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Abstract: This paper aims to assess the influence of canopy cover on lichen growth in boreal forests along a regional forest gradient. Biomass and area gain, and some acclimation traits, [...]

This study reports on mollusc grazing of two epiphytic cyanobacterial lichens (Pseudocyphellaria crocata and Lobaria pulmonaria) transplanted within three Picea abies-dominated boreal rain forest stands (clear-cut, young and old forests) in west central Norway. Grazing was particularly high in transplants located in the old forest and was almost absent in clear-cut transplants. Grazing marks were absent on natural thalli on nearby spruce twigs (required creeping distance for mollusc from the ground >4 m). Transplantation of lichens from twigs to artificial transplantation frames reduced the creeping distance to 1.2 m, and caused a significant increase in grazing damage in P. crocata. Given a paired choice under transplantation, molluscs consistently preferred P. crocata and avoided L. pulmonaria, implying species-specific differences in herbivore defence. Pseudocyphellaria crocata has a much lower content of the medullary depsidones stictic and constictic acid than L. pulmonaria. Heavy grazing occurred in the P. crocata thalli lowest in these two depsidones. The upper part of the medulla hosting the photobiont was the preferred fodder for grazing molluscs. Molluscs avoided the yellow soralia in P. crocata (localised pulvinic acid), suggesting a role for pulvinic acid in preventing grazing of detached soredia and early establishment stages. The preference of P. crocata for thin spruce twigs is probably a result of a lower grazing pressure on twigs compared to e.g. deciduous stems that frequently support the better defended L. pulmonaria. Ongoing climate changes with increased annual rainfall and milder winters have presumably increased mollusc grazing, particularly in SW parts of Norway which have more species of lichen-feeding molluscs than the boreal sites studied. These temperate areas lacking natural spruce populations have recently experienced reported extinctions of the poorly defended P. crocata from rocks and deciduous stems prone to mollusc grazing. Lichen-feeding molluscs have likely played a role in these extinctions, causing spruce twigs in Atlantic boreal forests to be a last strong foothold for P. crocata in Scandinavia. [ABSTRACT FROM AUTHOR]

The article talks about the difference between fibrils and main stems in the epiphytic macrolichen called Usnea longissima in terms of the secondary compounds and chlorophylls. The author said that Usnea longissima often exceed one meter in length. He also said that it belongs to the subgenus Dolichousnea. It was also reported that the fibrils of the lichens are photosynthetic organs.

Grazing marks from snails are frequently observed in populations of the old forest epiphyte Lobaria pulmonaria. However, grazing marks are more numerous in thalli from deciduous broad-leaved forests than in thalli from boreal Picea abies forests, due to higher populations of lichen-feeding molluscs in deciduous stands. Here we tested for deleterious effects of snails on the lichens by transplanting 600 more or less grazed L. pulmonaria thalli from deciduous forests to snail-free P. abies forests. Subsequent measurements showed that growth rates were as high in thalli with many grazing marks as those without, suggesting that growth of mature lobes of L. pulmonaria are not inhibited by the recorded grazing pressure imposed by lichen feeding snails. [ABSTRACT FROM AUTHOR]

Synthesis of the cortical anthraquinone pigment parietin (= physcion) was studied in acetone-rinsed, parietin-free Xanthoria parietina thalli. UV-B induced the synthesis, which increased linearly with UV-B (log-transformed) to the highest applied UV-B level (1.8 W m−2). At natural UV-B levels (0.75 W m−2), parietin resynthesis occurred at a constant pace (106 mg m−2 d−1) during a 14-d period at 220 µmol m−2 s−1 PAR. Under these conditions, 56% of the natural parietin content prior to extraction was resumed, accounting for 10% of total net carbon gain. In the presence of UV-B, the remaining results were consistent with the hypothesis assuming that photosynthates regulate the pace at which parietin is synthesized by the mycobiont. Resynthesis was rapid when photosynthesis was activated by light, or when certain carbohydrates were added exogenously. Additions of ribitol, the carbohydrate delivered from the photobiont, increased the parietin resynthesis substantially. Mannitol, the main fungal polyol, was significantly less effective. Furthermore, parietin resynthesis in X. parietina was depressed at high and low hydration when net photosynthesis is depressed. Therefore, the photobiont regulates the parietin resynthesis pace in its mycobiont partner by the delivery of photosynthates. In conclusion, both lichen bionts play important roles in the synthesis of parietin, which probably acts as a PAR- rather than a UV-B-screen. [ABSTRACT FROM AUTHOR]

Seed germination, growth and flowering of the arctic-alpine annual Koenigia islandica L. were studied in controlled environment. Intact (unabraded) seeds germinated poorely at temperatures up to 18 deg. C,with an optimum at 24 deg. C (89 % in 10 d). Scarified seeds germinated rapidly, and reached 100 % germination in 3 d at 21 deg. C, butno more than 40 % germination occurred at 9 and 12 deg. C. The seeds had no light requirement for germination, nor did fluctuating temperatures improve germination. Dry matter production was optimal at 12deg. C in both short day (SD) and long day (LD) conditions, but was markedly higher in LD than in SD at identical fluences at all temperatures except 21 deg. C where the plants showed symptoms of severe heat stress. The temperature compensation point for net productivity was estimated to 24 deg. C, and negative carbon balance at higher temperatures might be an important physiological mechanism limiting the distribution of K. islandica in Scandinavia. Flowering was extremely rapid and independent of daylength, even in a high-arctic population from 79 deg. N. In full summer daylight anthesis was reached 24 d after germination and seeds ripened after 36 d at 15 deg. C. Days to anthesis varied little across the temperature range from 6 to 21 deg. C, giving a linear decrease in the heat-sum requirement for the attainment of flowering with decreasing temperature. It is concluded that conservative seed germination strategy, tininess and rapid development, low temperature optima for growth and reproduction, and daylength indifference of flowering are important adaptations for success of an annual plant in high-arctic and high-alpine environments. Daylength neutrality has facilitated the wide latitudinal distribution of K. islandica, including the penetration of the species to the southern hemisphere. [ABSTRACT FROM AUTHOR]

Studies high-light damage in air-dry thalli of the old forest lichen Lobaria pulmonaria. Interactions of irradiance, exposure duration and high temperature; Susceptibility to light of hydrated thalli of lichens; Effects of desiccation on the lichen.

Two neighbouring even-aged 130-yr old Picea abies trees in a homogeneous stand can differ substantially with respect to their epiphytic vegetation. Sampled branches from the canopy of one tree harboured 781specimens of the old forest lichen Usnea longissima of which only 50could be seen from the ground, whereas no U. longissima were found on its nearest neighbour. Usnea longissima was most abundant on branchtips in lower parts of the canopy on branch segments having the highest biomass of other alectorioid species. Trees with and without U. longissima showed a different pattern in their mineral composition, suggesting that a tree-specific difference in nutritional status might contribute to explain the patchy distribution of this lichen within seemingly homogeneous stands. [ABSTRACT FROM AUTHOR]

This study aims to elucidate the connection between the phylogeny of epiphytic macrolichens and their chemical niches. We analyzed published floristic and environmental data from 90 canopies of Picea glauca x engelmannii across various forest settings in British Columbia. To explore the concordance between a principal coordinates analysis of the cladistic distance matrix and a global non-metric multidimensional scaling of the ecological distance matrix, we used Procrustean randomization tests. The findings uncover a robust association between large-scale macrolichen phylogeny and canopy throughfall chemistry. The high calcium-scores of the studied species effectively distinguished members of the Peltigerales from those of the Lecanorales, although parameters linked with Ca such as Mn, Mg, K, bark-, and soil-pH, may contribute to the niche partitioning along the oligotrophic-mesotrophic gradient. The substantial large-scale phylogenetic variation in the macrolichens' Ca-scores is consistent with an ancient adaptation to specialized chemical environments. Conversely, the minor variation in Ca-scores within families and genera likely stems from more recent adaptation. This study highlights crucial functional and chemical differences between members of the Lecanorales and Peltigerales. The deep phylogenetic connection to the chemical environment underscores the value of lichens as transferable bioindicators for the chemical environment and emphasizes the importance of elucidating the intricate interplay between chemical factors and lichen evolution., Competing Interests: Declarations. Conflict of interest: The authors declare that they have no conflict of interest. Ethical approval: Ethics approval was not required for this non-destructive study. Consent to participate: Not applicable. Consent for publication: Not applicable., (© 2024. The Author(s).)

The mat-forming fruticose lichens Cladonia stellaris and Cetraria islandica frequently co-occur on soils in sun-exposed boreal, subarctic, and alpine ecosystems. While the dominant reindeer lichen Cladonia lacks a cortex but produces the light-reflecting pale pigment usnic acid on its surface, the common but patchier Cetraria has a firm cortex sealed by the light-absorbing pigment melanin. By measuring reflectance spectra, high-light tolerance, photosynthetic responses, and chlorophyll fluorescence in sympatric populations of these lichens differing in fungal pigments, we aimed to study how they cope with high light while hydrated. Specimens of the two species tolerated high light equally well but with different protective mechanisms. The mycobiont of the melanic species efficiently absorbed excess light, consistent with a lower need for its photobiont to protect itself by non-photochemical quenching (NPQ). By contrast, usnic acid screened light at 450-700 nm by reflectance and absorbed shorter wavelengths. The ecorticate usnic species with less efficient fungal light screening exhibited a consistently lower light compensation point and higher CO 2 uptake rates than the melanic lichen. In both species, steady state NPQ rapidly increased at increasing light with no signs of light saturation. To compensate for less internal shading causing light fluctuations with a larger amplitude, the usnic lichen photobiont adjusted to changing light by faster induction and faster relaxation of NPQ rapidly transforming excess excitation energy to less damaging heat. The high and flexible NPQ tracking fluctuations in solar radiation probably contributes to the strong dominance of the usnic mat-forming Cladonia in open lichen-dominated heaths., (© 2024. The Author(s).)

Mat-forming lichens dominating high-latitudinal habitats vary in color and geometry. Widespread species are light greenish yellow (usnic acid) and reflect solar radiation, whereas melanic species absorbing most solar wavelengths are spatially more restricted. Color thereby influences lichens' energy budget and thus their hydration and photosynthetically active periods. By using well-defined cushions from early successional stages on glacier forelands - three melanic (m) and three usnic (u) mat-forming lichens with hair-like branches (Alectoria ochroleuca (u) , Gowardia nigricans (m) ), hollow terete branches (Cladonia uncialis (u) , Cetraria muricata (m) ), and flat branches (Flavocetraria nivalis (u) , Cetraria islandica (m) ) - we quantified hydration traits and analyzed how color and cushion size affect water loss rate (WLR) and duration of active periods. Main findings: 1) WLR declined with cushion size and was highest in melanic lichens. 2) Active periods were longer for usnic than for melanic lichens and increased with size in all groups. 3) Size, color, and taxon nested in color significantly influenced WLR and duration of active periods in linear mixed models. 4) Hair lichen cushions had shorter active periods than growth forms with terete or flat branches due to their more open canopy architecture and lower water holding capacity (WHC). 5) WHC measured for isolated branches highly underestimated WHC for intact cushions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Main Conclusion: During desiccation, both apparent electron transport rate (ETR app ) and photosynthetic CO 2 uptake peak when external water has evaporated. External water, causing suprasaturation, weakens the strong correlation between ETR app and CO 2 uptake. Lichens are poikilohydric organisms passively regulated by ambient conditions. In theory, apparent electron transport rate (ETR app ), estimated by photosystem II yield measured in light (Φ PSII ), is a proxy of photosynthetic CO 2 uptake. Hydration level, however, is a complicating factor, particularly during suprasaturation that strongly reduces CO 2 diffusion. Here, the cephalolichen Lobaria pulmonaria and two chlorolichens Parmelia sulcata and Xanthoria aureola were excessively hydrated before photosynthetic CO 2 uptake and Φ PSII using imaging fluorescence tools were simultaneously measured while drying at 200 µmol photons m -2  s -1 . CO 2 uptake peaked when hydration had declined to a level equivalent to their respective internal water holding capacity (WHC internal ) i.e., the water per thallus area after blotting external water. CO 2 uptake and ETR app in all species were highly correlated at hydration levels below WHC internal , but weaker at higher hydration (chlorolichens) or absent (cephalolichen). Yet, at a specimen level for the two chlorolichens, the correlation was strong during suprasaturation. The CO 2 uptake-ETR app relationship did not differ between measured species, but may vary between other lichens because the slope depends on cortical transmittance and fraction of electrons not used for CO 2 uptake. For new lichen species, calibration of ETR app against CO 2 uptake is therefore necessary. At intrathalline scales, Φ PSII during drying initially increased along thallus margins before reaching maximum values in central portions when hydration approached WHC internal . WHC internal represents the optimal hydration level for lichen photosynthesis. In conclusion, ETR app is an easily measured and reliable proxy of CO 2 uptake in thalli without external water but overestimates photosynthesis during suprasaturation.

Although water is essential for photosynthetic activation in lichens, rates of vapor uptake and activation in humid air, which likely influence their niche preferences and distribution ranges, are insufficiently known. This study simultaneously quantifies rehydration kinetics and PSII reactivation in sympatric, yet morphologically and functionally distinct cephalolichens (Lobaria amplissima, Lobaria pulmonaria, Lobaria virens). High-temporal resolution monitoring of rehydrating thalli by automatic weighing combined with chlorophyll fluorescence imaging of maximal PSII efficiency (F V /F M ) was applied to determine species-specific rates of vapor uptake and photosynthetic activation. The thin and loosely attached growth form of L. pulmonaria rehydrates and reactivates faster in humid air than the thick L. amplissima, with L. virens in between. This flexible hydration strategy is consistent with L. pulmonaria's wide geographical distribution stretching from rainforests to continental forests. By contrast, the thick and resupinate L. amplissima reactivates slowly in humid air but stores much water when provided in abundance. This prolongs active periods after rain, which could represent an advantage where abundant rain and stem flow alternates with long-lasting drying. Understanding links between morphological traits and functional responses, and their ecological implications for species at risk, is crucial to conservation planning and for modelling populations under various climate scenarios., (Copyright © 2020 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Main Conclusion: Photobiont type influences the relative humidity threshold at which photosystem II activates in green algal lichens. Water vapor uptake alone can activate photosynthesis in lichens with green algal photobionts. However, the minimum relative humidity needed for activation is insufficiently known. The objective of this study was to quantify the humidity threshold for photosystem II (PSII) activation in a range of chlorolichen species associated with photobionts from Trebouxiaceae, Coccomyxaceae and Trentepohliaceae. These lichens exhibit distribution, habitat and substrate patterns that are likely coupled to their efficiency in utilizing water vapor at lower levels of relative humidity (RH) for photosynthesis. Using chlorophyll fluorescence imaging during water uptake from humid air of 25 species of chlorolichens representing the above photobiont groups, we monitored PSII activation within controlled chambers with constant RH at five levels ranging from 75.6 to 95.4%. The results demonstrate clear photobiont-specific activation patterns: the trentepohlioid lichens activated PSII at significantly lower RH (75.6%) than trebouxioid (81.7%) and coccomyxoid (92.0%) lichens. These responses are consistent with a preference for warm and sheltered habitats for trentepohlioid lichens, with cool and moist habitats for the coccomyxoid lichens, and with a more widespread occurrence of the trebouxioid lichens. Within each photobiont group, lichen species exposed to marine aerosols in their source habitats seemed to be activated at lower RH than lichens sampled from inland sites. High osmolyte concentration may therefore play a role in lowering a photobiont's activation threshold. We conclude that photobiont type influences water vapor-driven photosynthetic activation of lichens, thereby shaping the ecological niches in which they occur.

Main Conclusion: Chlorophyll fluorescence, infrared gas exchange and photoinhibition data consistently show that vulpinic acid in L. vulpina functions as a strong blue light screening compound. The cortical lichen compounds, parietin, atranorin, usnic acid and melanins are known to screen photosynthetically active radiation (PAR), thereby protecting the underlying photobionts. The role of the toxic UV-/blue light-absorbing vulpinic acid in lichen cortices is poorly documented. By comparing controls with acetone-rinsed Letharia vulpina thalli (75% reduced vulpinic acid concentration), we aimed to test PAR screening by vulpinic acid. We exposed such thalli to blue, green and red irradiance, respectively, and recorded light quality-specific light saturation curves of CO 2 uptake, quantum yields of CO 2 uptake (QY CO2 ) and effective quantum yields of PSII (Φ PSII ). We also quantified light quality-dependent photoinhibition after 4-h exposure to 400 µmol photons m -2  s -1 . In controls, the greatest high light-induced reductions in CO 2 uptake and Φ PSII , as well as the strongest photoinhibition [lowered maximal quantum yield of PSII (F v /F m )], occurred in red light, followed by green, and was low in blue light. Removal of vulpinic acid significantly exacerbated photoinhibition, reduced Φ PSII , and increased QY CO2 in blue light. By contrast, acetone rinsing had no or weak effects in green and red lights. Comparing control with acetone-rinsed thalli, blue light screening was estimated at 69% using Φ PSII data and 49% using QY CO2 . To compensate for the 25% residual vulpinic acid left after rinsing, we repeated the screening estimation by comparing responses in blue and red lights. This resulted in 88% screening using Φ PSII data and 77% using QY CO2 . The consistent responses in all photosynthetic parameters support the hypothesis that vulpinic acid functions as a blue light screen in L. vulpina.

Lichen photobionts in situ have an extremely UV-B tolerant photosystem II efficiency (Fv/Fm). We have quantified the UV-B-screening offered by the mycobiont and the photobiont separately. The foliose lichens Nephroma arcticum and Umbilicaria spodochroa with 1: intact or 2: removed cortices were exposed to 0.7 Wm -2 UV-B BE for 4 h. Intact thalli experienced no reduction in Fv/Fm, whereas cortex removal lowered Fv/Fm in exposed photobiont layers by 22% for U. spodochroa and by 14% for N. arcticum. We also gave this UV-B dose to algal cultures of Coccomyxa and Trebouxia, the photobiont genera of N. arcticum and U. spodochroa, respectively. UV-B caused a 56% reduction in Fv/Fm for Coccomyxa, and as much as 98% in Trebouxia. The fluorescence excitation ratio (FER) technique comparing the fluorescence from UV-B or UV-A-excitation light with blue green excitation light using a Xe-PAM fluorometer showed that these photobiont genera did not screen any UV-B or UV-A The FER technique with a Multiplex fluorometer estimated the UV-A screening of isolated algae to be 13-16%, whereas intact lichens screened 92-95% of the UV-A. In conclusion, the cortex of N. arcticum and U. spodochroa transmitted no UV-B and little UV-A to the photobiont layer beneath. Thereby, the upper lichen cortex forms an efficient fungal solar radiation screen providing a high UV-B tolerance for studied photobionts in situ. By contrast, isolated photobionts have no UV-B screening and thus depend on their fungal partners in nature., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

Lichen photobionts in situ have an extremely UV-B tolerant photosystem II efficiency (Fv/Fm). We have quantified the UV-B-screening offered by the mycobiont and the photobiont separately. The foliose lichens Nephroma arcticum and Umbilicaria spodochroa with 1: intact or 2: removed cortices were exposed to 0.7 Wm -2 UV-B BE for 4 h. Intact thalli experienced no reduction in Fv/Fm, whereas cortex removal lowered Fv/Fm in exposed photobiont layers by 22% for U. spodochroa and by 14% for N. arcticum. We also gave this UV-B dose to algal cultures of Coccomyxa and Trebouxia, the photobiont genera of N. arcticum and U. spodochroa, respectively. UV-B caused a 56% reduction in Fv/Fm for Coccomyxa, and as much as 98% in Trebouxia. The fluorescence excitation ratio (FER) technique comparing the fluorescence from UV-B or UV-A-excitation light with blue green excitation light using a Xe-PAM fluorometer showed that these photobiont genera did not screen any UV-B or UV-A The FER technique with a Multiplex fluorometer estimated the UV-A screening of isolated algae to be 13-16%, whereas intact lichens screened 92-95% of the UV-A. In conclusion, the cortex of N. arcticum and U. spodochroa transmitted no UV-B and little UV-A to the photobiont layer beneath. Thereby, the upper lichen cortex forms an efficient fungal solar radiation screen providing a high UV-B tolerance for studied photobionts in situ. By contrast, isolated photobionts have no UV-B screening and thus depend on their fungal partners in nature., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

Lichens are considered freezing tolerant, although few species have been tested. Growth, a robust measure of fitness integrating processes in all partners of a lichen thallus, has not yet been used as a viability measure after freezing. We compared relative growth rates (RGR) after freezing with short-term viability measures of photo- and mycobiont functions in the coastal Lobaria virens and the widespread L. pulmonaria to test the hypothesis that low temperature shapes the coastal distribution of L. virens. Hydrated thalli from sympatric populations were subjected to freezing at -10, -20 and -40 °C for 5 h. The rate of cooling and subsequent warming was 5 °C h -1 . Short-term viability measures of photobiont (maximal photosystem II efficiency, effective PSII yield) and mycobiont viability (conductivity index), as well as subsequent RGR, were assessed. The exotherms showed that L. virens froze at -3 °C; L. pulmonaria, at -4 °C. Freezing significantly impaired short-term viability measures of both photo- and mycobiont, particularly in the coastal species. Lobaria pulmonaria grew 2.1 times faster than L. virens, but the short-term damage after one freezing event did not affect the long-term RGR in any species. Thereby, short-term responses were impaired by freezing, long-term responses were not. While the lacking RGR-responses to freezing suggest that freezing tolerance does not shape the coastal distribution of L. virens, the significant reported adverse short-term effects in L. virens may be aggravated by repeated freezing-thawing cycles in cold winters. In such a perspective, repeated freezing may eventually lead to reduced long-term fitness in L. virens., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Background and Aims: Understanding to what extent parasites affect host fitness is a focus of research on ecological interactions. Fungal parasites usually affect the functions of vascular plants. However, parasitic interactions comprising effects of fungal parasites on the fitness of lichen hosts are less well known. This study assesses the effects of the abundance of two highly specialized gall-forming fungi on growth of their two respective lichen hosts and tests whether these fungal parasites reduce lichen fitness., Methods: The relative biomass and thallus area growth rates, and change in specific thallus mass of Lobaria pulmonaria and L. scrobiculata were compared between lichens with and without galls of the lichenicolous fungi Plectocarpon lichenum and P. scrobiculatae, cultivated in a growth chamber for 14 d. By estimating the thallus area occupied by the galls, it was also assessed whether growth rates varied with effective photosynthetic lichen surface area., Key Results: Plectocarpon galls significantly reduced relative growth rates of the lichen hosts. Growth rates decreased with increasing cover of parasitic galls. The presence of Plectocarpon-galls per se, not the reduced photosynthetic thallus surface due to gall induction, reduced relative growth rates in infected hosts. Specific thallus mass in the hosts changed in species-specific ways., Conclusions: This study shows that specialized fungal parasites can reduce lichen fitness by reducing their growth rates. Higher parasite fitness correlated with lower host fitness, supporting the view that these associations are antagonistic. By reducing hosts' growth rates, these parasites in their symptomatic life stage may affect important lichen functions. This fungal parasite-lichen study widens the knowledge on the ecological effects of parasitism on autotrophic hosts and expands our understanding of parasitic interactions across overlooked taxonomic groups., (© The Authors 2017. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Lichens are hosts for a variety of lichenicolous fungi. By investigating two lichens with specialized parasites, we will test the hypothesis that these parasites reduce lichen fitness by increasing the palatability of their respective hosts. The palatability of Lobarina scrobiculata and Lobaria pulmonaria with or without galls of the lichenicolous fungi, Plectocarpon scrobiculatae and P. lichenum, respectively, were quantified in a feeding-preference experiment with grazing snails (Cepaea hortensis). We repeated the experiment for pairs with or without gall in which the carbon-based secondary compounds (CBSCs) had been reduced nondestructively by acetone rinsing. Lichens with galls had lower concentration of CBSCs than those without, but this contrast disappeared after acetone rinsing. In the lichen high in nitrogen (N) (the cyanolichen L. scrobiculata), the grazing was low, and the snails did not discriminate between specimens with and without Plectocarpon-galls. In L. pulmonaria low in N (green algae as main photobiont), the parasite reduced the lichen C : N ratio and the snails strongly preferred specimens with Plectocarpon-galls, regardless of whether CBSC concentration had been reduced or not. In conclusion, some lichen parasites can indirectly reduce lichen fitness by increasing its palatability and thus the grazing pressure from snails, whereas other parasites do not affect grazing preferences., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Lichens are useful monitoring organisms for heavy metal pollution. They are high in carbon based secondary compounds (CBSCs) among which some may chelate heavy metals and thus increase metal accumulation. This study quantifies CBSCs in four epiphytic lichens transplanted for 6months on stands along transects from a highway in southern Norway to search for relationships between concentrations of heavy metals and CBSCs along a gradient in heavy metal pollutants. Viability parameters and concentrations of 21 elements including nutrients and heavy metals in these lichen samples were reported in a separate paper. Medullary CBSCs in fruticose lichens (Ramalina farinacea, Usnea dasypoga) were reduced in the most polluted sites, but not in foliose ones (Parmelia sulcata, Lobaria pulmonaria), whereas cortical CBSC did not change with distance from the road in any species. Strong positive correlations only occurred between the major medullary compound stictic acid present in L. pulmonaria and most heavy metals, consistent with a chelating role of stictic acid, but not of other studied CBSCs or in other species. However, heavy metal chelating did not protect L. pulmonaria against damage because this species experienced the strongest reduction in viability in the polluted sites. CBSCs with an accumulation potential for heavy metals should be quantified in lichen biomonitoring studies of heavy metals because they, like stictic acid, could overshadow pollutant inputs in some species rendering biomonitoring data less useful. In the two fruticose lichen species, CBSCs decreased with increasing heavy metal concentration, probably because heavy metal exposure impaired secondary metabolism. Thus, we found no support for a heavy metal protection role of any CBSCs in studied epiphytic lichens. No intraspecific relationships occurred between CBSCs versus N or C/N-ratio. Interspecifically, medullary CBSCs decreased and cortical CBSCs increased with increasing C/N-ratio., (Copyright © 2015 Elsevier B.V. All rights reserved.)

This growth chamber experiment evaluates how temperature and humidity regimes shape soluble carbohydrate pools and growth rates in lichens with different photobionts. We assessed soluble carbohydrates, relative growth rates (RGRs) and relative thallus area growth rates (RTA GRs) in Parmelia sulcata (chlorolichen), Peltigera canina (cyanolichen) and Peltigera aphthosa (cephalolichen) cultivated for 14 d (150 μmol m(-2) s(-1) ; 12-h photoperiod) at four day : night temperatures (28 : 23°C, 20 : 15°C, 13 : 8°C, 6 : 1°C) and two hydration regimes (hydration during the day, dry at night; hydration day : night). The major carbohydrates were mannitol (cephalolichen), glucose (cyanolichen) and arabitol (chlorolichen). Mannitol occurred in all species. During cultivation, total carbohydrate pools decreased in cephalo-/cyanolichens, but increased in the chlorolichen. Carbohydrates varied less than growth with temperature and humidity. All lichens grew rapidly, particularly at 13 : 8°C. RGRs and RTA GRs were significantly higher in lichens hydrated for 24 h than for 12 h. Strong photoinhibition occurred in cephalo- and cyanolichens kept in cool dry nights, resulting in positive relationships between RGR and dark-adapted photosystem II (PSII) efficiency (Fv /Fm ). RGR increased significantly with the photobiont-specific carbohydrate pools within all species. Average RGR peaked in the chlorolichen lowest in total and photobiont carbohydrates. Nocturnal hydration improved recovery from photoinhibition and/or enhanced conversion rates of photosynthates into growth., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Photosynthesis was compared in two cyanobacterial lichens (Lobaria hallii and Peltigera praetextata) and two green algal lichens (Lobaria pulmonaria and Peltigera leucophlebia) exposed to red, green or blue light. Cyanolichens had substantially lower photosynthetic CO(2) uptake and O(2) evolution than the green algal lichens in blue light, but slightly higher photosynthesis in red and green light. The effective quantum yield of photosystem (PS) II (Φ(PSII)) decreased with increasing red and green light for all species, but in blue light this response occurred in green algal lichens only. Cyanolichen Φ(PSII) increased with increasing blue light at low irradiances, but decreased at stronger exposures. However, after adding red light the efficiency of blue light for photosynthetic O(2) evolution increased by 2.4 times. Because phycobilisomes associated with PSII have a low blue light absorption, our results are consistent with blue light absorption mainly by Chl in PSI. Thereby, unequal allocation of excitation energy between PSII and PSI results in low cyanolichen photosynthesis under blue light. This is new knowledge in the science of lichenology with important implications for e.g. the reliability of using Chl fluorometers with blue light for cyanolichens., (© The Author 2014. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Pendulous lichens dominate canopies of boreal forests, with dark Bryoria species in the upper canopy vs. light Alectoria and Usnea species in lower canopy. These genera offer important ecosystem services such as winter forage for reindeer and caribou. The mechanism behind this niche separation is poorly understood. We tested the hypothesis that species-specific sunscreening fungal pigments protect underlying symbiotic algae differently against high light, and thus shape the vertical canopy gradient of epiphytes. Three pale species with the reflecting pigment usnic acid (Alectoria sarmentosa, Usnea dasypoga, U. longissima) and three with dark, absorbing melanins (Bryoria capillaris, B. fremontii, B. fuscescens) were compared. We subjected the lichens to desiccation stress with and without light, and assessed their performance with chlorophyll fluorescence. Desiccation alone only affected U. longissima. By contrast, light in combination with desiccation caused photoinhibitory damage in all species. Usnic lichens were significantly more susceptible to light during desiccation than melanic ones. Thus, melanin is a more efficient light-screening pigment than usnic acid. Thereby, the vertical gradient of pendulous lichens in forest canopies is consistent with a shift in type and functioning of sunscreening pigments, from high-light-tolerant Bryoria in the upper to susceptible Alectoria and Usnea in the lower canopy.

Acetone-extractable carbon based secondary compounds (CBSCs) were quantified in two epiphytic lichens to study possible effects of external factors (season and aspect) on secondary chemistry and to relate defense investments to biomass growth and changes in specific thallus mass (STM). At the end of four separate annual cycles starting in each of the four seasons, the cyanolichen Lobaria scrobiculata and the cephalolichen Lobaria pulmonaria (green algae as the primary photobiont and with localized Nostoc in internal cephalodia) were monitored in their natural forest habitats and after being transplanted at three contrasting aspects in open sites. Season strongly influenced most CBSCs. Medullary CBSCs in both species were twice as high in summer as in winter. Aspect hardly affected major CBSCs, whereas transplantation from forest to clear-cut slightly reduced these compounds. No major CBSCs in any species showed a trade-off with growth rate. Dry matter- as well as thallus area-based medullary CBSC contents increased with STM. The cortical usnic acid strongly increased with growth rate and followed spatial, but not seasonal variations in light exposure. Maximal CBSC levels during seasons with most herbivores is consistent with the hypothesis inferring that herbivory is a major selective force for CBSCs. Lack of trade-off between growth and defence investments suggests that these two processes do not compete for photosynthates., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

This study investigates how hydration during light and dark periods influences growth in two epiphytic old forest lichens, the green algal Lobaria pulmonaria and the cyanobacterial L. scrobiculata. The lichens were cultivated in growth chambers for 14 days (200 μmol m(-1) s(-2); 12 h photoperiod) at four temperature regimes (25/20 °C, 21/16 °C, 13/8 °C, and 6/1 °C; day/night temperatures) and two hydration regimes (12 h day-time hydration; 12 h day-time + 12 h night-time hydration). Growth was highly dynamic, showing that short-term growth experiments in growth cabinets have a high, but largely unexplored potential in functional lichen studies. The highest measured growth rates were not far from the maximal dry matter gain estimated from published net photosynthetic CO2 uptake data. For the entire data set, photobiont type, temperature, hydration regime and specific thallus mass accounted for 46.6 % of the variation in relative growth rate (RGR). Both species showed substantially higher relative growth rates based on both biomass (RGR) and thallus area (RTAGR) when they were hydrated day and night compared to hydration in light only. Chronic photoinhibition was substantial in thalli hydrated only during the day time and kept at the highest and lowest temperature regimes, resulting in exponential increases in RGR with increasing maximal PSII efficiency (F v/F m) in both species. However, the depression in F v/F m was stronger for the cyanolichen than for the cephalolichen at extreme temperatures. The growth-stimulating effect of night-time hydration suggests that nocturnal metabolic activity improves recovery of photoinhibition and/or enhances the conversion rate of photosynthates into thallus extension.

Desiccation tolerance was quantified in four cyanolichens (Lobaria hallii, Lobaria retigera, Lobaria scrobiculata, Pseudocyphellaria anomala), one cephalolichen (Lobaria pulmonaria) and one chlorolichen (Platismatia glauca) from xeric and mesic, open and closed North American boreal forests. These sympatric epiphytes were exposed to 0%, 33%, 55% and 75% relative humidity with or without medium light (200 μmol m⁻² s⁻¹) for 7 d. Permanent and temporary photoinhibitory damage was recorded as viability measures. All species tolerated well the drying in darkness, but L. hallii and L. retigera, associated with a very humid climate, showed minor damage at the hardest drying (silica gel). Simultaneous exposure to medium light severely aggravated the drying damage at all relative humidity levels. Combined drying-light exposure was particularly devastating for the widespread chloro- and cephalolichens, whereas cyanolichens, including rare old forest species, were fairly resistant. The ability to recover after combined drying-light stress (this study) correlated positively with increasing species-specific water holding capacities (from the literature). Cyanolichens, depending on liquid water and large internal water storage, probably require strong drying-light resistance to handle long periods between hydration events, whereas chlorolichens can regularly maintain their photosynthetic apparatus during frequent and rapid activation by humid air on clear mornings., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)

Growth in two old forest lichens was studied to evaluate how temporal (seasonal) and spatial (aspect-wise) partitioning of biomass and area growth respond to seasonal changes in light and climate. We monitored relative growth rates during annual courses in the cephalolichen Lobaria pulmonaria and the cyanolichen Lobaria scrobiculata transplanted in boreal clear-cut to five fixed aspects in winter, spring, summer, and autumn. For each annual set, growth was quantified in January-March, April-June, July-September and October-December. Mean biomass and area increased in all seasons, but growth was highest in July-September. Mass growth did not follow area increment during a year. As a result, mass per area (specific thallus mass (STM)) declined (L. scrobiculata) or stayed constant (L. pulmonaria) in the dark, humid October-December season, whereas it strongly increased in the dry, sunny April-June season. Aspect influenced growth in species-specific ways. Seasonality in biomass growth mainly followed light availability, whereas area growth was strongest during humid seasons. The substantial STM changes across seasons, species, and aspects can be explained as passive responses to seasonal climate. However, as STM, according to the literature, is a driver of water storage, recorded changes probably improve fitness by prolonging hydration in places or during times with high evaporative demands., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)

The optimal defense theory (ODT) deals with defensive compounds improving fitness of a particular organism. It predicts that these compounds are allocated in proportion to the risk for a specific plant tissue being attacked and this tissue's value for plant fitness. As the benefit of defense cannot easily be measured in plants, the empirical evidence for ODT is limited. However, lichens are unique in the sense that their carbon-based secondary compounds can nondestructively be removed or reduced in concentration by acetone rinsing. By using such an extraction protocol, which is lethal to plants, we have tested the ODT by studying lichens instead of plants as photosynthetically active organisms. Prior to acetone rinsing, we found five times higher concentration of meta-scrobiculin in the reproductive parts (soralia) of Lobaria scrobiculata compared to somatic parts of this foliose epiphytic lichen species. At this stage, the lichen-feeding snail Cochlodina laminata avoided the soralia. However, after removal of secondary compounds, the snail instead preferred the soralia. In this way, we have successfully shown that grazing pattern inversely reflects the partitioning of the secondary compounds that have a documented deterring effect. Thus our study provides strong and novel evidence for the ODT.

Lichens, representing mutualistic symbioses between photobionts and mycobionts, often accumulate high concentrations of secondary compounds synthesized by the fungal partner. Light screening is one function for cortical compounds being deposited as crystals outside fungal hyphae. These compounds can non-destructively be extracted by 100% acetone from air-dry living thalli. Extraction of atranorin from Physcia aipolia changed the lichen colour from pale grey to green in the hydrated state, whereas acetone-rinsed and control thalli were all pale grey when dry. Removal of parietin from Xanthoria parietina changed the colour of desiccated thalli from orange to grey. Colour changes were quantified by reflectance measurements. By a new chlorophyll fluorescence method, screening was assessed as the decrease in incident irradiance (PAR) necessary to reach identical effective quantum yields of PSII (Phi(PSII)) in acetone-rinsed and control thalli. Thereby, we estimated a screening efficiency due to cortical atranorin crystals at 61, 38, and 40% of blue, green and red light, respectively, whereas parietin screened 81, 27 and 1% of these wavelength ranges. Removal of atranorin caused similar levels of increased photoinhibition for P. aipolia in blue, green and red light, whereas parietin-deficient thalli of X. parietina exhibited increased photoinhibition with decreasing wavelengths. Atranorin possibly prevents water from entering the spaces between the hyphae in the cortex. The air-filled cavities with white atranorin crystals reflect excess light, whereas the yellow compound parietin absorbs excess light. Thereby, both atranorin and parietin play significant photoprotective roles for symbiotic green algae, but with compound-specific screening mechanisms.

Some lichen compounds are known to deter feeding by invertebrate herbivores. We attempted to quantify the deterring efficiency of lichen compounds against a generalist vertebrate, the bank vole (Myodes glareolus). In two separate experiments, caged bank voles had the choice to feed on lichens with natural or reduced concentrations of secondary compounds. We rinsed air-dry intact lichens in 100% acetone to remove extracellular compounds non-destructively. In the first experiment, pairs of control and rinsed lichen thalli were hydrated and offered to the bank voles. Because the lichens desiccated fast, we ran a second experiment with pairs of ground control and compound-deficient thalli, each mixed with water to porridge. Eight and six lichen species were tested in the first and second experiment, respectively. In the first, bank voles preferred compound-deficient thalli of Cladonia stellaris and Lobaria pulmonaria, but did not discriminate between the other thallus pairs. This was likely a result of deterring levels of usnic and stictic acid in the control thalli. When lichens were served as porridge, significant preference was found for acetone-rinsed pieces of Cladonia arbuscula, C. rangiferina, Platismatia glauca, and Evernia prunastri. The increased preference was caused mainly by lower consumption of control thalli. Grinding and mixing of thallus structures prevented bank voles from selecting thallus parts with lower concentration of secondary compounds and/or strengthened their deterring capacity. We conclude that some lichen secondary compounds deter feeding by bank voles.

Relationships between thallus size and growth variables were analysed for the foliose Lobaria pulmonaria and the pendulous Usnea longissima with the aim of elucidating their morphogenesis and the factors determining thallus area (A) versus biomass (dry weight (DW) gain. Size and growth data originated from a factorial transplantation experiment that included three boreal climate zones (Atlantic, suboceanic and continental), each with three successional forest stands (clear-cut, young and old). When A was replaced by the estimated photobiont layer area in an area-DW scatterplot including all thalli (n = 1080), the two separate species clusters merged into one, suggesting similar allocation patterns between photobionts and mycobionts across growth forms. During transplantation, stand-specific water availability boosted area gain in foliose transplants, consistent with a positive role of water in fungal expansion. In pendulous lichens, A gain greatly exceeded DW gain, particularly in small transplants. The A gain in U. longissima increased with increasing DW:A ratio, consistent with a reallocation of carbon, presumably mobilized from the dense central chord. Pendulous lichens with cylindrical photobiont layers harvest light from all sides. Rapid and flexible three-dimensional A gain allows the colonization of spaces between canopy branches to utilize temporary windows of light in a growing canopy. Foliose lichens with a two-dimensional photobiont layer have more coupled A and DW gains.

THIS STUDY AIMS: (1) to quantify mollusc grazing on juvenile and mature thalli of the foliose epiphytic lichen Lobaria pulmonaria, and (2) to test the hypothesis inferring a herbivore defensive role of lichen depsidones in forests with indigenous populations of lichen-feeding molluscs. Lichens were transplanted in shaded and less shaded positions in each of two calcareous broadleaved deciduous forests, one poor in lichens, one with a rich Lobarion community. Preventing the access of molluscs significantly reduced the loss of juvenile L. pulmonaria, particularly in the naturally lichen-poor forest. Molluscs also severely grazed mature thalli in the lichen-poor forest, especially thalli placed under the more shading canopies. Furthermore, reducing the natural concentration of depsidones by pre-rinsing with acetone increased subsequent grazing significantly, showing that lichen depsidones function as herbivore defence in natural habitats. Our results suggest that mollusc grazing may play important roles in shaping the epiphytic vegetation in calcareous deciduous forests, and that recently established juvenile L. pulmonaria thalli seem to be particularly vulnerable.

Three epiphytic old forest lichens (Usnea longissima, Pseudocyphellaria crocata, and Lobaria pulmonaria) were transplanted along a natural shade-sun gradient comprising three successional stages in boreal spruce forests (dense young forest, open old forest, and clear-cut) for one summer. After harvest, extractable secondary compounds were analyzed by high-performance liquid chromatography, and the brown pigmentation in melanic species was quantified by reflectance measurements. Cortical compounds in all species increased from shady young forests to exposed clear-cuts. Usnic acid, the major cortical, secondary compound in U. longissima, showed consistently higher concentration in the clear-cut than in the two forested stands. Pseudocyphellaria crocata and L. pulmonaria, lacking extractable secondary compounds in the cortex, significantly increased their amounts of cortical melanins in well-lit stands. The medullary compounds showed more complex responses. Many were not influenced by environmental conditions during the transplantation, whereas the majority of those that responded showed the lowest concentration in clear-cut transplants. Only a few medullary compounds showed the highest concentration in the clear-cut, and at a low level of significance. The synthesis of UV-B-absorbing usnic acid and melanins seems to be part of an acclimation to increased light exposure. The medullary compounds in studied species barely function as solar screens despite their strong UV-B absorbance.

This study analysed relationships between secondary chemistry, lichen growth rates and external habitat factors for two groups of UV-B-absorbing secondary compounds in the lichen Lobaria pulmonaria in order to test some hypotheses on their formation and function. Medullary depsidones and cortical melanins were quantified in thalli transplanted to three successional forest stands (shaded young forest, open old forest, sun-exposed clear-cut area) and subjected to different watering regimes (spraying with water, water + nitrogen, no spraying). Growth rates were already known. The total concentration of all seven depsidones was constant across the entire range of growth rates and sun exposures, showing that these depsidones serve functions other than photoprotection. Thalli from the well-lit transplantation sites had the highest synthesis of melanins. Within each forest type there was a trade-off between growth and melanin synthesis. Melanins and photosynthetic acclimation enhanced survival on a subsequent exposure to high light intensity, despite excessive temperatures resulting from higher absorption of solar energy in melanic thalli relative to pale thalli. In conclusion, the highly responsive melanic pigments play a photoprotective role in light acclimation, whereas the constant amount of depsidones across a wide spectrum of growth ranges and irradiances is consistent with herbivore defence functions.

This study aims to assess the photoprotective potential of desiccation-induced curling in the light-susceptible old forest lichen Lobaria pulmonaria by using chlorophyll fluorescence imaging. Naturally curled thalli showed less photoinhibition-induced limitations in primary processes of photosynthesis than artificially flattened specimens during exposures to 450 micromol m-2 s-1 in the laboratory after both 12- (medium dose treatment) and 62-h duration (high dose treatment). Thallus areas shaded by curled lobes during light exposure showed unchanged values of measured chlorophyll fluorescence parameters (FV/FM, PhiPS II), whereas non-shaded parts of curled thalli, as well as the mean for the entire flattened thalli, showed photoinhibitory limitation after light treatments. Furthermore, the chlorophyll fluorescence imaging showed that the typical small-scale reticulated ridges on the upper side of L. pulmonaria caused a spatial, small-scale reduction in damage due to minor shading. Severe dry-state photoinhibition readily occurred in flattened and light-treated L. pulmonaria, although the mechanisms for such damage in a desiccated and inactive stage are not well known. Natural curling is one strategy to reduce the chance for serious photoinhibition in desiccated L. pulmonaria thalli during high light exposures.

Seasonal variation in maximal photochemical quantum yield (F(V)/F(M)) of photosystem II (PS II), light adapted quantum yield (Phi(II)) of PS II, non-photochemical quenching (NPQ), contents of chlorophylls, and xanthophyll cycle pigments (VAZ) was studied in Xanthoria parietina repeatedly sampled in one location in S Norway during one year. The seasonal course in the susceptibility to photoinhibition was evaluated as high light-induced changes (1,800 micromol photons m(-2) s(-1) for 24h) in F(V)/F(M), Phi(II), and NPQ, measured as the ability to recover after 2 and 20 h at low light in control thalli with a natural cortical parietin screen, and in thalli from which parietin had been removed prior to high light exposures. F(V)/F(M), Phi(II), chlorophyll content, and the conversion state of VAZ (DEPS) reached minimum in spring. At the same time, yearly maxima of VAZ content and NPQ were recorded. Thereafter, F(V)/F(M), Phi(II), and chlorophyll content increased gradually, reaching maximum values in late autumn. DEPS peaked already in summer. Similarly, VAZ and NPQ decreased from early summer until winter. All data show that the X. parietina photobiont acclimates to seasonal changes in solar radiation, consistent with the lichen's preference for well-lit habitats. However, a comparison with a study of seasonal acclimation in the X. parietina mycobiont shows that in order to understand the seasonal photobiont acclimation, one has to consider the seasonal variation in internal screening caused by the fungal regulation of the PAR-absorbing parietin. A joint effort of both bionts seems to be required to avoid serious photoinhibition.

This study aims to assess biomass and area growth of 600 thalli of the old forest lichen, Lobaria pulmonaria, transplanted to three successional boreal forest stands with (1) natural rainfall regime, (2) additional moistening during dry days, and (3) additional moistening with added nutrients. Mean biomass growth during 100 days varied from 8.3% in the dark young spruce forest to 23.1% in the clear-cut area, with the old forest in between (16.0%). Additional moistening did not enhance lichen growth, probably because the transplantation period was wet. Nutrient additions slightly increased area growth compared to artificial water additions only. Growth was determined by a combination of external (forest stand, site factors) and internal factors (chlorophyll content, biomass per area). Transplants acclimated to high light by increasing thickness and chlorophyll a/b-ratio. Some visible bleaching and a strong positive correlation between chlorophyll content per area and lichen growth in clear-cuts suggest some high light-induced chlorophyll degradation. We believe that biomass growth and natural occurrence of L. pulmonaria is controlled by a delicate balance between light availability and desiccation risk, and that the species is confined to old forests due to a physiological trade-off between growth potential and fatal desiccation damage, both of which increase with increasing light. The discrepancy between potential and realized ecological niches is probably caused by a long-term risk to be killed in open habitats by high light during long periods with no rain.

Lichens are well-suited organisms for experimental herbivory studies because their secondary compounds, assumed to deter grazing, can be non-destructively extracted. Thalli of 17 lichen species from various habitats were cut in two equal parts; compounds were extracted from one part by acetone, the other served as a control. These two pieces were offered as a paired choice to the generalist herbivore snail Cepaea hortensis. Control thalli of all lichens were consumed at a low rate regardless of their investments in acetone-extractable lichen compounds; naturally compound-deficient lichen species were not preferred compared to those with high contents. However, for extracted thalli, there was a highly significant positive correlation between rate of consumption and the extracted compound contents. These data imply that herbivore defence has evolved in different directions in different lichens. Studied members of Parmeliaceae, common in oligotrophic habitats, have high contents of carbon-rich acetone-soluble compounds; these lichens became highly palatable to snails subsequent to acetone rinsing. Extracted lichen compounds were applied to pieces of filter paper and fed to snails. Extracts from members of the Parmeliaceae significantly deterred feeding on paper. Such data suggest that generalist herbivores may have shaped evolution in the widespread and highly diverse Parmeliaceae towards high investments in lichen compounds. On the other hand, lichens belonging to the Physciaceae and Teloschistales, common in nutrient-enriched habitats, are deficient in, or have low concentrations of, lichen compounds. Such lichens did not become more palatable after acetone rinsing. The orange anthraquinone compound parietin, restricted to the Teloschistales, and which has previously been found to protect against excess light, did not deter grazing.