Lichen Species Richness Research Articles

Large beech (Fagus sylvatica) trees as ‘lifeboats’ for lichen diversity in central European forests

The forest management practices used in central Europe in the last several centuries have led to loss of lichen diversity that may be largely attributed to a loss of substrate variability and quantity. In an attempt to obtain information enabling us to mitigate this process, we surveyed affinity of lichen species to the substrates they currently occupy in six forest areas in the Czech Republic, located between 200 and 1000 m a.s.l. Tree bases and stems represented the most important substrate for lichen species, and especially so for threatened (i.e. red-listed) species. Lichen species richness per individual tree generally increased with stem diameter, especially for beech. Stems and tree bases of large-diameter beeches provide habitats that have enabled the survival of a crucial component of the red-listed lichen species in central Europe, far outweighing other tree species. The deciduous tree species that are commonly considered as favourable for lichen diversity (e.g. maples, ash, elms) were inhabited by only a few other lichen species additional to those associated with beech. This may be due to the low frequency of these tree species in most managed forests, and also some forest reserves, at the present time. Similarly, low incidence of dead wood in managed forests has likely limited its contribution to the lichen diversity, despite the high potential for lichen diversity associated with such substrates. It is thus apparent that bark of large-diameter live beech trees comprises a keystone habitat element in the provision of lichen diversity in central European forests.

Short-term response of Cladonia lichen communities to logging and fire in boreal forests

Fire is the major disturbance agent in boreal forests that initiates forest regeneration and succession, and plays a major role in determining the composition of terricolous lichen communities. In recent decades however, logging has become an important disturbance agent in boreal forests. In this context, growing concerns about logging effects on endangered species such as woodland caribou (Rangifer tarandus caribou) has emerged and drawn attention on how reindeer lichens (i.e. Cladonia arbuscula, Cladonia mitis, Cladonia rangiferina, Cladonia stellaris, and Cladonia stygia) respond to logging and fire given their importance as winter forage for caribou. We compared critical lichen habitat factors (i.e., forest floor thickness, tree height, and canopy closure) between fire and logged sites and evaluated how Cladonia lichen species richness, biomass, and ground cover were related to these habitat factors in the first decades following disturbance. We found no significant differences in habitat factors and no significant differences in lichen species richness or ground cover between logged and fire sites. However, Cladonia lichen biomass was significantly higher following logging. These results support the hypothesis that Cladonia lichens can persist following logging and suggest that forestry practices that include in their toolbox winter harvesting or machinery traffic restricted to specific trails are likely to preserve undisturbed forest floor habitat conditions with source populations of lichens.

Corticolous lichens as environmental indicators of natural sulphur emissions near the sulphur mine El Vinagre (Cauca, Colombia)

AbstractThe aim of this study was to evaluate the spatial effect of a natural source of sulphur pollution on the species diversity, richness and distribution of corticolous lichens in a páramo zone at the mine ‘El Vinagre’ (Puracé, Cauca, Colombia). Three zones at different distances from the pollution source were established: zone 1 with a high degree of contamination, a potentially mildly affected or transitional zone 2, and a zone 3 free of disturbance. In each zone, 10 phorophytes ofWeinmannia microphylla(Cunoniaceae) were sampled, and all lichens in a 150 cm vertical transect 50 cm above the ground were collected and identified. Phorophyte parameters were measured (bark pH and diameter at breast height) and the samples were georeferenced. In order to evaluate the impact on lichens, non-metric multidimensional scaling (NMS) and indicator species analysis were carried out. ANOVA and Spearman correlations were performed to assess the relationships of environmental and tree variables between zones and with lichen community structure. The index of atmospheric purity (IAP) and the environmental rating factor (FCA) were evaluated for the three zones. In total, 104 lichen species were recorded, of which 72 were identified to species, 17 to genus, and four to family; 11 samples could not be identified. NMS clustered samples according to zone and the main axis which were correlated with bark pH and distance from pollution source. We found eight indicator species characterizing different zones, and four marginally significant indicator species. Using the IAP, we established three isocontamination areas, with zones 2 and 3 classified as more or less pristine zones and zone 1 as a polluted zone (supported by bark pH as a proxy for current pollution). Diversity was lowest in zone 1, closest to the pollution source, and lichen species composition differed between zones. Differences between lichens in zones 2 and 3 appear mostly unrelated to the current pollution source and might be more related to historical differences in impact from a 20-year-old pollution source.

A Pine Is a Pine and a Spruce Is a Spruce--The Effect of Tree Species and Stand Age on Epiphytic Lichen Communities.

With an increasing demand for forest-based products, there is a growing interest in introducing fast-growing non-native tree species in forest management. Such introductions often have unknown consequences for native forest biodiversity. In this study, we examine epiphytic lichen species richness and species composition on the trunks of non-native Pinus contorta and compare these to the native Pinus sylvestris and Picea abies in managed boreal forests in northern Sweden across a chronosequence of age classes. Overall, we recorded a total of 66,209 lichen occurrences belonging to 57 species in the 96 studied forest stands. We found no difference in species richness of lichens between stands of P. contorta and P. sylvestris, but stands of P. abies had higher total species richness. However, species richness of lichens in stands of P. abies decreased with increasing stand age, while no such age effect was detected for P. contorta and P. sylvestris. Lichen species composition progressively diverged with increasing stand age, and in 30-year-old stands all three tree species showed species-specific assemblages. Epiphytic lichen assemblages in stands of 30-year-old P. contorta were influenced by greater basal area, canopy closure, and average diameter at breast height, P. abies stands by higher branch density and canopy closure, and stands of P. sylvestris by greater bark crevice depth. Differences in lichen species richness and composition were mainly explained by canopy closure and habitat availability, and the greater canopy closure in mature P. abies stands promoted the colonization and growth of calicioid lichen species. Our results indicate that the non-native P. contorta have similar species richness as the native P. sylvestris. The main difference in lichen species richness and composition is between P. abies and Pinus spp. in managed forests of boreal Sweden.

Lichen species richness is highest in non-intensively used grasslands promoting suitable microhabitats and low vascular plant competition

Lichens are very sensitive to habitat changes and their species richness is likely to decline under intensive land use. Currently, a comprehensive study analyzing lichen species richness in relation to land-use types, extending over different regions and including information on habitat variables, is missing for temperate grasslands. In three German regions we studied lichen species richness in 490 plots of 16 m2 representing different land-use types, livestock types, and habitat variables. Due to the absence of low-intensity pastures and substrates such as woody plants, deadwood and stones, there were no lichens in the 78 plots in Schorfheide-Chorin. In the two other regions, the richness of lichen species was 45 % higher in pastures than in meadows, and 77 % higher than in mown pastures, respectively. Among the pastures, the richness of all lichen species was on average 10 times higher in sheep-grazed pastures than in the ones grazed by cattle or horses. On average, the richness of all lichen species increased by 3.3 species per additional microhabitat. Furthermore, the richness of corticolous lichens increased by 1.2 species with 10 % higher cover of woody plants, lignicolous lichen species richness increased by 4.8 species with 1 % higher cover of deadwood, and saxicolous lichen species richness increased by 1.0 species with 1 % higher cover of stones. Our findings highlight the importance of low-intensity land use for lichen conservation. In particular, the degradation of grasslands rich in microhabitats and the destruction of lichen substrates by intensification, and conversion of unfertilized pastures formerly grazed at low intensity to meadows should be avoided to maintain lichen diversity.

Effects of Management on Lichen Species Richness, Ecological Traits and Community Structure in the Rodnei Mountains National Park (Romania).

Lichens are valuable bio-indicators for evaluating the consequences of human activities that are increasingly changing the earth’s ecosystems. Since a major objective of national parks is the preservation of biodiversity, our aim is to analyse how natural resource management, the availability of lichen substrates and environmental parameters influence lichen diversity in Rodnei Mountains National Park situated in the Eastern Carpathians. Three main types of managed vegetation were investigated: the transhumance systems in alpine meadows, timber exploitation in mixed and pure spruce forests, and the corresponding conserved sites. The data were sampled following a replicated design. For the analysis, we considered not only all lichen species, but also species groups from different substrates such as soil, trees and deadwood. The lichen diversity was described according to species richness, red-list status and substrate-specialist species richness. The variation in species composition was related to the environmental variables. Habitat management was found to negatively influence species richness and alter the lichen community composition, particularly for threatened and substrate-specialist species. It reduced the mean level of threatened species richness by 59%, when all lichen species were considered, and by 81%, when only epiphytic lichens were considered. Management-induced disturbance significantly decreased lichen species richness in forest landscapes with long stand continuity. The diversity patterns of the lichens indicate a loss of species richness and change in species composition in areas where natural resources are still exploited inside the borders of the national park. It is thus imperative for protected areas, in particular old-growth forests and alpine meadows, to receive more protection than they have received in the past to ensure populations of the characteristic species remain viable in the future.

A survey method for assessing the richness of epiphytic lichens using growth forms

The species richness of epiphytic lichens is continuously decreasing by degradation and loss of habitat. Considering that taxonomic identification of all species is time and resource consuming, rapid assessment methods to extrapolate the total number of species are needed for practical conservation. This paper describes an alternative method using the correlation between lichens growth forms and species richness. The study was conducted in 406 forest stands located in Central Spain, covering a wide range of mediterranean-climate ecosystem regions, management intensity levels, canopy cover conditions, and tree sizes. The presence/absence of epiphytic lichens was determined in 6090 trees, which were dominated by oak species (Quercus ilex, Q. faginea, and Q. pyrenaica). In all type of forests, the diversity of growth forms was positively correlated with the total epiphytic lichen richness. In all cases, species richness increased in non-managed forest stands with dense canopies. Thus, we propose the use of lichen growth forms as a helpful surrogate of species richness to detect potentially conservation priority areas in the Mediterranean region.

Macrolichen substrate selection: Patterns among aspen, non-aspen hardwood, and conifer-dominated forests in the Wasatch Mountains, Utah

Abstract Lichens in montane, forested ecosystems of the Intermountain West and central Rocky Mountains tend to be understudied. Focusing on forested ecosystems in Utah, specifically Santaquin and Payson Canyons, we describe lichen community diversity and composition and their relation to percent aspen cover, and contrast lichens in aspen, non-aspen hardwood and coniferous environments. We collected 351 macrolichens, a total of 34 species and 15 genera, from 22 circular plots with a 34.7 m radius. We found three major gradients in lichen community composition in our study area. First, many species of Xanthomendoza had strong positive correlations with increasing aspen cover, while several Melanohalea species declined with greater aspen cover. Xanthomendoza galericulata was a significant indicator for aspen forests, but this lichen was not limited to aspen as it colonizes various barks flexibly. Second, corticolous and saxicolous taxa positively associated with a trend in lichen species richness, while terr...

Contrasting patterns of lichen functional diversity and species richness across an elevation gradient

Major environmental gradients co‐vary with elevation and have been a longstanding natural tool allowing ecologists to study global diversity patterns at smaller scales, and to make predictions about the consequences of climate change. These analyses have traditionally studied taxonomic diversity, but new functional diversity approaches may provide a deeper understanding of the ecological mechanisms driving species assembly. We examined lichen taxonomic and functional diversity patterns on 195 plots (200 m²) together with forest structure along an elevational gradient of 1000 m in a temperate low mountain range (Bohemian Forest, Germany). Along this elevation gradient temperature decreased and precipitation increased, two macroclimatic variables critical for lichens. Elevation was more important than forest structure in driving taxonomic and functional diversity. While species richness increased with elevation, functional diversity decreased and revealed that community patterns shift with elevation from random to clustered, reflecting selection for key shared traits. Higher elevations favored species with a complex growth form (which takes advantage of high moisture) and asexual reproductive mode (facilitating establishment under low temperature conditions). Our analysis highlights the need to examine alternative forms of diversity and opens the avenue for community predictions about climate change. For a regional scenario with increasing temperature and decreasing availability of moisture, we expect a loss of specialized species with a complex growth form and those with vegetative organs at higher elevations in low mountain ranges in Europe.

Lichen traits and species as indicators of vegetation and environment

Abstract Lichens in the Arctic play important ecological roles. They also face the threats of increasing fire and shrub and tree expansion, exacerbated or caused by climate change. These forces may lead to changes not only in lichen community composition but also in the abundance, diversity and distribution of lichen functional traits. We sought to connect landscape-scale patterns of lichen community composition and traits to environmental gradients to both monitor lichen communities and clarify community-trait-environment relationships. We measured lichens throughout one of the largest and most remote U.S. National Parks within the Arctic. We then analyzed lichen community composition and species richness within ecologically informative lichen trait groups along environmental and vascular vegetation gradients. Macrolichen species richness in 0.4 ha plots averaged 41 species with a total landscape level observed gamma diversity of 262 macrolichen species. Jackknife estimators placed the landscape level ma...

Species richness and vertical stratification of epiphytic lichens in subtropical primary and secondary forests in southwest China

An entire-tree investigation was conducted in two primary and three secondary forest types in the subtropical Ailao Mountains of southwest China to determine whether species richness and vertical stratification of epiphytic lichens responded to forest type and host attributes. Lichen species number, composition and cover differed significantly among forest types and tree species, while tree diameter and tree height had a modest influence. Epiphytic lichen species and functional groups showed clear vertical stratification. Epiphytic lichens were richer in canopies than on trunks and exhibited a great preference for the intermediate zones of trees, while five lichen groups showed distinct vertical diversification. The stratification patterns are clearly related to forest type and may reflect the microclimatic requirements of individual species, e.g. light availability and humidity. (C) 2015 Elsevier Ltd and The British Mycological Society. All rights reserved.

The effects of rock climbing on cliff‐face vegetation

AbstractQuestionWhat are the effects of rock climbing on diversity, abundance and composition on cliff‐face vegetation along environmental and use gradients typical of modern climbing?LocationNew River Gorge National River, WV, US.MethodsWe compared species richness, abundance and composition of vascular plants, bryophytes and lichens on 79 pre‐established rock climbs and 32 unclimbed ‘control’ sites across potential climb‐use intensity and cliff structure using linear models, residual analysis and NMDS ordination.ResultsDifferences in species richness and abundance associated with potential climb‐use intensity and cliff structure were variable across taxonomic groups. Linear models indicated that cliff angle was the strongest explanatory variable of species richness and abundance for all three taxonomic groups. Once the effects of biophysical variables were modelled, analysis of the residuals indicated that potential climbing‐use intensity had a small but negative effect on species richness and abundance of vascular plants (range 3–6%), no effect on bryophytes (0%) and a substantial effect on lichens (range 10–12%). Similarly, NMDS ordination indicated that cliff angle, canopy height and aspect were the primary drivers of species composition. We observed no change in community composition due to climbing.ConclusionsOther studies have observed significant impacts of climbing on vegetation at the base of cliffs. Here, we observe that potential climbing‐use intensity has some impact on species richness and abundance of vascular plants and more significant impacts on lichens, however cliff angle is a fundamental control on cliff vegetation. Because of the challenge in sampling steep and overhanging cliffs, we recommend that climbers be directly involved in the assessment, monitoring and management of cliff resources in order to ensure that the full range of biophysical conditions of cliff ecosystems are considered.

Genotype variation in bark texture drives lichen community assembly across multiple environments.

A major goal of community genetics is to understand the influence of genetic variation within a species on ecological communities. Although well-documented for some organisms, additional research is necessary to understand the relative and interactive effects of genotype and environment on biodiversity, identify mechanisms through which tree genotype influences communities, and connect this emerging field with existing themes in ecology. We employ an underutilized but ecologically significant group of organisms, epiphytic bark lichens, to understand the relative importance of Populus angustifolia (narrowleaf cottonwood) genotype and environment on associated organisms within the context of community assembly and host ontogeny. Several key findings emerged. (1) In a single common garden, tree genotype explained 18-33% and 51% of the variation in lichen community variables and rough bark cover, respectively. (2) Across replicated common gardens, tree genotype affected lichen species richness, total lichen cover, lichen species composition, and rough bark cover, whereas environment only influenced composition and there were no genotype by environment interactions. (3) Rough bark cover was positively correlated with total lichen cover and richness, and was associated with a shift in species composition; these patterns occurred with variation in rough bark cover among tree genotypes of the same age in common gardens and with increasing rough bark cover along a -40 year tree age gradient in a natural riparian stand. (4) In a common garden, 20-year-old parent trees with smooth bark had poorly developed lichen communities, similar to their 10-year-old ramets (root suckers) growing in close proximity, while parent trees with high rough bark cover had more developed communities than their ramets. These findings indicate that epiphytic lichens are influenced by host genotype, an effect that is robust across divergent environments. Furthermore, the response to tree genotype is likely the result of genetic variation in the timing of the ontogenetic shift from smooth to rough bark allowing communities on some genotypes to assemble faster than those on other genotypes. Organisms outside the typical sphere of community genetics, such as lichens, can help address critical issues and connect plant genotype effects to long-established streams of biological research, such as ontogeny and community assembly.

Different patterns in species richness and community composition between trees, plants and epiphytic lichens in semi-natural pastures under agri-environment schemes

Agri-environment schemes (AES) have been established to counteract negative effects of agricultural intensification on e.g. semi-natural pastures and meadows. The efficiency of most AESs have, however, been poorly evaluated. We evaluated the success of a Swedish AES for the management of semi-natural pastures by comparing species richness and composition of vascular plants (except trees), epiphytic lichens and trees among pastures receiving higher (high value pastures) and lower levels of AES paymens (general value pastures). There was no difference in the number of tree species among high and general value pastures, even though AES regulations allow a maximum of 60 and 100 trees/ha in general and high value pastures, respectively. High value pastures had, however, a higher number of plant and epiphytic lichen species than common value pastures. Moreover, a higher number of pasture specialist plant species were indicative of high value pastures than of general value pastures. No lichen species indicating high value pastures are associated with habitats with low canopy cover (such as e.g. pastures). Finally, tree identity was an important factor for explaining the number and composition of epiphytic lichen species. Our study highlights that species groups can respond differently to agri-environment schemes and other conservation measures. Even though the effects are the desired on the diversity of one assessed taxon, this is not always the case for non-target organism groups.

Effects of human trampling on abundance and diversity of vascular plants, bryophytes and lichens in alpine heath vegetation, Northern Sweden.

This study investigated the effects of human trampling on cover, diversity and species richness in an alpine heath ecosystem in northern Sweden. We tested the hypothesis that proximity to trails decreases plant cover, diversity and species richness of the canopy and the understory. We found a significant decrease in plant cover with proximity to the trail for the understory, but not for the canopy level, and significant decreases in the abundance of deciduous shrubs in the canopy layer and lichens in the understory. Proximity also had a significant negative impact on species richness of lichens. However, there were no significant changes in species richness, diversity or evenness of distribution in the canopy or understory with proximity to the trail. While not significant, liverworts, acrocarpous and pleurocarpous bryophytes tended to have contrasting abundance patterns with differing proximity to the trail, indicating that trampling may cause shifts in dominance hierarchies of different groups of bryophytes. Due to the decrease in understory cover, the abundance of litter, rock and soil increased with proximity to the trail. These results demonstrate that low-frequency human trampling in alpine heaths over long periods can have major negative impacts on lichen abundance and species richness. To our knowledge, this is the first study to demonstrate that trampling can decrease species richness of lichens. It emphasises the importance of including species-level data on non-vascular plants when conducting studies in alpine or tundra ecosystems, since they often make up the majority of species and play a significant role in ecosystem functioning and response in many of these extreme environments.

Elevational gradients of terricolous lichen species richness in the Western Himalaya

Elevation confers limitations on distribution of organisms through correlated variations in temperature, moisture, radiations and precipitation. The elevation gradients of terricolous lichen species richness in Garhwal, western Himalaya were assessed using generalized additive models, in order to compare distribution patterns of different growth-forms, photobiont types, and dominant families. A total of 148 terricolous lichen species belonging to 42 genera and 19 families were recorded. The total species richness showed unimodal relationship with elevation, where the highest species richness was observed at mid elevations (3,200 m). The species richness of lichens with green algae (chlorolichens) and of lichens with cyanobacteria (cyanolichens) also exhibited significant unimodal elevational patterns with cyanolichens peaked at somewhat lower (2,800–2,900 m) elevation than chlorolichens (3,200 m). Growth forms showed statistically significant relationship of species richness to elevation, with crustose and squamulose lichens reaching their maxima at higher elevation than foliose, fruticose and dimorphic terricolous lichens. Unimodal pattern of species richness was also followed by six dominant families, with these families reaching maximum richness at different elevations. Elevational variation in topography, climate, and competition from vascular plant communities, together with the tolerance of specific growth forms to zoo-anthropogenic pressures, shape the distribution of terricolous lichens in the Garhwal Himalaya.

How do soil factors determine vegetation structure and species richness in post-smelting dumps?

This study aims to determine the effects of extreme heavy metal contamination and other soil factors on the structure and species richness of pioneer communities on artificial slag-soil substrates. Lichens, bryophytes, and vascular plants were examined in study plots and chemical properties of corresponding substrate samples were analyzed. Species responses to soil factors were identified by detrended correspondence analysis with interpretation of ordination axes and identification of significant soil parameters using linear mixed-effects models. The interrelationships between species richness and soil factors were evaluated by generalized linear mixed-effects model with Poisson error distribution. Three different responses to substrate contamination were recognized: species most tolerant and favored by strong contamination, less resistant species appearing only on less contaminated dumps and cryptogams indifferent to heavy metal contamination and abundant on all dumps. The species richness of lichens and vascular plants respond differently in relation to heavy metal contamination. Our results indicate that certain cryptogamic species are indifferent to enormous contamination and well adapted to slag substrate. In contrast, highly elevated levels of heavy metals negatively affect vascular plant species richness. Assemblages of pioneer cryptogams are an important element in the natural regeneration of post-smelting wastes and should be taken into account when reclamation activities are planned.

Influence of Time since Fire and Micro-Habitat Availability on Terricolous Lichen Communities in Black Spruce (Picea mariana) Boreal Forests

Terricolous lichens are an important component of boreal forest ecosystems, both in terms of function and diversity. In this study, we examined the relative contribution of microhabitat characteristics and time elapsed since the last fire in shaping terricolous lichen assemblages in boreal forests that are frequently affected by severe stand-replacing fires. We sampled 12 stands distributed across five age classes (from 43 to >200 years). In each stand, species cover (%) of all terricolous lichen species and species richness were evaluated within 30 microplots of 1 m2. Our results show that time elapsed since the last fire was the factor that contributed the most to explaining terricolous lichen abundance and species composition, and that lichen cover showed a quadratic relationship with stand age. Habitat variables such as soil characteristics were also important in explaining lichen richness. These results suggest that the presence of suitable substrates is not sufficient for the conservation of late-successional terricolous lichen communities in this ecosystem, and that they also need relatively long periods of times for species dispersal and establishment.

Epiphytic lichens onJuniperus communis– an unexplored component of biodiversity in threatened alvar grassland

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 juniperJuniperus communisand 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 size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.

Impacts of nitrogen deposition on herbaceous ground flora and epiphytic foliose lichen species in southern Ontario hardwood forests

In this study 70 sugar maple (Acer saccharum Marsh.) dominated plots in Ontario, Canada were sampled in the spring of 2009 and 2010 and herbaceous plant and epiphytic foliose lichen species data were compared against modeled N and S deposition data, climate parameters and measured soil and plant/lichen S and N concentration. Herbaceous plant species richness was positively correlated with temperature and indices of diversity (Shannon Weiner and Simpson's Index) were positively correlated with soil pH but not N or S deposition or standardized foliar N scores. Herbaceous community composition was strongly controlled by traditional factors, but there was a small and significant influence of atmospheric S and N deposition. Epiphytic lichen species richness exhibited a strong negative relationship with standardized foliar N score and only one lichen species (Phaeophyscia rubropulchra) was observed at sites with a standardized foliar N score of 0.76.