a1.
*B. pycnidiata (Czarnota et Coppins) Czarnota et Guz.-Krzem. — on bark of Alnus glutinosa; 2 (LE L-21828; GenBank accession number: OQ238791). New to North-Western European Russia. In European Russia the nearest locality is known in the Orel Region (Muchnik, 2016). Distribution in Fennoscandia and Baltic countries: Norway, Finland (Westberg et al, 2021), Lithuania (Motiejünaite, 2017), and Estonia (Randlane et al, 2021).
The species is distinguished by having abundant whitish long-necked pycnidia, similar to Fellhanera subtilis (Vezda) Diederich et Serus., on finely granular to slightly coralloid thallus, which resembles thallus of Bacidina modesta (Zwackh ex Vain.) S. Ekman (Czarnota, Coppins, 2006).
Biatora efflorescens (Hedl.) Räsänen — on bark of Acer platanoides and Salix fragilis; 4, 10.
Buellia griseovirens (Turner et Borrer ex Sm.) Almb. — on bark of Alnus glutinosa and Salix sp.; 2, 9, a4.
B. schaereri De Not. — on bark of Alnus glutinosa; 2.
Caloplaca cerina (Hedw.) Th. Fr. — on bark of Populus tremula and Salix fragilis; 6, 11.
Candelariella aurella (Hoffm.) Zahlbr. — on concrete; a6.
C. efflorescens R. C. Harris et W. R. Buck — on bark of Alnus glutinosa, Betula sp., Picea glauca, Prunus padus, and Salix spp.; 1-3, 5, 6, 10, 11. On bark of Ulmus sp. near the garden, 17 IX 2004, Kurbatova, LE L-3797 (sub Parmelia sulcata).
C. lutella (Vain.) Räsänen — on bark of Salix sp.; 9. On bark of S. mirsinifolia in willow thickets, 17 IX 2004, Kurbatova, LE L-3807 (sub Myriolecis hagenii).
C. reflexa (Nyl.) Lettau — on bark of Picea glauca (branches); 3.
C. vitellina (Hoffm.) Müll. Arg. — on bark of Hippophae rhamnoides and Salix spp., on granite; 1, 5, 6, 8-11. On bark of Salix mirsinifolia in willow thickets, 17 IX 2004, Kurbatova, LE L-3804 (sub Athallia pyracea).
C. xanthostigma (Ach.) Lettau — on bark of Salix spp.; 5, 9, 10. The specimen published by Kataeva (2005) as Candelariella xanthostigma (LE L-3804) refers to C. vitellina.
Catillaria nigroclavata (Nyl.) Schuler — on bark of Salix sp.; 5.
Catinaria atropurpurea (Schaer.) Vezda et Poelt — on wood of Salix fragilis; 1.
Cetraria sepincola (Ehrh.) Ach. — on bark of Alnus glutinosa; 2. On bark of A. glutinosa in black alder forest near the brook (Kataeva, 2005, LE L-3769).
R Chaenotheca brachypoda (Ach.) Tibell — on bark of Alnus glutinosa and Salix fragilis; 10, a4.
C. furfuracea (L.) Tibell — on bark of Alnus glutinosa and Salix fragilis; 2, 10.
R iC. hispidula (Ach.) Zahlbr. — on bark of Alnus glutinosa; 2.
C. trichialis (Ach.) Th. Fr. — on bark of Alnus glutinosa and Salix fragilis; 2, 4, 10.
Cladonia chlorophaea (Flörke ex Sommerf.) Spreng. — on bark of Alnus glutinosa and Salix fragilis; 2, 10. On soil in the meadow (Kataeva, 2005, LE L-3780). Specimens contain fumarpro-tocetraric acid complex and fatty acid.
C. coniocraea (Flörke) Spreng. — on bark of Alnus glutinosa and Salix fragilis; 2, 10.
C. fimbriata (L.) Fr. — on bark of Alnus glutinosa; 2. On bark of Salix sp. in black alder forest with willow (Kataeva, 2005, LE L-3779). The specimen LE L-3779 contains fumarproto-cetraric acid complex.
Evernia prunastri (L.) Ach. — on bark of Alnus glutinosa, Picea glauca, Prunus padus, and Salix spp.; 2, 3, 5, 10, 11, a4. On bark of Prunus padus on the glade and on bark of old Salix sp. in black alder forest (Kataeva, 2005).
*Fellhanera bouteillei (Desm.) Vezda (Fig. 3A) — abundant on twigs and needles of Picea glauca; 3 (LE L-21829). New to North-Western European Russia. Nearest localities in European Russia known from Kaliningrad (Himelbrant et al, 2022) and Moscow (Golubkova, 1959) regions. Distribution in Fennoscandia and Baltic countries: Norway, Sweden, Finland (Westberg et al, 2021), Lithuania (Motiejünaite, 2017), and Estonia (Randlane et al, 2021).
Crustose lichen, characterized by entirely sorediate thallus, pinkish apothecia, and 1-sep-tate ascospores (Cannon et al, 2022).
RFlavoparmelia caperata (L.) Hale (Fig. 3B) — well developed on bark of Picea glauca (branches); 3 (LE L-21834).
Graphis scripta (L.) Ach. — on bark of Prunus padus; 2.
#Heterocephalacria physciacearum (Diederich) Millanes et Wedin — on thallus of Phy-scia aipolia growing on bark of Salix sp.; 5.
Hypocenomyce scalaris (Ach.) M. Choisy — on bark of Alnus glutinosa; 2.
Hypogymnia physodes (L.) Nyl. — on bark of Alnus glutinosa, Betula sp., Picea glauca, Prunus padus, and Salix spp.; 2-6, 10. On bark of various deciduous trees in different habitats [Kataeva, 2005, LE L-3785 (sub Tuckermannopsis chlorophylla)].
Hypogymnia tubulosa (Schaer.) Hav. — on bark of Alnusglutinosa, Betula sp., Piceaglauca, and Salix spp.; 2-6, 10. On bark of Prunuspadus on the glade [Kataeva, 2005, as Hypogymnia tubulosa f. farinosa (Hillm.) Rassad., LE L-3810].
R sHypotrachyna revoluta (Flörke) Hale (Fig. 3C) — well developed on bark of Picea glauca (branches); 3 (LE L-21830).
#Illosporiopsis christiansenii (B. L. Brady et D. Hawksw.) D. Hawksw. — on thalli of Phy-scia spp. growing on bark of Salix spp.; 1, 5, 9, 10.
Lecania cyrtella (Ach.) Th. Fr. — on bark of Acerplatanoides, Betula sp., Populus tremula, and Salix spp.; 1, 4, 6, 8-11. The specimen published by Kataeva (2005) as Lecania cyrtella (LE L-3805) refers to Lecanora albellula.
L. cyrtellina (Nyl.) Sandst. — on fruit body of polypore growing on old Salix sp.; 5.
L. dubitans (Nyl.) A. L. Sm. — on bark of Salix fragilis; 1.
L. naegelii (Hepp) Diederich et van den Boom — on bark of Acer platanoides, Alnus glutinosa, Picea glauca, Populus tremula, Prunus padus, and Salix spp.; 1-6, 8-11. On bark of Salix sp. in swampy willow thickets, 24 VI 2004, Kataeva, LE L-3773 (as Lecania fuscella).
*L. olivacella (Nyl.) Zahlbr. (Fig. 3D) — on rusty iron; 6 (LE L-21831). New to Russia. Distribution in Fennoscandia and Baltic countries: Norway and Sweden (Westberg et al., 2021). The rare but widespread in Europe species; also known from Austria, France, Great Britain, Italy, and Spain (Atienza, Segarra, 1999; Nimis et al., 2018; Cannon et al, 2021).
This normally saxicolous crustose lichen is characterized by persistent thalline margin of apothecia with a rich photobiont layer, flat to moderately convex disc, 1-septate ascospores, and areolate to nearly squamulose thallus with no pruina (Cannon et al, 2021).
Lecanora aitema (Ach.) Hepp — on bark of Alnus glutinosa, Picea glauca, and Salix fragilis; 1, 3, 4, 10, 11.
L. albellula (Nyl.) Th. Fr. — on bark of Alnus glutinosa and Salix sp.; 2, 4, 9. On bark of Salix spp. in willow thickets, 24 VI 2004, Kataeva, LE L-3785 (sub Tuckermannopsis chlorophylla); on bark of Alnus incana in gray alder stand, 24 VI 2004, Kataeva, LE L-3786 (sub Rinodina septen-trionalis); on bark of Alnus glutinosa in black alder forest, 24 VI 2004, Kataeva, LE L-3793 (as Lecanora saligna); on bark of Salix fragilis, 17 IX 2004, Kurbatova, LE L-3805 (as Lecania cyrtella, sub Polycauliona polycarpa).
L. allophana Nyl. — on bark of Populus tremula and Salix fragilis; 6, 10, 11. The specimen published by Kataeva (2005) as Lecanora allophana (LE L-3791) refers to L. chlarotera.
L. argentata (Ach.) Malme — on bark of Salix fragilis; 1.
L. carpinea (L.) Vain. — on bark of Alnus glutinosa, Betula sp., Populus tremula, Prunus padus, Salix spp., and Sorbus aucuparia; 1, 2, 4-6, 8-11. On bark of Salix sp. in willow thickets (Kataeva, 2005, LE L-3795); same substrate and biotope, 17 IX 2004, Kataeva, L-3809 (as Lecanora leptyrodes).
L. chlarotera Nyl. — on bark of Acer platanoides, Prunus padus, Salix spp., and Sorbus aucuparia; 1, 2, 4, 8, 9. On bark of Prunus padus on the glade, 24 VI 2004, Kataeva, LE L-3791 (as Lecanora allophana) and Salix sp. in swampy willow thickets, 24 VI 2004, Kataeva, LE L-3794 (as Lecanora populicola).
L. pulicaris (Pers.) Ach. — on bark of Alnus glutinosa, Betula sp., and Picea spp.; 2-4, 6, 7.
L. symmicta (Ach.) Ach. — on bark of Alnus glutinosa, Betula sp., Picea spp., Prunus padus, and Salix spp.; 1-7, 9-11, a4. On bark of Alnus spp., Prunus padus, and Salix spp. in different habitats [Kataeva, 2005, LE L-3786 (sub Rinodina septentrionalis), L-3805, L-3808 (sub Polycauliona polycarpa)].
L. umbrina (Ach.) A. Massal. — on bark of Acer platanoides, Alnus glutinosa, Betula sp., Hippophae rhamnoides, Picea abies, Prunus padus, and Salix spp.; 1, 2, 4-11.
L. varia (Hoffm.) Ach. — on bark of Salix fragilis; 1.
Lecidea nylanderi (Anzi) Th. Fr. — on bark of Alnus glutinosa; 2.
Lecidella elaeochroma (Ach.) M. Choisy — on bark of Picea glauca, Populus tremula, Prunus padus, and Salix spp.; 1-3, 5, 6, 9, 11.
L. euphorea (Flörke) Hertel — on bark of Populus tremula; 6.
L. flavosorediata (Vezda) Hertel et Leuckert — on bark of Alnus glutinosa; a5.
*L. subviridis Tensberg — on bark of Alnus glutinosa; 2 (LE L-21833). The specimen contains atranorin, thiophanic acid, cf. arthothelin, cf. expallens-unknown. New to North-Western European Russia. In European Russia the species is known also from the Kaliningrad Region (Himelbrant et al, 2022). Distribution in Fennoscandia and Baltic countries: Norway, Sweden (Westberg et al, 2021), Latvia (Motiejünaite et al., 2016), Lithuania (Motiejünaite, 2017), and Estonia (Randlane et al, 2021).
The species is characterized by thin, sometimes almost leprose, pale yellow-green thallus, and specific secondary metabolites (Zdunczyk, Kukwa 2014).
Lepraria elobata Tonsberg — on bark of Alnus glutinosa; 2.
L. incana (L.) Ach. — on bark of Alnus glutinosa and Salix fragilis; 2, 10. The specimen published by Kataeva (2005) from bark of Alnus glutinosa, willow thickets, 24 VI 2004, Kataeva (LE L-3784), is undeterminable (poor material).
R' iLeptogium saturninum (Dicks.) Nyl. (Fig. 3E) — small thallus (ca. 1 cm in diam.) on bark of very old Salix sp. (Fig. 3F); 5.
Leptorhaphis atomaria (Ach.) Szatala — on bark of Populus tremula; 6.
Melanelixia glabratula (Lamy) Sandler et Arup — on bark of Prunus padus; 2.
M. subaurifera (Nyl.) O. Blanco et al. — on bark of Alnus glutinosa, Betula sp., Hippophae rhamnoides, Picea glauca, Prunus padus, and Salix spp.; 2, 3, 5, 6, 8-11, a4.
Melanohalea exasperata (De Not.) O. Blanco et al. — on bark of Alnus glutinosa and Salix spp.; 2, 5, 9, 11.
M. exasperatula (Nyl.) O. Blanco et al. — on bark of Alnus glutinosa, Picea spp., Prunus padus, and Salix spp.; 2-5, 7, 10, 11. On bark of Alnus incana, Prunus padus, and Salix sp. in different habitats [Kataeva, 2005, as Melanelia exasperatula (Nyl.) Essl., LE L-3775].
M. olivacea (L.) O. Blanco et al. — on bark of Alnus glutinosa, Betula sp., and Salix sp.; 2, 4-6.
Mycobilimbia epixanthoides (Nyl.) Vitik. et al. — on mosses growing on bark of Alnus glutinosa; 2.
+Mycocalicium subtile (Pers.) Szatala — on wood of Alnus glutinosa; a3.
Myriolecis dispersa (Pers.) Sliwa et al. — on concrete; 6, a6.
M. hagenii (Ach.) Sliwa et al. — on bark of Alnus glutinosa and Salix spp.; 1, 4, 5, 9-11. On bark of Betula sp., Prunus padus, and Salix sp. [Kataeva, 2005, as Lecanora hagenii (Ach.) Ach., LE L-3807]; on bark of Alnus incana in gray alder stand, 24 VI 2004, Kataeva, LE L-3786 (sub Rinodina septentrionalis).
M. sambuci (Pers.) Clem. — on bark of Ulmus sp. in the garden, 17 IX 2004, Kurbatova, LE L-3800 (sub Phaeophyscia orbicularis).
M. semipallida (H. Magn.) Sliwa et al. — on concrete; a6.
Naetrocymbe punctiformis (Pers.) R. C. Harris — on bark of Alnus glutinosa and Salix sp.; 4, 5.
+Naevia punctiformis (Ach.) A. Massal. — on bark of Alnusglutinosa and Salix sp.; 4, 5.
Pachyphiale fagicola (Hepp) Zwackh — on bark of Salix sp.; 5.
Parmelia sulcata Taylor — on bark of Alnus glutinosa, Betula sp., Picea spp., Populus tremula, and Salix spp., on treated wood; 1-7, 9-11, a4. On bark of various deciduous trees in different habitats [Kataeva, 2005, LE L-3797, L-3785 (sub Tuckermannopsis chlorophylla)].
Parmeliopsis ambigua (Wulfen) Nyl. — on bark of Alnus glutinosa; 2, a4.
Peltigera didactyla (With.) J. R. Laundon — on clayey soil on the shore of the Gulf of Finland (Kataeva, 2005, LE L-3776).
P. neopolydactyla (Gyeln.) Gyeln. — on bark of Salix fragilis; 10.
P. praetextata (Flörke ex Sommerf.) Zopf — on bark of Salix fragilis and on mosses growing on granite boulder; 6, 10.
Phaeophyscia nigricans (Flörke) Moberg — on bark of Populus tremula and Salix spp., on wood of Salix fragilis; 1, 5, 6, 9, 11.
P. orbicularis (Neck.) Moberg — on bark of Alnus glutinosa, Betula sp., Hippophae rham-noides, Picea abies, Populus tremula, Prunus padus, and Salix spp., on wood of Salix fragilis, and on concrete; 1, 4, 5-11, a6. On bark of Ulmus sp. near the gardens (Kataeva, 2005, LE L-3800).
P. sciastra (Ach.) Moberg — on granite; a1.
Phlyctis argena (Spreng.) Flot. — on bark of Alnus glutinosa, Picea glauca, Prunus padus, and Salix spp.; 2, 3, 9, 10.
Physcia adscendens H. Olivier — on bark of Alnus glutinosa, Hippophae rhamnoides, Picea spp., Prunus padus, and Salix spp., on wood of Salix fragilis; 1-3, 5, 7-11. On bark of Salix sp. in swampy willow thickets (Kataeva, 2005, LE L-3781).
P. aipolia (Ehrh. ex Humb.) Fürnr. — on bark of Alnus glutinosa, Betula sp., and Salix spp.; 1, 2, 4-6, 8-11. On bark of S. mirsinifolia in willow thickets, 17 IX 2004, Kurbatova, LE L-3807 (sub Myriolecis hagenii).
P. alnophila (Vain.) Loht. et al. — on bark of Alnus glutinosa and Salix spp.; 1, 2, 5, 9, 11.
P. caesia (Hoffm.) Fürnr. — on granite; 6.
P. dubia (Hoffm.) Lettau — on bark of Betula sp.; 6. On bark of Salix sp. in swampy willow thickets, 24 VI 2004, Kataeva, LE L-3781 (sub Physcia adscendens); on bark of Ulmus sp. in the garden, 17 IX 2004, Kurbatova, LE L-3800 (sub Phaeophyscia orbicularis).
P. stellaris (L.) Nyl. — on bark of Alnus glutinosa, Hippophae rhamnoides, and Salix spp.; 2, 4, 5, 8, 9, 11. On bark of Salix sp. and Ulmus sp. in different habitats (Kataeva, 2005, LE L-3798).
P. tenella (Scop.) DC. — on bark of Alnus glutinosa, Betula sp., Hippophae rhamnoides, Picea spp., and Salix spp., on treated wood; 1-11. On bark of various deciduous trees in different habitats [Kataeva, 2005, LE L-7807, L-3781 (sub Physcia adscendens)].
Physconia detersa (Nyl.) Poelt — on bark of Salix fragilis; 10.
P. distorta (With.) J. R. Laundon — on bark of Picea glauca and Salix fragilis; 3, 10, 11.
P. enteroxantha (Nyl.) Poelt — on bark of Betula sp.; 6.
Placynthiella dasaea (Stirt.) Tonsberg — on wood of Alnus glutinosa and Salix fragilis; 11, a4.
Platismatia glauca (L.) W. L. Culb. et C. F. Culb. — on bark of Alnus glutinosa and Salix fragilis; 10, a4. On bark of Alnus glutinosa in black alder forest and Salix sp. in swampy willow thickets (Kataeva, 2005, LE L-3782).
Polycauliona candelaria (L.) Fröden et al. — on bark of Salix sp.; 5.
P. polycarpa (Hoffm.) Fröden et al. — on bark of Alnus glutinosa, Betula sp., Hippophae rhamnoides, Picea abies, Prunus padus, and Salix spp., on treated wood; 1, 2, 4-11. On bark of various deciduous trees in different habitats [Kataeva, 2005, LE L-3797 (sub Parmelia sulcata)
and L-3804 (sub Athalliapyracea)]; on bark of Salix sp. in willow thickets, 17 IX 2004, Kurba-tova, LE L-3805 [as Caloplaca lobulata (Flörke) Hellb.].
Pseudevernia furfuracea (L.) Zopf — on bark of Alnusglutinosa and Prunuspadus; 2, a4.
Pseudoschismatomma rufescens (Pers.) Ertz et Tehler — on bark of Salix fragilis; 1.
R sPunctelia jeckeri (Roum.) Kalb (Fig. 3G) — well developed on bark of Picea glauca branches (Fig. 3H); 3 (LE L-21832).
Ramalina farinacea (L.) Ach. — on bark of Alnus glutinosa, Picea glauca, Prunus padus, and Salix spp.; 1-3, 9-11. On bark of old Salix sp. in black alder forest (Kataeva, 2005, LE L-3770); on bark of Salix sp. in swampy willow thickets, 24 VI 2004, Kataeva, LE L-3778 (as Ramalina pollinaria).
R. fraxinea (L.) Ach. — on bark of Prunus padus; a4.
Rinodina exigua (Ach.) Gray — on bark of Alnus glutinosa on the shore of the Gulf of Finland (Kataeva, 2005), on bark of Salix fragilis in willow thickets, 17 IX 2004, Kurbatova, LE L-3805 (sub Polycauliona polycarpa). The specimen published by Kataeva (2005) as Rinodina exigua (LE L-3801) refers to R. septentrionalis.
R. pyrina (Ach.) Arnold — on bark of Hippophae rhamnoides and Salix spp.; 1, 8, 9, 11.
R. septentrionalis Malme — on bark of Salix sp.; 5, 8. On bark of Alnus incana (Kataeva, 2005, LE L-3786); on bark of Salix sp. in swampy willow thickets, 24 VI 2004, Kataeva, LE L-3788 (as Rinodina laevigata); on bark of Salix mirsinifolia in willow thickets, 17 IX 2004, Kurbatova, LE L-3801 (as Rinodina exigua) and L-3808 (sub Lecanora symmicta).
Ropalospora viridis (Tonsberg) Tonsberg — on bark of Alnus glutinosa and Prunus padus; 2.
Scoliciosporum chlorococcum (Graewe ex Stenh.) Vezda — on bark of Populus tremula and Salix sp.; 5, 6, 9. On bark of Alnus incana, 24 VI 2004, Kataeva, LE L-3772 [as Bacidia subin-compta (Nyl.) Arnold].
S. sarothamni (Vain.) Vezda — on bark of Alnus glutinosa, Betula sp., Hippophae rhamnoides, Picea spp., Populus tremula, Prunus padus, and Salix spp., on wood of S. fragilis; 1-11. On bark of Ulmus sp. in a garden, 17 IX 2004, Kataeva, LE L-3797 (sub Parmelia sulcata); on bark of Salix sp. in willow thickets, 17 IX 2004, Kataeva, LE L-3806 (as Scoliciosporum umbrinum); same substrate and biotope, 24 VI 2004, Kataeva, LE L-3785 (sub Tuckermannopsis chloro-hylla); on bark of Padus avium in open place, 24 VI 2004, Kataeva, LE L-3771 (as Scoliciosporum umbrinum).
RScytinium subtile (Schrad.) Otalora et al. — on bark of Salix fragilis; 10.
Steinia geophana (Nyl.) Stein — on clayey soil on the shore of the Gulf of Finland, 24 VI 2004, Kataeva, LE L-3777 [as Mycobilimbia hypnorum (Lib.) Kalb et Hafellner].
+Stenocybe pullatula (Ach.) Stein — on bark of Alnus glutinosa; 2, 4.
Strangospora pinicola (A. Massal.) Körb. — on bark of Alnus incana, 6 VIII 2004, Kataeva, LE L-2069 (as Strangospora deplanata).
Thelenella pertusariella (Nyl.) Vain. — on bark of Prunus padus; 1.
Toniniopsis separabilis (Nyl.) Gerasimova et A. Beck — on bark of Acer platanoides and Salix fragilis; 1, 4. The specimen published by Kataeva (2005) as Bacidia subincompta (Nyl.) Arnold (LE L-3772) refers to Scoliciosporum chlorococcum.
#Tremella lichenicola Diederich — on thallus of Violella fucata growing on bark of Alnus glutinosa; 2.
Tuckermannopsis chlorophylla (Willd. ex Humb.) HaLE — on bark of Alnus glutinosa, Picea glauca, Prunus padus, and Salix sp.; 2, 3, 5, a4. On bark of Alnus glutinosa in black alder forest and Salix sp. in swampy willow thickets [Kataeva, 2005, as Cetraria chlorophylla (Willd.) Vain., LE L-3785].
Usnea hirta (L.) F. H. Wigg. — on bark of Salix sp.; 5.
U. subfloridana Stirt. — on bark of Prunuspadus and Salix sp.; 5, a4.
Verrucaria muralis Ach. — on concrete; 6.
Violella fucata (Stirt.) T. Sprib. — on bark of Alnusglutinosa; 2.
Vulpicida pinastri (Scop.) J.-E. Mattsson et M. J. Lai — on bark of Alnus glutinosa, Picea glauca, and Prunus padus; 2, 3, a4. On bark of various deciduous trees in different habitats (Ka-taeva, 2005, LE L-3783).
Xanthoria parietina (L.) Th. Fr. — on bark of Alnus glutinosa, Betula sp., Hippophae rhamnoides, Picea spp., Populus trémula, Prunus padus, and Salix spp., on wood of S. fra-gilis; 1-11. On bark of various deciduous trees in different habitats (Kataeva, 2005, LE L-3802).
Taxa excluded from lichen flora of Strelninsky Bereg Protected Area
Calogaya lobulata (Flörke) Arup et al. — reported by Kataeva (2005) as Caloplaca lo-bulata (Flörke) Hellb., the specimen belongs to Polycaulionapolycarpa (LE L-3805).
Lecania fuscella (Schaer.) A. Massal. — reported by Kataeva (2005), the specimen belongs to L. naegelii (LE L-3773).
Lecanora intumescens (Rebent.) Rabenh. — reported by Kataeva (2005), the specimen LE L-3792 is poorly developed for correct identification, but does not belong to L. intu-mescens.
L. leptyrodes (Nyl.) Degel. — reported by Kataeva (2005), the specimen belongs to L. carpinea (LE L-3809).
L. populicola (DC.) Duby — reported by Kataeva (2005), the specimen belongs to L. chlarotera (LE L-3794).
L. saligna (Schrad.) Zahlbr. — reported by Kataeva (2005), the specimen belongs to L. albellula (LE L-3793).
Bryobilimbia hypnorum (Lib.) Fryday et al. — reported by Kataeva (2005) as Myco-bilimbia hypnorum (Lib.) Kalb et Hafellner, the specimen belongs to Steinia geophana (LE L-3777).
Ramalina pollinaria (Westr.) Ach. — reported by Kataeva (2005), the specimen belongs to R. farinacea (LE L-3778).
Rinodina laevigata (Ach.) Malme — reported by Kataeva (2005), the specimen belongs to R. septentrionalis (LE L-3788).
Scoliciosporum umbrinum (Ach.) Arnold — reported by Kataeva (2005), the specimens belong to S. sarothamni (LE L-3771, L-3806).
Strangospora deplanata (Almq.) Clauzade et Cl. Roux — reported by Kataeva (2005), the specimen belongs to S. pinicola (LE L-2069).
Fig. 3. The lichens of Strelninsky Bereg and their habitats. A — Fellhanera bouteillei on needles of Picea glauca (LE L-21829); B — Flavoparmelia caperata on twig of Picea glauca (LE L-21834); C — Hypotrachyna revoluta on twig of Picea glauca (LE L-21830); D — Lecania olivacella on rusty iron (LE L-21831); E — Leptogium saturninum on bark of old Salix sp. (field record, SA 5); F — Habitat of Leptogium saturninum: old willow tree in thickets (SA 5); G — Punctelia jeckeri on twig of Picea glauca (LE L-21832); H — Picea glauca trees on a meadow. Scale bars: A-C, E, G — 1 cm; D — 1 mm.
Discussion
The total revealed lichen diversity of Strelninsky Bereg Protected Area includes 130 species, 123 of them are lichenized, four are lichenicolous, and three are non-li-chenized saprobic fungi. The lichen biota of Strelninsky Bereg is quite rich, despite extremely small size of the area, and is richer than a number of compatible small territories within the city limits (Himelbrant et al., 2007; Atlas..., 2016; Stepanchikova et al., 2020; unpublished authors' data). For example, the list of lichens and allied fungi of Petrovsky Pond Protected Area (ca. 0.03 km2) counts 30 species only, Elagin Island Protected Area (ca. 1 km2) — 87 species, Piskarevsky Park (ca. 1.2 km2) — 91 species, Polezhaevsky Park (ca. 1.5 km2) — 108 species, Dubki Park (0.6 km2) — 117 species, proposed protected area Gagarka (ca. 0.3 km2) — 121 species. Obviously, the diversity of lichens and allied fungi depends not only on the size of the area, but more on the diversity and condition of substrates and biotopes, as well as on the position of the area within the city.
In the result of the revision of the herbarium specimens in LE, we accepted the reports of 25 lichen species out of 41 which were published (Kataeva, 2005). Moreover, some additional species were identified in herbarium specimens. Altogether, 39 species were correctly published or collected before our study.
In May 2022, we found 124 species in Strelninsky Bereg. In course of this inventory, we did not record only six species which existed here in 2004: Athallia cerinel-loides, Myriolecis sambuci, Peltigera didactyla, Rinodina exigua, Steinia geophana, and Strangospora pinicola. They all are widely distributed in the Leningrad Region and St. Petersburg, but are small-sized and normally not abundant, so they could be just overlooked.
Most common species registered in all 11 SA were Physcia tenella, Scoliciosporum sarothamni, and Xanthoria parietina; Lecania naegelii, Lecanora symmicta, L. umbrina, Parmelia sulcata, and Polycauliona polycarpa found in 10 SA. Controversially, 51 species were found in our investigation in only one SA or additional plot.
An average amount of species per SA was 33.1+3.7, with minimum 12 (young shadowy spruce alley, SA 7), and maximum 56 (black alder forest with bird cherry, SA 2). This average number is similar with territories along the northern shore of the Gulf of Finland within St. Petersburg, where an average is 33.3+1.3 (Stepanchikova et al., 2020), but significantly lower than in more remote territories of the Leningrad Region or even St. Petersburg with very rich lichen diversity. For example, in the proposed protected area Motornoe-Zaostrovje (LR) an average amount of species per standard SA was 58.6+3.1 (Stepanchikova et al., 2022), in Konevets Island (LR) — 47.7+2.6 (Himelbrant et al., 2018), in proposed protected area Pukhtolova Gora (SPb) — 49.7+5.5 (Stepanchikova et al., 2021), and in the valley of the Smorodinka River (LR) — 42.9+4.0 (Stepanchikova et al., 2013).
Altogether 78 species (60.0% of the lichen biota revealed in 2022) were recorded in communities dominated by willows, of them 60 species (46.2%) inhabited old-growth willow stands of Salix fragilis and S. myrsinifolia, and 51 species (39.2%) were found in
willow thickets. Black alder forests which are typical natural communities of the study area were also comparatively rich in lichens: 73 species (56.2%) were recorded there. In young birch and aspen stand (SA 6), 37 species (28.5%) were recorded. Planted spruces (alley and single trees) brought to the lichen biota 32 species (24.6%). Concrete constructions and granite boulders were very poor in lichens (8 species altogether).
The majority of lichens in Strelninsky Bereg Protected Area were epiphytic (104 species, 80.0% of the lichen biota revealed in 2022), which is typical for the region. The richest phorophytes were Salix spp. (71 species) and Alnusglutinosa (58 species). Other phorophytes are arranged below in descending order of the number of species: Picea glauca and Prunuspadus (28 species each), Betula sp. (20 species), Populus tremula (15 species), Hippophae rhamnoides and Picea abies (11 species each), Acer platanoides (7 species), and Sorbus aucuparia (2 species). Very few species were found on stones (12 species, 9.2%), wood (11 species, 8.5%). Lichenicolous fungi were represented by 4 species. Some species were recorded also on mosses, iron, fruit bodies of polypores, and epiphytic algae. Terricolous lichens were not observed during the last investigation, though several species were recorded before by Kataeva (2005).
We found four species, previously unknown in St. Petersburg and the Leningrad Region: Lecania olivacella, which is also new to Russia, Bacidinapycnidiata, Fellhanera bouteillei, and Lecidella subviridis, new to North-Western European Russia. However, these new records were important, but not the main result of the study, which brought us additionally two "discoveries".
One "discovery" concerns a species which had not been recorded in St. Petersburg for a long time: surprisingly, we found a small thallus of Leptogium saturninum on bark of very old willow tree in swampy willow thickets with tall grasses (SA 5; Fig. 3E, 3F). This corticolous cyanolichen is one of indicator species of biologically valuable forests in North-Western European Russia (Vyyavlenie... , 2009). The species was listed in the Red Data Book of St. Petersburg as regionally extinct (Krasnaya..., 2018); its last collections within St. Petersburg were made in 1927 by M. M. Gollerbakh in Du-dergof (LE L-10634) and Pavlovsk (Rassadina, 1930; LE L-10635). Moreover, the nearest known actual localities in the Leningrad Region are remote from the city: near Mattia (Kingisepp District) 67 km from St. Petersburg, Lisinskoe (Tosno District) 28 km from St. Petersburg, and Orekhovo (Priozersk District) 34 km from St. Petersburg (unpublished authors' data). Leptogium saturninum as a cyanolichen known as very sensitive to atmospheric pollution (Richardson, Cameron, 2004), so its reappearance could reflect a decline of pollution in the city suburbs. It is not a unique case within the limits of St. Petersburg — single finds of the other sensitive species, such as Acolium inquinans (Sm.) A. Massal., Nephroma parile (Ach.) Ach., Schismatomma pericleum (Ach.) Branth et Rostr., Sclerophora pallida (Pers.) Y. J. Yao et Spooner, Scytinium teretiusculum (Wallr.) Otalora et al., are known from Oranienbaum Park in Lomonosov, as well as Sclerophora coniophaea (Norman) J. Mattsson et Middelb. from Dubki Park in Sestroretsk (Krasnaya., 2018).
Another "discovery" of our investigation was an unusual lichen community on branches, twigs, and needles of planted Picea glauca (SA 3; Fig. 3H). This community included 28 species, most interesting of them were Fellhanera bouteillei (already mentioned above as new for the region), Flavoparmelia caperata, Hypotrachyna revoluta, and Punctelia jeckeri. Three last foliose species were known from St. Petersburg in 1920s and early 1930s from old parks of the Petrodvorets District, faced to the Gulf of Finland (LE; Rassadina, 1930; Minyaev, 1936; Krasnaya..., 2018), but were not registered there later despite intensive investigations. The four species mentioned are currently known as suboceanic; around Baltic Sea, they are distributed mainly in the southwest and west, with eastern parts of their ranges lying in Lithuania, Kaliningrad Region of Russia, and Sweden (Faltynowicz, Kossowska, 2016; Motiejunaite, 2017; Himelbrant et al., 2020, 2022; Westberg et al, 2021; Printzen et al, 2022). During the previous lichen inventory in 2004, none of these species was found in Strelninsky Bereg (Kataeva, 2005), as well as Picea glauca was not recorded there in course of the comprehensive complex inventory (Strelninsky., 2005). Considering this, the trees of Picea glauca were planted not more than 18 years ago (though at least ten years ago or more, given their twisted shape). The age of the spruces now is about 35 years. Obviously, when the trees were brought from the Kaliningrad Region or some European country, their epiphytic lichens came together with phorophytes, and then survived. Surprisingly, all four "alien" lichen species did not decline, on the contrary, they actively colonized young branches. However, they were absent in the neighboring willow thickets and black alder stands, probably due to lack of favorable substrate. In our opinion, long-term successful surviving of four introduced lichen species usually occurring in the European regions with milder climate could be a possible result of modern climate change in the region of St. Petersburg and in the World. This is in a good concordance with the historical fact of early 20th century warming (1920-s and 1930-s) (Hegerl et al., 2018), when Flavoparmelia caperata, Hypotrachyna revoluta, and Punctelia jeckeri were registered in parks of the Petrodvorets District of St. Petersburg. During next decades with colder climate these species were not recorded in the region. Based on this data, we report here a successful occasional reintroduction of lichen species as the first recordable change in the regional lichen diversity due to modern climate change. The lower level of atmospheric pollution in St. Petersburg in last decades might be the other reason for the species to come back. However, at least Flavoparmelia caperata and Punctelia jeckeri are not highly sensitive to air quality, e. g., they occur on bark of trees growing along highways in the Kaliningrad Region (Himelbrant et al, 2020, 2022).
Altogether, nine lichen species red-listed in St. Petersburg (Krasnaya., 2018) were found in the study area: Anaptychia ciliaris, Arthonia helvola, Chaenotheca brachypoda, C. hispidula, Flavoparmelia caperata, Hypotrachyna revoluta, Lepto-gium saturninum, Punctelia jeckeri, and Scytinium subtile. The lichens Arthonia helvola, Chaenotheca brachypoda, C. hispidula, and Leptogium saturninum are indicator species, whereas Hypotrachyna revoluta and Punctelia jeckeri are habitat specialists
of biologically valuable forests in North-Western European Russia (Vyyavlenie..., 2009). The red-listed and indicator species were found on bark of old trees of Alnus glutinosa and Salix spp. in swampy or very wet habitats, except for the reintroduced species discussed before.
The lichen biota of Strelninsky Bereg Protected Area seems to be quite rich, including a number of red-listed and indicator species. However, the most interesting part of lichen diversity is represented by the successfully reintroduced species and the species coming back to St. Petersburg, which makes the territory extremely valuable for the monitoring studies of the lichen diversity in the future and reflects the processes taking place in the regional biota.
Acknowledgments
The authors are grateful to the Directorate of Nature Protected Areas of St. Petersburg, as well as to V. N. Khramtsov and E. A. Volkova (Komarov Botanical Institute of RAS, St. Petersburg) for support in organization of field studies. We thank G. A. Isachenko (St. Petersburg State University) for consultations on regional climate changes. The laboratory and herbarium studies were supported by the Ministry of Science and Higher Education of the Russian Federation under Agreement No. 075-15-2021-1056. Authors are grateful to anonymous reviewers for valuable comments and corrections.
References / Литература
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