According to available published, herbarium and recent inventory data at least 48 (and potentially over 80) species of lichenized and allied fungi can inhabit burnt areas and/or charred wood in Estonia. Most of these are widespread epiphytes and terrestrial lichens but the list also includes all five species considered ‘fire-dependent’ in Fennoscandia. Future studies should assess the specificity of species for charred wood, and analyse the features that make it a quality habitat for lichen species.

Kruustük, Kerttu

Lõhmus, Piret

Journals from University of Tartu

Folia Cryptog. Estonica, Fasc. 47: 37–41 (2010)INTRODUCTIONBurnt wood (trees) can be found abundantly in many regions and vegetation types all over the world where intensive burns take place, but also as solitary and partly burnt trees, stumps and other woody items in forests, mires and anthropogenic habitats. In the boreal vegeta-tion zone, wildfires have been a major natural disturbance regime historically, later comple-mented by slash-and-burn techniques of early farming (Esseen et al., 1992). In such areas, charred wood provides specific habitat for sev-eral organisms, including lichens. For taxa as some Hypocenomyce species (H. anthracophila, H. castaneocinerea, H. friesii, H. scalaris) and Hertelidea botryosa (Timdal, 1984; Esseen et al., 1992) fire is nessessary for colonization and establishment, and they have been termed ‘fire-dependent’ by Esseen et al. (1997). Other organisms (‘fire related’ sensu Esseen et al., 1997) are not restricted to but have a preference for post-fire successional stands; for example, some lecideoid lichens usually inhabit burnt ground soon after fire (Ahti, 1977).Despite the widespread occurrence of burnt areas and substrata in the boreal zone, very few studies of their lichen richness and composi-tion have been carried out. This is particularly unfortunate because several species of fire-created habitats are suspected to be threatened nowadays – fire management has dramatically reduced the frequency and extent of fires in northern Europe, including Estonia, during the last 100 years (Esseen et al., 1997; Lõhmus et al., 2004). Three of four investigations known to us (Nguyen-Xuan et al., 2000; Uotila & Kouki 2005; Ruokolainen & Salo, 2006) explore lichen communities on burnt ground. Only Johansson et al. (2006) explored also charred wood finding 32 lichen taxa eight years post fire. Altogether, these four studies list 67 taxa, mostly mac-rolichens. In contrast, the catalogue of lichen-forming and lichenicolous fungi of Fennoscandia (Santesson et al., 2004) only notes six taxa as inhabitants of burnt places (Cladonia cariosa, Peltigera didactyla) or charred wood (the four ‘fire-dependent’ taxa listed above).In Estonia, burnt wood is one of the least studied lichen substrata (Lõhmus, 2003). The published data only include nine species re-ported on burnt wood in quantitative lichen studies (Lõhmus 2003) and additional four taxa (Hypocenomyce anthracophila, H. friesii, H. scalaris, Hertelidea botryosa) mentioned in the key books of the lichen biota (Trass & Rand-lane 1994; Randlane & Saag 2004). Because the total number of such species is obviously much higher, we (1) present a preliminary list of lichens growing in burnt areas and/or on charred wood in Estonia based on available published, herbarium and recent inventory data, and (2) assess the need for future studies on fire-dependent taxa in this paper.Lichens on burnt wood in Estonia: a preliminary assessmentPiret Lõhmus & Kerttu KruustükDepartment of Botany, Institute of Ecology and Earth Sciences, University of Tartu, 40 Lai St., EE-51005 Tartu, Estonia E-mail: piret.lohmus@ut.eeAbstract: According to available published, herbarium and recent inventory data at least 48 (and potentially over 80) species of lichenized and allied fungi can inhabit burnt areas and/or charred wood in Estonia. Most of these are widespread epiphytes and terrestrial lichens but the list also includes all five species considered ‘fire-dependent’ in Fennoscandia. Future studies should assess the specificity of species for charred wood, and analyse the features that make it a quality habitat for lichen species.Kokkuvõte: Samblikud põlenud puidul Eestis: esialgne hinnangAvaldatud, hiljutiste inventuuride ja herbaariumi andmete põhjal tehti kindlaks, et põlendikke, põlenud kasvupindu ja/või söestunud puitu võib Eestis asustada vähemalt 48 (potentsiaalselt üle 80) lihheniseerunud ja neile lähedase seene liiki. Enamik neist on laialt levinud puudel ja maapinnal kasvavad samblikud, kuid nimekiri sisaldab ka viit Fennoskandias „põlengust sõltuvaks” peetavat liiki. Edasised uuringud on vajalikud, et hinnata, millised liigid vajavad kasvupinnana söestunud puitu ning selgitada omadusi, mis muudavad antud kasvupinna samblikele väärtuslikuks.38 Folia Cryptog. EstonicaMATERIALS AND METHODSTo compile the list of lichens (i.e. lichen-forming, lichenicolous and allied fungi) found from burnt substrata in Estonia, we used two datasets. First, the database of the Estonian lichenologi-cal herbarium at the University of Tartu (TU; http://elurikkus.ut.ee/, accessed December 18, 2009) was scanned using key words “põlendik” (burnt area), “söestunud” (charred), “tulease” (fireplace) and “põlenud” (burned). The result-ing habitat and substratum data were critically checked (in doubtful cases inspecting also the specimens in TU), i.e. whether the specimen had been collected from charred wood or bark and/or in a burnt area (in forests or bogs). The second dataset includes the results of recent fixed-area fixed-effort surveys in 2-ha plots in Estonia (see Lõhmus & Lõhmus, 2009 for details). The 92 plots represented, in a balanced design, 4 site types and 4 management types (old growth, mature commercial forest, clear-cut with and without retention trees). In each plot, lichens were searched and recorded from all substrata, including charred wood. Finally, the nine species noted to inhabit burnt wood by Lõhmus (2003) were also included in the list according to the raw data of that study.The nomenclature follows Randlane et al. (2008); species found from fewer than six locali-ties in Estonia are considered as ‘nationally rare’ and red listed status of the species is reported based on Randlane et al. (2008).RESULTSAltogether, 84 herbarium records and 72 original-survey records (from 15 plots) were ac-cepted for inclusion into the list. Those records represent a total of 48 lichen taxa (including 11 microlichens) found in burnt areas and/or on burnt substrata in forests and semi-open habitats in Estonia (Table 1). Seven species (Chaenotheca ferruginea, Cladonia coniocraea, C. digitata, Hypocenomyce anthracophila, H. scalaris, Micarea melaena, Trapeliopsis granu-losa) appeared frequent (5–20 records). The list includes one nationally rare species (Hypoceno-myce castaneocinerea), but no red listed taxa. Most records were from burnt stumps, snags and ground, but some were also from logs and live trees. Among tree species, records from Scots pine (Pinus sylvestris) comprised 60%; ad-ditional 34% were from unspecified tree species (possibly including deciduous trees also) and the rest were from Norway spruce (Picea abies). Based on the data from other studies of boreal burnt areas, 34 additional lichen-forming spe-cies could potentially inhabit burnt areas and/or charred wood in Estonia (see the footnote of Table 1).DISCUSSIONOur preliminary assessment shows that at least 48 lichen-forming and allied fungi can inhabit burnt substrata and/or areas in Estonia. Poten-tially (considering studies from the boreal zone) this number exceeds 80 species. Most of these are common and widespread epiphytic, epixylic and epigaeic species that inhabit forest and semi-open areas in Estonia (Randlane & Saag, 1999; Lõhmus, 2003). Therefore, charred wood is a generally suitable substratum for the colo-nization of many species after fire. This concurs with the findings by Romagni and Gries (2000) who showed that, by creating more free space and eliminating competition, severe fire may positively affect the (re-)colonization of epiphytic lichens in oak woodlands.Almost half of the species on the Estonian list (46%; especially Cladonia spp.) overlap with those known from burnt areas or charred wood in Fennoscandia (see Introduction and Table 1). Most importantly, all ‘fire-dependent’ lichens distinguished by Esseen et al. (1992) also oc-curred on charred wood in Estonia, although only two such species (H. anthracophila and H. scalaris) were frequent. Considering that the list of species obviously varies because of the small samples, it seems that lichen composition on burnt substrata does not vary considerably across northern Europe. However, the reasons of rarity of some ‘fire-dependent’ species deserve further study. The preliminary list does not allow to assess the specificity of species for charred wood in Estonia, and datasets representative of all main woody substrata should be collected for that purpose. More generally, it is unknown which features of charred wood, in addition to its being a competitor-free surface for colonization, make it a quality habitat for lichen species. One pos-sibility is that, because of its dark colour, burnt wood may create warm microhabitats necessary for thermophilous species, which obviously is 39SpeciesHabitat Burnt substrata Occurrence in other studies1Burnt areaForest (Semi)open habitat Bryoria sp. 1 tree* ACetraria aculeata (Schreb.) Fr. 1 groundChaenotheca ferruginea (Turner & Borrer) Mig. 6 stump, logChaenotheca trichialis (Ach.) Th. Fr. 1 stumpChaenothecopsis pusilla (Ach.) A. F. W. Schmidt + 1 stumpCladina arbuscula (Wallr.) Hale & W. L. Culb. 1 stump CDCladina mitis (Sandst.) Hustich 1 1 log, stump ABDCladina rangiferina (L.) Nyl. 1 1 log, stump ABDCladonia bacilliformis (Nyl.) Glück 1 1 log, stumpCladonia botrytes (K. G. Hagen) Willd. 1 1 log, stump ADCladonia cenotea (Ach.) Schaer. 1 3 log, stump BDCladonia chlorophaea (Flörke ex Sommerf.) Spreng. 1 ground DCladonia coniocraea (Flörke) Spreng. 1 2 2 log, snag*, stump ACladonia cornuta (L.) Hoffm. 1 ground BCDCladonia deformis (L.) Hoffm. 1 2 ground, stump BCDCladonia digitata (L.) Hoffm. 5 2 log, snag*, stump DCladonia fimbriata (L.) Fr. 1 1 2 log, stump Cladonia gracilis (L.) Willd. 2 1 stump ABDCladonia macilenta Hoffm. 1 snagCladonia polydactyla (Flörke) Spreng. 1 logCladonia squamosa Hoffm. 1 groundHertelidea botryosa (Fr.) Printzen & Kantvilas 4 snag, stumpHypocenomyce anthracophila (Nyl.) P. James & Gotth. Schneid. 1 13 1 snag*, stump, (tree)Hypocenomyce caradocensis (Leigth. ex Nyl.) P. James & Gotth. Schneid. 2 stump, tree*Hypocenomyce castaneocinerea (Räsänen) Timdal 1 snagHypocenomyce friesii (Ach.) P. James & Gotth. Schneid. 4 snag, treeHypocenomyce scalaris (Ach.) M. Choisy 19 1 log, snag, stump, treeHypocenomyce sp. 3 stump, treeHypogymnia farinacea Zopf 1 snagHypogymnia physodes (L.) Nyl. 1 stump ADImshaugia aleurites (Ach.) S. L. F. Meyer 1 logLecanora sp. 1 treeLepraria incana (L.) Ach. 2 stumpMicarea melaena (Nyl.) Hedl. 11 snag*, stumpMicarea prasina Fr. 1 stumpTable 1. Occurrence (no. of records) of lichen-forming and allied fungi (marked +) on burnt sub-strata and/or charred wood found at different habitats in Estonia. (Semi)open habitats include clear-cuts, retention-cuts and alvars. Substrata marked with * refer to charred bark. Other stud-ies: A – Johansson et al. (2006); B – Nguyen-Xuan et al. (2000); C – Ruokolainen & Salo (2006); D – Uotila & Kouki (2005)40 Folia Cryptog. EstonicaTable 1 (continued)SpeciesHabitat Burnt substrata Occurrence in other studies1Burnt areaForest (Semi)open habitat Ochrolechia androgyna (Hoffm.) Arnold 1 tree rootsParmeliopsis ambigua (Wulfen) Nyl. 1 1 stump DParmeliopsis hyperopta (Ach.) Arnold 1 stump DPeltigera canina (L.) Willd. 1 fine woody debrisPeltigera degenii Gyeln. 1 groundPeltigera didactyla (With.) J. R. Laundon 1 ground APeltigera lepidophora (Nyl. ex Vain.) Bitter 2 groundPeltigera neckeri Hepp ex Müll. Arg. 1 fine woody debrisPeltigera polydactylon (Neck.) Hoffm. 2 1 log, groundPeltigera praetextata (Flörke ex Sommerf.) Zopf 1 groundPeltigera rufescens (Weiss) Humb. 2 groundPlacynthiella icmalea (Ach.) Coppins & P. James 1 1 1 stump, logTrapeliopsis flexuosa (Fr.) Coppins & P. James 2 1 1 stump, log ATrapeliopsis granulosa (Hoffm.) Lumbsch 1 3 1ground, log, stump, tree BDUsnea hirta (L.) F. H. Wigg. 1 treeVulpicida pinastri (Scop.) J.-E. Mattsson & M. J. Lai 1 stump ACDmore important in northern areas. Another problem with the current studies appears to be their short time frame. ‘Fire-dependent’ Hypocenomyce species have been observed on burnt pine snags as long as nearly 300 years post burning (Esseen et al., 1992). Therefore, the first decade after burning – the main target of the studies performed – does not necessarily reflect the importance of charred wood for its specific species.ACKNOWLEDGEMENTSWe thank Asko Lõhmus, Tiina Randlane and anonymous referee for valuable corrections and comments on the manuscript. The study was supported by the Estonian Science Foun-dation (grant 7987), the Estonian Ministry of Education and Science (target-financing project 0180012s09) and the European Union through the European Regional development Fund (Cen-tre of Excellence FIBIR).REFERENCESAhti, T. 1977. Lichens of the boreal coniferous zone. In: Seaward, M. R. D. (ed.), Lichen Ecology, pp. 145–181. Academic Press, London.Esseen, P.-A., Ehnström, B., Ericson, L. & Sjöberg, K. 1992. Boreal forests – The Focal Habitats of Fennoscandia. In: Hansson, L. (ed.), Ecological Principles of Nature Conservation, pp.253–325. Elsevier Applied Sciences.1Additional species found in studies A–D and known to occur in Estonia: Baeomyces carneus Flörke (study D), Caloplaca holocarpa (Hoffm. ex Ach.) A. E. Wade (A), Candelariella xanthostigma (Ach.) Lettau (A), Catillaria nigroclavata (Nyl.) Schuler (A), Cetraria islandica (L.) Ach. (C), Cladina stellaris (Opiz) Brodo (CD), Cladonia borealis S. Stenroos (D), C. carneola (Fr.) Vain. (D), C. cervicornis (Ach.) Flot. (D), C. coccifera (L.) Willd. (D), C. crispata (Ach.) Flot. (BD), C. furcata (Huds.) Schrad. (D), C. grayi G. Merr. ex Sandst. (D), C. macrophylla (Schaer.) Stenh. (D), C. phyllophora Hoffm. (D), C. pleurota (Flörke) Schaer. (D), C. pyxidata (L.) Hoffm. (D), C. sulphurina (Michx.) Fr. (D), C. turgida Hoffm. (D), C. uncialis (L.) F. H. Wigg. (D), Evernia mesomorpha Nyl. (A), Flavoparmelia caperata (L.) Hale (A), Melanelia subaurifera (Nyl.) Essl. (A), Micarea misella (Nyl.) Hedl. (A), Nephroma arcticum (L.) Torss. (D), Ochrolechia arborea (Kreyer) Almb. (A), Parmelia sulcata Taylor (A), Peltigera neopolydactyla (Gyeln.) Gyeln. (A), Physcia adscendens (Fr.) H. Olivier (A), Placynthiella dasaea (Stirt.) Tonsberg (A), Platismatia glauca (L.) W. L. Culb. & C. F. Culb. (D), Pseudevernia furfuracea (L.) Zopf (D), Scoliciosporum chlorococcum (Stenh.) Vezda (A), Stereocaulon paschale (L.) Hoffm. (D).41Esseen, P.-A., Ehnström, B., Ericson, L. & Sjöberg, K. 1997. Boreal forests. Ecol. Bull. 46: 16–47.Johansson, P., Wetmore, C.M.,  Carlson, D.J., Reich, P.B. & Thor, G. 2006. Habitat preference, growth form, vegetative dispersal and population size of lichens along a wildfire severity gradient. Bryolo-gist 104: 527–540. Lõhmus, A., Kohv, K., Palo, A., & Viilma, K. 2004. Loss of old-growth, and the minimum need for strictly protected forests in Estonia. Ecol. Bull. 51: 401–411.Lõhmus, P. 2003. Composition and substrata of forest lichens in Estonia: a meta- analysis. Folia Cryptog. Estonica 40: 19–38.Lõhmus, P. & Lõhmus, A. 2009. The importance of representative inventories for lichen conservation assessments: the case of Cladonia norvegica and C. parasitica. Lichenologist 41: 61–67.Nguyen-Xuan, T., Bergeron, Y., Simard, D., Fyles J. W. & Pare, D. 2000. The importance of forest floor disturbance in the early regeneration patterns of the boreal forest of western and central Quebec: a wildfire versus logging comparison. Can. J. For. Res. 30: 1353–1364.Randlane, T. & Saag, A. (eds) 1999. Second checklist of lichenized, lichenicolous and allied fungi of Estonia. Folia Cryptog. Estonica 35: 1–132.Randlane, T. & Saag, A. (eds) 2004. Eesti pisisamb-likud (Microlichens of Estonia).  Tartu Ülikooli Kirjastus, Tartu. 582 pp. (In Estonian).  Randlane, T., Saag, A. & Suija, A. 2008. Lichenized, lichenicolous and allied fungi of Estonia. – http://www.ut.ee/ial5/lich/e_liigid/samblik_e_2008.htmlRandlane, T., Jüriado, I., Suija, A., Lõhmus, P. & Leppik, E. 2008. Lichens in the new Red List of Estonia. Folia Cryptog. Estonica 44: 113–120.Romagni, J. & Gries, C. 2000. Post-fire recoloniza-tion of dominant epiphytic lichen species on Quercus hypoleucoides (Fagaceae). Am. J. Bot. 87: 1815–1820.Ruokolainen, L. & Salo, K. 2006. The succession of boreal forest vegetation during ten years after slash-burning in Koli National Park, eastern Finland. Ann. Bot. Fenn. 43: 363–378.Santesson, R., Moberg, R., Nordin, A., Tønsberg, T. & Vitikainen, O. 2004. Lichen-forming and Li-chenicolous Fungi of Fennoscandia. Museum of Evolution, Uppsala University, Uppsala. 359 pp.Timdal, E. 1984. The genus Hypocenomyce (Lecano-rales, Lecideaceae) with special emphasis on the Norwegian and Swedish species. Nord. J. Bot. 4: 83–108.Trass, H. & Randlane, T. (eds) 1994. Eesti suursamb-likud (Macrolichens of Estonia). Tartu. 399 pp. (In Estonian).Uotila, A. & Kouki, J. 2005. Understorey vegetation in spruce-dominated forests in eastern Finland and Russian Karelia: Successional patterns after anthropogenic and natural disturbances. Forest Ecol. Manag. 215: 113–137.42 Folia Cryptog. Estonica

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Lichens on burnt wood in Estonia: a preliminary assessment

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Lichens on burnt wood in Estonia: a preliminary assessment

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