lichens and their substrate preferences on the pakri

14
Folia Cryptog. Estonica, Fasc. 48: 45–58 (2011) INTRODUCTION The Pakri Peninsula is situated in the northwest- ern part of the Estonian mainland, Harjumaa County, between the Lahepere and Paldiski Bays of the Baltic Sea (59°19–23’N 24°02–10’E) (Fig. 1). The length of the peninsula is 12 km, the width is 5 km, and the area in total is c. 40 km 2 . The climate of the Pakri Peninsula resembles that of the western and northwestern coasts of Estonia and the climate of our islands: the aver- age annual temperature is 5,9 °C; the coldest month is February, with the average tempera- ture of -3,8 °C, and the average temperature of the warmest month July is 16,6 °C. The mean precipitation is 567 mm, dropping to 356 mm in the warm period (from April to October). The average annual sunshine duration is 1800 hours, and the snow cover duration amounts to 90–100 days (EMHI, Kliima). The Pakri Peninsula is a plateau which is bordered by the Ordovician and Cambrian limestone outcrop. The highest part of the lime- stone bank on the Pakri Cape is 24,8 m above the sea level. The relief of the peninsula is flat, with some ice-edge formations, like ridges and Lichens and their substrate preferences on the Pakri Peninsula (Northwest Estonia) Ljudmilla Martin 1 , Tiina Randlane 2 & Jüri Martin 1 1 Euroacademy, Mustamäe tee 4, 10621 Tallinn, Estonia. E-mail: [email protected] 2 Institute of Ecology and Earth Sciences, University of Tartu, Lai 38/40, 51005 Tartu, Estonia. E-mail: [email protected] Abstract: A checklist of lichenized and lichenicolous fungi of the Pakri Peninsula is presented and an overview of substrate preferences of recorded taxa is compiled. e list includes 243 species, eight of them appear new to Estonia: Catillaria ato- marioides, Lecanora persimilis, Micarea lithinella, Opegrapha calcarea, Staurothele rupifraga, elidium olivaceum, T. papulare and Verrucaria hochstetteri. Verrucaria maculiformis, which was considered extinct in Estonia, is re-found, and Staurothele caesia, which has earlier been recorded as dubious for Estonia, is verified hereby. e greatest diversity of species is recorded among epilithic taxa, altogether with 121 species of which 63 species occurred on calcareous stones, 60 on granite, 18 on concrete and 12 on sandstone. Kokkuvõte: Pakri poolsaare (Loode-Eesti) samblikud ja nende substraadieelistused Esitatakse Pakri poolsaare lihheniseerunud ja lihhenikoolsete seente nimekiri ning käsitletakse nende liikide substraadieelistusi. Nimekiri sisaldab 243 liiki, neist kaheksa (Catillaria atomarioides, Lecanora persimilis, Micarea lithinella, Opegrapha calcarea, Staurothele rupifraga, elidium olivaceum, T. papulare ja Verrucaria hochstetteri) on Eestile uued. Üks samblik (Verrucaria maculiformis), mida varem peeti Eestis tõenäoliselt hävinuks, on taasleitud ning teise, eelnevalt kaheldavalt määratud liigi (Staurothele caesia) leidumine siin on nüüd tõestatud. Samblike suurim liigirikkus tuvastati epiliitide hulgas – kokku leiti erinevatelt kivisubstraatidelt 121 liiki, neist 63 liiki lubjakivilt, 60 graniidilt, 18 betoonilt and 12 liivakivilt. Fig. 1. Location of the study area. TALLINN

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Folia Cryptog. Estonica, Fasc. 48: 45–58 (2011)

INTRODUCTION

The Pakri Peninsula is situated in the northwest-ern part of the Estonian mainland, Harjumaa County, between the Lahepere and Paldiski Bays of the Baltic Sea (59°19–23’N 24°02–10’E) (Fig. 1). The length of the peninsula is 12 km, the width is 5 km, and the area in total is c. 40 km2.The climate of the Pakri Peninsula resembles that of the western and northwestern coasts of Estonia and the climate of our islands: the aver-age annual temperature is 5,9 °C; the coldest month is February, with the average tempera-ture of -3,8 °C, and the average temperature of the warmest month July is 16,6 °C. The mean precipitation is 567 mm, dropping to 356 mm in the warm period (from April to October). The average annual sunshine duration is 1800 hours, and the snow cover duration amounts to 90–100 days (EMHI, Kliima).

The Pakri Peninsula is a plateau which is bordered by the Ordovician and Cambrian limestone outcrop. The highest part of the lime-stone bank on the Pakri Cape is 24,8 m above the sea level. The relief of the peninsula is flat, with some ice-edge formations, like ridges and

Lichens and their substrate preferences on the Pakri Peninsula (Northwest Estonia)

Ljudmilla Martin1, Tiina Randlane2 & Jüri Martin1

1Euroacademy, Mustamäe tee 4, 10621 Tallinn, Estonia. E-mail: [email protected]

2Institute of Ecology and Earth Sciences, University of Tartu, Lai 38/40, 51005 Tartu, Estonia. E-mail: [email protected]

Abstract: A checklist of lichenized and lichenicolous fungi of the Pakri Peninsula is presented and an overview of substrate preferences of recorded taxa is compiled. The list includes 243 species, eight of them appear new to Estonia: Catillaria ato-marioides, Lecanora persimilis, Micarea lithinella, Opegrapha calcarea, Staurothele rupifraga, Thelidium olivaceum, T. papulare and Verrucaria hochstetteri. Verrucaria maculiformis, which was considered extinct in Estonia, is re-found, and Staurothele caesia, which has earlier been recorded as dubious for Estonia, is verified hereby. The greatest diversity of species is recorded among epilithic taxa, altogether with 121 species of which 63 species occurred on calcareous stones, 60 on granite, 18 on concrete and 12 on sandstone.

Kokkuvõte: Pakri poolsaare (Loode-Eesti) samblikud ja nende substraadieelistusedEsitatakse Pakri poolsaare lihheniseerunud ja lihhenikoolsete seente nimekiri ning käsitletakse nende liikide substraadieelistusi. Nimekiri sisaldab 243 liiki, neist kaheksa (Catillaria atomarioides, Lecanora persimilis, Micarea lithinella, Opegrapha calcarea, Staurothele rupifraga, Thelidium olivaceum, T. papulare ja Verrucaria hochstetteri) on Eestile uued. Üks samblik (Verrucaria maculiformis), mida varem peeti Eestis tõenäoliselt hävinuks, on taasleitud ning teise, eelnevalt kaheldavalt määratud liigi (Staurothele caesia) leidumine siin on nüüd tõestatud. Samblike suurim liigirikkus tuvastati epiliitide hulgas – kokku leiti erinevatelt kivisubstraatidelt 121 liiki, neist 63 liiki lubjakivilt, 60 graniidilt, 18 betoonilt and 12 liivakivilt.

Fig. 1. Location of the study area.

TALLINN

46 Folia Cryptog. Estonica

moraines (Fig.2). The surface covering is mainly gravel and slag, the thickness of which ranges from some centimeters to some meters. The bedrock of the soil includes Cambrian sands and clay, sorted out gravel, clay and sands (Kink, 1996). Lithologically different types and formations of calcareous substrate are repre-sented, e.g. limestone and dolomites from Väo Formation (Lasnamäe Stage), Kõrgekalda For-mation (Uhaku Stage) and glauconite limestone from Toila Formation (Volkhov Stage) (Raukas, 1988). The Pakri Peninsula is a vivid example of the northern coast of the North-Estonian Klint (North-Estonian limestone escarpment) with its peculiar landscape.

Different vegetation types are found on the peninsula. The so-called klint forest is an unu-sual habitat for temperate broad-leaved tree spe-cies (Acer platanoides L., Fraxinus excelsior L., Tilia cordata Mill., Quercus robur L., and Padus avium Mill.) in northern Europe; it occurs as a narrow stripe on the talus slopes (Paal, 1998; Jüriado et al., 2009). Already Lippmaa (1935) singled out the area with this special vegeta-tion as “Estonia clivosa”. In the lower coastal areas alder trees (Alnus incana (L.) Moench) pre-dominate (Tamm et al., 1997). Coniferous trees, mainly Pinus sylvestris L., are found in small numbers on the eastern coast, on drifted sand. Alvar forests represent one of the rarest forest types in Europe – their distribution is limited to limestone areas with very thin soil layer ranging from a couple of cm to 30 cm; they spread only in western and northwestern Estonia and southern Sweden (incl. Gotland and Öland) (Laasimer, 1965; Meier & Paal 2009). Other forest types, with the soil turning marshy with over moisture can also be found on the peninsula (Truus &

Kannukene, 1996).The wetter the soil, the wider is the extent of rich paludified forest types (e.g. Filipendula site type with Alnus glutinosa, Betula pubescens Ehrh., and Fraxinus excelsior). The vascular flora of the Pakri Peninsula includes 542 taxa, 177 out of them have been found be-fore 1970s (Jõe & Ramst, 1996; Kukk & Kull, 2005). Calciphilous plant species which prefer calcareous soil dominate in the flora. Several new species (Crambe maritima L., Dactylorhiza maculata (L.) Soó, Daphne mezereum L., Pul-satilla pratensis (L.) Mill.,) were recorded only recently, in 2009, in the Pakrineeme research area, where altogether 167 vascular plant spe-cies were registered (Martin et al., 2009).

Pakri Landscape Reserve was founded in 1998 to protect the local landscape, with its fauna and flora. The Reserve is situated in Harjumaa, on the territory of Paldiski city, con-taining predominantly the coastal areas of the Pakri Peninsula, northern parts of Suur-Pakri (Stora-Rågö) and Väike-Pakri (Lilla-Rågo) Is-lands, also, the sea between them together with Kappa and Bjargranne islets, plus the southern part of Väike-Pakri Island. The total area of the Reserve is 1450 ha (Vabariigi Valitsuse määrus 97, 1998).

During the World War 2nd and thereafter, there were great changes in the land use on the Peninsula. For more than 50 years, between 1939–1995, the peninsular territory had been closed for economic activities and civilian visits. Therefore only scarce data about the lichen biota of the Pakri Penisnula were known earlier. The aim of the present study was to compile a check-list of lichenized, lichenicolous and allied fungi of this remarkable area, and give an overview of substrate preferences of recorded taxa.

Fig. 2. Plateau of the Pakri Peninsula (left), and coastal terraces of limestone pebbles.

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MATERIAL AND METHODS

The list of taxa is based on the herbarium speci-mens which are stored in the herbaria of the following institutions: H – Finnish Museum of Natural History, University of Helsinki (Finland), ICEB – Euroacademy (Tallinn, Estonia), TAAM – Institute of Agricultural and Environmental Sciences, Estonian University of Life Sciences (Tartu, Estonia), TAM – Estonian Museum of Natural History (Tallinn, Estonia), and TU – Natural History Museum, University of Tartu (Tartu, Estonia).

The list of earlier collections from the Pakri peninsula was compiled using the database of Estonian lichens eSamba (http://www.ut.ee/ial5/lich/baasid/esamba.html) which mainly accumulates data from TU and TAM. The data-base was scanned using key words “Pakri Penin-sula” and “Paldiski”, resulting in 112 references to the herbarium specimens representing 70 species. Information of lichen specimens collect-ed by the following persons was included: Ernst Häyren (collecting performed in 1924), Inga Jüriado (1997, 2001), Eugen Niclasen (1910), Erast Parmasto (1995), Taimi Piin (1960), Silvi Pärn-Eilart (1960), Jaak Ruubel (1932), Veli Räsänen (1929), Mari Sarv (1995), and Heljo Ting (1960). Furthermore, fresh material of more than 600 lichen samples was collected by Ljudmilla Martin and Jüri Martin in 1999 in the central part of the peninsula; in 2009–2010 on its northeastern coast; and by Tatjana Shadrina in 2005 on concrete in various localities of the peninsula. The fresh material was mainly identi-fied by the first author.

The data about red-listed species originate from Randlane et al. (2008), and the data about protected species from Riigi Teataja (Vabariigi Valitsuse määrus 195, 2004; Keskkonnaministri määrus 51, 2004). A rare species is defined here as a species with ten or less recorded localities in Estonia (Randlane & Saag, 1999); the infor-mation about rarity comes from the databases eSamba (http://www.ut.ee/ial5/lich/baasid/esamba.html) and eElurikkus (http://elurikkus.ut.ee/). The nomenclature follows Randlane et al. (2009).

RESULTS AND DISCUSSION

The list of taxa

In total 243 species of lichenized and licheni-colous fungi are listed on the Pakri Peninsula

(Appendix), eight of them appear new to Estonia. Among recorded taxa, 233 are lichenized and 10 belong to the group of lichenicolous fungi. One species which was considered regionally extinct is re-found, and another species which has earlier been recorded as dubious in Estonia is verified hereby.

The oldest specimens available from this area were collected by Niclasen in 1910 (one species – Physcia tenella (Scop.) DC.), by Häyren in 1924 (two species) and by Räsänen in 1929 (25 species); Räsänen identified also the samples collected by Ruubel in 1932. During the whole soviet period, a few specimens were collected by Piin, Pärn-Eilart and Ting in 1960 close to the road to Paldiski city; only in 1990ies the penin-sula was re-opened for civil public. The following taxa are recorded according to the old collections exclusively, and were not re-found during recent field work: Arctoparmelia incurva (Pers.) Hale, Cetraria ericetorum Opiz, Diploschistes musco-rum (Scop.) R. Sant., Flavocetraria nivalis (L.) Kärnefelt & A. Thell, Gyalecta jenensis (Batch) Zahlbr., Phaeophyscia ciliata (Hoffm.) Moberg, Schaereria fuscocinerea (Nyl.) Clauzade & Cl. Roux, Solorina saccata (L.) Ach., Staurothele frustulenta Vain., Umbilicaria torrefacta (Lightf.) Schrad., and Usnea hirta (L.) F.H. Wigg. (Appen-dix). Among them, three species (Arctoparmelia incurva, Flavocetraria nivalis and Solorina sac-cata) are assigned to the red-listed categories in the recent Red List of Estonia (Randlane et al., 2008) and two species (Gyalecta jenensis and Staurothele frustulenta) appear rare in Estonia (Randlane & Saag, 1999).

Species new to Estonia

Catillaria atomarioides (Müll. Arg.) Kilias – Harjumaa Co., Pakri Peninsula, 59°23’03’’N 24°04’11’’E, on a granite boulder on the marine terrace, leg. Ljudmilla & Jüri Martin 10.07.2009 (ICEB). The species is characterized by a dark brown or greenish black true exciple, and it differs from the similar C. chalybeia (Borrer) A. Massal. by the colourless hypothecium. On the British Isles it is frequent on siliceous rocks, but has earlier been much overlooked (Smith et al., 2009). In Fennoscandia it is recorded in Norway and Sweden (Santesson et al., 2004).

leCanora persimilis (Th. Fr.) Nyl. – Harjumaa Co., Pakri Peninsula, 59°23’02’’N 24°04’11’’E, on dry twigs of Rosa sp. in a coastal meadow, leg. Ljudmilla & Jüri Martin 08.07.2009 (ICEB). A

48 Folia Cryptog. Estonica

member of the Lecanora dispersa group, being most similar to L. sambuci (Pers.) Nyl. which differs in having 12 or more spored asci, or to L. hagenii (Ach.) Ach. which often has pruinose disc and white or grey lecanorine apothecial margin. L. persimilis has epruinose disc, often brown-ish biatorine apothecial margin and 8 spores in asci. The taxon is common on branches and twigs of deciduous trees and shrubs with a neutral bark (Śliwa, 2007). In Fennoscandia it is recorded in Norway and Sweden (Santesson et al., 2004); in the Baltic states – in Lithuania (Motiejunaite, 2011).

miCarea lithinella (Nyl.) Hedl. – Harjumaa Co., Pakri Peninsula, 59°22’59’’N 24°04’20’’E, on the side surface of granite gravel in the old quarry on the plateau, leg. Ljudmilla & Jüri Martin 22.08.2009 (ICEB). The species is considered a pioneer colonizer of shaded stones, widespread but overlooked (Smith et al., 2009). In Fennos-candia it is recorded in Finland, Norway and Sweden (Santesson et al., 2004), in the Baltic states – in Lithuania (Motiejunaite, 2011).

opegrapha CalCarea Turn. ex Sm. – Harjumaa Co., Pakri Peninsula, 59°23’07’’N 24°04’35’’E, on limestone gravel on the sea terrace, leg. Ljudmilla & Jüri Martin 08.07.2009 (two speci-mens, ICEB). Another saxicolous Opegrapha species, O. rupestris Pers., which has earlier been recorded in Estonia, differs by K+ brown-red reaction on the exciple while O. calcarea has K+ olive-green exciple reaction (Smith et al., 2009). In Fennoscandia it is known in Norway and Sweden (Santesson et al., 2004).staurothele rupifraga (A. Massal.) Arnold – Harjumaa Co., Pakri Peninsula, 59°23’06’’N 24°04’23’’E, on limestone gravel on the sea ter-race, leg. Ljudmilla & Jüri Martin 08.07.2009 (four specimens, ICEB); 59°22’47’’N 24°05’11’’E, on a pile of limestone gravel on the forest edge, on the edge of the plateau, leg. Ljudmilla & Jüri Martin 29.06.2009 (two specimens, ICEB). The species is easily recognized by the perithecia im-mersed in pits of the substrate, by 4(-5)-spored asci, and by dark brown, often opaque as-cospores (Orange, 2008; Smith et al., 2009). In Fennoscandia it is recorded in Finland (Pykälä, 2010b), Norway and Sweden (Santesson et al., 2004).

thelidium olivaCeum (Fr.) Körb. – Harjumaa Co., Pakri Peninsula, 59°22’47’’N 24°05’11’’E, on

a pile of limestone gravel on the forest edge, on the edge of the plateau, leg. Ljudmilla & Jüri Martin 29.06.2009, det. Juha Pykälä 15.10.2010 (ICEB). The species is characterized by dark olive-brown diffuse to determinate thal-lus, relatively small semi-immersed to almost superficial perithecia (0.2–0.3 mm in diam.), and black involucrellum, contiguous with the exciple. Ascospores are (0–)1-septate, ellipsoidal to subcylindrical, 15–25 × 6–12 μm (Pykälä, 2007; McCarthy, 2011). In Fennoscandia it is recorded in Finland (Pykälä, 2007), and in the Baltic states – in Latvia (Piterans, 2001); known also in Denmark (Søchting & Alstrup, 2008).thelidium papulare (Fr.) Arnold – Harjumaa Co., Pakri Peninsula, 59°22’47’’N 24°05’11’’E, on a pile of limestone gravel on the forest edge, on the edge of the plateau, leg. Ljudmilla & Jüri Martin 29.06.2009 (two specimens, ICEB). Character-two specimens, ICEB). Character- specimens, ICEB). Character-ized by the relatively large perithecia (0.4–0.8 mm in diam.), well-developed involucrellum, and 3-(5)-septate, large (36–50 × 14–19 μm) ascospores. The species is often found on wet calcareous rocks (Orange, 2008; Smith et al., 2009). In Fennoscandia it is recorded in Finland (Pykälä, 2010a), Norway and Sweden (Santesson et al., 2004).

verruCaria hoChstetteri Fr. – Harjumaa Co., Pakri Peninsula, 59°23’07’’N 24°04’35’’E, on li-59°23’07’’N 24°04’35’’E, on li-mestone gravel on the sea terrace, leg. Ljudmilla & Jüri Martin 08.07.2009 (ICEB); 59°22’47’’N 24°05’11’’E, on a pile of limestone gravel on the forest edge, on the edge of the plateau, leg. Ljudmilla & Jüri Martin 29.06.2009 (ICEB). The species is characterized by immersed thal-lus, large perithecia sunken in pits, absence of involucrellum and large (29–35 × 16–20 μm) ascospores (Orange, 2008; Smith et al., 2009). In Fennoscandia it is recorded in Finland (Pykälä 2010a), Norway and Sweden (Santesson et al., 2004).

Protected, red-listed and rare species

Out of 243 species of lichenized and licheni-colous fungi recorded on the Pakri Peninsula (Appendix), 11 species are included in the so-called red-listed categories of the latest Red List of Estonia (Randlane et al., 2008): one species (Arctoparmelia incurva) belongs to the category Endangered; four (Arthonia lapidicola (Taylor) Branth. & Rostr., Leptogium schraderi (Bernh.) Nyl., L. subtile (Schrad.) Torss., Opegrapha

49

rupestris) – to the category Vulnerable; three (Cladonia pocillum (Ach.) Grognot, Flavocetraria nivalis, Solorina saccata) – to the category Near Threatened; two (Leptogium gelatinosum (With.) J.R. Laundon, Melanelia sorediata (Ach.) Goward & Ahti) – to the category Data Deficient; and one species (Verrucaria maculiformis Kremp.) belonged to the category Regionally Extinct, be-ing thus re-found hereby. One of the red-listed lichens, Solorina saccata, is assigned to the legally protected species in Estonia – to the pro-tection category II (Vabariigi Valitsuse määrus nr 195, 2004). About half of the recorded taxa, 127 species, are placed in the Red List category Least Concerned, while 105 species have not been evaluated against the criteria proposed by IUCN guidelines for the system of red list categories yet. Among the not-evaluated taxa, however, many species appear rare in Estonia; therefore the estimation of ‘rare’ versus ‘com-mon’ taxa was performed (Randlane & Saag, 1999. The number of rare taxa (70 species), with ten or less recorded localities in Estonia, forms almost one third of all recorded species on the peninsula) (Appendix). One of the rare lichens, Staurothele caesia (Arnold) Arnold, has earlier been recorded as dubious in Estonia (Randlane & Saag, 2004) as the according herbarium ma-terial had not been studied, and the taxon was included in the list on literature data only (Brut-tan, 1870). Now the occurrence of this species in Estonia is verified.

Lichens on different substrates

The following main substrate groups were considered: deciduous trees, coniferous trees, lignum, calcareous stone, concrete, granite, sandstone, soil, and lichens (Fig. 3).

The greatest diversity of species is recorded among epilithic taxa, altogether with 121 spe-cies of which 63 species occurred on calcareous stones, 60 on granite, 18 on concrete and 12 on sandstone. Calcareous substrate includes rock fragments and pebbles which occur in three different ecotopes on the Pakri peninsula: (1) limestone pebbles and rocks on the plateau and in quarries, (2) limestone outcrops on the slope of the klint under the canopy of deciduous trees, and (3) limestone pebbles of coastal terraces. The first ecotope is the most rich in species, with Arthonia lapidicola, Aspicilia moenium (Vain.) G. Thor & Timdal, Lecanora dispersa (Pers.)

Sommerf., Lecidella stigmatea (Ach.) Hertel & Leuckert and Verrucaria muralis Ach. dominat-ing on pebbles and rocks on the plateau and in quarries. Several common lichens, e.g. Aspicilia calcarea (L.) Mudd, Caloplaca citrina (Hoffm.) Th. Fr., Lecanora dispersa, Rinodina immersa (Körb.) Arnold etc. are abundant on limestone pebbles of the coastal terraces. Some species are adapted to the specific habitats, e.g. Acarospora heppii (Nägeli ex Hepp) Nägeli was found in the pits and fissures of the unstable limestone pebbles on the marine terraces close to the coastline while Gyalecta subclausa Anzi was found in the localities outside the direct influence of the sea, on the lower surface of loose limestone rocks, or on the surface covered with mosses or other stones. Lichen diversity was separately recorded on concrete, an artificial substrate which is most similar to limestone by its chemical and physi-cal properties. On the Pakri peninsula, many military structures of concrete were built during the last 100 years, and debris and fragments of these are still exposed. The species richness of lichens is smaller on concrete compared with limestone, and only common species (e.g. As-picilia moenium, Caloplaca citrina, C. decipiens (Arnold) Blomb. & Forssell, Candelariella aurella (Hoffm.) Zahlbr., Lecanora albescens (Hoffm.) Branth & Rostr., Protoblastenia rupestris (Scop.) J. Steiner etc.) were listed on this substrate. Granite substrate includes erratic boulders and pebbles (including crystalline pebbles of granite, gneiss and schist). Acarospora fuscata (Nyl.) Arnold, Amandinea punctata (Hoffm.) Coppins & Scheid., Aspicilia cinerea (L.) Körb., Cande-

Fig. 3. Distribution of lichenized and licheni-colous fungi recorded on the Pakri Peninsula between the main substrate groups.

no of taxa

50 Folia Cryptog. Estonica

lariella vitellina (Hoffm.) Müll. Arg., Catillaria chalybeia (Borrer) A. Massal., Lecanora rupicola (L.) Zahlbr., Lecidea lapicida var. pantherina Ach., Parmelia saxatilis (L.) Ach., and Xanthopar-melia conspersa (Ach.) Hale are common taxa on granite.The species composition of granite boulders located on the plateau and on marine terraces differs with regard to rare species. For example, Acarospora smaragdula (Wahlenb.) A. Massal., Ramalina subfarinacea (Nyl. ex Cromb.) Nyl., and Sarcogyne privigna (Ach.) A. Massal. grow on the boulders on the plateau while Amandinea cacuminum (Th. Fr.) H. Mayrhofer & Sheard, Buellia aethalea (Ach.) Th. Fr., and Pseudosagedia chlorotica (Ach.) Hafellner & Kalb were found on the boulders on marine terraces. Lecidella scabra (Taylor) Hertel & Leuckert was recorded only on the pebbles of sandstone on marine terraces.

Epiphytic lichen biota includes 81 species on the Pakri peninsula. One of the main factors, affecting the occurrence of epiphytic lichen species, is the acidity of the phorophyte bark. Coniferous trees (and birch) have normally acid and deciduous trees have subneutral bark in unpolluted areas (Wirth, 1995). In the study area coniferous trees are represented by two species, Pinus sylvestris (hereafter ‘pine’) and Juniperus communis L. Pines grow here in two separate localities: on the plateau in south-eastern part of the peninsula, and on the coastal sandy plain in the north-eastern part of the peninsula. Altogether 31 lichen species were recorded on pines. Common acidic and acidophilic lichens, e.g. Hypogymnia tubulosa (Schaer.) Hav., Par-meliopsis ambigua (Wulfen) Nyl., Platismatia glauca (L.) W.L. Culb. & C.F. Culb., species of Bryoria and Usnea are absent on the Pakri pen-insula (U. hirta was found once in 1929). Buellia schaereri De Not., Dimerella pineti (Ach.) Vězda, Hypogymnia physodes (L.) Nyl., Lecanora sym-micta (Ach.) Ach. and Parmelia sulcata Taylor are common here on pine trunks; these species have wider amplitude to acidity of substrate (Wirth, 1995). Subneutrophilic and neutrophilic species Lecidella elaeochroma (Ach.) M. Choisy, Physcia adscendens (Fr.) H. Olivier, P. tenella, Ramalina fastigiata (Pers.) Ach., and Xanthoria parietina (L.) Th. Fr. were recorded on pines on the plateau, indicating a change in the chemical composition and properties of the tree bark. On the pines growing on coastal plain, on the con-

trary, typical species of acidic bark, e.g. Micarea prasina Fr., Ochrolechia pallescens (L.) A. Mas-sal., Scoliciosporum chlorococcum (Stenh.) Vězda and Vulpicida pinastri (Scop.) J.-E. Mattsson & M.J. Lai, were found. 63 species of epiphytic lichens were recorded on the phorophytes with neutral or subneutral bark. The greatest diver-sity of species occurred on Fraxinus excelsior growing on the plateau (27), and on Acer plata-noides growing on the coastal plain (22). Leca-nia naegelii (Hepp) Diederich & Van den Boom, Lecanora carpinea (L.) Vain., L. rugosella Zahlbr., Lecidella elaeochroma, L. euphorea (Flörke) Her-tel, species of the genera Physcia, Physconia, and Xanthoria were abundant on different deciduous trees and shrubs. Anaptychia ciliaris (L.) Körb. and Ramalina fraxinea (L.) Ach. grew only on Fraxinus excelsior on the plateau.

Thirty four species were recorded among epigeic lichens. Species of Cetraria, Cladina, Cladonia, Peltigera and Stereocaulon grow on the coastal plains in areas with sandy soil, among the dis-persed graminaceous vegetation. Collema tenax (Sw.) Ach., Leptogium lichenoides (L.) Zahlbr., L. schraderi, L. subtile, Mycobilimbia species, Rino-dina conradii Körb., Peltigera rufescens (Weiss) Humb. and Verrucaria xyloxena Norman were found on the plateau, on the calcareous alvar soil among mosses.

ACKNOWLEDGEMENTS

We are grateful to Olle Hints (Institute of Geol-ogy, Tallinn University of Technology) for the assistance in determination the rock samples; to Tatjana Shadrina, MSc, for the permission to use her data about lichens on concrete, and to Elena Andrianova, MSc (both from Euroa-cademy), for the help in collecting and sorting material. Authors thank Juha Pykälä (Finnish Environmental Institute) for identifying some Thelidium and Verrucaria samples. The research was partly financed by Euroacademy grant EU KK d 01, and Baltic Gas, Ltd., contract on stra-tegic environmental assessment.

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52 Folia Cryptog. Estonica

Num

ber o

f spe

cies

on

diffe

rent

subs

trat

es22

136

376

731

65

419

92

1758

923

4612

1810

34Ab

roth

allu

s cae

rules

cens

rN

EA

Acar

ospo

ra fu

scata

fqN

EAB

Acar

ospo

ra g

lauc

ocar

pafq

NE

ABAc

aros

pora

hep

pii

rN

EB

Acar

ospo

ra sm

arag

dula

rN

EA

Acar

ospo

ra v

eron

ensis

fqN

EB

BB

Acro

cord

ia ca

vata

fqN

EA

Acro

cord

ia g

emm

ata

fqLC

ACA

Aman

dine

a ca

cum

inum

r

NE

BAm

andi

nea

coni

ops

fqN

EB

Aman

dine

a pu

ncta

tafq

LCAB

BB

Anap

tych

ia ci

liaris

fqLC

AAr

ctopa

rmeli

a in

curv

afq

ENA

Arth

onia

disp

ersa

rN

EB

Arth

onia

gla

ucom

aria

rN

EB

Arth

onia

lapi

dico

lar

VU

AA

Arth

onia

vin

osa

fqLC

BAs

picil

ia ca

esioc

iner

ear

NE

BAs

picil

ia ca

lcare

afq

NE

ABC

Aspi

cilia

cine

rea

fqLC

BAB

Aspi

cilia

cont

orta

ssp.

cont

orta

rN

EAs

picil

ia co

ntor

ta ss

p. h

offm

anni

ana

fqLC

ABA

AAs

picil

ia m

oeni

umfq

NE

ABAB

Athe

lia a

rach

noid

eafq

LCA

Bacid

ia b

aglie

ttoan

afq

NE

A

Spec

ies o

f lic

heni

zed

and

liche

nico

lous

fung

i

Frequency in Estonia

Category of red list 2008

Subs

trat

es

trees

wood

stone

slichens

soil / plant debris

Acer platanoides

Alnus incana

Betula sp.

Fraxinus excelsior

Juniperus communis

Malus sp.

Pinus sylvestris

Populus tremula

Rosa sp.

Salix caprea

Sorbus aucuparia

Tilia cordata

Ulmus glabra

limestone pebbles

limestone rock

granite pebbles

granite boulders

sandstone pebbles

concrete

Appen

dix

. Th

e list

of

lich

eniz

ed a

nd l

ich

enic

olou

s fu

ngi

in

th

e Pakri

Pen

insu

la,

an

d t

hei

r su

bst

rate

s. A

bbre

viati

ons

of,

(a)

freq

uen

cy o

f sp

ecie

s in

E

ston

ia:

r –

rare

(1–1

0 l

ocaliti

es), f

q –

fre

qu

ent

(11 o

r m

ore

loca

liti

es); (

b)

cate

gori

es o

f th

e R

ed l

ist

of E

ston

ia:

RE

– R

egio

nally

Ext

inct

, E

N –

En

dan

-ge

red, V

U –

Vu

lner

able

, N

T –

Nea

r Th

reate

ned

, D

D –

Data

Defi

cien

t, L

C –

Lea

st C

once

rned

, N

E –

Not

Eva

luate

d;

(c) ec

otop

es: A

– p

late

au

, B

– c

oast

al

terr

ace

s, C

– s

lope

(klin

t).

53Ba

cidia

bec

khau

siifq

NE

BBa

cidia

frax

inea

fqLC

ACC

Buell

ia a

etha

lear

NE

BBu

ellia

disc

iform

isfq

LCB

Buel

lia g

riseo

vire

nsfq

LCAB

BBu

ellia

scha

erer

ifq

NE

ABC

alici

um g

lauc

ellum

fqLC

BC

alop

laca

alo

ciza

rN

EAB

Cal

opla

ca ce

rina

fqLC

AC

alop

laca

citri

nafq

LCAB

AC

alop

laca

cren

ulat

ella

rN

EA

Cal

opla

ca d

ecip

iens

fqLC

AC

AC

alop

laca

flav

orub

escen

sfq

LCA

BC

alop

laca

hol

ocar

pafq

LCA

AC

alop

laca

lacte

afq

LCAB

AC

alop

laca

saxi

cola

fqLC

AA

Cal

opla

ca v

aria

bilis

fqN

EAB

Cal

opla

ca v

itelli

nula

fqN

EB

BC

ande

larie

lla a

urell

afq

LCAB

AC

ande

larie

lla v

itelli

nafq

LCAB

ABB

Can

dela

riella

xan

thos

tigm

afq

LCA

Cat

illar

ia a

phan

ar

NE

BC

atill

aria

ato

mar

ioid

esN

ewN

EB

Cat

illar

ia ch

alyb

eiafq

NE

ABB

Cat

illar

ia le

nticu

laris

rN

EAB

Cet

raria

acu

leata

fqLC

BC

etra

ria er

iceto

rum

fqLC

AC

etra

ria is

land

ica

fqLC

ABC

haen

othe

ca fe

rrug

inea

fqLC

AC

ladi

na a

rbus

cula

fqLC

BC

ladi

na m

itis

fqLC

BC

ladi

na ra

ngife

rina

fqLC

ABC

lado

nia

ceno

tea

fqLC

BC

lado

nia

chlo

roph

aea

fqLC

BC

lado

nia

coni

ocra

eafq

LCB

Cla

doni

a co

rnut

afq

LCB

Cla

doni

a cr

ispat

afq

LCAB

Cla

doni

a di

gita

ta

fqLC

BC

lado

nia

fimbr

iata

fqLC

AC

lado

nia

furc

ata

fq

LCAB

Cla

doni

a ph

yllop

hora

fqLC

BC

lado

nia

pocil

lum

fqN

TAB

54 Folia Cryptog. Estonica

Cla

doni

a py

xida

tafq

LCAB

Cla

doni

a su

bula

tafq

LCB

Cla

doni

a su

lphu

rina

fqLC

BC

lado

nia

sym

phyc

arpi

afq

LCAB

Cla

uzad

ea m

ontic

ola

fqLC

ABA

Clio

stom

um g

riffith

iifq

NE

BCC

olle

ma

fusc

ovire

nsfq

NE

ABA

Col

lema

polyc

arpo

nr

NE

AC

ollem

a te

nax

fqN

EA

ABC

ortic

ifrag

a fu

ckeli

ir

NE

ABD

imer

ella

pine

ti fq

NE

BAB

Dip

losch

istes

mus

coru

mfq

NE

AD

iplo

tom

ma

phar

cidiu

mfq

NE

CB

Elix

ia fl

exell

ar

NE

AEn

doco

ccus

exer

rans

rN

EB

Ever

nia

prun

astri

fqLC

AB

Flav

ocet

raria

niv

alis

fqN

TA

Gra

phis

scrip

tafq

LCB

Gya

lecta

jene

nsis

rN

EC

Gya

lecta

subc

laus

ar

NE

BH

ymen

elia

carn

ulos

ar

NE

ABH

ymen

elia

epul

otica

rN

EAB

Hym

eneli

a he

tero

mor

pha

rN

EAB

Hym

eneli

a rh

odop

isr

NE

AH

ypoc

enom

yce s

cala

ris

fqLC

BH

ypog

ymni

a ph

ysod

esfq

LCAB

BA

Leca

nia

cyrt

ella

fqLC

BAB

BB

Leca

nia

fusce

llar

NE

ALe

cani

a na

egel

iifq

NE

BA

BA

BLe

cano

ra a

lbesc

ens

fqLC

ABA

Leca

nora

carp

inea

fqLC

BA

BB

AB

B

Spec

ies o

f lic

heni

zed

and

liche

nico

lous

fung

i

Frequency in Estonia

Category of red list 2008

Subs

trat

es

trees

wood

stone

s

lichens

soil / plant debris

Acer platanoides

Alnus incana

Betula sp.

Fraxinus excelsior

Juniperus communis

Malus sp.

Pinus sylvestris

Populus tremula

Rosa sp.

Salix caprea

Sorbus aucuparia

Tilia cordata

Ulmus glabra

limestone pebbles

limestone rock

granite pebbles

granite boulders

sandstone pebbles

concrete

55Le

cano

ra ch

laro

tera

fqLC

BA

BLe

cano

ra cr

enul

ata

fqN

EAB

ALe

cano

ra d

isper

safq

LCAB

ALe

cano

ra ex

palle

nsfq

LCB

CAB

BLe

cano

ra h

agen

iifq

LCA

Leca

nora

heli

copi

sfq

LCB

Leca

nora

intri

cata

fqN

EB

Leca

nora

per

prui

nosa

rN

EA

Leca

nora

per

simili

sN

ewN

EB

Leca

nora

pin

iper

dar

NE

ALe

cano

ra p

olyt

ropa

fqLC

BB

Leca

nora

pul

icar

isfq

LCB

AB

Leca

nora

rugo

sella

fqLC

BA

AA

BLe

cano

ra ru

pico

lafq

LCB

ABLe

cano

ra sa

ligna

fqLC

ALe

cano

ra su

bint

ricat

ar

NE

ALe

cano

ra su

lphu

rea

fqLC

BLe

cano

ra sy

mm

icta

fqLC

BB

BB

AAB

BB

Leca

nora

var

iafq

LCB

ALe

cano

ra x

anth

osto

ma

fqN

EAB

Lecid

ea ‘h

ypop

ta’r

NE

ALe

cidea

fusco

atra

fqN

EAB

Lecid

ea la

picid

a va

r. pa

nthe

rina

fqLC

AB

Lecid

ella

carp

athi

car

NE

BLe

cidell

a ela

eoch

rom

afq

LCC

CB

BA

AA

BA

Lecid

ella

euph

orea

fqLC

BA

ALe

cidell

a sca

bra

rN

EB

Lecid

ella

stigm

atea

fqLC

ABB

Lepr

aria

ebur

nea

fqN

EC

Lepr

aria

lobi

fican

sfq

LC

CLe

ptog

ium

gela

tinos

umr

DD

ALe

ptog

ium

imbr

icatu

mr

NE

BLe

ptog

ium

lich

enoi

des

fqLC

AA

ABLe

ptog

ium

schr

ader

ir

VU

ALe

ptog

ium

subt

iler

VU

A Li

chen

ocon

ium

erod

ens

fqN

EB

Mela

nelia

exas

pera

ta

fqLC

BM

elane

lia fu

ligin

osa

fqLC

BA

BM

elane

lia so

redi

ata

fqD

DB

Mela

nelia

suba

urife

rafq

LCA

AA

Mica

rea

deni

grat

afq

NE

BM

icare

a lit

hine

llaN

ewN

EA

56 Folia Cryptog. Estonica

Mica

rea

pras

ina

s. la

t.fq

LCB

Mue

llere

lla li

chen

icola

rN

EAB

Mue

llere

lla p

ygm

aea

rN

EB

Myc

obili

mbi

a m

icroc

arpa

rN

EA

Myc

obili

mbi

a te

tram

era

fqN

EAB

Myc

omicr

othe

lia co

nfus

ar

NE

AN

eofu

scelia

loxo

des

fqLC

BN

eofu

scelia

pul

lafq

LCA

Och

rolec

hia

palle

scens

rN

EB

Ope

grap

ha ca

lcare

aN

ewN

EB

Ope

grap

ha ru

fesce

nsfq

LCBC

Ope

grap

ha ru

pestr

isr

VU

ABO

pegr

apha

var

iafq

LCC

Ope

grap

ha v

ulga

tafq

LCB

Pach

yphi

ale f

agico

lafq

NE

BA

APa

rmeli

a sa

xatil

isfq

LCAB

Parm

elia

sulca

tafq

LCB

CB

ABB

BA

APe

ltige

ra ca

nina

fqLC

AAB

Pelti

gera

did

acty

lafq

LCA

AAB

Pelti

gera

mem

bran

acea

fqLC

BPe

ltige

ra n

ecke

rifq

LCAB

Pelti

gera

pol

ydac

tylo

nfq

LCC

APe

ltige

ra p

raet

exta

tafq

LCA

APe

ltige

ra ru

fesce

nsfq

LCA

AAB

Pert

usar

ia co

ccod

esfq

LCA

Pert

usar

ia le

iopl

aca

fqLC

BA

APh

aeoc

alici

um p

opul

neum

rN

EC

Phae

ophy

scia

cilia

tafq

LCA

Phae

ophy

scia

nigr

icans

fqLC

APh

aeop

hysci

a or

bicu

laris

fqLC

AB

AA

Phlyc

tis a

rgen

afq

LCA

AA

Spec

ies o

f lic

heni

zed

and

liche

nico

lous

fung

i

Frequency in Estonia

Category of red list 2008

Subs

trat

es

trees

wood

stone

s

lichens

soil / plant debris

Acer platanoides

Alnus incana

Betula sp.

Fraxinus excelsior

Juniperus communis

Malus sp.

Pinus sylvestris

Populus tremula

Rosa sp.

Salix caprea

Sorbus aucuparia

Tilia cordata

Ulmus glabra

limestone pebbles

limestone rock

granite pebbles

granite boulders

sandstone pebbles

concrete

57Ph

yscia

ads

cend

ens

fqLC

BAB

AA

BA

Phys

cia a

ipol

iafq

LCA

Phys

cia ca

esia

fqLC

AA

Phys

cia d

ubia

fqLC

APh

yscia

stell

aris

fqLC

BB

BPh

yscia

tene

llafq

LCAB

AA

Phys

coni

a di

stort

a fq

LCB

BAB

BPl

acyn

thiel

la ic

mal

eafq

LCB

BPl

acyn

thiu

m n

igru

mfq

LCC

ABPo

lybla

stia

albi

dar

NE

ABPo

lybla

stia

cupu

laris

rN

EA

Porp

idia

sore

dizo

des

rN

EB

Porp

idia

tube

rcul

osa

rN

EB

BPr

otob

laste

nia

calv

ar

NE

APr

otob

laste

nia

rupe

stris

fqLC

ABC

APr

otop

arm

elia

badi

afq

NE

BPr

otop

arm

eliop

sis m

ural

isfq

LCAB

Pseu

deve

rnia

furfu

race

afq

LCA

Pseu

dosa

gedi

a ch

loro

tica

rN

EB

Ram

alin

a fa

rinac

eafq

LCAB

BAB

ARa

mal

ina

fasti

giat

afq

LCAB

ARa

mal

ina

fraxi

nea

fqLC

ARa

mal

ina

subf

arin

acea

rLC

ARh

izoc

arpo

n di

stinc

tum

fqN

EB

Rhiz

ocar

pon

geog

raph

icum

fqN

EB

Rhiz

ocar

pon

gran

der

NE

BRh

izoc

arpo

n po

lycar

pum

fqN

EAB

BRh

izoc

arpo

n re

ductu

mfq

NE

BRi

nodi

na b

ischo

ffii

fqN

EAB

Rino

dina

conr

adii

rN

EB

Rino

dina

exig

uafq

LCAB

Rino

dina

imm

ersa

fqN

EAB

Rino

dina

pyr

ina

fqLC

ACB

Rino

dina

soph

odes

fqLC

BB

Sarc

ogyn

e priv

igna

rN

EA

Sarc

ogyn

e reg

ular

isfq

NE

BSc

haer

eria

fusco

ciner

eafq

NE

ASc

hism

atom

ma

peric

leum

fqN

EC

Scol

icio

spor

um c

hlor

ococ

cum

fqLC

BB

Scol

icios

poru

m u

mbr

inum

fqN

EB

ABB

Solo

rina

sacc

ata

rN

TC

Spha

erell

othe

cium

pro

pinq

uellu

mfq

NE

B

58 Folia Cryptog. Estonica

Stau

roth

ele ca

esia

rN

EB

Stau

roth

ele fr

ustu

lenta

rN

EC

Stau

roth

ele ru

pifra

gaN

ewN

EAB

Sten

ocyb

e pul

latu

lafq

LCB

Ster

eoca

ulon

tom

ento

sum

fqLC

BSy

zygo

spor

a ph

yscia

cear

umfq

NE

BTe

phro

mela

atra

fqLC

ABB

Theli

dium

dec

ipien

sr

NE

ABTh

elidi

um in

cava

tum

rN

EB

Theli

dium

min

utul

um.

rN

EA

Theli

dium

oliv

aceu

mN

ewN

EA

Theli

dium

pap

ular

eN

ewN

EAB

Trap

eliop

sis fl

exuo

safq

LCB

Trap

eliop

sis g

ranu

losa

fqLC

BU

mbi

licar

ia to

rrefa

ctafq

LCB

Usn

ea h

irta

fqLC

AVe

rruc

aria

calci

seda

fqN

EB

Verr

ucar

ia d

olos

ar

NE

ABVe

rruc

aria

fusce

llar

NE

BVe

rruc

aria

hoc

hste

tteri

New

NE

ABVe

rruc

aria

mac

ulifo

rmis

rR

EB

Verr

ucar

ia m

ural

isfq

LCAB

ABVe

rruc

aria

nig

resce

nsfq

LCAB

CVe

rruc

aria

xylo

xena

rN

EAB

Vulp

icida

pin

astri

fqLC

BB

Xant

hopa

rmeli

a co

nspe

rsafq

LCAB

Xant

hopa

rmeli

a so

mlo

ënsis

fqLC

AXa

ntho

ria ca

ndela

riafq

LCB

Xant

horia

par

ietin

a fq

LCB

ABB

AB

BA

Xant

horia

pol

ycar

pafq

LCB

BB

B

Spec

ies o

f lic

heni

zed

and

liche

nico

lous

fung

i

Frequency in Estonia

Category of red list 2008

Subs

trat

es

trees

wood

stone

s

lichens

soil / plant debris

Acer platanoides

Alnus incana

Betula sp.

Fraxinus excelsior

Juniperus communis

Malus sp.

Pinus sylvestris

Populus tremula

Rosa sp.

Salix caprea

Sorbus aucuparia

Tilia cordata

Ulmus glabra

limestone pebbles

limestone rock

granite pebbles

granite boulders

sandstone pebbles

concrete