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Lichens have long been considered as composite organisms composed of algae and/or cyanobacteria hosted by a fungus in a mutualistic relationship. Other organisms have been gradually discovered within the lichen thalli, such as multiple algal species, yeasts, or even viruses. Of pivotal relevance is the existence of the lichen microbiome, which is a community of microorganisms that can be found living together on the lichen surface. This community performs a growing number of functions. In this entry, we explore the journey of lichens being considered from a dual partnership to a multi-species symbiotic relationship.

While restrictions imposed by the COVID-19 pandemic have strongly limited and affected the work of scientists and communicators, the pandemic has also encouraged the development of new ways of networking and public engagement. People have had to resort to virtual events, with a subsequent proliferation of webinars, online meetings, and digital resources. In this situation we have had to find new ways of measuring the impact of these activities. Using the activities of the Italian Lichen Society (Società Lichenologica Italiana, SLI), Google Trends and colleagues’ contributions, we evaluated the performances and impacts of virtual tools on lichenological literacy. We compared the relative success of virtual and in-person events and the effort required; we evaluated followers’ appreciation of various categories of posts on SLI Facebook page; and designed a questionnaire to collate information on individual experiences of in-person and virtual events linked to lichens. As expected, online events generally required less effort to put together and deploy than in-person events and engaged more people, especially when recorded and made available online for a long time. Using online searches for the word “lichens” we found an association with national events, and there was a notable increase in membership of SLI over the last 10 years, demonstrating an increasing interest by people in lichens. Without excluding the positive effects of in-person experiences, we believe that online events offer a powerful tool to help increase interest in, and knowledge about lichens. This interest may help to mitigate the impact of anthropogenic activities on this sensitive component of the ecosystem and help human-lichen relationships. Supplementary Information The online version contains supplementary material available at 10.1007/s13199-021-00780-6.

Climate change and atmospheric nitrogen (N) deposition on drylands are greatly threatening these especially vulnerable areas. Soil biocrust-forming lichens in drylands can provide early indicators of these disturbances and play a pivotal role, as they contribute to key ecosystem services. In this study, we explored the effects of different long-term water availability regimes simulating climate changes and their interaction with N addition on the physiological response of the soil lichen Cladonia rangiferina. Three sets of this lichen were subjected to control, reduced watering, and reduced watering and N addition (40 kg NH4NO3 ha−1 year−1) treatments for 16 months. Finally, all samples were subjected to daily hydration cycles with N-enriched water at two levels (40 and 80 kg NH4NO3 ha−1 year−1) for 23 days. We found that reduced watering significantly decreased the vitality of this lichen, whereas N addition unexpectedly helped lichens subjected to reduced watering to cope with stress produced by high temperatures. We also found that long-term exposure to N addition contributed to the acclimation to higher N availability. Overall, our data suggest that the interactions between reduced watering and increased N supply and temperature have an important potential to reduce the physiological performance of this soil lichen.

Forests host most terrestrial biodiversity and their sustainable management is crucial to halt biodiversity loss. Although scientific evidence indicates that sustainable forest management (SFM) should be assessed by monitoring multi-taxon biodiversity, most current SFM criteria and indicators account only for trees or consider indirect biodiversity proxies. Several projects performed multi-taxon sampling to investigate the effects of forest management on biodiversity, but the large variability of their sampling approaches hampers the identification of general trends, and limits broad-scale inference for designing SFM. Here we address the need of common sampling protocols for forest structure and multi-taxon biodiversity to be used at broad spatial scales. We established a network of researchers involved in 41 projects on forest multi-taxon biodiversity across 13 European countries. The network data structure comprised the assessment of at least three taxa, and the measurement of forest stand structure in the same plots or stands. We mapped the sampling approaches to multi-taxon biodiversity, standing trees and deadwood, and used this overview to provide operational answers to two simple, yet crucial, questions: what to sample? How to sample? The most commonly sampled taxonomic groups are vascular plants (83% of datasets), beetles (80%), lichens (66%), birds (66%), fungi (61%), bryophytes (49%). They cover different forest structures and habitats, with a limited focus on soil, litter and forest canopy. Notwithstanding the common goal of assessing forest management effects on biodiversity, sampling approaches differed widely within and among taxonomic groups. Differences derive from sampling units (plots size, use of stand vs. plot scale), and from the focus on different substrates or functional groups of organisms. Sampling methods for standing trees and lying deadwood were relatively homogeneous and focused on volume calculations, but with a great variability in sampling units and diameter thresholds. We developed a handbook of sampling methods (SI 3) aimed at the greatest possible comparability across taxonomic groups and studies as a basis for European-wide biodiversity monitoring programs, robust understanding of biodiversity response to forest structure and management, and the identification of direct indicators of SFM., This review was funded by the EU Framework Programme Horizon 2020 through the COST Association: COST Action CA18207: BOTTOMSUP- Biodiversity Of Temperate forest Taxa Orienting Management Sustainability by Unifying Perspectives.

Increasing economic growth and industrial development in China is starting to impact even remote areas such as the Shennongjia nature reserve, where nitrogen pollution is becoming a major environmental threat. The epiphytic lichen flora is particularly rich in this area and is one of the components of this habitat most sensitive to nitrogen pollution. Since lichens represent an important food resource for the endangered monkey species Rhinopithecus roxellana, a reduction in lichen availability would have harmful consequences for the conservation of its habitat in the Shennongjia Mountains. To investigate the effects of increased nitrogen availability on the local lichen communities, so far scarcely considered, we conducted a one-year field experiment measuring growth, survival, and phosphomonoesterase activity of the widespread species Usnea luridorufa in response to nitrogen (up to 50 kg N ha−1 year−1 deposition) and phosphorus supply. Growth and survival of thalli and propagules of U. luridorufa decreased when treated with N deposition >12.05 kg N ha−1 year−1 and >2.14 kg N ha−1 year−1, respectively. The important role of phosphorus availability in relation to nitrogen supply was demonstrated by the increase in phosphomonoesterase activity with increasing nitrogen availability until a nitrogen toxicity threshold was reached. However, the high concentration of phosphorus in rainwater showed that phosphorus is not a limiting nutrient in the area. The results make a contribution to the knowledge of the negative effects of increased N deposition in the Shennongjia forest ecosystem.

An ecological biogeographical perspective provides an understanding of the factors that shape the geographical distribution of organisms, their biodiversity and ecological speciation. Focusing on members of the lichen genus Solenopsora, which are strongly linked to a Mediterranean-type climate, we aimed to depict their environmental niches in the Apennine Peninsula. This area represents their ecological optima, as well as the biogeographical centre of distribution. On the basis of occurrences of Solenopsora congeners, we identified the key ecological factors that shape their environmental niches. Applying an ensemble approach, which merges the results of Random Forest, GLM and MaxEnt algorithms, suitability maps were developed. These are mainly influenced by geological substratum, temperature and precipitation. Occurrence of Solenopsora taxa seems to be mainly governed by low variability in diurnal temperature and tolerance to dryness, with precipitation in the range of 0–20 mm in the driest month and a minimum temperature of >5 °C in the coldest month. The sensitivity to diurnal temperature, an important indicator for climate change, suggests that the taxa confined to Mediterranean bioclimatic types (i.e. Solenopsora grisea, S. marina, S. olivacea subsp. olbiensis and S. olivacea subsp. olivacea) might be good indicators of climatic stability. The geological substratum was a strong limiting factor and separated the taxa into three groups: those growing on calcareous, siliceous and ultramafic substrata. Limited co-occurrence of species confined to one of the three categories suggests that the level of niche differentiation is on a microhabitat level. Accounting for ecological requirements, the taxa differ in their tolerance to sub-optimal conditions. The ecological niches of a sister subspecies pair with different reproduction strategies, fertile S. olivacea subsp. olivacea and sorediate S. olivacea subsp. olbiensis, overlap strongly. However, habitat suitability for S. olivacea subsp. olbiensis is greater in areas with higher precipitation in the driest month, whereas S. olivacea subsp. olivacea is more restricted to warmer and drier areas. We also report new regional records for Italy and the first records of S. cesatii in Serbia and Ukraine, and of S. liparina in Serbia.

Book review

The lichen Cladonia portentosa is generally considered to be sensitive to increased environmental nitrogen (N) deposition. However, the presence of this lichen in impacted environments suggests that it can cope with prolonged exposure to high N availability. To test the tolerance of this species to N, photosynthetic parameters, carbon and N concentrations and isotopic signature, chitin concentration, surface pH and extracellular enzymatic activity were measured in samples exposed for 11 years to different N doses and forms at the Whim bog N manipulation experimental site (United Kingdom). The results showed that C. portentosa is tolerant to long-term exposure to wet N deposition, maintaining its functionality with almost unaltered physiological parameters. The comparison of the proteome of short- and long-term exposed samples showed similar changes in protein expression suggesting that mechanisms to cope with N are not dependent on the exposure time even after more than a decade. Since empirical N Critical Loads are based on the response of sensitive components of the ecosystem, like C. portentosa, its capacity to cope with short- and long-term exposure to N needs to be recognized and taken into account when setting them, likewise, the significance of the form of N., Investigador - Fundação para Ciência e Tecnologia (FCT) contract and the FCT projects IF/00964/2013 and UID/BIA/00329/2013 (2015–2018).

Ammonia and ammonium have received less attention than other forms of air pollution, with limited progress in controlling emissions at UK, European and global scales. By contrast, these compounds have been of significant past interest to science and society, the recollection of which can inform future strategies. Sal ammoniac ( nūshādir , nao sha ) is found to have been extremely valuable in long-distance trade ( ca AD 600–1150) from Egypt and China, where 6–8 kg N could purchase a human life, while air pollution associated with nūshādir collection was attributed to this nitrogen form. Ammonia was one of the keys to alchemy—seen as an early experimental mesocosm to understand the world—and later became of interest as ‘alkaline air’ within the eighteenth century development of pneumatic chemistry. The same economic, chemical and environmental properties are found to make ammonia and ammonium of huge relevance today. Successful control of acidifying SO 2 and NO x emissions leaves atmospheric NH 3 in excess in many areas, contributing to particulate matter (PM 2.5 ) formation, while leading to a new significance of alkaline air, with adverse impacts on natural ecosystems. Investigations of epiphytic lichens and bog ecosystems show how the alkalinity effect of NH 3 may explain its having three to five times the adverse effect of ammonium and nitrate, respectively. It is concluded that future air pollution policy should no longer neglect ammonia. Progress is likely to be mobilized by emphasizing the lost economic value of global N emissions ($200 billion yr −1 ), as part of developing the circular economy for sustainable nitrogen management. This article is part of a discussion meeting issue ‘Air quality, past present and future’.

Review of the book "Sailing School Navigating Science and Skill, 1550-1800", from Margaret E. Schotte. With a style enjoyable to scholars and enthusiasts, Margaret Schotte analyzes archival records from six different countries to trace the evolution of a new paradigm of knowledge transfer, education and practice of navigation.

Soils are being degraded at an alarming rate and thereby also crucial ecosystem goods and services. Nitrogen (N) enrichment is a major driver of this degradation. While the negative impacts of N enrichment on vegetation are well known globally, those on various ecological interactions, and on ecosystem functioning, remain largely unknown. Because Mediterranean ecosystems are N limited, they are good model systems for evaluating how N enrichment impacts not only vegetation but also ecological partnerships and ecosystem functioning. Using a 7-year N-manipulation (dose and form) field experiment running in a Mediterranean Basin maquis located in a region with naturally low ambient N deposition (

Although it is generally accepted that δ15N in lichen reflects predominating N isotope sources in the environment, confirmation of the direct correlation between lichen δ15N and atmospheric δ15N is still missing, especially under field conditions with most confounding factors controlled. To fill this gap and investigate the response of lichens with different tolerance to atmospheric N deposition, thalli of the sensitive Evernia prunastri and the tolerant Xanthoria parietina were exposed for ten weeks to different forms and doses of N in a field manipulation experiment where confounding factors were minimized. During this period, several parameters, namely total N, δ15N and chlorophyll a fluorescence, were measured. Under the experimental conditions, δ15N in lichens quantitatively responded to the δ15N of released gaseous ammonia (NH3). Although a high correlation between the isotopic signatures in lichen tissue and supplied N was found both in tolerant and sensitive species, chlorophyll a fluorescence indicated that the sensitive species very soon lost its photosynthetic functionality with increasing N availability. The most damaging response to the different N chemical forms was observed with dry deposition of NH3, although wet deposition of ammonium ions had a significant observable physiological impact. Conversely, there was no significant effect of nitrate ions on chlorophyll a fluorescence, implying differential sensitivity to dry deposition versus wet deposition and to ammonium versus nitrate in wet deposition. Evernia prunastri was most sensitive to NH3, then NH4+, with lowest sensitivity to NO3−. Moreover, these results confirm that lichen δ15N can be used to indicate the δ15N of atmospheric ammonia, providing a suitable tool for the interpretation of the spatial distribution of NH3 sources in relation to their δ15N signal.

Symbiosis, mainly due to the advances in -omics technology and to the microbiome revolution, is being increasingly acknowledged as fundamental to explain any aspect of life existence. Previously considered an exception, a peculiar characteristic of few systems like lichens, corals and mycorrhizas, symbiosis is nowadays recognized as the rule, with the microbiome being part of all living entities and systems. However, our knowledge of the ecological meaning and functioning of many symbiotic systems is still limited. Here, we discuss a new, integrative approach based on current findings that looks at commonalities among symbiotic systems to produce theoretical models and conceptual knowledge that would allow a more efficient exploitation of symbiosis-based biotechnologies. The microbiome recruitment and assemblage processes are indicated as one of the potential targets where a holistic approach could bring advantages. Finally, we reflect on the potential socio-economic and environmental consequences of a symbiotic view of the world, where co-dependence is the matrix of life.

The interpretation of lichen bioaccumulation data is of paramount importance in environmental forensics and decision-making processes. By implementing basic ideas underlying previous interpretative scales, new dimensionless, species-independent &ldquo, bioaccumulation scales&rdquo, for native and transplanted lichens are proposed. Methodologically consistent element concentration datasets were populated with data from biomonitoring studies relying on native and transplanted lichens. The scale for native lichens was built up by analyzing the distribution of ratios between element concentration data and species-specific background concentration references (B ratios), herein provided for Flavoparmelia caperata and Xanthoria parietina (foliose lichens). The scale for transplants was built up by analyzing the distribution of ratios between element concentration in exposed and unexposed samples (EU ratio) of Evernia prunastri and Pseudevernia furfuracea (fruticose lichens). Both scales consist of five percentile-based classes, namely, &ldquo, Absence of&rdquo, &ldquo, Low&rdquo, Moderate&rdquo, High&rdquo, and &ldquo, Severe&rdquo, bioaccumulation. A comparative analysis of extant interpretative tools showed that previous ones for native lichens suffered from the obsolescence of source data, whereas the previous expert-assessed scale for transplants failed in describing noticeable element concentration variations. The new scales, based on the concept that pollution can be quantified by dimensionless ratios between experimental and benchmark values, overcome most critical points affecting the previous scales.

Lichens and mosses represent the macroscopic components of biological soil crusts (biocrusts). Their ability to exploit short periods of water availability and reversibly deactivate metabolism is crucial for their growth and survival. In this work we investigated photosynthesis reactivation, respectively after long (65–66 days) and short (15 days) dry periods, in lichen and moss species widespread in two Mediterranean environments (Portugal and Italy). Chlorophyll a fluorescence emission of the samples was investigated and the parameter Fv/Fm, an indicator of vitality of photosynthetic organisms, was used as a proxy for photosynthesis reactivation. The fruticose lichens Cladonia convoluta and C. rangiformis, and the moss Pleurochaete squarrosa, typical of Mediterranean environments, showed a significantly slower reactivation of photosynthetic activity when subjected to a longer period of drought. Conversely, the alien invasive moss Campylopus introflexus was not affected by prolonged dry conditions. The study showed that drought duration influences the reactivation of photosynthetic activity in terricolous lichens and mosses forming biocrusts in re-hydration cycles. These results indicate the likelihood of a reduction in biocrust productivity as a consequence of climate change in Mediterranean drylands.

A comprehensive biomonitoring programme should integrate several methods distributed along the biomonitoring chain, allowing to detect exposure, threads and impacts. In the case of a municipal solid waste incinerator (MSWI), biomonitoring of air pollution can contribute to source attribution, detection of ongoing processes and assessment of environmental effects. Three different methods were used to assess the biological effects of air pollution around a MSWI using lichens as biomonitors: (1) lichen diversity; (2) bioaccumulation of trace elements; and (3) physiological status (photosynthetic efficiency, cell membrane damage, viability). The first method takes into account the native lichen flora, while the other two were applied to thalli of the lichen Evernia prunastri transplanted for 6 months in the study area. Lichen diversity and physiological parameters reflected the effects of air pollution around the incinerator and the surrounding industrial area. High frequencies of non-nitrophilous species corresponded to sites with higher environmental quality, while high frequencies of nitrophilous species corresponded to sites with higher level of eutrophication. Transplanted samples showed increased cell membrane damage and reduced vitality respect to control samples. Bioaccumulation of trace elements pointed at the atmospheric origin of Hg depositions in the area. These results suggest that an integrated use of lichen-based methods along the biomonitoring chain can provide useful biological outputs for decision-makers to establish correct sustainable waste management policies.

Ajuts: This manuscript resulted from the 1st "Committee on Air Pollution Effects Research on Mediterranean Ecosystems" (CAPERmed) Conference celebrated in Lisbon, 2014 (http://capermed.weebly.com/). All attendants to this conference are, therefore, greatly acknowledged. Fundação para a Ciência e Tecnologia (FCT) is acknowledged through the Post Doc grant SFRH/BPD/85419/2012 to Teresa Dias and the Investigador grant to Silvana Munzi. ADM, EP and PS thank the LIFE10 ENV/FR/208 project MOTTLES. EP thanks also the Fondazione Cassa di Risparmio di Firenze (2013/7956). ROH is supported by a Juan de la Cierva Fellowship (IJCI-2014-21252) of the Spanish Ministry of Economy and Competitiveness. The authors declare no conflicts of interest. Mediterranean Basin ecosystems, their unique biodiversity, and the key services they provide are currently at risk due to air pollution and climate change, yet only a limited number of isolated and geographically-restricted studies have addressed this topic, often with contrasting results. Particularities of air pollution in this region include high O3 levels due to high air temperatures and solar radiation, the stability of air masses, and dominance of dry over wet nitrogen deposition. Moreover, the unique abiotic and biotic factors (e.g., climate, vegetation type, relevance of Saharan dust inputs) modulating the response of Mediterranean ecosystems at various spatiotemporal scales make it difficult to understand, and thus predict, the consequences of human activities that cause air pollution in the Mediterranean Basin. Therefore, there is an urgent need to implement coordinated research and experimental platforms along with wider environmental monitoring networks in the region. In particular, a robust deposition monitoring network in conjunction with modelling estimates is crucial, possibly including a set of common biomonitors (ideally cryptogams, an important component of the Mediterranean vegetation), to help refine pollutant deposition maps. Additionally, increased attention must be paid to functional diversity measures in future air pollution and climate change studies to establish the necessary link between biodiversity and the provision of ecosystem services in Mediterranean ecosystems. Through a coordinated effort, the Mediterranean scientific community can fill the above-mentioned gaps and reach a greater understanding of the mechanisms underlying the combined effects of air pollution and climate change in the Mediterranean Basin.

In this paper, we investigated whether growth form and substrate in lichens influence their physiological responses along an aridity gradient. Thalli of the foliose lichen Parmotrema perlatum and the fruticose lichen Ramalina canariensis were transplanted in selected rural/forested sites of Southern Portugal characterized by a different aridity index. Physiological parameters including photosynthetic performances, assimilation pigments, ergosterol content and sample viability were measured prior to exposure (winter) and after 6-month exposure (summer). Photosynthetic performances were also investigated in common native foliose and fruticose epiphytic lichens and in fruticose terricolous species. Both transplanted and native lichens showed a decrease in photosynthetic performances in summer and lower performances in sites classified as drier and higher performances in humid forested sites. No relevant differences occurred between epiphytic foliose and fruticose growth forms. However, terricolous fruticose samples showed a significant difference in humid and drier sites and between winter and summer, probably due to microclimatic conditions similarly to other biological crusts.

Main conclusion: Different nitrogen forms affect different metabolic pathways in lichens. In particular, the most relevant changes in protein expression were observed in the fungal partner, with NO3−mostly affecting the energetic metabolism and NH4+affecting transport and regulation of proteins and the energetic metabolism much more than NO3−did.Excess deposition of reactive nitrogen is a well-known agent of stress for lichens, but which symbiont is most affected and how, remains a mystery. Using proteomics can expand our understanding of stress effects on lichens. We investigated the effects of different doses and forms of reactive nitrogen, with and without supplementary phosphorus and potassium, on the proteome of the lichen Cladonia portentosa growing in a ‘real-world’ simulation of nitrogen deposition. Protein expression changed with the nitrogen treatments but mostly in the fungal partner, with NO3− mainly affecting the energetic metabolism and NH4+ also affecting the protein synthesis machinery. The photobiont mainly responded overexpressing proteins involved in energy production. This suggests that in response to nitrogen stress, the photobiont mainly supports the defensive mechanisms initiated by the mycobiont with an increased energy production. Such surplus energy is then used by the cell to maintain functionality in the presence of NO3−, while a futile cycle of protein production can be hypothesized to be induced by NH4+ excess. External supply of potassium and phosphorus influenced differently the responses of particular enzymes, likely reflecting the many processes in which potassium exerts a regulatory function.

Summary 1. In this work, we hypothesized that in urban areas with reduced pollution levels, local climatic conditions can be an important driver of lichen diversity. Thus, lichen functional groups could be used to study the effects of the urban heat island, characterized by higher temperature and lower humidity in urban than in peri-urban areas. To test this hypothesis, we sampled functional groups of epiphytic lichens in 48 forest remnants of a Mediterranean urban area (Almada, Portugal). 2. Of all tested functional groups (including those most closely associated with air pollution), groups related to water requirements showed the most significant correlations with surrounding artificial areas, roads and forest. These results suggest that the current major driver of lichen diversity is the climate within the urban area rather than air pollution, as frequently assumed. 3. The analysis of local climate using meteorological data of temperature and humidity confirmed the existence of urban heat island in the study area. 4. Synthesis and applications. The analysis of lichen functional groups gave an integrated response to the climatic modifications occurring in urban areas, namely to the urban heat island, suggesting that lichens can be used as a tool to evaluate the impact of urban areas on local climate and the effectiveness of mitigation and adaptation strategies if pollution levels are low.

Environmental cerium (Ce) levels are likely to increase in the near future and monitoring of its biological effects will therefore be necessary. The aim of this study was to test if treatment of the lichen Xanthoria parietina with Ce-containing solutions (0.1 mM, 1 mM, 10 mM and 100 mM) causes Ce bioaccumulation (both extra- and intra-cellularly) as well as physiological (sample viability, membrane lipids peroxidation, photosynthetic performance, water-soluble proteins content) and ultrastructural alterations. The results showed that treatment with Ce solutions induces Ce bioaccumulation, both extra-cellularly and intra-cellularly, which in turn causes an acute toxicity, evident as decreased sample viability, marked decrease in the photosynthetic performance and important changes in the ultrastructure.

Increased atmospheric nitrogen (N) deposition is known to alter ecosystem carbon source-sink dynamics through changes in soil CO2 fluxes. However, a limited number of experiments have been conducted to assess the effects of realistic N deposition in the Mediterranean Basin, and none of them have explored the effects of N addition on soil respiration (R s ). To fill this gap, we assessed the effects of N supply on R s dynamics in the following two Mediterranean sites: Capo Caccia (Italy), where 30kgha-1year-1 was supplied for 3years, and El Regajal (Spain), where plots were treated with 10, 20, or 50kg Nha-1year-1 for 8years. Results show a complex, non-linear response of soil respiration (R s ) to N additions with R s overall increasing at Capo Caccia and decreasing at El Regajal. This suggests that the response of R s to N addition depends on dose and duration of N supply, and the existence of a threshold above which the N introduced in the ecosystem can affect the ecosystem's functioning. Soil cover and seasonality of precipitations also play a key role in determining the effects of N on R s as shown by the different responses observed across seasons and in bare soil vs. the soil under canopy of the dominant species. These results show how increasing rates of N addition may influence soil C dynamics in semiarid ecosystems in the Mediterranean Basin and represent a valuable contribution for the understanding and the protection of Mediterranean ecosystems., This research was also financially supported by a research grant of the Spanish Ministerio de Economía y Competitividad (BIOMOD CGL-2013-44661-R). Comunidad de Madrid also funded this work through REMEDINAL-3 project (S2013/MAE-2719)

The mechanisms of nitrogen (N) tolerance in lichens are not yet fully understood. Here, we investigated how the increase of chitin content is related with N excess at inter- and intra-specific levels, by using species with differing ecological N tolerances (the tolerant Xanthoria parietina and Parmotrema hypoleucinum and the sensitive Evernia prunastri and Usnea sp.) and thalli of X. parietina and P. hypoleucinum from sites with different availabilities of N of agricultural origin (livestock), as confirmed by lichen N content and δ15N. Nitrogen, chitin (N-containing compound), and ergosterol contents were measured in lichen thalli. Nitrogen and chitin contents were higher in tolerant species than those in sensitive ones (inter-specific level) and in thalli collected from the N-polluted site than in thalli from the clean site (intra-specific level). We suggest that chitin contributes to N stress tolerance in lichens, and that excess N can be partially stored as chitin (non-toxic form) in the cell walls of tolerant species.

We tested whether freezing of air-dried lichen thalli alter physiological parameters commonly used as valuable stress markers in laboratory and field ecophysiological studies, namely integrity of cell membranes (measured as electrolyte leakage), assimilation pigments (chlorophyll a, chlorophyll b, chlorophyll a+b, total carotenoids content), chlorophyll integrity (OD435/OD415) and chlorophyll a fluorescence emission (expressed as F V/F M, the potential quantum yield of primary photochemistry and PIABS, a global indicator of the photosynthetic performance). Thalli of the lichen Evernia prunastri (L.) Ach. were air-dried, stored in the freezer at ca. −18°C under dark conditions and analysed after short-term storage (15, 30 and 90 days). These periods are compatible with the needs of a relatively rapid data evaluation in biomonitoring studies. After freezing, the investigated parameters showed that lichens remained healthy and suitable for later physiological measurements without biasing data quality.

Atmospheric dust pollution, especially particulate matter below 2.5 μm, causes 3.3 million premature deaths per year worldwide. Although pollution sources are increasingly well known, the role of ecosystems in mitigating their impact is still poorly known. Our objective was to investigate the role of forests located in the surrounding of industrial and urban areas in reducing atmospheric dust pollution. This was tested using lichen transplants as biomonitors in a Mediterranean regional area with high levels of dry deposition. After a multivariate analysis, we have modeled the maximum pollution load expected for each site taking into consideration nearby pollutant sources. The difference between maximum expected pollution load and the observed values was explained by the deposition in nearby forests. Both the dust pollution and the ameliorating effect of forested areas were then mapped. The results showed that forest located nearby pollution sources plays an important role in reducing atmospheric dust pollution, highlighting their importance in the provision of the ecosystem service of air purification.

This study aimed at investigating the cellular distribution of Hg in the lichens Cladonia arbuscula subsp. mitis and Peltigera rufescens treated with Hg²(+) and at testing if Hg treatment affects selected physiological parameters. In both species, increasing Hg accumulation under increasing Hg supply in the treatment solutions was found. P. rufescens showed a higher intracellular accumulation. Photosynthetic parameters were negatively affected in both species, as indicated by the decrease in photosynthetic pigments content, photosynthetic efficiency and chlorophyll integrity. Cell membranes of both species endured damage as indicated by the increase in the concentration of products of lipid peroxidation and decrease in ergosterol content. Nevertheless, differences between the two species were found, suggesting a differential sensitivity to Hg.

The present work aims at testing if exposure time and dose play a role in the response of lichen species to nitrogen (N) pollution. To this purpose, samples of the N-sensitive Evernia prunastri and the N-tolerant Xanthoria parietina were treated for 5 weeks either with solutions of NH 4 NO 3 0.05 and 1 M, or (NH 4 ) 2 SO 4 0.025 and 0.5 M. Photosynthetic efficiency was measured as an indicator of sample vitality. The results showed that the lowest concentrations were ineffective at the beginning, but after several supplies both compounds inhibited photosynthetic activity of E. prunastri . The highest concentrations had a deleterious effect, but with a temporal trend. For X. parietina no effect was found for the lowest concentrations, while the same trend shown by E. prunastri was instead observed following treatments with the highest concentrations. It was concluded that the response of lichens to N supply is not only species-specific, but also time- and dose-dependent. The results give a clue on field studies on the relationships between lichens and N pollution.

This study was carried out to investigate changes in lichen photosynthetic pigments induced by different combinations of light irradiance and water availability during a Mediterranean summer. To this purpose, thalli of three epiphytic lichens with a markedly different ecology concerning photo-hygrophytism, namely Evernia prunastri (hygro-mesophytic), Flavoparmelia caperata (mesophytic) and Xanthoria parietina (xero-mesophytic), were transplanted for 30 days to N- and S-facing cardinal exposures in central Italy. To investigate the effect of thallus hydration, at each cardinal exposure 50% of thalli were hydrated daily with deionised water. The results showed that changes in the concentrations of photosynthetic pigments are species-specific and consist in a general depression of photosynthetic pigments only in the hygro-mesophytic species E. prunastri. The pattern of photosynthetic pigments was also investigated in spontaneous samples along contrasting aspects. In this case, X. parietina from S-facing slopes, adapted to direct solar influx, showed higher pigment content than N-facing thalli; F. caperata and E. prunastri avoid direct extreme solar radiation and assimilation pigments were influenced more by the shadowing of tree canopies than by the cardinal exposure. The influence of drought on lichen photosynthetic pigments in the Mediterranean area is discussed.

In the present study we report some preliminary results concerning the evaluation of antiproliferative activity on murine myeloma cells (P3X63-Ag8.653) of crude extracts of two common lichen species, Evernia prunastri and Xanthoria parietina.

The results of antiproliferative activity against murine myeloma cells (line P3X63-Ag8.653) of water, ethanol, and methanol crude extracts of three lichen species belonging to the genus Peltigera (P. canina, P. elisabethae, and P. praetextata) are reported. Cell proliferation was evaluated by means of the 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide test, which is commonly used to assess the activity of living cells through mitochondrial dehydrogenases. The results showed that water extracts were the most effective in reducing cell proliferation; however, 40 μl of all Peltigera extracts, irrespective of water, ethanol, or methanol, induced a consistent reduction in cell proliferation. It is suggested that this important biological activity is due to antioxidant properties, possibly linked with phenolic compounds. A possible role of quaternary ammonium compounds is also discussed.

This study investigated the relationship between nitrogen and carbon in the thalli of the lichen Evernia prunastri, with an emphasis on the role of the C/N balance in the response of this species to N excess. For this purpose, thalli collected at a remote site were sprayed for 5 weeks with NH4NO3 (0.05 and 1 M) and (NH4)2SO4 (0.025 and 0.5 M) solutions, and later analyzed for their C and N content. The results showed that the total N content of thalli increased and the total C content decreased along with increasing N concentration in the treatment solutions. Consequently, the C/N ratio also significantly decreased in treated samples. The Fv/Fm ratio, taken as a vitality index, turned out to be positively correlated with C content and negatively correlated with N content. It is suggested that N stress is caused by the lack of compensation for elevated ammonium levels with higher C assimilation. It is concluded that E. prunastri is a suitable bioindicator of N pollution and that the C/N ratio deserves further investigation as a promising tool for early warning of N excess.

This study investigated the influence of angular exposure and distance from vehicular traffic on the diversity of epiphytic lichens and the bioaccumulation of traffic-related elements in a town of central Italy. An Index of Lichen Diversity (ILD) was calculated on the street-facing and the opposite side of road-lining trees and in a urban park 250 m away, and the content of selected trace elements (Al, Ba, Ce, Cd, Cr, Cu, Fe, Mn, Ni, Pb, Sb, V, and Zn) was determined in samples of the lichen Punctelia borreri (Sm.) Krog growing on tree bark, both on the exposed and opposite sides. ILD increases with distance from traffic emissions. However, at the site with vehicle traffic, non-nitrophilous lichens decreased while nitrophilous ones increased. The concentration of the traffic-related elements Ba, Cr, Cu, Mn, Sb, and Zn accumulated in thalli of P. borreri was higher on roadside trees than in trees from the urban park. ILD was not affected by the angular exposure to the road and the bioaccumulation of traffic-related elements was similar in lichens from the side of the bole exposed to traffic emissions and particulate resuspension and from the opposite side. The angular exposure in respect to the traffic source does not influence trace element accumulation. These results are important when using lichens for biomonitoring purposes, both for planning future studies and for the reliability of the interpretation of past surveys that do not report information about the angular exposure of the collected lichen material.

In this paper we tested if treating the lichen Xanthoria parietina with Sb-containing solutions causes Sb bioaccumulation as well as physiological and ultrastructural changes. Total and intracellular antimony content in Sb-treated samples increased progressively with increasing concentration in the treatment solutions. Incubation of X. parietina thalli with Sb at concentrations as low as 0.1 mM caused a decrease in sample viability, measured as intensity of respiratory activity, and damage to cell membranes, expressed in terms of membrane lipid peroxidation, as well as ultrastructural changes such as plasmolysis, impairment of the thylakoid system of the alga and cytoplasmic lipid droplets. The photosynthetic system hardly responded, at least under the tested experimental conditions.

To investigate the mechanisms of nitrogen (N) tolerance in lichens, we examined the physiological responses to increased N availability in different functional groups. Thalli of the nitrophytic Xanthoria parietina (L.) Th.Fr. previously grown both in an N-poor environment (~2 kg N ha–1 year–1) and in an N-rich environment (~52 kg N ha–1 year–1) were compared with the oligotrophic species Evernia prunastri (L.) Ach. and Usnea sp. Lichens were submitted to ammonium treatments. Maximum PSII efficiency, redistribution of the ions between the intra- and extracellular compartments and potassium and magnesium concentrations were the parameters used to check for the effects of N supply. The buffering capacity of the lichen extracts was also determined in untreated lichen thalli to check if different lichen behaviours were due to their ability to maintain the pH. The results showed a more similar response between X. parietina from the N-poor environment and the N-sensitive species than between X. parietina from the N-poor and N-rich environments, suggesting that X. parietina achieved N-tolerance after long-term exposure to N-rich environment. These results are important in understanding the effects of chronic ammonium pollution on one of the most sensitive components of the ecosystem, linking physiological response and ecological consequences.

This study investigated the physiological response of the epiphytic lichen Evernia prunastri to ecologically relevant concentrations of nitrogen compounds. Lichen samples were sprayed for 4 weeks either with water or 50, 150 and 500 μM NH 4 Cl. The integrity of cell membranes and chlorophyll a fluorescence emission ( F V / F M and PI ABS ) were analyzed. No membrane damage occurred after the exposure period. F V / F M , a classical fluorescence indicator, decreased during the second week of treatment with 500 μM NH 4 Cl and the third week with 50 and 150 μM NH 4 Cl. PI ABS , an overall index of the photosynthetic performance, was more sensitive and decreased already during the first week with 500 μM NH 4 Cl and the second week with 150 μM NH 4 Cl. Since E. prunastri has been exposed to ammonium loads corresponding to real environmental conditions, these findings open the way to an effective use of this species as early indicators of environmental nitrogen excess.