Inga Jüriado | University of Tartu (original) (raw)

Papers by Inga Jüriado

Research paper thumbnail of Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape

Biodiversity and Conservation, 2011

ABSTRACT Changes in the forest management practices have strongly influenced the distribution of ... more ABSTRACT Changes in the forest management practices have strongly influenced the distribution of species inhabiting old-growth forests. The epiphytic woodland lichen Lobaria pulmonaria is frequently used as a model species to study the factors affecting the population biology of lichens. We sampled 252 L. pulmonaria individuals from 12 populations representing three woodland types differing in their ecological continuity and management intensity in Estonia. We used eight mycobiont-specific microsatellite loci to quantify genetic diversity among the populations. We calculated the Sørensen distance to estimate genetic dissimilarity among individuals within populations. We revealed that L. pulmonaria populations have significantly higher genetic diversity in old-growth forests than in managed forests and wooded meadows. We detected a significant woodland-type-specific pattern of genetic dissimilarity among neighbouring L. pulmonaria individuals, which suggests that in wooded meadows and managed forests dominating is vegetative reproduction. The vegetative dispersal distance between the host trees of L. pulmonaria was found to be only 15–30m. Genetic dissimilarity among individuals was also dependent on tree species and trunk diameter. Lobaria pulmonaria populations in managed forests included less juveniles compared to old-growth forests and wooded meadows, indicating that forest management influences life stage structure within populations. We conclude that as intensive stand management reduces the genetic diversity of threatened species in woodland habitats, particular attention should be paid to the preservation of remnant populations in old-growth habitats. Within managed habitats, conservation management should target on maintenance of the stand’s structural diversity and availability of potential host trees. KeywordsGenetic dissimilarity–Genetic distance–Epiphytic lichen–Microsatellites–Managed forest–Old-growth forest–Population genetics–Wooded meadow

Research paper thumbnail of Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Nordic Journal of Botany

Semi-natural grasslands, among them thin-soil calcareous grasslands (alvars), have great conserva... more Semi-natural grasslands, among them thin-soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red-listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its siz...

Research paper thumbnail of Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Biodiversity and Conservation, 2009

Tree and stand level variables affecting the species richness, cover and composition of epiphytic... more Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens.

Research paper thumbnail of Biogeographical determinants of lichen species diversity on islets in the West‐Estonian Archipelago

Journal of Vegetation Science, 2006

Questions: Do islet area, number of biotopes and distance from the mainland shape lichen species ... more Questions: Do islet area, number of biotopes and distance from the mainland shape lichen species richness on islets? Are there any species-or substrate group-specific trends associated with these factors? Location: Islets of the west Estonian Archipelago, Estonia. Methods: A species list was compiled for each of the 32 islets and the relative abundance of each species was estimated. The lichens were divided into seven groups according to their substrate preferences. Generalized linear model (GLIM) analysis was applied to test the effect of the islet traits on the number of lichen species on the islets and in the substrate groups. The probability of presence/absence and abundance of the most frequent species according to the islet traits were tested with GLIM and general linear mixed model.

Research paper thumbnail of Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions

The Lichenologist, 2009

The species richness and composition of lichens was quantified for four temperate broad-leaved tr... more The species richness and composition of lichens was quantified for four temperate broad-leaved tree species (Acer platanoides, Fraxinus excelsior, Tilia cordata and Ulmus glabra) in boreo-nemoral forests on the talus slope of the North-Estonian limestone escarpment (North-Estonian Klint). Thirteen study sites were distributed along the klint on a west to east gradient. The effects of tree and stand characteristics and geographical location of a stand on composition and diversity of epiphytic lichens were evaluated by multivariate analyses (DCA, CCA, pCCA) and by general linear mixed models (GLMM). Tree level variables (e.g. bark pH, bryophytes cover and host tree species) explained the largest fraction of the variation in lichen species composition. However, species richness and composition were significantly influenced also by the unique habitat conditions of klint forest (length of the forest fragment, proximity of the stand to the sea and height of the escarpment). A significant correlation between stand locality and lichen diversity on the tree bole was found which is most likely related to local air pollution gradients caused by alkaline cement dust and acid pollutants in the north-eastern part of Estonia.

Research paper thumbnail of Influence of tree stand age and management on the species diversity in Estonian eutrophic alvar and boreo-nemoral Pinus sylvestris forests

Scandinavian Journal of Forest Research, 2005

Research paper thumbnail of Epiphytic and epixylic species diversity in Estonian natural forests

Biodiversity and Conservation

Research paper thumbnail of Epiphytic and epixylic lichen species diversity in Estonian natural forests

Biodiversity and Conservation, 2003

Research paper thumbnail of Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape

Biodiversity and Conservation, 2011

ABSTRACT Changes in the forest management practices have strongly influenced the distribution of ... more ABSTRACT Changes in the forest management practices have strongly influenced the distribution of species inhabiting old-growth forests. The epiphytic woodland lichen Lobaria pulmonaria is frequently used as a model species to study the factors affecting the population biology of lichens. We sampled 252 L. pulmonaria individuals from 12 populations representing three woodland types differing in their ecological continuity and management intensity in Estonia. We used eight mycobiont-specific microsatellite loci to quantify genetic diversity among the populations. We calculated the Sørensen distance to estimate genetic dissimilarity among individuals within populations. We revealed that L. pulmonaria populations have significantly higher genetic diversity in old-growth forests than in managed forests and wooded meadows. We detected a significant woodland-type-specific pattern of genetic dissimilarity among neighbouring L. pulmonaria individuals, which suggests that in wooded meadows and managed forests dominating is vegetative reproduction. The vegetative dispersal distance between the host trees of L. pulmonaria was found to be only 15–30m. Genetic dissimilarity among individuals was also dependent on tree species and trunk diameter. Lobaria pulmonaria populations in managed forests included less juveniles compared to old-growth forests and wooded meadows, indicating that forest management influences life stage structure within populations. We conclude that as intensive stand management reduces the genetic diversity of threatened species in woodland habitats, particular attention should be paid to the preservation of remnant populations in old-growth habitats. Within managed habitats, conservation management should target on maintenance of the stand’s structural diversity and availability of potential host trees. KeywordsGenetic dissimilarity–Genetic distance–Epiphytic lichen–Microsatellites–Managed forest–Old-growth forest–Population genetics–Wooded meadow

Research paper thumbnail of Lichens in the new Red List of Estonia

Abstract: The compilation of the current Red List of Estonia took place during 2006–2008; the IUC... more Abstract: The compilation of the current Red List of Estonia took place during 2006–2008; the IUCN system of categories
and criteria (vers. 6.1), which is accepted worldwide, was applied. Out of the 1019 lichenized, lichenicolous and closely allied
fungal species recorded in Estonia in 2006, 464 species (45.5%) were evaluated while 555 species remained not estimated
– in the category Not Evaluated (NE). Of the evaluated species, 213 were assigned to the so-called red-listed categories:
Regionally Extinct (RE), Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT) and Data
Deficient (DD). 113 of them were classified as threatened (belonging to the categories CR, EN, VU). 251 species were
assigned to the category Least Concerned (LC). The full enumeration of the red-listed lichens of Estonia with appropriate
category and criteria is presented.

Research paper thumbnail of New or interesting lichens and lichenicolous fungi found during the 5th IAL Symposium in Estonia

Abstract: Altogether 74 species of lichens and lichenicolous fungi found during IAL5 (Estonia, Au... more Abstract: Altogether 74 species of lichens and lichenicolous fungi found during IAL5 (Estonia, August 2004) are listed; 30
of them are new to the country. New lichens: Absconditella celata, A. delutula, A. sphagnorum, Bacidia adastra, B. caligans, Bacidina
chloroticula, Cladonia monomorpha, Dirina massiliensis f. sorediata, Lecanora subcarpinea, Lecidella flavosorediata, Micarea anterior, M. lutulata,
M. micrococca, Ramonia aff. nigra, Rinodina degeliana, Thelocarpon lichenicola, Trapelia involuta, Verrucaria bryoctona, V. dolosa. New
lichenicolous fungi: Arthonia digitatae, A. galactinaria, Cercidospora macrospora, Cornutispora lichenicola, Marchandiomyces aurantiacus,
Merismatium heterophractum, Phoma epiphyscia, Pronectria xanthoriae, Telogalla olivieri, Trichonectria anisospora, T. hirta.

Research paper thumbnail of Threatened forest lichen Lobaria pulmonaria – its past, present and future in Estonia

Abstract. Lobaria pulmonaria is a conspicuous and widely known epiphytic lichen species which mai... more Abstract. Lobaria pulmonaria is a conspicuous and widely known epiphytic
lichen species which mainly occurs on mature hardwood trees in old forests.
The distribution of L. pulmonaria has become highly fragmented, showing
a decline across Europe. Lobaria pulmonaria is unevenly distributed among
ca. 600 localities in Estonia. Most localities are in north-eastern and southwestern
Estonia (50% and 21% of records respectively). 86% of L. pulmonaria
localities are in nature reserves, national parks or woodland key habitats
(WKH). In 81% of historical localities known until 1992, L. pulmonaria could
be considered extinct. 6% of L. pulmonaria localities have become destroyed
in the period from 1993 to 2010 due to clear cutting; in 19% of localities the
species could be considered endangered, as its neighbourhood is affected by
forest management activities. 40% of destroyed localities are located in forests
designated as WKHs in north-eastern Estonia, and most of the endangered
populations are also within WKHs. In forest management and conservation
planning, WKHs should be preserved with surrounding buffer areas in order
to avoid abrupt changes in environmental condition and natural disturbance
regimes. There is also a necessity to develop specific management methods
to recover reforested wooded meadows, to ensure that species of semi-open
habitats are retained, e.g. by reducing the density of spruce.

Research paper thumbnail of Distribution and habitat ecology of the threatened forest lichen Lobaria pulmonaria in Estonia

Abstract: Lobaria pulmonaria is a widely used as an indicator species of undisturbed old-growth f... more Abstract: Lobaria pulmonaria is a widely used as an indicator species of undisturbed old-growth forests ecosystems, but the knowledge about its habitat ecology is still highly fragmented. To quantify the distribution, habitat preference and host tree specificity of Lobaria pulmonaria we utilised data from databases and field surveys in Estonia. The number of L. pulmonaria localities is the highest in the densely forested regions, concentrated mainly in small forest patches defined as ‘ecologically highly valuable’. The species grows mostly on deciduous trees, particularly on aspen (Populus tremula). L. pulmonaria is most common in oligo-mesotrophic boreal, eutrophic boreo-nemoral and in eutrophic paludifying forests, and prefers forests with an average age of trees more than 100 years. In addition, we found that younger stands could be suitable habitats for L. pulmonaria if the structure of the stand is comparable to mature stands. In spite of the many localities of L. pulmonaria in Estonia, the species is still threatened because (1) the rotation period of tree stands is short, (2) it is abundant in forest types which are rare or under strong economic pressure, (3) and it prefers host trees which have a restricted distribution in Estonia or are not favoured in forest management practice.

Research paper thumbnail of Habitat conditions and host tree properties affect the occurrence, abundance and fertility of the endangered lichen Lobaria pulmonaria in wooded meadows of Estonia

Abstract: We assessed multiple environmental factors that might influence the population vitality... more Abstract: We assessed multiple environmental factors that might influence the population vitality of
the epiphytic lichen Lobaria pulmonaria at the individual tree and habitat levels in partially overgrown
wooded meadows in Estonia. A total of 301 trees of four species were sampled at nine study plots,
using a stratified factorial scheme, 151 colonized by L. pulmonaria and 150 not colonized by L.
pulmonaria forming the control group. We used the Generalized Linear Models (GLZ) to identify a
complex of factors which predicts the probability of the lichen occurring on tree trunks and the
presence of apothecia on its individuals. We employed the General Linear Mixed Model (GLMM) to
study the relationship between cover of L. pulmonaria and environmental factors. The occurrence
probability of L. pulmonaria on tree trunks increased with increasing light availability and height of
deciduous shrubs near the trunk, and decreased with increasing distance to the nearest colonized tree.
The host tree species and its trunk properties were also of importance, particularly the facilitating effect
of the cover of bryophytes upon L. pulmonaria. The probability of occurrence of apothecia increased
with maximum values of bark pH and cover of L. pulmonaria on the trunk.We conclude that partially
overgrown wooded meadows are suitable habitats for L. pulmonaria. However, to maintain the vitality
of these populations, a specificmanagement scheme, preventing development of a dense stand, should
be applied. Management requirements would include 1) selective cutting of overgrowing coniferous
trees (particularly spruce), 2) preservation of adult and younger potential host trees within 10–20 m of
colonized trees, 3) preservation of scattered deciduous shrubs in the vicinity of the host trees.

Research paper thumbnail of Changes in stand structure due to the cessation of traditional land use in wooded meadows impoverish epiphytic lichen communities

Abstract: Wooded meadows with a history of traditional land use over thousands of years support a... more Abstract: Wooded meadows with a history of traditional land use over thousands of years support a
great diversity of various taxa. Today, however, high-species-rich communities in wooded meadows
are threatened because of the cessation of traditional management in large areas. We studied lichen
communities on 136 deciduous trees (Betula spp., Fraxinus excelsior and Quercus robur) in 12 wooded
meadows in three regions of Estonia, and assessed the effect of habitat change due to the abandonment
of traditional management on epiphytic lichen species composition, considering factors on three
spatial scales: regional, habitat and individual tree. The variation partitioning approach in partial
Canonical Correspondence Analysis (pCCA) revealed that most of the variation in species composition
is described by the species of host tree and tree bark pH. Other tree level variables, foremost tree
diameter, described as much of the compositional variation as geographic location (region) or
environmental conditions in wooded meadows. Of the environmental factors studied, woodland
canopy cover is the strongest predictor of the change in epiphytic lichen species composition from the
community type of semi-open wooded meadows to species-poor communities characteristic of
secondary forest. General Linear Model (GLM) analysis of the abundance of the 35 most frequently
observed lichen species revealed that more than half of them (21) are influenced by site openness
(canopy cover and/or undergrowth density), showing that increasing canopy cover has a negative effect
on the abundance of epiphytic lichen species characteristic of traditionally managed semi-open
wooded meadows. The results emphasize that the preservation of large old deciduous trees of various
species and the maintenance of the semi-open structure of stands are vitally important for the
protection of epiphytic lichen communities in wooded meadows.

Research paper thumbnail of The conservation of ground layer lichen communities in alvar grasslands and the relevance of substitution habitats

Semi-natural calcareous grasslands (alvars) are biodiversity hotspots in Northern Europe, particu... more Semi-natural calcareous grasslands (alvars) are biodiversity hotspots in Northern Europe, particularly for herb
layer plants. In the last century, traditional management has ceased, and the area of grasslands has declined
due to extensive encroachment. We were interested in the drivers of ground layer (alias terricolous or epigeic)
lichen communities. Our survey consisted of 86 habitat fragments in western Estonia, covering four types of
historic alvar grasslands and three types of alvar-like habitats. We found that the ground lichen communities
were primarily soil-type-specific, but were also affected by historic disturbances and land use change. In
contrast to knowledge about herb layer communities, for which shrub encroachment has been shown to be
main driver, the increased density of the herb layer and the reduced diversity of microhabitats were major
drivers for the ground layer lichen community. These drivers caused a decrease in species richness, but only
within the species of conservation value, and also led to a shift in the composition of lichen growth form from
the dominance of squamulose and crustose towards fruticose lichens. We conclude that the traditional practice
of restoring alvars by cutting shrubs is insufficient to maintain ground layer lichen biodiversity. Alvar
maintenance practices should include grazing, which creates various small-scale ground disturbances and
increases microhabitat heterogeneity. Alvar-like habitats originating from large-scale historic disturbances
appeared to be suitable for calcicolous epigeic lichens, and can therefore be considered to be temporary
substitution habitats, i.e. refugia for the regional species pool.

Research paper thumbnail of Functional ecology of rare and common epigeic lichens in alvar grasslands

The autecology of rare species can be derived using similarities among functional traits and envi... more The autecology of rare species can be derived using similarities among functional traits and
environmental conditions observed for common species, i.e. we employed the ‘matching
analogy approach’ with the analytical scheme ‘common species / driver / trait /
driver/rare species’.We addressed the driveretrait relationship for common epigeic lichens
of thin-soil calcareous grasslands, which are endangered by cessation of traditional land use.
Common lichens were suppressed by encroaching herbs and shrubs, and were supported by
ground disturbances. The lichens of open low-productivity alvars are predominantly calciphilous,
epibryic, crustose-squamulose, sexually reproducing and contain UV-protective pigments.
Lichens of encroached alvars are soil-type generalists, fruticose, reproduce vegetatively
and contain herbivore-deterring compounds. Rare lichens resemble the species of open lowproductivity
alvars, except their more limited niche space, i.e. they are restricted to arcticalpine
habitats. The conservation practices on alvars should support the formation of crustforming
communities by suppressing the growth of shrubs and herbs, and by promoting
recurrent small-scale soil disturbances.

Research paper thumbnail of Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions

Lichenologist, 2009

The species richness and composition of lichens was quantified for four temperate broad-leaved tr... more The species richness and composition of lichens was quantified for four temperate broad-leaved tree species (Acer platanoides, Fraxinus excelsior, Tilia cordata and Ulmus glabra) in boreo-nemoral forests on the talus slope of the North-Estonian limestone escarpment (North-Estonian Klint). Thirteen study sites were distributed along the klint on a west to east gradient. The effects of tree and stand characteristics and geographical location of a stand on composition and diversity of epiphytic lichens were evaluated by multivariate analyses (DCA, CCA, pCCA) and by general linear mixed models (GLMM). Tree level variables (e.g. bark pH, bryophytes cover and host tree species) explained the largest fraction of the variation in lichen species composition. However, species richness and composition were significantly influenced also by the unique habitat conditions of klint forest (length of the forest fragment, proximity of the stand to the sea and height of the escarpment). A significant correlation between stand locality and lichen diversity on the tree bole was found which is most likely related to local air pollution gradients caused by alkaline cement dust and acid pollutants in the north-eastern part of Estonia.

Research paper thumbnail of Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions – ERRATUM

Research paper thumbnail of Biogeographical determinants of lichen species diversity on islets in the West-Estonian Archipelago

Journal of Vegetation Science, 2006

AbstractQuestions: Do islet area, number of biotopes and distance from the mainland shape lichen ... more AbstractQuestions: Do islet area, number of biotopes and distance from the mainland shape lichen species richness on islets? Are there any species- or substrate group-specific trends associated with these factors?Do islet area, number of biotopes and distance from the mainland shape lichen species richness on islets? Are there any species- or substrate group-specific trends associated with these factors?Location: Islets of the west Estonian Archipelago, Estonia.Islets of the west Estonian Archipelago, Estonia.Methods: A species list was compiled for each of the 32 islets and the relative abundance of each species was estimated. The lichens were divided into seven groups according to their substrate preferences. Generalized linear model (GLIM) analysis was applied to test the effect of the islet traits on the number of lichen species on the islets and in the substrate groups. The probability of presence/absence and abundance of the most frequent species according to the islet traits were tested with GLIM and general linear mixed model.A species list was compiled for each of the 32 islets and the relative abundance of each species was estimated. The lichens were divided into seven groups according to their substrate preferences. Generalized linear model (GLIM) analysis was applied to test the effect of the islet traits on the number of lichen species on the islets and in the substrate groups. The probability of presence/absence and abundance of the most frequent species according to the islet traits were tested with GLIM and general linear mixed model.Results: The lichen flora of the islets consisted of 326 taxa, the number of lichen species per islet varied from 2 to 197. Total number of species per islet and within the substrate groups was positively correlated with islet area and with number of biotopes, and negatively correlated with distance from the mainland; however, these relationships varied among the substrate groups. Although individual lichen species showed variation in responses, general trends in island biogeography were evident.The lichen flora of the islets consisted of 326 taxa, the number of lichen species per islet varied from 2 to 197. Total number of species per islet and within the substrate groups was positively correlated with islet area and with number of biotopes, and negatively correlated with distance from the mainland; however, these relationships varied among the substrate groups. Although individual lichen species showed variation in responses, general trends in island biogeography were evident.Conclusions: The distribution pattern of lichens on the studied islets follows the theory of island biogeography: the number of species per islet depends on isolation, area and biotope diversity. Species specific traits, such as dispersal strategy and growth form, as well as availability of a particular substrate are important for formation of the lichen flora on islets.The distribution pattern of lichens on the studied islets follows the theory of island biogeography: the number of species per islet depends on isolation, area and biotope diversity. Species specific traits, such as dispersal strategy and growth form, as well as availability of a particular substrate are important for formation of the lichen flora on islets.

Research paper thumbnail of Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape

Biodiversity and Conservation, 2011

ABSTRACT Changes in the forest management practices have strongly influenced the distribution of ... more ABSTRACT Changes in the forest management practices have strongly influenced the distribution of species inhabiting old-growth forests. The epiphytic woodland lichen Lobaria pulmonaria is frequently used as a model species to study the factors affecting the population biology of lichens. We sampled 252 L. pulmonaria individuals from 12 populations representing three woodland types differing in their ecological continuity and management intensity in Estonia. We used eight mycobiont-specific microsatellite loci to quantify genetic diversity among the populations. We calculated the Sørensen distance to estimate genetic dissimilarity among individuals within populations. We revealed that L. pulmonaria populations have significantly higher genetic diversity in old-growth forests than in managed forests and wooded meadows. We detected a significant woodland-type-specific pattern of genetic dissimilarity among neighbouring L. pulmonaria individuals, which suggests that in wooded meadows and managed forests dominating is vegetative reproduction. The vegetative dispersal distance between the host trees of L. pulmonaria was found to be only 15–30m. Genetic dissimilarity among individuals was also dependent on tree species and trunk diameter. Lobaria pulmonaria populations in managed forests included less juveniles compared to old-growth forests and wooded meadows, indicating that forest management influences life stage structure within populations. We conclude that as intensive stand management reduces the genetic diversity of threatened species in woodland habitats, particular attention should be paid to the preservation of remnant populations in old-growth habitats. Within managed habitats, conservation management should target on maintenance of the stand’s structural diversity and availability of potential host trees. KeywordsGenetic dissimilarity–Genetic distance–Epiphytic lichen–Microsatellites–Managed forest–Old-growth forest–Population genetics–Wooded meadow

Research paper thumbnail of Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Nordic Journal of Botany

Semi-natural grasslands, among them thin-soil calcareous grasslands (alvars), have great conserva... more Semi-natural grasslands, among them thin-soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red-listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its siz...

Research paper thumbnail of Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Biodiversity and Conservation, 2009

Tree and stand level variables affecting the species richness, cover and composition of epiphytic... more Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens.

Research paper thumbnail of Biogeographical determinants of lichen species diversity on islets in the West‐Estonian Archipelago

Journal of Vegetation Science, 2006

Questions: Do islet area, number of biotopes and distance from the mainland shape lichen species ... more Questions: Do islet area, number of biotopes and distance from the mainland shape lichen species richness on islets? Are there any species-or substrate group-specific trends associated with these factors? Location: Islets of the west Estonian Archipelago, Estonia. Methods: A species list was compiled for each of the 32 islets and the relative abundance of each species was estimated. The lichens were divided into seven groups according to their substrate preferences. Generalized linear model (GLIM) analysis was applied to test the effect of the islet traits on the number of lichen species on the islets and in the substrate groups. The probability of presence/absence and abundance of the most frequent species according to the islet traits were tested with GLIM and general linear mixed model.

Research paper thumbnail of Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions

The Lichenologist, 2009

The species richness and composition of lichens was quantified for four temperate broad-leaved tr... more The species richness and composition of lichens was quantified for four temperate broad-leaved tree species (Acer platanoides, Fraxinus excelsior, Tilia cordata and Ulmus glabra) in boreo-nemoral forests on the talus slope of the North-Estonian limestone escarpment (North-Estonian Klint). Thirteen study sites were distributed along the klint on a west to east gradient. The effects of tree and stand characteristics and geographical location of a stand on composition and diversity of epiphytic lichens were evaluated by multivariate analyses (DCA, CCA, pCCA) and by general linear mixed models (GLMM). Tree level variables (e.g. bark pH, bryophytes cover and host tree species) explained the largest fraction of the variation in lichen species composition. However, species richness and composition were significantly influenced also by the unique habitat conditions of klint forest (length of the forest fragment, proximity of the stand to the sea and height of the escarpment). A significant correlation between stand locality and lichen diversity on the tree bole was found which is most likely related to local air pollution gradients caused by alkaline cement dust and acid pollutants in the north-eastern part of Estonia.

Research paper thumbnail of Influence of tree stand age and management on the species diversity in Estonian eutrophic alvar and boreo-nemoral Pinus sylvestris forests

Scandinavian Journal of Forest Research, 2005

Research paper thumbnail of Epiphytic and epixylic species diversity in Estonian natural forests

Biodiversity and Conservation

Research paper thumbnail of Epiphytic and epixylic lichen species diversity in Estonian natural forests

Biodiversity and Conservation, 2003

Research paper thumbnail of Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape

Biodiversity and Conservation, 2011

ABSTRACT Changes in the forest management practices have strongly influenced the distribution of ... more ABSTRACT Changes in the forest management practices have strongly influenced the distribution of species inhabiting old-growth forests. The epiphytic woodland lichen Lobaria pulmonaria is frequently used as a model species to study the factors affecting the population biology of lichens. We sampled 252 L. pulmonaria individuals from 12 populations representing three woodland types differing in their ecological continuity and management intensity in Estonia. We used eight mycobiont-specific microsatellite loci to quantify genetic diversity among the populations. We calculated the Sørensen distance to estimate genetic dissimilarity among individuals within populations. We revealed that L. pulmonaria populations have significantly higher genetic diversity in old-growth forests than in managed forests and wooded meadows. We detected a significant woodland-type-specific pattern of genetic dissimilarity among neighbouring L. pulmonaria individuals, which suggests that in wooded meadows and managed forests dominating is vegetative reproduction. The vegetative dispersal distance between the host trees of L. pulmonaria was found to be only 15–30m. Genetic dissimilarity among individuals was also dependent on tree species and trunk diameter. Lobaria pulmonaria populations in managed forests included less juveniles compared to old-growth forests and wooded meadows, indicating that forest management influences life stage structure within populations. We conclude that as intensive stand management reduces the genetic diversity of threatened species in woodland habitats, particular attention should be paid to the preservation of remnant populations in old-growth habitats. Within managed habitats, conservation management should target on maintenance of the stand’s structural diversity and availability of potential host trees. KeywordsGenetic dissimilarity–Genetic distance–Epiphytic lichen–Microsatellites–Managed forest–Old-growth forest–Population genetics–Wooded meadow

Research paper thumbnail of Lichens in the new Red List of Estonia

Abstract: The compilation of the current Red List of Estonia took place during 2006–2008; the IUC... more Abstract: The compilation of the current Red List of Estonia took place during 2006–2008; the IUCN system of categories
and criteria (vers. 6.1), which is accepted worldwide, was applied. Out of the 1019 lichenized, lichenicolous and closely allied
fungal species recorded in Estonia in 2006, 464 species (45.5%) were evaluated while 555 species remained not estimated
– in the category Not Evaluated (NE). Of the evaluated species, 213 were assigned to the so-called red-listed categories:
Regionally Extinct (RE), Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT) and Data
Deficient (DD). 113 of them were classified as threatened (belonging to the categories CR, EN, VU). 251 species were
assigned to the category Least Concerned (LC). The full enumeration of the red-listed lichens of Estonia with appropriate
category and criteria is presented.

Research paper thumbnail of New or interesting lichens and lichenicolous fungi found during the 5th IAL Symposium in Estonia

Abstract: Altogether 74 species of lichens and lichenicolous fungi found during IAL5 (Estonia, Au... more Abstract: Altogether 74 species of lichens and lichenicolous fungi found during IAL5 (Estonia, August 2004) are listed; 30
of them are new to the country. New lichens: Absconditella celata, A. delutula, A. sphagnorum, Bacidia adastra, B. caligans, Bacidina
chloroticula, Cladonia monomorpha, Dirina massiliensis f. sorediata, Lecanora subcarpinea, Lecidella flavosorediata, Micarea anterior, M. lutulata,
M. micrococca, Ramonia aff. nigra, Rinodina degeliana, Thelocarpon lichenicola, Trapelia involuta, Verrucaria bryoctona, V. dolosa. New
lichenicolous fungi: Arthonia digitatae, A. galactinaria, Cercidospora macrospora, Cornutispora lichenicola, Marchandiomyces aurantiacus,
Merismatium heterophractum, Phoma epiphyscia, Pronectria xanthoriae, Telogalla olivieri, Trichonectria anisospora, T. hirta.

Research paper thumbnail of Threatened forest lichen Lobaria pulmonaria – its past, present and future in Estonia

Abstract. Lobaria pulmonaria is a conspicuous and widely known epiphytic lichen species which mai... more Abstract. Lobaria pulmonaria is a conspicuous and widely known epiphytic
lichen species which mainly occurs on mature hardwood trees in old forests.
The distribution of L. pulmonaria has become highly fragmented, showing
a decline across Europe. Lobaria pulmonaria is unevenly distributed among
ca. 600 localities in Estonia. Most localities are in north-eastern and southwestern
Estonia (50% and 21% of records respectively). 86% of L. pulmonaria
localities are in nature reserves, national parks or woodland key habitats
(WKH). In 81% of historical localities known until 1992, L. pulmonaria could
be considered extinct. 6% of L. pulmonaria localities have become destroyed
in the period from 1993 to 2010 due to clear cutting; in 19% of localities the
species could be considered endangered, as its neighbourhood is affected by
forest management activities. 40% of destroyed localities are located in forests
designated as WKHs in north-eastern Estonia, and most of the endangered
populations are also within WKHs. In forest management and conservation
planning, WKHs should be preserved with surrounding buffer areas in order
to avoid abrupt changes in environmental condition and natural disturbance
regimes. There is also a necessity to develop specific management methods
to recover reforested wooded meadows, to ensure that species of semi-open
habitats are retained, e.g. by reducing the density of spruce.

Research paper thumbnail of Distribution and habitat ecology of the threatened forest lichen Lobaria pulmonaria in Estonia

Abstract: Lobaria pulmonaria is a widely used as an indicator species of undisturbed old-growth f... more Abstract: Lobaria pulmonaria is a widely used as an indicator species of undisturbed old-growth forests ecosystems, but the knowledge about its habitat ecology is still highly fragmented. To quantify the distribution, habitat preference and host tree specificity of Lobaria pulmonaria we utilised data from databases and field surveys in Estonia. The number of L. pulmonaria localities is the highest in the densely forested regions, concentrated mainly in small forest patches defined as ‘ecologically highly valuable’. The species grows mostly on deciduous trees, particularly on aspen (Populus tremula). L. pulmonaria is most common in oligo-mesotrophic boreal, eutrophic boreo-nemoral and in eutrophic paludifying forests, and prefers forests with an average age of trees more than 100 years. In addition, we found that younger stands could be suitable habitats for L. pulmonaria if the structure of the stand is comparable to mature stands. In spite of the many localities of L. pulmonaria in Estonia, the species is still threatened because (1) the rotation period of tree stands is short, (2) it is abundant in forest types which are rare or under strong economic pressure, (3) and it prefers host trees which have a restricted distribution in Estonia or are not favoured in forest management practice.

Research paper thumbnail of Habitat conditions and host tree properties affect the occurrence, abundance and fertility of the endangered lichen Lobaria pulmonaria in wooded meadows of Estonia

Abstract: We assessed multiple environmental factors that might influence the population vitality... more Abstract: We assessed multiple environmental factors that might influence the population vitality of
the epiphytic lichen Lobaria pulmonaria at the individual tree and habitat levels in partially overgrown
wooded meadows in Estonia. A total of 301 trees of four species were sampled at nine study plots,
using a stratified factorial scheme, 151 colonized by L. pulmonaria and 150 not colonized by L.
pulmonaria forming the control group. We used the Generalized Linear Models (GLZ) to identify a
complex of factors which predicts the probability of the lichen occurring on tree trunks and the
presence of apothecia on its individuals. We employed the General Linear Mixed Model (GLMM) to
study the relationship between cover of L. pulmonaria and environmental factors. The occurrence
probability of L. pulmonaria on tree trunks increased with increasing light availability and height of
deciduous shrubs near the trunk, and decreased with increasing distance to the nearest colonized tree.
The host tree species and its trunk properties were also of importance, particularly the facilitating effect
of the cover of bryophytes upon L. pulmonaria. The probability of occurrence of apothecia increased
with maximum values of bark pH and cover of L. pulmonaria on the trunk.We conclude that partially
overgrown wooded meadows are suitable habitats for L. pulmonaria. However, to maintain the vitality
of these populations, a specificmanagement scheme, preventing development of a dense stand, should
be applied. Management requirements would include 1) selective cutting of overgrowing coniferous
trees (particularly spruce), 2) preservation of adult and younger potential host trees within 10–20 m of
colonized trees, 3) preservation of scattered deciduous shrubs in the vicinity of the host trees.

Research paper thumbnail of Changes in stand structure due to the cessation of traditional land use in wooded meadows impoverish epiphytic lichen communities

Abstract: Wooded meadows with a history of traditional land use over thousands of years support a... more Abstract: Wooded meadows with a history of traditional land use over thousands of years support a
great diversity of various taxa. Today, however, high-species-rich communities in wooded meadows
are threatened because of the cessation of traditional management in large areas. We studied lichen
communities on 136 deciduous trees (Betula spp., Fraxinus excelsior and Quercus robur) in 12 wooded
meadows in three regions of Estonia, and assessed the effect of habitat change due to the abandonment
of traditional management on epiphytic lichen species composition, considering factors on three
spatial scales: regional, habitat and individual tree. The variation partitioning approach in partial
Canonical Correspondence Analysis (pCCA) revealed that most of the variation in species composition
is described by the species of host tree and tree bark pH. Other tree level variables, foremost tree
diameter, described as much of the compositional variation as geographic location (region) or
environmental conditions in wooded meadows. Of the environmental factors studied, woodland
canopy cover is the strongest predictor of the change in epiphytic lichen species composition from the
community type of semi-open wooded meadows to species-poor communities characteristic of
secondary forest. General Linear Model (GLM) analysis of the abundance of the 35 most frequently
observed lichen species revealed that more than half of them (21) are influenced by site openness
(canopy cover and/or undergrowth density), showing that increasing canopy cover has a negative effect
on the abundance of epiphytic lichen species characteristic of traditionally managed semi-open
wooded meadows. The results emphasize that the preservation of large old deciduous trees of various
species and the maintenance of the semi-open structure of stands are vitally important for the
protection of epiphytic lichen communities in wooded meadows.

Research paper thumbnail of The conservation of ground layer lichen communities in alvar grasslands and the relevance of substitution habitats

Semi-natural calcareous grasslands (alvars) are biodiversity hotspots in Northern Europe, particu... more Semi-natural calcareous grasslands (alvars) are biodiversity hotspots in Northern Europe, particularly for herb
layer plants. In the last century, traditional management has ceased, and the area of grasslands has declined
due to extensive encroachment. We were interested in the drivers of ground layer (alias terricolous or epigeic)
lichen communities. Our survey consisted of 86 habitat fragments in western Estonia, covering four types of
historic alvar grasslands and three types of alvar-like habitats. We found that the ground lichen communities
were primarily soil-type-specific, but were also affected by historic disturbances and land use change. In
contrast to knowledge about herb layer communities, for which shrub encroachment has been shown to be
main driver, the increased density of the herb layer and the reduced diversity of microhabitats were major
drivers for the ground layer lichen community. These drivers caused a decrease in species richness, but only
within the species of conservation value, and also led to a shift in the composition of lichen growth form from
the dominance of squamulose and crustose towards fruticose lichens. We conclude that the traditional practice
of restoring alvars by cutting shrubs is insufficient to maintain ground layer lichen biodiversity. Alvar
maintenance practices should include grazing, which creates various small-scale ground disturbances and
increases microhabitat heterogeneity. Alvar-like habitats originating from large-scale historic disturbances
appeared to be suitable for calcicolous epigeic lichens, and can therefore be considered to be temporary
substitution habitats, i.e. refugia for the regional species pool.

Research paper thumbnail of Functional ecology of rare and common epigeic lichens in alvar grasslands

The autecology of rare species can be derived using similarities among functional traits and envi... more The autecology of rare species can be derived using similarities among functional traits and
environmental conditions observed for common species, i.e. we employed the ‘matching
analogy approach’ with the analytical scheme ‘common species / driver / trait /
driver/rare species’.We addressed the driveretrait relationship for common epigeic lichens
of thin-soil calcareous grasslands, which are endangered by cessation of traditional land use.
Common lichens were suppressed by encroaching herbs and shrubs, and were supported by
ground disturbances. The lichens of open low-productivity alvars are predominantly calciphilous,
epibryic, crustose-squamulose, sexually reproducing and contain UV-protective pigments.
Lichens of encroached alvars are soil-type generalists, fruticose, reproduce vegetatively
and contain herbivore-deterring compounds. Rare lichens resemble the species of open lowproductivity
alvars, except their more limited niche space, i.e. they are restricted to arcticalpine
habitats. The conservation practices on alvars should support the formation of crustforming
communities by suppressing the growth of shrubs and herbs, and by promoting
recurrent small-scale soil disturbances.

Research paper thumbnail of Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions

Lichenologist, 2009

The species richness and composition of lichens was quantified for four temperate broad-leaved tr... more The species richness and composition of lichens was quantified for four temperate broad-leaved tree species (Acer platanoides, Fraxinus excelsior, Tilia cordata and Ulmus glabra) in boreo-nemoral forests on the talus slope of the North-Estonian limestone escarpment (North-Estonian Klint). Thirteen study sites were distributed along the klint on a west to east gradient. The effects of tree and stand characteristics and geographical location of a stand on composition and diversity of epiphytic lichens were evaluated by multivariate analyses (DCA, CCA, pCCA) and by general linear mixed models (GLMM). Tree level variables (e.g. bark pH, bryophytes cover and host tree species) explained the largest fraction of the variation in lichen species composition. However, species richness and composition were significantly influenced also by the unique habitat conditions of klint forest (length of the forest fragment, proximity of the stand to the sea and height of the escarpment). A significant correlation between stand locality and lichen diversity on the tree bole was found which is most likely related to local air pollution gradients caused by alkaline cement dust and acid pollutants in the north-eastern part of Estonia.

Research paper thumbnail of Diversity of epiphytic lichens in boreo-nemoral forests on the North-Estonian limestone escarpment: the effect of tree level factors and local environmental conditions – ERRATUM

Research paper thumbnail of Biogeographical determinants of lichen species diversity on islets in the West-Estonian Archipelago

Journal of Vegetation Science, 2006

AbstractQuestions: Do islet area, number of biotopes and distance from the mainland shape lichen ... more AbstractQuestions: Do islet area, number of biotopes and distance from the mainland shape lichen species richness on islets? Are there any species- or substrate group-specific trends associated with these factors?Do islet area, number of biotopes and distance from the mainland shape lichen species richness on islets? Are there any species- or substrate group-specific trends associated with these factors?Location: Islets of the west Estonian Archipelago, Estonia.Islets of the west Estonian Archipelago, Estonia.Methods: A species list was compiled for each of the 32 islets and the relative abundance of each species was estimated. The lichens were divided into seven groups according to their substrate preferences. Generalized linear model (GLIM) analysis was applied to test the effect of the islet traits on the number of lichen species on the islets and in the substrate groups. The probability of presence/absence and abundance of the most frequent species according to the islet traits were tested with GLIM and general linear mixed model.A species list was compiled for each of the 32 islets and the relative abundance of each species was estimated. The lichens were divided into seven groups according to their substrate preferences. Generalized linear model (GLIM) analysis was applied to test the effect of the islet traits on the number of lichen species on the islets and in the substrate groups. The probability of presence/absence and abundance of the most frequent species according to the islet traits were tested with GLIM and general linear mixed model.Results: The lichen flora of the islets consisted of 326 taxa, the number of lichen species per islet varied from 2 to 197. Total number of species per islet and within the substrate groups was positively correlated with islet area and with number of biotopes, and negatively correlated with distance from the mainland; however, these relationships varied among the substrate groups. Although individual lichen species showed variation in responses, general trends in island biogeography were evident.The lichen flora of the islets consisted of 326 taxa, the number of lichen species per islet varied from 2 to 197. Total number of species per islet and within the substrate groups was positively correlated with islet area and with number of biotopes, and negatively correlated with distance from the mainland; however, these relationships varied among the substrate groups. Although individual lichen species showed variation in responses, general trends in island biogeography were evident.Conclusions: The distribution pattern of lichens on the studied islets follows the theory of island biogeography: the number of species per islet depends on isolation, area and biotope diversity. Species specific traits, such as dispersal strategy and growth form, as well as availability of a particular substrate are important for formation of the lichen flora on islets.The distribution pattern of lichens on the studied islets follows the theory of island biogeography: the number of species per islet depends on isolation, area and biotope diversity. Species specific traits, such as dispersal strategy and growth form, as well as availability of a particular substrate are important for formation of the lichen flora on islets.