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Although it is generally accepted that δ 15 N in lichen reflects predominating N isotope sources in the environment, confirmation of the direct correlation between lichen δ 15 N and atmospheric δ 15 N 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, δ 15 N and chlorophyll a fluorescence, were measured. Under the experimental conditions, δ 15 N in lichens quantitatively responded to the δ 15 N of released gaseous ammonia (NH 3 ). 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 NH 3 , 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 NH 3 , then NH 4 + , with lowest sensitivity to NO 3 - . Moreover, these results confirm that lichen δ 15 N can be used to indicate the δ 15 N of atmospheric ammonia, providing a suitable tool for the interpretation of the spatial distribution of NH 3 sources in relation to their δ 15 N signal., (Copyright © 2018. Published by Elsevier B.V.)

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 δ 15 N. 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.

Reactive nitrogen (Nr) is an important driver of global change, causing alterations in ecosystem biodiversity and functionality. Environmental assessments require monitoring the emission and deposition of both the amount and types of Nr. This is especially important in heterogeneous landscapes, as different land-cover types emit particular forms of Nr to the atmosphere, which can impact ecosystems distinctively. Such assessments require high spatial resolution maps that also integrate temporal variations, and can only be feasibly achieved by using ecological indicators. Our aim was to rank land-cover types according to the amount and form of emitted atmospheric Nr in a complex landscape with multiple sources of N. To do so, we measured and mapped nitrogen concentration and isotopic composition in lichen thalli, which we then related to land-cover data. Results suggested that, at the landscape scale, intensive agriculture and urban areas were the most important sources of Nr to the atmosphere. Additionally, the ocean greatly influences Nr in land, by providing air with low Nr concentration and a unique isotopic composition. These results have important consequences for managing air pollution at the regional level, as they provide critical information for modeling Nr emission and deposition across regional as well as continental scales., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Despite reductions in atmospheric sulfur (S) concentrations due to abatement policies in some countries, modeling the dispersion of this pollutant and disentangling anthropogenic sources from natural ones is still of great concern. Lichens have been used as biomonitors of the impacts of S for over 40 years, but their potential as source-tracers of specific sources, including natural ones, remains unexplored. In fact, few attempts have been made to try to distinguish and spatially model different sources of S using lichens. We have measured S concentrations and isotopic values in lichens within an industrial coastal region where different sources of S, natural and anthropogenic, interplay. We detected a prevailing influence of natural sea-originated S that mixed with anthropogenic sources of S. We were then able to disentangle the sources of S, by removing the ocean influence on S isotopic values, enabling us to model the impact of different anthropogenic sources on S deposition and highlighting the potential use of lichens to evaluate the weight of different types of anthropogenic sources.

One of the main drawbacks of using lichens to monitor atmospheric PAHs has been reported as the inexistence of studies aiming to translate PAH values in lichens into the atmospheric equivalents ones, in order to use this information for regulatory purposes. In this work, PAH concentrations in lichens were compared with PAH concentrations measured in a conventional active sampler in an outdoor environment for a 9-month span. Significant positive correlations between HMW-PAHs, Σ16 EPA-PAHs, and BaP equivalent concentrations in lichens and those in air (TSP) were found. Concentrations of Σ16 EPA-PAHs in lichens and air showed a seasonal variation, with highest values during winter and lowest values during summer. Meteorological variables - temperature, atmospheric pressure, relative humidity, and wind speed - showed to significantly influence PAH concentrations in both lichens and air. Based on the significant linear correlations, equations for translating PAH concentrations measured in lichens into equivalent ones for air were proposed for the first time, allowing a broader use of lichens' information regarding PAHs in monitoring schemes and decision-making., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The aim of this study was to validate lichens as biomonitors of PAH atmospheric deposition; for that, an inter-comparison between the PAH profile and concentrations intercepted in lichens with those of air, soil and pine needles was performed. The study was conducted in a petro-industrial area and the results showed that PAH profiles in lichens were similar to those of the air and pine needles, but completely different from those of soils. Lichens accumulated higher PAH concentrations when compared to the other environmental compartments and its concentrations were significantly and linearly correlated with concentrations of PAHs in soil; we showed that a translation of the lichen PAHs concentrations into regulatory standards is possible, fulfilling one of the most important requirements of using lichens as biomonitors. With lichens we were then able to characterize the air PAHs profile of urban, petro-industrial and background areas., (Copyright (c) 2009 Elsevier Ltd. All rights reserved.)

The stable isotopic composition of water is routinely used as a tracer to study water exchange processes in vascular plants and ecosystems. To date, no study has focussed on isotope processes in non-vascular, poikilohydric organisms such as lichens and bryophytes. To understand basic isotope exchange processes of non-vascular plants, thallus water isotopic composition was studied in various green-algal lichens exposed to desiccation. The study indicates that lichens equilibrate with the isotopic composition of surrounding water vapour. A model was developed as a proof of concept that accounts for the specific water relations of these poikilohydric organisms. The approach incorporates first their variable thallus water potential and second a compartmentation of the thallus water into two isotopically distinct but connected water pools. Moreover, the results represent first steps towards the development of poikilohydric organisms as a recorder of ambient vapour isotopic composition.

The aim of this study was to compare the performance of two lichen species--Xanthoria parietina and Ramalina canariensis--as biomonitors of the toxic organic compounds dioxins and furans (PCDD/Fs). For that purpose, the concentrations and profiles of PCDD/Fs found in samples of these two lichen species were compared. Results showed that R. canariensis presented higher concentrations than X. parietina and that the PCDD/F homologue profiles were substantially different between these species. Xanthoria parietina appeared to be a more efficient interceptor of more chlorinated PCDD/Fs and also of particles, whereas R. canariensis mainly reflected less chlorinated PCDD/Fs. Results also showed that the PCDD/F profile of X. parietina differed from the one found in other foliose and fruticose lichen species. Despite the differences observed between the profiles of R. canariensis and X. parietina, the calibration of PCDD/F concentrations between the two species was achieved, allowing the biomonitoring of PCDD/Fs at a regional scale using both species simultaneously.

PAHs are toxic compounds emitted by several anthropogenic sources, which have a great impact on human health. We show, for the first time, how spatial models based on PAHs intercepted by lichens can be used for fingerprinting multisource atmospheric pollution in a regional area. Urban-industrial areas showed the highest atmospheric deposition of PAHs followed by urban > industrial > agricultural > forest Multivariate analysis of lichen data showed, for the first time, a clear distinction between various sources of PAHs in the same area: urban are dominated by 4-ring PAHs, forest by 3-ring PAHs, and industrial by 5- and 6-ring PAHs or by 2-ring PAHs (petrogenic or pyrogenic, respectively). Heavy metals were also used for supporting the fingerprinting of PAH sources, reinforcing the industrial origin of 5- and 6-ring PAHs and revealing their particular nature. The spatial structure of the models for different PAHs seems to be dependent on the following factors: size and hydrophilic character of different PAHs, type of emission sources (point or nonpoint), and dispersion associated with particulates of different sizes. Based on the long-term integration of PAHs in lichens, these spatial models will significantly improve our knowledge on the impact of PAH chronic-exposure to humans and ecosystems.

With the aim of determining the main drivers of changes in nitrophytic and oligotrophic macro-lichen communities in an industrial region with a Mediterranean climate, we considered both land-cover types and atmospheric pollutants. We determined the relation between the abundance of nitrophytic and oligotrophic species with environmental factors considering the distance of influence of land-cover types. The results showed that oligotrophic species decreased in the proximity of artificial areas, barren land and agricultural areas, associated with higher concentrations of NO2 and Zn, and Ti, probably dust of industrial and agricultural origin. Nitrophytic species were positively related to all the mentioned land-cover types, and with higher concentrations of Fe and N. Magnesium, probably from ocean aerosols, was negatively related to oligotrophic species and positively to nitrophytic.

The study presents first data on the delta(18)O performance of poikilohydrous lichen ground cover, and its potential impact on the isotopic composition of water fluxes arising from subjacent soil layers. As a model organism, the globally distributed lichen Cladina arbuscula was studied under laboratory conditions as well as in the field. During a desiccation experiment, delta(18)O of the lichen's thallus water and of its respired CO(2) became enriched by approximately 7 per thousand and followed a similar enrichment pattern to that expected from homoiohydrous, vascular plants. However, the observed degree of enrichment was lower in comparison to vascular plants due to (i) the lichen's inherent lower evaporative resistances; and (ii) a stronger effect of the more depleted surrounding water vapour. In lichens growing in their natural habitat, this specific pattern may show substantial variations depending on prevailing microclimatic conditions. Within a field study, thallus water delta(18)O of lichens principally proved to become more depleted when close to equilibration with the surroundings. It thereby strongly depended on the absorption of surrounding water vapour. Moreover, the results indicate that lichen mats substantially reduce evaporation rates arising from subjacent soil layers, and may alter the isotopic signal of vapour diffusing away from these layers into more depleted values.

The objective of this work was to determine the impact of neighbourhood land-cover in epiphytic lichen diversity. We used geostatistics to analyse the spatial structure of lichen-indicators (number of lichen species and Lichen Diversity Value) and correlate them to land-cover considering different distances from the observed data. The results showed that lichen diversity was influenced by different environmental factors that act in the same territory but impact lichens at different distances from the source. The differences in the distance of influence of the several land-cover types seem to be related to the size of pollutants/particles that predominantly are dispersed by each land-cover type. We also showed that a local scale of analysis gives a deeper insight into the understanding of lichen richness and abundance in the region. This work highlighted the importance of a multiple spatial scale of analysis to deeply interpret the relation between lichen diversity and the underling environmental factors.

The aim of this study was to compare the physiological responses to increased nitrogen (N) supply between the nitrophytic lichen Xanthoria parietina (L.) Th. Fr. and the acidophytic lichen Evernia prunastri (L.) Ach. The two lichens were exposed to a weekly dosage of 0.05, 0.1, 0.2, 0.6 or 2.4 g N m(-2) for 2 months, administered as NH(4)NO(3) dissolved in artificial rainwater (1 l m(-2)). After the treatments, in vivo chlorophyll a fluorescence was determined to assess vitality; concentrations of total N, ammonium, nitrate and dominant amino acids, including glutamate, glutamine and arginine, were quantified in order to follow changes in N status; and the polyols ribitol, arabitol and mannitol were quantified to follow changes in the lichens' carbon (C) status. The uptake of N was quantified by labelling the fertiliser with (15)N in the ammonium position; chlorophyll a was used as an indirect marker for algal activity, and ergosterol as an indirect marker of fungal activity. Nitrogen uptake was higher in E. prunastri than in X. parietina, although the latter species may have used the mannitol reserves to obtain C skeletons and energy for N assimilation. Chlorophyll a and ergosterol concentrations remained unaltered in X. parietina irrespective of N dosage while ergosterol decreased with increasing N uptake in E. prunastri. The latter species had accumulated a large pool of ammonium at the highest N dosage, whilst in X. parietina a significant nitrate pool was instead observed. Taken together, these short-term responses to high N supply observed in the two lichens, and the differences between them, can partly explain the higher tolerance of X. parietina towards increased atmospheric N levels.

The study presents first data on the δ18O performance of poikilohydrous lichen ground cover, and its potential impact on the isotopic composition of water fluxes arising from subjacent soil layers. As a model organism, the globally distributed lichen Cladina arbuscula was studied under laboratory conditions as well as in the field. During a desiccation experiment, δ18O of the lichen's thallus water and of its respired CO2 became enriched by ∼7‰ and followed a similar enrichment pattern to that expected from homoiohydrous, vascular plants. However, the observed degree of enrichment was lower in comparison to vascular plants due to (i) the lichen's inherent lower evaporative resistances; and (ii) a stronger effect of the more depleted surrounding water vapour. In lichens growing in their natural habitat, this specific pattern may show substantial variations depending on prevailing microclimatic conditions. Within a field study, thallus water δ18O of lichens principally proved to become more depleted when close to equilibration with the surroundings. It thereby strongly depended on the absorption of surrounding water vapour. Moreover, the results indicate that lichen mats substantially reduce evaporation rates arising from subjacent soil layers, and may alter the isotopic signal of vapour diffusing away from these layers into more depleted values. [ABSTRACT FROM AUTHOR]

Variations in stable carbon isotope discrimination (Δ) were investigated across the thalli of several lichen species possessing different photobiont associations. Lichens containing (i) green algae (phycobiont), (ii) green algae in association with cyanobacteria confined in cephalodia, or (iii) cyanobacteria (cyanobiont) as the photobiont partner were studied. Carbon isotope discrimination was analysed in different thallus sections, which varied in distance from the margin and in age. The marginal thallus region is considered to be youngest, while the central region is thought to be oldest. This analysis showed a clear variation in Δ across the thallus related to distance from the growing margin. In most of the species examined, the highest Δ values were found in marginal regions (younger), while the central and basal regions (older) showed significantly lower Δ. To investigate the effects of the historical increase in atmospheric CO2 concentration and the concurrent decrease in the 13C content of atmospheric CO2 on the Δ of lichens, experiments were carried out on herbarium samples of tobaria pulmonaria collected from the mid 19th Century to 1953. The results obtained showed a pattern of variation of Δ consistent with that of freshly collected samples; A decreased substantially with increasing distance from the thallus margin, irrespective of the collection date. Moreover, no consistent variation of discrimination was found among different collection dates. These results demonstrate that the observed variation in Δ is caused by age-related changes in the physiological behaviour of different thallus sections, and that the past 150 years of increasing CO concentration have not had significant effects on Δ in L. pulmonaria. Photosynthesis measurements, chlorophyll analysis and observations using optical microscopy, performed on freshly collected lichens, showed significant changes in morphological and physiological characteristics across the thallus. Particularly, remarkable variations in thickness were found across the thallus. These anatomical changes may be responsible for the variation in Δ, through variations in CO2 transfer resistance and, consequently, in CO2 availability across the thallus. The lack of age-dependent variation in Δ in cyanobiont lichens is possibly attributable to the operation of a CO2 -concentrating mechanism and, therefore, to a more constant CO2 environment across the thallus in this lichen group. [ABSTRACT FROM AUTHOR]

Abstract: The effects of land-use on species and communities are usually studied by considering high-intensity land-use gradients. However, in many Mediterranean Europe areas, the traditional land-use regime is of low-intensity, including low-impact forestry practices and extensive livestock pasture. This low-intensity land-use is associated with ‘High Nature Value Farmlands’, which occupy more than 50% of total farmland area in Mediterranean areas. The objective of this work was to determine the effect of traditional low-intensity land-use on epiphytic lichen communities in Mediterranean cork-oak woodlands. We intend to investigate the value of lichen communities as early indicators of the effects of land-use changes. For that, lichens were sampled along a gradient of low-intensity land-use that included forestry and pasture activities. As lichen diversity variables we considered: (i) total species richness, (ii) total species frequency (LDV), and (iii) richness and frequency of functional groups dividing species according to their eutrophication-tolerance. We found an increase of total species richness with an increase of land-use intensity which is opposite to expectations and which was shown for first time in lichens. The increase in total species richness was due to an increase of the number of eutrophication tolerant species accompanied by the maintenance of the sensitive one when we analyzed the lichen-data at the functional response group level. These results fit with the intermediate disturbance hypothesis that predicts higher diversity under an intermediate disturbance due to the coexistence of species with contrasting ecological requirements. Traditionally managed cork-oak woodlands with low-intensity land-use can thus support a high diversity of lichen species. An important outcome was that we confirmed the importance of using ecological indicators based on functional-groups for monitoring changes in ecosystems and, since lichens responded to low levels of land-use intensity, we have also shown the possibility of using lichens as early indicators of the impacts of low intensity land-use. [Copyright &y& Elsevier]