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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.

Lichens occur in most terrestrial ecosystems; they are often present as minor contributors, but in some forests, drylands and tundras they can make up most of the ground layer biomass. As such, lichens dominate approximately 8% of the Earth's land surface. Despite their potential importance in driving ecosystem biogeochemistry, the influence of lichens on community processes and ecosystem functioning have attracted relatively little attention. Here, we review the role of lichens in terrestrial ecosystems and draw attention to the important, but often overlooked role of lichens as determinants of ecological processes. We start by assessing characteristics that vary among lichens and that may be important in determining their ecological role; these include their growth form, the types of photobionts that they contain, their key functional traits, their water-holding capacity, their colour, and the levels of secondary compounds in their thalli. We then assess how these differences among lichens influence their impacts on ecosystem and community processes. As such, we consider the consequences of these differences for determining the impacts of lichens on ecosystem nutrient inputs and fluxes, on the loss of mass and nutrients during lichen thallus decomposition, and on the role of lichenivorous invertebrates in moderating decomposition. We then consider how differences among lichens impact on their interactions with consumer organisms that utilize lichen thalli, and that range in size from microfauna (for which the primary role of lichens is habitat provision) to large mammals (for which lichens are primarily a food source). We then address how differences among lichens impact on plants, through for example increasing nutrient inputs and availability during primary succession, and serving as a filter for plant seedling establishment. Finally we identify areas in need of further work for better understanding the role of lichens in terrestrial ecosystems. These include understanding how the high intraspecific trait variation that characterizes many lichens impacts on community assembly processes and ecosystem functioning, how multiple species mixtures of lichens affect the key community- and ecosystem-level processes that they drive, the extent to which lichens in early succession influence vascular plant succession and ecosystem development in the longer term, and how global change drivers may impact on ecosystem functioning through altering the functional composition of lichen communities., (© 2016 Cambridge Philosophical Society.)

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.)

While there has been much interest in the relationships between traits of primary producers and composition of associated invertebrate consumer communities, our knowledge is largely based on studies from vascular plants, while other types of functionally important producers, such as lichens, have rarely been considered. To address how physiological traits of lichens drive community composition of invertebrates, we collected thalli from 27 lichen species from southern Norway and quantified the communities of associated springtails, mites, and nematodes. For each lichen species, we measured key physiological thallus traits and determined whether invertebrate communities were correlated with these traits. We also explored whether invertebrate communities differed among lichen groups, categorized according to nitrogen-fixing ability, growth form, and substratum. Lichen traits explained up to 39% of the variation in abundances of major invertebrate groups. For many invertebrate groups, abundance was positively correlated with lichen N and P concentrations, N:P ratio, and the percentage of water content on saturation (WC), but had few relationships with concentrations of carbon-based secondary compounds. Diversity and taxonomic richness of invertebrate groups were sometimes also correlated with lichen N and N:P ratios. Nitrogen-fixing lichens showed higher abundance and diversity of some invertebrate groups than did non-N-fixing lichens. However, this emerged in part because most N-fixing lichens have a foliose growth form that benefits invertebrates, through, improving the microclimate, independently of N concentration. Furthermore, invertebrate communities associated with terricolous lichens were determined more by their close proximity to the soil invertebrate pool than by lichen traits. Overall, our results reveal that differences between lichen species have a large impact on the invertebrate communities that live among the thalli. Different invertebrate groups show contrasting responses to traits that are indicative of thallus quality (nutrient concentrations), and thallus growth form is often an important determinant of the invertebrate community. Given the large diversity of lichen traits and growth forms that occur in many ecosystems, lichen-invertebrate communities may be an important contributor to overall community diversity in boreal forests.

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.)

In the long-term absence of major disturbances ecosystems enter a state of retrogression, which involves declining soil fertility and consequently a reduction in decomposition rates. Recent studies have looked at how plant traits such as specific leaf mass and amounts of secondary compounds respond to declining soil fertility during retrogression, but there are no comparable studies for lichen traits despite increasing recognition of the role that lichens can play in ecosystem processes. We studied a group of 30 forested islands in northern Sweden differing greatly in fire history, and collectively representing a retrogressive chronosequence, spanning 5000 years. We used this system to explore how specific thallus mass (STM) and carbon based secondary compounds (CBSCs) change in three common epiphytic lichen species (Hypogymnia phsyodes, Melanohalea olivacea and Parmelia sulcata) as soil fertility declines during this retrogression. We found that STMs of lichens increased sharply during retrogression, and for all species soil N to P ratio (which increased during retrogression) was a strong predictor of STM. When expressed per unit area, medullary CBSCs in all species and cortical CBSCs in P. sulcata increased during retrogression. Meanwhile, when expressed per unit mass, only cortical CBSCs in H. physodes responded to retrogression, and in the opposite direction. Given that lichen functional traits are likely to be important in driving ecological processes that drive nutrient and carbon cycling in the way that plant functional traits are, the changes that they undergo during retrogression could potentially be significant for the functioning of the ecosystem.

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.

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.

Lichens produce various carbon-based secondary compounds (CBSCs) in response to abiotic conditions and herbivory. Although lichen CBSCs have received considerable attention with regard to responses to UV-B exposure, very little is known about intra-specific variation across environmental gradients and their role in protection against herbivory in the Antarctic. Here we report on the variation in CBSCs of two widely distributed and common Antarctic lichens, Usnea antarctica and Umbilicaria antarctica, between sites with different solar exposure (NW–SE) and along natural nitrogen (N) gradients which are associated with changing lichen-invertebrate associations on Signy Island (South Orkney Islands, maritime Antarctic). Fumarprotocetraric and usnic acid concentrations in Usnea showed no relationships with solar exposure, lichen-N or associated invertebrate abundance. However, fumarprotocetraric acid concentration was 13 times higher at inland sites compared to coastal sites along the N-gradients. Gyrophoric acid concentration in Umbilicaria was 33% lower in sun-facing (northerly exposed) habitats compared to more shaded (south-facing) rocks and declined with elevation. Gyrophoric acid concentration was positively correlated with the abundance and species richness of associated microarthropods, similar to the patterns found with lichen N. This initial investigation indicates that there can be large intraspecific variation in lichen CBSC concentrations across relative short distances (< 500 m) on Signy Island and raises further questions regarding current understanding of the role of CBSCs in Antarctic lichens in relation to biotic and abiotic pressures. [ABSTRACT FROM AUTHOR]

Foundation species provide habitat and modify the availability of resources to other species. In nature, multiple foundation species may occur in mixture, but little is known on how their interactions shape the community assembly of associated species. Lichens provide both structural habitat and resources to a variety of associated organisms and thereby serve as foundation species. In this study, we use mat-forming lichens and their associated micro-arthropods as a miniature ecosystem to study potential synergies between foundation species diversity and the abundance and functional diversity of higher trophic levels. We created lichen patches with monocultures and mixtures of up to four species, and extracted Collembola (identified to species level), Oribatida, Mesostigmata, Pseudoscorpiones, and Araneae with Tullgren apparatuses after 106 days of incubation within a natural lichen mat. We found that different lichen species supported different arthropod abundances. For 19 out of a total of 55 lichen mixtures and arthropod groups, we found non-additive, synergistic effects on arthropod abundance, although the specific lichen mixture causing synergistic effects differed with arthropod group. In addition, synergistic effects on arthropod abundance were more common for arthropod groups at lower trophic levels. The functional diversity of lichen mixtures explained patterns in Collembola abundance, but in the opposite direction than hypothesized because synergistic responses were more frequent in functionally similar lichen mixtures. Finally, we found few effects of lichen mixture identity or diversity on the functional diversity of Collembola communities. When applied to large-scale ecosystems, our results suggest that understanding interactions between coexisting foundation species and identifying those species that drive synergistic effects of foundation species on consumer biota, is likely to be of importance to biodiversity conservation and restoration efforts. [ABSTRACT FROM AUTHOR]

Aim: Functional traits offer a window into how organisms are adapted, and might acclimate, to environmental pressures. Despite being important in ecosystem function, lichens are underrepresented in trait‐based research; understanding how lichen functional traits vary with climate and habitat availability will be useful in predicting how communities will respond to climate change, for example, in wetter and warmer boreal and arctic ecosystems. Here, we assess the influence of macroclimate and forest availability on the spatial distribution of lichen traits across Norway. Location: Norwegian mainland. Taxon: Lichens. Methods: We used relative trait frequency (RTF) data from LIAS gtm, a database combining trait information from LIAS (A Global Information System for Lichenized and Non‐Lichenized Ascomycetes) and GBIF (Global Biodiversity Information Facility) species observations. The 20 traits included photobiont types, growth forms, cortical features and reproductive modes. Nonparametric multiplicative regression (NPMR) models were used to explore the relationships between the environmental predictors of precipitation, temperature and forest availability. Results: All traits showed significant relationships with the three environmental predictors. Photobiont type and reproductive mode traits produced the strongest models and revealed ecologically meaningful biogeographical patterns. Trebouxioid species peaked in colder, drier upland regions, while trentepohlioid lichens displayed an affinity for wetter and warmer climates and had a western and southern distribution. Cyanolichens increased with increasing precipitation and were strongly coastal. Sorediate and isidiate lichens were positively related to temperature, the former also increasing with forest cover. The above responses were consistent with the physiological and habitat requirements of the associated lichens. The remaining traits had weaker responses. Main conclusions: Discrete traits (i.e. photobiont type and reproductive mode) with relatively low ecological plasticity reflect clear functional environmental responses at the large scale. By contrast, growth form and thallus structural features—proxies for continuous variables—are too variable within each given category to show observable distribution patterns. [ABSTRACT FROM AUTHOR]

Question: Cold environments are stressful for vascular plants, and stress-tolerant non-vascular photoautotrophs, e.g. bryophytes and lichens, become relatively more important as competition from vascular plants decreases towards higher elevations. Under increasingly stressful climatic conditions, species assembly of vascular plants is commonly driven more by environmental filtering, and abiotic constraints may lead to increased similarity between species and thus low functional diversity. Because bryophytes and lichens are less constrained by harsh environments, environmental filtering may be less strong. Instead, reduced competition from vascular plants can potentially free up niche space for non-vascular vegetation. Therefore, we hypothesized that functional diversity of vascular plants, bryophytes and lichens are likely to show contrasting responses to elevation. Location: Finse Alpine Research Centre, Southern Norway. Methods: We utilized measurements of species abundance and functional traits of the three groups along a 500-m elevational gradient in alpine southern Norway and calculated multi-trait and single-trait functional dispersion. Results: Functional diversity of vascular plants declined with elevation, indicating increased environmental filtering. By contrast, functional diversity of lichens and bryophytes increased along the same gradient, suggesting they are less exposed to environmental filtering, in line with our hypothesis. Instead, they likely benefit from the lower abundance of vascular plants at higher elevation. Conclusions: Our findings suggest that different photoautotroph groups vary in how they respond to the same environmental gradient, which may contribute to contrasting community assembly processes across groups. These divergent responses likely occur because non-vascular vegetation differs from vascular plants in terms of nutrient acquisition and water economy strategies, meaning that they respond differently to the same factors. This highlights the need to explicitly consider bryophytes and lichens in community-level studies whenever these groups are abundant. [ABSTRACT FROM AUTHOR]

Lichen heaths are declining in abundance in alpine and Arctic areas partly due to an increasing competition with shrubs. This shift in vegetation types might have important consequences for the microclimate and climate on a larger scale. The aim of our study is to measure the difference in microclimatic conditions between lichen heaths and shrub vegetation during the growing season. With a paired plot design, we measured the net radiation, soil heat flux, soil temperature and soil moisture on an alpine mountain area in southern Norway during the summer of 2018 and 2019. We determined that the daily net radiation of lichens was on average 3.15 MJ (26 %) lower than for shrubs during the growing season. This was mainly due to a higher albedo of the lichen heaths but also due to a larger longwave radiation loss. Subsequently, we estimate that a shift from a lichen heath to shrub vegetation leads to an average increase in atmospheric heating of 3.35 MJ d -1 during the growing season. Surprisingly, the soil heat flux and soil temperature were higher below lichens than below shrubs during days with high air temperatures. This implies that the relatively high albedo of lichens does not lead to a cooler soil compared to shrubs during the growing season. We predict that the thicker litter layer, the presence of soil shading and a higher evapotranspiration rate at shrub vegetation are far more important factors in explaining the variation in soil temperature between lichens and shrubs. Our study shows that a shift from lichen heaths to shrub vegetation in alpine and Arctic areas will lead to atmospheric heating, but it has a cooling effect on the subsurface during the growing season, especially when air temperatures are relatively high. [ABSTRACT FROM AUTHOR]

Vascular plants and lichens often produce a diversity of carbon‐based secondary compounds (CBSCs) to protect them against biotic and abiotic stresses. These compounds play important but often compound‐specific roles in community and ecosystem processes by affecting herbivore and decomposer activity. However, our understanding of what drives community‐level CBSCs among ecosystems or across environmental gradients is limited.We measured concentrations and compositions of CBSCs for all dominant vascular plant and lichen species present across a 500‐m alpine elevational gradient. These measurements were combined with data on species composition and abundance to obtain whole‐community measures of plant and lichen CBSCs across the gradient.At the whole community level, plant CBSCs had the lowest concentrations while lichen CBSCs had the highest concentrations at the highest elevations. Further, plant CBCSs shifted from those associated with herbivore defence towards those protecting against light and oxidative stress as elevation increased, while lichen CBSCs showed the opposite pattern.Synthesis. Our findings that individual compounds show contrasting responses to the same environmental gradient highlight the importance of studying qualitative as well as quantitative changes in CBSCs. Further, the divergent responses between vascular plants and lichens reveal that in systems where both groups are abundant, they need to be considered simultaneously to better understand how future environmental changes may impact on ecosystem‐level processes. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Lichen heaths are declining in abundance in alpine and arctic areas partly due to an increasing competition with shrubs. This shift in vegetation types might have important consequences for the microclimate and climate on a larger scale. The aim of our study is to measure the difference in microclimatic conditions between lichen heaths and shrub vegetation during the growing season. With a paired plot design, we measured the net radiation, soil heat flux, soil temperature, and soil moisture on an alpine mountain area in south Norway during the summer of 2018 and 2019. We determined that the daily net radiation of lichens was on average 3.15 MJ (26 %) lower than for shrubs during the growing season. This was mainly due to a higher albedo of the lichen heaths, but also due to a larger longwave radiation loss. Subsequently, we estimate that a shift from a lichen heath to shrub vegetation leads to an average increase in atmospheric heating of 3.35 MJ per day during the growing season. Surprisingly, the soil heat flux and soil temperature were higher below lichens than below shrubs during days with high air temperatures. This implies that the relatively high albedo of lichens does not lead to a cooler soil compared to shrubs during the growing season. We hypothesize that the thicker litter layer, the presence of soil shading, and a higher evapotranspiration rate at shrub vegetation are far more important factors in explaining the variation in soil temperature between lichens and shrubs. Our study shows that a shift from lichen heaths to shrub vegetation in alpine and arctic areas will lead to atmospheric heating, but has a cooling effect on the subsurface during the growing season, especially when air temperatures are relatively high. [ABSTRACT FROM AUTHOR]

Lichens and bryophytes are abundant primary producers in high latitude and high elevation ecosystems, and they play an important role in ecosystem processes such as decomposition and nutrient cycling. Despite their importance, little is known about the decomposability of lichens and bryophytes either among or within species, at the whole community level, or how this decomposability is affected by their functional traits. Here, we studied decomposability of lichens and bryophytes at the community‐level and individual species‐level (using 21 species and genera) collected from an elevational gradient in alpine Norway. In order to isolate the elevation effect on litter quality, we used a standardized laboratory bioassay to measure decomposability. In contrast to our expectations, we found that community‐level decomposability of lichens and bryophytes increased with elevation and thus decreasing temperature. In contrast, phosphorus release from the litter decreased with elevation while nitrogen release was unresponsive. Decomposability was explained by nutrient concentrations, litter pH and primary producer group identity (lichens versus bryophytes) at both the individual species and community levels. Species turnover (changes in species composition and abundance) was the main driver of decomposability across elevation at the community level, despite some of the traits explaining decomposability showing high intraspecific variability. Our study highlights the importance of among‐species variation in determining lichen and bryophyte decomposability. Further, the higher decomposability that we found for higher elevations suggests that global warming might result in a shift towards slower decomposable lichen and bryophyte species. [ABSTRACT FROM AUTHOR]

Lichen heaths are declining in abundance while shrubs are increasing their range in alpine and arctic areas due to climate change. This can have a large impact on the surface albedo of these areas. The aim of this article is to quantify the difference in albedo between lichen heaths and shrub-dominated vegetation and the variability within lichen heaths. Several environmental conditions that can influence the albedo measurements are considered. We measured the albedo of twenty lichen- and shrub-dominated plots on an alpine mountain area in southern Norway in the summer of 2018 with two radiometers using a paired plot design. With this design, we ensured similar weather conditions, aspects, and zenith angles between the paired lichen- and shrubdominated plots. In addition, we collected patches of Cladonia stellaris and Flavocetraria nivalis to measure their albedo. The average difference in albedo between the lichen- and shrub-dominated plots is 0.124. The albedo of the lichen-dominated plots varies between 0.227 and 0.284, and that of the shrub-dominated plots varies between 0.115 and 0.148. This variation in albedo is explained by differences in aspect and vegetation composition. Further studies should focus on the consequences of this decrease in albedo for the microclimate in alpine and arctic areas. [ABSTRACT FROM AUTHOR]

Across environmental gradients, community‐level functional traits of plants can change due to species turnover, intraspecific variation and their covariation. Studies on vascular plants suggest that species turnover is the main driver of trait variation across gradients, although intraspecific variation can also be important. However, there is limited knowledge about whether this holds for non‐vascular primary producers such as lichens and bryophytes. We hypothesized that intraspecific variation is more important for non‐vascular than for vascular primary producers because they lack specialized structures to maintain homeostasis and should therefore be more responsive to extrinsic factors.To assess the relative importance of species turnover versus intraspecific variation for vascular plants, lichens and bryophytes, we estimated species abundance and measured chemical (tissue nitrogen (N) and phosphorous (P) content, N:P ratio and pH) and non‐chemical (specific leaf or thallus area, dry matter content and water holding capacity) functional traits along an elevational gradient in alpine southern Norway. We calculated community‐weighted mean traits and quantified the relative contribution of species turnover, intraspecific variation and their covariation to total trait variation across the gradient.We found mixed support for our hypothesis: the contribution of intraspecific variation to total trait variation for N and N:P was higher in lichens than in vascular plants and bryophytes, but in general the contribution of intraspecific variation differed among functional traits and producer groups. Nutrient variables (N, P and N:P) were significantly impacted by intraspecific variation for vascular plants and lichens but not for bryophytes. Non‐chemical traits and pH were mainly driven by species turnover effects in all primary producer groups.Our results highlight that while nearly all studies on primary producer trait variation across environments have focused on vascular plants, trait variation of other largely neglected but ecologically important producer groups, such as lichens and bryophytes, may show very different responses to the same environmental factors. In order to fully understand how future environmental changes impact on community‐ and ecosystem‐level processes, traits of primary producers other than vascular plants—and their within‐species variation—need to be considered in systems where these groups are abundant. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

The article discusses research which examined whether thalli infected with lichenicolous fungus Plectocarpon lichenum have lower synthesis of lichen produce carbon-based secondary compounds (CBSC). Topics discussed include the reduction in the total CBSC concentration in healthy thalli with acetone treatment and adverse influence of Plectocarpon lichenum on its lichen host by reducing CBSC and growth.

A limited understanding of how variation in the species composition among communities (i.e., β-diversity) changes along natural environmental gradients, and the mechanisms responsible, inhibits our ability to understand large-scale biodiversity change resulting from either natural or anthropogenic drivers. Therefore, our aim was to test key drivers of β-diversity patterns along a strong, natural environmental gradient for seven widely different organisms groups, that is, root-associated fungi, litter fungi, soil nematodes, vascular plants, epiphytic lichens, beetles, and spiders. Using previously published community-level data from boreal-forested islands, we calculated α-diversity and β-diversity for each of the seven organism groups. Out of several available environmental variables, we identified four variables, that is, ecosystem age, total C storage, net primary productivity (NPP), and N-to-P ratio, as potential predictors of variation in β-diversity. We found that ecosystem age was the variable with the highest overall importance. We then used two different methods to quantify the relative importance of stochastic and deterministic processes underlying patterns in β-diversity along the ecosystem age gradient, and our detailed knowledge based on prior data collection in the study system to mechanistically explain amonggroup differences in these patterns. We found divergent responses in β-diversity along the age gradient for the seven different organism groups, due to among-group differences in the relative importance of deterministic vs. stochastic community assembly, and attributed these results to reliance on resources from different energy channels that are not always related to NPP. Our results highlight the necessity to consider the importance of taxon-specific resources, and not only NPP, to obtain an understanding of βdiversity patterns among organism groups and ecosystems, as well as large-scale patterns in biodiversity. They therefore also suggest that management and protection of β-biodiversity in the landscape requires explicit consideration of a wide range of habitats. [ABSTRACT FROM AUTHOR]

Abstract: Phenolic compounds have been shown in several studies to have important ‘carryover effects’ on litter decomposition, microbial nutrient immobilization and nutrient availability. These effects arise in part because of the adverse effect they have on the feeding activities of litter-feeding invertebrates such as micro-arthropods that drive decomposition processes. However, the interactive effects of phenolic compounds and soil micro-arthropods on litter decomposition are poorly understood. Phenolic compounds can easily be removed by acetone rinsing from living lichens, allowing us to specifically test the role that phenolic compounds (and their removal) have in controlling the effects of micro-arthropods on the decomposition of their litter. We performed a litter-bag experiment aimed at exploring how lichen litter mass loss and nutrient release during decomposition was affected by phenolics (by using acetone rinsed and non-rinsed lichen material) and micro-arthropod activity (by using different mesh sizes to allow or exclude entry by micro-arthropods) for each of six contrasting lichen species (Cladonia rangiferina, Cladonia stellaris, Evernia prunastri, Hypogymnia physodes, Pseudevernia furfuracea and Usnea dasypoga). Both the removal of phenolic compounds and the presence of micro-arthropods accelerated mass and nutrient release overall, but not for either of the two Cladonia species. Removal of phenolics also had an overall positive effect on the effects of arthropods on the loss of P, but not mass and N, from the decomposing lichens. Further, for U. dasypoga, but not the other species, natural levels of phenolic compounds deterred micro-arthropods from accelerating mass loss, and the removal of these compounds enabled micro-arthropods to enhance its decomposition. Our findings that lichen phenolic compounds can sometimes interact with micro-arthropods to influence lichen litter mass loss and nutrient release during decomposition assists our understanding of how lichens and their consumers may impact on organic matter dynamics, biochemical nutrient cycling and other related ecosystem processes. [Copyright &y& Elsevier]

Abstract: Epiphytic lichens can contribute significantly to ecosystem nutrient input because they efficiently accumulate atmospheric mineral nutrients and, in the case of cyanolichens, also fix nitrogen. The rate at which carbon and other nutrients gained by lichens enters the ecosystem is determined by lichen litter decomposability and by invertebrate consumption of lichen litter. In turn, these processes are driven by the secondary compounds present in senesced lichens. Therefore, we explored how lichen palatability and concentrations of secondary compounds change with tissue senescence for Lobaria pulmonaria, a green-algal lichen with cyanobacterial cephalodia, and Lobaria scrobiculata, a cyanobacterial lichen. During senescence both lichens lost 38–48 % of their stictic acid chemosyndrome, while m-scrobiculin and usnic acid in L. scrobiculata remained unchanged. Snails preferred senesced rather than fresh L. pulmonaria, while senesced L. scrobiculata were avoided. This provides evidence that species with labile secondary compounds will have higher turnover rates, through consumption and decomposition, than those producing more stable secondary compounds. [Copyright &y& Elsevier]

Abstract: We quantified carbon based secondary compounds (CBSC) in 182 Lobaria pulmonaria thalli sampled in a range of forest types along an elevation gradient in southern Norway to assess the importance of external and internal factors as a source of variation for the CBSC content. The best multiple regression model for CBSC content included altitude, soil pH, dry weight (DW) per area and thallus size. Total CBSC content decreased with altitude and increased with pH. L. pulmonaria maximizes its CBSC content in sites with long gastropod grazing seasons (lowland forests) and high gastropod abundance (calcareous soils). Secondly, we assessed grazing damage in the L. pulmonaria thalli to test the hypothesis that natural populations of gastropods respond to the CBSC level in situ. We found a significantly negative correlation between the stictic acid concentration and grazing damage from gastropods, which is consistent with an evolutionary arms race between lichens and gastropods. This relationship became stronger when using the tree trunk as a factor in the ANCOVA, consistent with spatial variation in gastropod occurrences. [Copyright &y& Elsevier]

The article focuses on the study which investigates whether the palatability to lichenivorous gastropods differs with the two chemical races of Lobaria pulmonaria. It says that high performance liquid chromatography (HPLC), online ultraviolet (UV)-spectra , and co-chromatography of norstictic acid were performed. It shows that 28 of the 39 thalli from L. pulmonaria belonged to chemical race I, while all of the chemical race II thalli were less or more grazed.

Thalli of Lobaria pulmonaria of different sizes were collected from the trunks of two specimens of Populus tremula. The secondary lichen compounds, stictic, constictic, norstictic, peristictic, cryptostictic and methyl stictic acid were quantified by HPLC and ranked in order of decreasing concentration. There was a highly significant positive correlation between thallus size and the total concentration of secondary compounds, as well as the total content per unit area for specimens from the two sampled trees. According to hypotheses inferring a herbivore deterrent role of secondary lichen compounds, small, juvenile thalli should be more susceptible to herbivores than larger thalli. Thus herbivory might limit survival of young specimens in habitats rich in lichen-feeding molluscs and thereby reduce reproductive success. [ABSTRACT FROM AUTHOR]

Community ecology has experienced a major transition, from a focus on patterns in taxonomic composition, to revealing the processes underlying community assembly through the analysis of species functional traits. The power of the functional trait approach is its generality, predictive capacity such as with respect to environmental change, and, through linkage of response and effect traits, the synthesis of community assembly with ecosystem function and services. Lichens are a potentially rich source of information about how traits govern community structure and function, thereby creating opportunity to better integrate lichens into 'mainstream' ecological studies, while lichen ecology and conservation can also benefit from using the trait approach as an investigative tool. This paper brings together a range of author perspectives to review the use of traits in lichenology, particularly with respect to European ecosystems from the Mediterranean to the Arctic-Alpine. It emphasizes the types of traits that lichenologists have used in their studies, both response and effect, the bundling of traits towards the evolution of life-history strategies, and the critical importance of scale (both spatial and temporal) in functional trait ecology. [ABSTRACT FROM AUTHOR]