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Maritime Antarctica's harsh abiotic conditions forged simple terrestrial ecosystems, mostly constituted of bryophytes, lichens, and vascular plants. Though biotic interactions are, together with abiotic factors, thought to help shape this ecosystem, influencing species' distribution and, indirectly, mediating their response to climate, the importance of these interactions is still fairly unknown. We modeled current and future abundance patterns of bryophytes, lichens, and vascular plants, accounting for biotic interactions and abiotic drivers, along a climatic gradient in maritime Antarctica. The influence of regional climate and other drivers was modeled using structural equation models, with and without biotic interactions. Models with biotic interactions performed better; the one offering higher ecological support was used to estimate current and future spatial distributions of vegetation. Results suggest that plants are confined to lower elevations, negatively impacting bryophytes and lichens, whereas at higher elevations both climate and other drivers influence bryophytes and lichens. Our findings strongly support the use of biotic interactions to predict the spatial distribution of Antarctic vegetation., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Lichens are a mutualistic symbiosis between a fungus and one or more photosynthetic partners. They are photosynthetically active during desiccation down to relative water contents (RWCs) as low as 30% (on dry mass). Experimental evidence suggests that during desiccation, the photobionts have a higher hydration level than the surrounding fungal pseudo-tissues. Explosive cavitation events in the hyphae might cause water movements towards the photobionts. This hypothesis was tested in two foliose lichens by measurements of ultrasonic acoustic emissions (UAEs), a method commonly used in vascular plants but never in lichens, and by measurements of PSII efficiency, water potential, and RWC. Thallus structural changes were characterized by low-temperature scanning electron microscopy. The thalli were silent between 380% and 30% RWCs, when explosive cavitation events should cause movements of liquid water. Nevertheless, the thalli emitted UAEs at ~5% RWC. Accordingly, the medullary hyphae were partially shrunken at ~15% RWC, whereas they were completely shrunken at <5% RWC. These results do not support the hypothesis of hyphal cavitation and suggest that the UAEs originate from structural changes at hyphal level. The shrinking of hyphae is proposed as an adaptation to avoid cell damage at very low RWCs., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Despite being one of the most pristine regions in the world, Antarctica is currently also one of the most vulnerable to climate change. Antarctic vegetation comprises mostly lichens and bryophytes, complemented in some milder regions of Maritime Antarctica by two vascular plant species. Shifts in the spatial patterns of these three main vegetation groups have already been observed in response to climate change, highlighting the urgent need for the development of comprehensive large-scale ecological models of the effects of climate change. Besides climate, Antarctic terrestrial vegetation is also strongly influenced by non-climatic microscale conditions related to abiotic and biotic factors. Nevertheless, the quantification of their importance in determining vegetation patterns remains unclear. The objective of this work was to quantify the importance of abiotic and biotic microscale conditions in determining the spatial cover patterns of the major functional types, lichens, vascular plants and bryophytes, explicitly determining the likely confinement of each functional type to the microscale conditions, i.e., their ecological niche. Microscale explained >60 % of the spatial variation of lichens and bryophytes and 30 % of vascular plants, with the niche analysis suggesting that each of the three functional types may be likely confined to specific microscale conditions in the studied gradient. Models indicate that the main microscale ecological filters are abiotic but show the potential benefits of including biotic variables and point to the need for further clarification of vegetation biotic interactions' role in these ecosystems. Altogether, these results point to the need for the inclusion of microscale drivers in ecological models to track and forecast climate change effects, as they are crucial to explain present vegetation patterns in response to climate, and for the interpretation of ecological model results under a climate change perspective., 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 © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Lichens are increasingly recognised as valuable bioindicators for environmental heavy metal pollution due to their sensitivity to spatial and temporal variations in pollution levels and their ability to adapt to diverse and often harsh habitats. This review initially examines the mechanisms of metal absorption in lichens, including particulate entrapment, ion exchange, and intracellular absorption, as well as their physiological responses to abiotic stressors such as heavy metal exposure and desiccation. In the latter part, we compile and synthesise evidence showing that secondary metabolites in lichens are significantly influenced by metal concentrations, with varying impacts across different species. Although extensive research has addressed the broader physiological effects of heavy metal hyperaccumulation in lichens, there remains a significant gap in understanding the direct or indirect influences of heavy metals on the lichen microbiome, possibly mediated by changes in secondary metabolite production. Our review integrates these aspects to propose new research directions aimed at elucidating the mechanisms underlying physiological responses such as resilience and adaptability in lichens. Overall, this review highlights the dynamic interplay between microbiome composition, secondary metabolite variation, and metal accumulation, suggesting that these factors collectively contribute to the physiological responses of lichens in polluted environments., (© 2024. The Author(s), under exclusive licence to Springer Nature Switzerland AG.)

The fruticose lichen Stereocaulon vesuvianum is among the most abundant and widespread lichens in upland Britain. It typically produces cephalodia (nodules) that contain the cyanobacterium Stigonema, which can fix atmospheric nitrogen. However, over much of England, Wales, and southern Scotland S. vesuvianum no longer produces cephalodia and does not fix nitrogen, a morphological change linked to elevated atmospheric nitrogen deposition. This provided a unique opportunity to compare the 15 N natural abundance signatures in N 2 -fixing and non-N 2 -fixing lichen populations, keeping in mind that fixed nitrogen has a 15 N content close to that of atmospheric N 2 while, in comparison, several components of atmospheric combined N (e.g. nitrate and ammonium in precipitation) tend to be 15 N depleted. We found that in N 2 -fixing samples, there was a steep gradient in 15 N relative abundance in the terminal 15 mm of thallus branches (pseudopodetia), changing from 15 N depleted tissues at 10-15 mm below the tips to values close to that of atmospheric N 2 at the apices while in non-N 2 -fixing samples thallus branches were uniformly 15 N depleted. The 15 N gradient in N 2 -fixing material could not be explained by the presence of cephalodia since these are more abundant towards branch bases. The data provide the first evidence in lichens of translocation of recently fixed N to sink regions of active growth and production of asexual reproductive propagules, bringing lichens into line with N source-sink relationships in N 2 -fixing plant symbioses., (© 2024 The Author(s). Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

Lichens are important components of high-latitude boreal and Arctic habitats. While stress tolerant, they are among the most sensitive ecosystem components to climate change, in particular, an increase in ultraviolet light (UV) arising from polar ozone depletion and deforestation. This study is the first to explore the effects of UV-B on gene expression in lichens to predict metabolic pathways involved in tolerance. Using transcriptome profiling and bioinformatic analyses, here we studied the effects of UV-B on gene expression in lichens using Lobaria pulmonaria (L.) Hoff. as a model species. UV-B exposure causes significant browning of the upper cortex of the thallus, which correlates to an increased expression of biosynthetic gene clusters involved in the synthesis of eu- and allomelanins and melanin precursors. Based on transcriptome analyses, we suggest that the biosynthesis of melanins and other secondary metabolites, such as naphthalene derivates, tropolones, anthraquinones, and xanthones, is a trade-off that lichens pay to protect essential metabolic processes such as photosynthesis and respiration. Expression profiles of general stress-associated genes, in particular, related to reactive oxygen species scavenging, protection of proteins, and DNA repair, clearly indicate that the mycobiont is the more UV-B-responsive and susceptible partner in lichen symbiosis. Our findings demonstrate that UV-B stress activates an intricate gene network involved in tolerance mechanisms of lichen symbionts. Knowledge obtained here may enable the prediction of likely effects on lichen biodiversity caused by climate change and pollution., (© 2024 Scandinavian Plant Physiology Society.)

Soil salinization is a major environmental threat to the entire terrestrial ecosystem. Lichens arose from the symbiosis of fungi and algae or cyanobacteria. They have a high tolerance to various extreme environments, including adaptation to saline-alkali habitats. Thus, lichens are pioneer species on saline-alkali soil. However, the separate resilience of the two symbiotic partners under saline-alkali conditions remains insufficiently understood. In this study, two representative symbiotic algae, Diplosphaera chodatii and Trebouxia jamesii, were studied for their physiological response to the saline-alkali stress by adjusting different concentrations of NaHCO 3 , together with their respective symbiotic fungi Endocarpon pusillum (terricolous lichen) and Umbilicaria muhlenbergii (saxicolous lichen). The results indicate that cell growth rate and biomass in all four cultures decreased in alkali-alkaline substrate, while cellular activities and ultrastructure were affected to a distinct extent. Compared with the symbiotic fungi, the algae were found to be more active in coordinating oxidative stress and lipid peroxidation damage under the saline-alkali stress. The antioxidant system of the alga was especially shown as a key adaptive trait and it provides an important strategy for species survival and persistence in arid saline-alkali desert. The specific survival ability of the lichen symbiosis relies on the stress resilience advantages of the symbiotic partners in combination. Our study provided new insights into understanding the adaptation of lichen symbiosis to desert saline-alkali soil, and the potential of lichen symbiotic algae in the future desert ecological restoration., (© 2024 Scandinavian Plant Physiology Society.)

The analysis of the interaction between main bionts (mycobiont and photobiont) in the lichen symbiosis delivers substantial information about their preferences in the selection of symbiotic partners, and their ecological preferences. The selectivity in the Parmelia genus has been defined as strong so far. However, data on this lichen genus, which includes several widely distributed species, are biogeographically limited. Therefore, using specialization indicators and extended sampling, in this study, we estimated the interactions between the main bionts of selected Parmelia spp., using two levels of estimation (species/OTU and haplotype). A comparison of mycobiont-photobiont interactions at different levels showed that considering only mycobiont species and Trebouxia OTUs, greater specialization is found, while Parmelia species studied in this work present a more generalistic strategy in photobiont choice when haplotypes are considered. Despite the uneven sampling of Parmelia species, the interpretation of specialization within species and individuals of the genus leads to a more precise and accurate interpretation of their adaptation strategies. Furthermore, the data from P. sulcata indicate the existence of a different pool of compatible haplotypes in some geographical regions compared to neighboring areas. This observation suggests the potential influence of climatic factors., (© 2024. The Author(s).)

Ecophysiological studies of lichens in tropical Asia are rare, and additional studies can increase the understanding of lichen life in this region. The main aim of this study was to observe the relationships between water availability and photosynthetic parameters, as well as hydration trait parameters, in macrolichens during the rainy and dry seasons in a tropical forest. A total of 11 lichen species growing in a lower montane rainforest in Thailand were collected and studied. The results clearly showed that the specific thallus mass (STM), net photosynthetic rate (Pn), the potential quantum yield of primary photochemistry (Fv/Fm), chlorophyll content, and carotenoid content of almost all lichens were lower in the dry season than in the rainy season. Field measurements in the dry season revealed that only the foliose chlorolichen Parmotrema tinctorum was metabolically active and exhibited slight carbon assimilation. In the rainy season, all lichens started their photosynthesis in the early morning, reached maximal values, declined, and ceased when the thalli desiccated. The photosynthetically active period of the lichens was approximately 2-3 h in the morning, and the activities of the cyanolichens ended approximately 30 min after the chlorolichens. The hydration trait parameters, including the STM, maximal water content (WC max ), and water holding capacity (WHC), were greater in the cyanolichens. In addition, the maximal Pn (Pn max ) and optimal water content (WC opt ) for Pn were also greater in the cyanolichens, but the maximal Fv/Fm (Fv/Fm max ) was lower. The cyanolichens compensated for their inability to use humid air to restore photosynthesis by having higher water content and storage, higher photosynthetic rates, and longer photosynthetically active periods. This study provides additional insights into lichen ecophysiology in tropical forests that can be useful for lichen conservation., (© 2024. The Author(s) under exclusive licence to The Botanical Society of Japan.)

Black-and-white snub-nosed monkeys (Rhinopithecus bieti) rely on behavioral and dietary flexibility to survive in temperate latitudes at high-elevation habitats characterized by climate and resource seasonality. However, little is known about how elevation influences their behavioral and dietary flexibility at monthly or seasonal scales. We studied an isolated R. bieti population at Mt. Lasha in the Yunling Provincial Nature Reserve, Yunnan, China, between May 2008 and August 2016 to assess the impacts of elevation on feeding behavior and diet. Across our sample, R. bieti occupied elevations between 3031 and 3637 m above mean sea level (amsl), with a 315.1 m amsl range across months and a 247.3 m amsl range across seasons. Contrary to expectations, individuals spent less time feeding when ranging across higher elevations. Lichen consumption correlated with elevation use across months and seasons, with individuals spending more time feeding on this important resource at higher elevations. Leaf consumption only correlated with elevation use during the spring. Our results suggest that R. bieti do not maximize their food intake at higher elevations and that monthly and seasonal changes in lichen and leaf consumption largely explain variation in elevation use. These findings shed light on the responses of R. bieti to environmental change and offer insight into strategies for conserving their habitats in the face of anthropogenic disturbance., (© 2024 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

The primary purpose of the study, as part of the planned conservation work, was to uncover all aspects of autochthonous biofilm pertaining to the formation of numerous deterioration symptoms occurring on the limestone Rožanec Mithraeum monument in Slovenia. Using state-of-the-art sequencing technologies combining mycobiome data with observations made via numerous light and spectroscopic (FTIR and Raman) microscopy analyses pointed out to epilithic lichen Gyalecta jenensis and its photobiont, carotenoid-rich Trentepohlia aurea, as the origin of salmon-hued pigmented alterations of limestone surface. Furthermore, the development of the main deterioration symptom on the monument, i.e., biopitting, was instigated by the formation of typical endolithic thalli and ascomata of representative Verrucariaceae family (Verrucaria sp.) in conjunction with the oxalic acid-mediated dissolution of limestone. The domination of lichenized fungi, as the main deterioration agents, both on the relief and surrounding limestone, was additionally supported by the high relative abundance of lichenized and symbiotroph groups in FUNGuild analysis. Obtained results not only upgraded knowledge of this frequently occurring but often overlooked group of extremophilic stone heritage deteriogens but also provided a necessary groundwork for the development of efficient biocontrol formulation applicable in situ for the preservation of similarly affected limestone monuments., (© 2024. The Author(s).)

A core vulnerability in symbioses is the need for coordination between the symbiotic partners, which are often assumed to be closely physiologically integrated. We critically re-examine this assumed integration between symbionts in lichen symbioses, recovering a long overlooked yet fundamental physiological asymmetry in carbon balance. We examine the physiological, ecological, and transcriptional basis of this asymmetry in the lichen Evernia mesomorpha . This carbon balance asymmetry depends on hydration source and aligns with climatic range limits. Differences in gene expression across the E. mesomorpha symbiosis suggest that the physiologies of the primary lichen symbionts are decoupled. Furthermore, we use gas exchange data to show that asymmetries in carbon balance are widespread and common across evolutionarily disparate lichen associations. Using carbon balance asymmetry as an example, we provide evidence for the wide-ranging importance of physiological asymmetries in symbioses.

Background and Aims: There is growing interest in the functional ecology of poikilohydric non-vascular photoautotrophs (NVPs), including 'cryptogamic' bryophytes and lichens. These organisms are structurally important in many ecosystems, contributing substantially to ecosystem function and services, while also being sensitive to climate change. Previous research has quantified the climate change response of poikilohydric NVPs using predictive bioclimatic models with standard climate variables including precipitation totals and temperature averages. This study aimed for an improved functional understanding of their climate change response based on their growth rate sensitivity to moisture and light., Methods: We conducted a 24-month experiment to monitor lichen hydration and growth. We accounted for two well-known features in the ecology of poikilohydric NVPs, and exemplified here for a structurally dominant lichen epiphyte, Lobaria pulmonaria: (1) sensitivity to multiple sources of atmospheric moisture including rain, condensed dew-formation and water vapour; and (2) growth determined by the amount of time hydrated in the light, driving photosynthesis, referred to as the Iwet hypothesis., Key Results: First, we found that even within an oceanic high-rainfall environment, lichen hydration was better explained by vapour pressure deficit than precipitation totals. Second, growth at a monthly resolution was positively related to the amount of time spent hydrated in the light, and negatively related to the amount of time spent hydrated in the dark., Conclusions: Using multimodel averaging to project growth models for an ensemble of future climate change scenarios, we demonstrated reduced net growth for L. pulmonaria by the late 21st century, explained by extended climate dryness and lichen desiccation for periods when there is otherwise sufficient light to drive photosynthesis. The results further emphasize a key issue of photoperiodism when constructing functionally relevant models to understand the risk of climate change, especially for poikilohydric NVPs., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Ecological interactions and symbiosis between algae and fungi are ancient, widespread, and diverse with many independent origins. The heterotrophic constraint on fungal nutrition drives fungal interactions with autotrophic organisms, including algae. While ancestors of modern fungi may have evolved as parasites of algae, there remains a latent ability in algae to detect and respond to fungi through a range of symbioses that are witnessed today in the astounding diversity of lichens, associations with corticoid and polypore fungi, and endophytic associations with macroalgae. Research into algal-fungal interactions and biotechnological innovation have the potential to improve our understanding of their diversity and functions in natural systems, and to harness this knowledge to develop sustainable and novel approaches for producing food, energy, and bioproducts., Competing Interests: Declaration of Competing Interest The author declares that he has no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The Lobaria pulmonaria holobiont comprises algal, fungal, cyanobacterial and bacterial components. We investigated L. pulmonaria's bacterial microbiome in the adaptation of this ecologically sensitive lichen species to diverse climatic conditions. Our central hypothesis posited that microbiome composition and functionality aligns with subcontinental-scale (a stretch of ~1100 km) climatic parameters related to temperature and precipitation. We also tested the impact of short-term weather dynamics, sampling season and algal/fungal genotypes on microbiome variation. Metaproteomics provided insights into compositional and functional changes within the microbiome. Climatic variables explained 41.64% of microbiome variation, surpassing the combined influence of local weather and sampling season at 31.63%. Notably, annual mean temperature and temperature seasonality emerged as significant climatic drivers. Microbiome composition correlated with algal, not fungal genotype, suggesting similar environmental recruitment for the algal partner and microbiome. Differential abundance analyses revealed distinct protein compositions in Sub-Atlantic Lowland and Alpine regions, indicating differential microbiome responses to contrasting environmental/climatic conditions. Proteins involved in oxidative and cellular stress were notably different. Our findings highlight microbiome plasticity in adapting to stable climates, with limited responsiveness to short-term fluctuations, offering new insights into climate adaptation in lichen symbiosis., (© 2024 The Author(s). Environmental Microbiology Reports published by John Wiley & Sons Ltd.)

Lichens are significant components of the biological soil crust communities in gypsum ecosystems and are involved in several processes related to ecosystem functioning, such as water and nutrient cycles or protection against soil erosion. Although numerous studies centered on lichen taxonomy and ecology have been performed in these habitats, global information about lichen species from gypsum substrates or their distributional ranges at a global scale is lacking. Thus, we compiled a global data set of recorded lichen species growing on gypsum. This review is based on systematic searches in two bibliographic databases (Web of Science and the more specialized database Mattick's Literature Index) using various keywords related to the substrate or ecology (i.e., gypsum, gypsiferous, semiarid, saxicolous, terricolous). In addition, we revised lichen literature from countries with gypsum soils using Mattick's, Hamburg University's Worldwide checklist, and different national lichen checklists. Ultimately the review includes a total of 321 studies. This data set included 6114 specimen records belonging to 336 recorded lichen species from 26 countries throughout the world. The results showed large differences in the number of species recorded among countries, reflecting differences in the sampling effort. We provide a table with the number of studies and species in relation to gypsum surface in order to account for the bias produced by sampling effort. The number of studies carried out per country was not related to the gypsum surface but probably to other factors, such as accessibility to field sampling, economic or political factors, or the presence of a wider community of lichenologists. Thus, Spain and Germany hosted the highest number of recorded species (160 and 114 species, respectively). Outside the European continent, only a few countries had a large number of species: Morocco (46), United States (42), and Iran (37). Remarkably, countries from the southern hemisphere (i.e., Australia, Chile, Namibia, and South Africa) showed a low number of studies from gypsum lands, supporting the stated biases observed in sampling efforts among countries. Considering the most studied countries, the results show that Teloschistaceae was the most represented family in gypsum ecosystems followed by Verrucariaceae and Cladoniaceae. Regarding particular species, Psora decipiens and Squamarina lentigera were some of the most widespread and abundant species in these habitats. This data set constitutes a basic and first step toward a much more comprehensive database, to be periodically updated in future releases, which also serves to identify countries or territories where future studies should be accomplished. There are no copyright restrictions on the data; please cite this data paper if the data are used in publications and teaching events., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

This study explores and compares the limits for photosynthesis in subzero temperatures of six Antarctic lichens: Sphaerophorus globosus, Caloplaca regalis, Umbilicaria antarctica, Pseudephebe minuscula, Parmelia saxatilis and Lecania brialmontii combining linear cooling and chlorophyll fluorescence methods. The results revealed triphasic S-curves in the temperature response of the maximum quantum yield (F V /F M ) and effective quantum yield of photosystem II (Φ PSII ) for all species. All investigated species showed a high level of cryoresistance with critical temperatures (T c ) below -20 °C. However, record low T c temperatures have been discovered for L. brialmotii (-54 °C for F V /F M and -40 °C for Φ PSII ) and C. regalis (-52 °C for F V /F M and -38 °C for Φ PSII ). Additionally, the yield differentials (F V /F M  - Φ PSII ) in functions of temperature revealed one or two peaks, with the larger one occurring for temperatures below -20 °C for the above-mentioned species. Finally, Kautsky kinetics were measured and compared at different temperatures (20 °C, 10 °C, 0 °C and -10 °C and then -10 °C after 1 h of incubation). This research serves as a foundation for further developing investigations into the biophysical mechanisms by which photosynthesis is carried out at subzero temperatures., 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 © 2024 Elsevier GmbH. All rights reserved.)

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).)

Most epiphytic lichens demonstrate high specificity to a habitat type, and sensitive hygrophilous species usually find shelter only in close-to-natural forest complexes. Some of them are considered as old-growth forest and/or long ecological continuity indicators. To evaluate general links between the narrow ecological range and physiological traits, two distinct sets of model lichens, i.e., old-growth forest ( Cetrelia cetrarioides (Duby) W.L. Culb. & C.F. Culb., Lobaria pulmonaria (L.) Hoffm., Menegazzia terebrata (Hoffm.) A. Massal.), and generalist ( Flavoparmelia caperata (L.) Hale, Hypogymnia physodes (L.) Nyl., Parmelia sulcata Taylor) ones, were examined in terms of sensitivity to long-term desiccation stress (1-, 2-, and 3-month) and photosynthesis activation rate upon rehydration. Desiccation tolerance and response rate to rehydration are specific to a given ecological set of lichens rather than to a particular species. Noticeable delayed and prompt recovery of high photosynthetic activity of photosystem II (PSII) characterize these sets, respectively. At the same time, although a decrease in the potential quantum yield of PSII in lichen thalli with a relative water content (RWC) at the level of 25% was observed, the efficiency remained at a very high level for all species, regardless of habitat preferences. Among the examined lichens, the fluorescence emission parameters for F. caperata were the fastest toward equilibrium upon rehydration, both after a shorter and a longer period of desiccation stress. In contrast to generalist lichens, retrieving of photosynthesis after 3-month desiccation failed in old-growth forest lichens. In the long term, prolonged rainless periods and unfavorable water balance in the environment predicted in the future may have a severely limiting effect on hygrophilous lichens during growing season (also in the sense of species associations) and, at the same time, promote the development of generalists.

Biological soil crusts (biocrusts) are widely distributed in global drylands and have multiple significant roles in regulating dryland soil and ecosystem multifunctionality. However, maps of their distribution over large spatial scales are uncommon and sometimes unreliable, because our current remote sensing technology is unable to efficiently discriminate between biocrusts and vascular plants or even bare soil across different ecosystem and soil types. The lack of biocrust spatial data may limit our ability to detect risks to dryland function or key tipping points. Here, we indirectly mapped biocrust distribution in China's drylands using spatial prediction modeling, based on a set of occurrences of biocrusts (379 in total) and high-resolution soil and environmental data. The results showed that biocrusts currently cover 13.9 % of China's drylands (or 5.7 % of China's total area), with moss-, lichen-, and cyanobacterial-dominated biocrusts each occupying 5.7 % to 10.7 % of the region. Biocrust distribution is mainly determined by soil properties (soil type and contents of gravel and nitrogen), aridity stress, and altitude. Their most favorable habitat is arenosols with low contents of gravel and nitrogen, in climate with a drought index of 0.54 and an altitude of about 500 m. By 2050, climate change will lead to a 5.5 %-9.0 % reduction in biocrust cover. Lichen biocrusts exhibit a high vulnerability to climate change, with potential reductions of up to 19.0 % in coverage. Biocrust cover loss is primarily caused by the combined effects of the elevated temperature and increased precipitation. Our study provides the first high-resolution (250 × 250 m) map of biocrust distribution in China's drylands and offers a reliable approach for mapping regional or global biocrust colonization. We suggest incorporating biocrusts into Earth system models to identify their significant impact on global or regional-scale processes under climate change., 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 © 2023 Elsevier B.V. All rights reserved.)

Desiccation tolerance (DT) is relatively frequent in non-vascular plants and green algae. However, it is poorly understood how successive dehydration/rehydration (D/R) cycles shape their transcriptomes and proteomes. Here, we report a comprehensive analysis of adjustments on both transcript and protein profiles in response to successive D/R cycles in Coccomyxa simplex (Csol), isolated from the lichen Solorina saccata. A total of 1833 transcripts and 2332 proteins were differentially abundant as a consequence of D/R; however, only 315 of these transcripts/proteins showed similar trends. Variations in both transcriptomes and proteomes along D/R cycles together with functional analyses revealed an extensive decrease in transcript and protein levels during dehydration, most of them involved in gene expression, metabolism, substance transport, signalling and folding catalysis, among other cellular functions. At the same time, a series of protective transcripts/proteins, such as those related to antioxidant defence, polyol metabolism and autophagy, was upregulated during dehydration. Overall, our results show a transient decrease in most cellular functions as a result of drying and a gradual reactivation of specific cell processes to accommodate the hydration status along successive D/R cycles. This study provides new insights into key mechanisms involved in the DT of Csol and probably other dehydration-tolerant microalgae. In addition, functionally characterising the high number of genes/proteins of unknown functions found in this study may lead to the discovery of new DT mechanisms., (© 2023. The Author(s).)

Epiphytic lichens constitute an important component of biodiversity in both deforested and forest ecosystems. Widespread occurrence is the domain of generalist lichens or those that prefer open areas. While, many stenoecious lichens find shelter only in a shaded interior of forests. Light is one of the factors known to be responsible for lichen distribution. Nevertheless, the effect of light intensity on photosynthesis of lichen photobionts remain largely unknown. We investigated photosynthesis in lichens with different ecological properties in relation to light as the only parameter modified during the experiments. The aim was to find links between this parameter and habitat requirements of a given lichen. We applied the methods based on a saturating light pulse and modulated light to perform comprehensive analyses of fast and slow chlorophyll fluorescence transient (OJIP and PSMT) combined with quenching analysis. We also examined the rate of CO 2 assimilation. Common or generalist lichens, i.e. Hypogymnia physodes, Flavoparmelia caperata and Parmelia sulcata, are able to adapt to a wide range of light intensity. Moreover, the latter species, which prefers open areas, dissipates the excess energy most efficiently. Conversely, Cetrelia cetrarioides considered an old-growth forest indicator, demonstrates definitely lower range of energy dissipation than other species, although it assimilates CO 2 efficiently both at low and high light. We conclude that functional plasticity of the thylakoid membranes of photobionts largely determines the dispersal abilities of lichens and light intensity is one of the most important factors determining the specificity of a species to a given habitat., (© 2023. The Author(s).)

Extreme climatic phenomena such as heat waves, heavy rainfall and prolonged droughts are one of the main problems associated with ongoing climate change. The global increase in extreme rainfalls associated with summer heatwaves are projected to increase in amplitude and frequency in the near future. However, the consequences of such extreme events on lichens are largely unknown. The aim was to determine the effect of heat stress on the physiology of lichen Cetraria aculeata in a metabolically active state and to verify whether strongly melanised thalli are more resistant than poorly melanised thalli. In the present study, melanin was extracted from C. aculeata for the first time. Our study showed that the critical temperature for metabolism is around 35 °C. Both symbiotic partners responded to heat stress, manifested by the decreased maximum quantum yield of PSII photochemistry, high level of cell membrane damage, increased membrane lipid peroxidation and decreased dehydrogenase activity. Highly melanised thalli were more sensitive to heat stress, which excludes the role of melanins as compounds protecting against heat stress. Therefore, mycobiont melanisation imposes a trade-off between protection against UV and avoidance of damage caused by high temperature. It can be concluded that heavy rainfall during high temperatures may significantly deteriorate the physiological condition of melanised thalli. However, the level of membrane lipid peroxidation in melanised thalli decreased over time after exposure, suggesting greater efficiency of antioxidant defence mechanisms. Given the ongoing climate changes, many lichen species may require a great deal of plasticity to maintain their physiological state at a level that ensures their survival., (© 2023. The Author(s).)

Nonvascular photoautotrophs (NVP), including bryophytes, lichens, terrestrial algae, and cyanobacteria, are increasingly recognized as being essential to ecosystem functioning in many regions of the world. Current research suggests that climate change may pose a substantial threat to NVP, but the extent to which this will affect the associated ecosystem functions and services is highly uncertain. Here, we propose a research agenda to address this urgent question, focusing on physiological and ecological processes that link NVP to ecosystem functions while also taking into account the substantial taxonomic diversity across multiple ecosystem types. Accordingly, we developed a new categorization scheme, based on microclimatic gradients, which simplifies the high physiological and morphological diversity of NVP and world-wide distribution with respect to several broad habitat types. We found that habitat-specific ecosystem functions of NVP will likely be substantially affected by climate change, and more quantitative process understanding is required on: (1) potential for acclimation; (2) response to elevated CO 2 ; (3) role of the microbiome; and (4) feedback to (micro)climate. We suggest an integrative approach of innovative, multimethod laboratory and field experiments and ecophysiological modelling, for which sustained scientific collaboration on NVP research will be essential., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

Arctic ecosystems are increasingly exposed to extreme climatic events throughout the year, which can affect species performance. Cryptogams (bryophytes and lichens) provide important ecosystem services in polar ecosystems but may be physiologically affected or killed by extreme events. Through field and laboratory manipulations, we compared physiological responses of seven dominant sub-Arctic cryptogams (three bryophytes, four lichens) to single events and factorial combinations of mid-winter heatwave (6°C for 7 days), re-freezing, snow removal and summer nitrogen addition. We aimed to identify which mosses and lichens are vulnerable to these abiotic extremes and if combinations would exacerbate physiological responses. Combinations of extremes resulted in stronger species responses but included idiosyncratic species-specific responses. Species that remained dormant during winter (March), irrespective of extremes, showed little physiological response during summer (August). However, winter physiological activity, and response to winter extremes, was not consistently associated with summer physiological impacts. Winter extremes affect cryptogam physiology, but summer responses appear mild, and lichens affect the photobiont more than the mycobiont. Accounting for Arctic cryptogam response to multiple climatic extremes in ecosystem functioning and modelling will require a better understanding of their winter eco-physiology and repair capabilities., (© 2023 The Authors. Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

Forest vegetation is key for buffering microclimatic factors and regulating atmospheric deposition. Epiphytic lichens are sensitive to these factors and can indicate the overall health status of the ecosystem. Specifically, the analysis of morpho-anatomical variations allows us to understand the degree of tolerance or sensitivity of these organisms exposed to agricultural crops and how vegetation might buffer this response. We analyzed variations in vegetative and reproductive characters and injuries in thalli of Parmotrema pilosum as a response to distance to crops and forest cover. The study was conducted in forest patches of the Espinal in central Argentina, an ecosystem threatened by agricultural activity. We selected 10 sites with different forest cover areas and two collection points differing in distance to crops: sites adjacent to (0 m) and far from (150 m) crops. We collected five thalli from each collection point and analyzed variations in morpho-anatomical characters at macro- and microscopic levels. We found a lower number of algae and a higher proportion of simple cilia in individuals at points adjacent to crops. At points with low forest cover, a thinner upper cortex was observed, whereas at points with greater forest cover, an increase of necrosis and greater presence of apothecia were detected. Bleaching was the most frequent injury at sites adjacent to crops, decreasing with increasing forest cover. Conservation and reforestation of Espinal forest patches would promote the propagation of lichens affected by agricultural practices., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

To create more resilient cities, it is important that we understand the effects of the global change drivers in cities. Biodiversity-based ecological indicators (EIs) can be used for this, as biodiversity is the basis of ecosystem structure, composition, and function. In previous studies, lichens have been used as EIs to monitor the effects of global change drivers in an urban context, but only in single-city studies. Thus, we currently do not understand how lichens are affected by drivers that work on a broader scale. Therefore, our aim was to quantify the variance in lichen biodiversity-based metrics (taxonomic and trait-based) that can be explained by environmental drivers working on a broad spatial scale, in an urban context where local drivers are superimposed. To this end, we performed an unprecedented effort to sample epiphytic lichens in 219 green spaces across a continental gradient from Portugal to Estonia. Twenty-six broad-scale drivers were retrieved, including air pollution and bio-climatic variables, and their dimensionality reduced by means of a principal component analysis (PCA). Thirty-eight lichen metrics were then modelled against the scores of the first two axes of each PCA, and their variance partitioned into pollution and climate components. For the first time, we determined that 15% of the metric variance was explained by broad-scale drivers, with broad-scale air pollution showing more importance than climate across the majority of metrics. Taxonomic metrics were better explained by air pollution, as expected, while climate did not surpass air pollution in any of the trait-based metric groups. Consequently, 85% of the metric variance was shown to occur at the local scale. This suggests that further work is necessary to decipher the effects of climate change. Furthermore, although drivers working within cities are prevailing, both spatial scales must be considered simultaneously if we are to use lichens as EIs in cities at continental to global scales., 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 Authors. Published by Elsevier Ltd.. All rights reserved.)

Poikilohydric autotrophs are the main colonizers of the permanent ice-free areas in the Antarctic tundra biome. Global climate warming and the small human footprint in this ecosystem make it especially vulnerable to abrupt changes. Elucidating the effects of climate change on the Antarctic ecosystem is challenging because it mainly comprises poikilohydric species, which are greatly influenced by microtopographic factors. In the present study, we investigated the potential effects of climate change on the metabolic activity and net primary photosynthesis (NPP) in the widespread lichen species Usnea aurantiaco-atra. Long-term monitoring of chlorophyll a fluorescence in the field was combined with photosynthetic performance measurements in laboratory experiments in order to establish the daily response patterns under biotic and abiotic factors at micro- and macro-scales. Our findings suggest that macroclimate is a poor predictor of NPP, thereby indicating that microclimate is the main driver due to the strong effects of microtopographic factors on cryptogams. Metabolic activity is also crucial for estimating the NPP, which is highly dependent on the type, distribution, and duration of the hydration sources available throughout the year. Under RCP 4.5 and RCP 8.5, metabolic activity will increase slightly compared with that at present due to the increased precipitation events predicted in MIROC5. Temperature is highlighted as the main driver for NPP projections, and thus climate warming will lead to an average increase in NPP of 167-171% at the end of the century. However, small changes in other drivers such as light and relative humidity may strongly modify the metabolic activity patterns of poikilohydric autotrophs, and thus their NPP. Species with similar physiological response ranges to the species investigated in the present study are expected to behave in a similar manner provided that liquid water is available., 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 Authors. Published by Elsevier B.V. All rights reserved.)

Lecideoid lichens as dominant vegetation-forming organisms in the climatically harsh areas of the southern part of continental Antarctica show clear preferences in relation to environmental conditions (i.e. macroclimate). 306 lichen samples were included in the study, collected along the Ross Sea coast (78°S-85.5°S) at six climatically different sites. The species compositions as well as the associations of their two dominant symbiotic partners (myco- and photobiont) were set in context with environmental conditions along the latitudinal gradient. Diversity values were nonlinear with respect to latitude, with the highest alpha diversity in the milder areas of the McMurdo Dry Valleys (78°S) and the most southern areas (Durham Point, 85.5°S; Garden Spur, 84.5°S), and lowest in the especially arid and cold Darwin Area (~ 79.8°S). Furthermore, the specificity of mycobiont species towards their photobionts decreased under more severe climate conditions. The generalist lichen species Lecanora fuscobrunnea and Lecidea cancriformis were present in almost all habitats, but were dominant in climatically extreme areas. Carbonea vorticosa, Lecidella greenii and Rhizoplaca macleanii were confined to milder areas. In summary, the macroclimate is considered to be the main driver of species distribution, making certain species useful as bioindicators of climate conditions and, consequently, for assessing the consequences of climate change., (© 2021. The Author(s).)

Toxin-antitoxin (TA) systems are genetic modules composed of a toxin interfering with cellular processes and its cognate antitoxin, which counteracts the activity of the toxin. TA modules are widespread in bacterial and archaeal genomes. It has been suggested that TA modules participate in the adaptation of prokaryotes to unfavorable conditions. The Bosea sp. PAMC 26642 used in this study was isolated from the Arctic lichen Stereocaulon sp. There are 12 putative type II TA loci in the genome of Bosea sp. PAMC 26642. Of these, nine functional TA systems have been shown to be toxic in Escherichia coli The toxin inhibits growth, but this inhibition is reversed when the cognate antitoxin genes are coexpressed, indicating that these putative TA loci were bona fide TA modules. Only the BoVapC1 (AXW83&#95;01405) toxin, a homolog of VapC, showed growth inhibition specific to low temperatures, which was recovered by the coexpression of BoVapB1 (AXW83&#95;01400). Microscopic observation and growth monitoring revealed that the BoVapC1 toxin had bacteriostatic effects on the growth of E. coli and induced morphological changes. Quantitative real time polymerase chain reaction and northern blotting analyses showed that the BoVapC1 toxin had a ribonuclease activity on the initiator tRNA fMet , implying that degradation of tRNA fMet might trigger growth arrest in E. coli Furthermore, the BoVapBC1 system was found to contribute to survival against prolonged exposure at 4°C. This is the first study to identify the function of TA systems in cold adaptation., (© 2021 Jeon et al.; Published by Cold Spring Harbor Laboratory Press for the RNA Society.)

Understanding the species richness and β-diversity patterns along elevation gradients can aid in formulating effective conservation strategies particularly in areas where local anthropogenic stresses and climate change are quite significant as in the Himalaya. Thus, we studied macrolichen richness and β-diversity along elevational gradients at three sites, namely Kashmir (2200 to 3800 m a.m.s.l), Uttarakhand (2000-3700 m a.m.s.l) and Sikkim (1700 to 4000 m a.m.s.l) which cover much of the Indian Himalayan Arc. In all, 245 macrolichen species belonging to 77 genera and 26 families were collected from the three sites. Only 11 species, 20 genera and 11 families were common among the three transects. Despite the differences in species composition, the dominant functional groups in the three sites were the same: foliose, fruticose and corticolous forms. The hump-shaped elevation pattern in species richness was exhibited by most of the lichen groups, though an inverse hump-shaped pattern was also observed in certain cases. β-diversity (β sor ) based on all pairs of comparisons along an elevation gradient varied from 0.48 to 0.58 in Kashmir, 0.03 to 0.63 in Uttarakhand and 0.46 to 0.77 in Sikkim. The contribution of turnover to β-diversity was more than nestedness at all the three transects. Along elevation β-diversity and its components of turnover and nestedness varied significantly with elevation. While species turnover increased significantly along the elevation in all the three transects, nestedness decreased significantly in Kashmir and Sikkim transects but increased significantly in the Uttarakhand transect. Except for the Kashmir Himalayan elevation transect, stepwise β-diversity and its components of turnover and nestedness did not vary significantly with elevation. The present study, the first of its kind in the Himalayan region, clearly brings out that macrolichen species richness, β-diversity, and its components of turnover and nestedness vary along the elevation gradients across the Himalayan Arc. It also highlights that contribution of turnover to β-diversity is higher in comparison to nestedness at all the three transects. The variations in species richness and diversity along elevation gradients underpin the importance of considering elevational gradients in planning conservation strategies., (© 2021. The Author(s).)

Greatly simplified ecosystems are often neglected for biodiversity studies. However, these simplified systems dominate in many regions of the world, and a lack of understanding of what shapes species occurrence in these systems can have consequences for biodiversity and ecosystem services at a massive scale. In Fennoscandia, ~90% of the boreal forest (~21Mha) is structurally simplified with little knowledge of how forest structural elements shape the occurrence and diversity of for example epiphytic lichens in these managed forests. One form of structural simplification is the reduction of the number and frequency of different tree species. As many lichen species have host tree preferences, it is particularly likely that this simplification has a huge effect on the lichen community in managed forests. In a 40-70 years old boreal forest in Sweden, we therefore related the occurrence and richness of all observed epiphytic lichens to the host tree species and beta and gamma lichen diversity at the forest stand level to the stand's tree species composition and stem diameter. Picea abies hosted the highest lichen richness followed by Pinus sylvestris, Quercus robur, Alnus glutinosa, Betula spp., and Populus tremula. However, P. tremula hosted twice as many uncommon species as any of the other tree species. Stand level beta and gamma diversity was twice as high on stands with four compared to one tree species, and was highest when either coniferous or deciduous trees made up 40-50% of the trees. The stem diameter was positively related to lichen richness at the tree and stand level, but negatively to beta diversity. For biodiversity, these findings imply that leaving a few trees of a different species during forest thinning is unlikely as effective as combining life-boat trees for endangered species with an even tree species mixture., Competing Interests: The authors have declared that no competing interests exist.

Mutualistic symbiosis, we now know, is a ubiquitous phenomenon in the natural world. And, in every case, there was an initial "genesis" - a "how" process that may have been at once unique to each situation and perhaps also shared a common set of facilitators. However, a full explanation of symbiogenesis also requires an answer to the "why" question, for natural selection is a stringent economizer. Something as contrarian as mutualistic cooperation between "differently named" organisms must also provide functional advantages for the participants that will be favored by natural selection (differential survival and reproduction). Enter the "Synergism Hypothesis" - the thesis that synergistic functional effects of various kinds are a common cause of cooperative relationships of all kinds in nature, including symbioses. When different organisms have complementary capabilities that are mutually beneficial and cannot otherwise be attained, the benefits derived from symbiotic cooperation will outweigh the costs. Among the many documented cases of symbiogenesis over time, lichens provide perhaps the most familiar, well-studied example, while the eukaryotes are often cited as a game-changer. The answer to the "why" question was, in each case, determinative for symbiogenesis., (Copyright © 2021. Published by Elsevier B.V.)

We studied the biodiversity of Asterochloris photobionts found in Bolivian lichens to better understand their global spatial distribution and adaptation strategies in the context of a worldwide phylogeny of the genus. Based on nuclear ITS rDNA, the chloroplast rbcL gene and the actin type I gene we reconstructed a phylogenetic tree that recovered nine new Asterochloris lineages, while 32 Bolivian photobiont samples were assigned to 12 previously recognized Asterochloris lineages. We also show that some previously discovered Asterochloris photobiont species and lineages may occur in a broader spectrum of climatic conditions, and mycobiont species and photobionts may show different preferences along an altitude gradient. To reveal general patterns of of mycobiont specificity towards the photobiont in Asterochloris, we tested the influence of climate, altitude, geographical distance and effects of symbiotic partner (mycobiont) at the species level of three genera of lichen forming fungi: Stereocaulon, Cladonia and Lepraria. Further, we compared the specificity of mycobionts towards Asterochloris photobionts in cosmopolitan, Neotropical, and Pantropical lichen forming fungi. Interestingly, cosmopolitan species showed the lowest specificity to their photobionts, but also the lowest haplotype diversity. Neotropical and Paleotropical mycobionts, however, were more specific.

Actinobacteria that inhabit lichen symbionts are considered a promising yet previously underexplored source of novel compounds. Here, for the first time, we conducted a comprehensive investigation with regard to strain isolation and identification of lichen-associated actinobacteria from Tibet Plateau, antimicrobial activity screening, biosynthetic genes detection, bioactive metabolites identification and activity prediction. A large number of culturable actinomycetes were isolated from lichens around Qinghai Lake, in Qinghai-Tibet Plateau. Twenty-seven strains with distinct morphological characteristics were preliminarily studied. 16S rRNA gene identification showed that 13 strains were new species. The PCR-screening of specific biosynthetic genes indicated that these 27 isolates had abundant intrinsic biosynthetic potential. The antimicrobial activity experiment screened out some potential biological control antagonistic bacteria. The metabolites of 13 strains of Streptomyces with antibacterial activity were analyzed by LC-HRMS, and further 18 compounds were identified by NMR and / or LC-HRMS. The identified compounds were mainly pyrrolidine and indole derivatives, as well as anthracyclines. Seven compounds were identified with less biological activity, then predicted and evaluated their biological activity. The predicted results showed that compound 2 had excellent inhibitory activity on HIV-1 reverse transcriptase. Overall, the results indicate actinobacteria isolated from unexploited plateau lichen are promising sources of biological active metabolite, which could provide important bioactive compounds as potential antibiotic drugs., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

Trebouxia sp. (TR9) and Coccomyxa simplex (Csol) are desiccation-tolerant lichen microalgae with different adaptive strategies in accordance with the prevailing conditions of their habitats. The remodelling of cell wall and extracellular polysaccharides depending on water availability are key elements in the tolerance to desiccation of both microalgae. Currently, there is no information about the extracellular proteins of these algae and other aero-terrestrial microalgae in response to limited water availability. To our knowledge, this is the first report on the proteins associated with the extracellular polymeric substances (EPS) of aero-terrestrial microalgae subjected to cyclic desiccation/rehydration. LC-MS/MS and bioinformatic analyses of the EPS-associated proteins in the two lichen microalgae submitted to four desiccation/rehydration cycles allowed the compilation of 111 and 121 identified proteins for TR9 and Csol, respectively. Both sets of EPS-associated proteins shared a variety of predicted biological functions but showed a constitutive expression in Csol and partially inducible in TR9. In both algae, the EPS-associated proteins included a number of proteins of unknown functions, some of which could be considered as small intrinsically disordered proteins related with desiccation-tolerant organisms. Differences in the composition and the expression pattern between the studied EPS-associated proteins would be oriented to preserve the biochemical and biophysical properties of the extracellular structures under the different conditions of water availability in which each alga thrives.

Mycobionts of many lichen genera appear to demonstrate strong selectivity in the choice of algal partner. The biological properties of a photobiont and its availability in an environment significantly determine the habitat requirements of lichens. Flexibility in photobiont choice extends the ecological amplitude of lichens; therefore, it may constitute an important adaptive strategy for colonization of extreme habitats. The photobiont inventory of the three epigeic lichens most resistant to soil pollution, i.e., Cladonia cariosa, C. rei , and the hyperaccumulator Diploschistes muscorum , was examined to verify whether and to what extent algal composition depends on the type of habitat and substrate enrichment with heavy metals. Photobionts Asterochloris and Trebouxia were identified in the studied lichen species; however, the presence of Trebouxia was directly related to anthropogenic sites with technogenic substrates, and the proportion of lichen specimens with these algae clearly depended on the level of heavy-metal soil pollution and the habitat type. The total number of algal haplotypes increased with increasing soil pollution, and the richness was associated more with soil pollution than with a given lichen species. Additionally, a large number of lichen individuals bearing multiple algal genotypes at polluted sites were recorded. Although Cladonia lichens were previously thought to be restricted to Asterochloris , they are able to start the relichenization process with Trebouxia under specific habitat conditions and to establish a stable association with these algae when colonization of disturbed sites takes place. Comparative analysis of the internal transcribed spacer (ITS) rDNA sequences revealed as many as 13 haplotypes of Trebouxia , and phylogenetic analysis grouped them into two different clades. Such a high level of genetic diversity indicates that Trebouxia is well adapted to metal pollution and could be an alternative photosynthetic partner for certain lichens, especially in polluted sites.

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.)

This study analyses the interactions among crustose and lichenicolous lichens growing on gypsum biocrusts. The selected community was composed of Acarospora nodulosa, Acarospora placodiiformis, Diploschistes diacapsis, Rhizocarpon malenconianum and Diplotomma rivas-martinezii. These species represent an optimal system for investigating the strategies used to share phycobionts because Acarospora spp. are parasites of D. diacapsis during their first growth stages, while in mature stages, they can develop independently. R. malenconianum is an obligate lichenicolous lichen on D. diacapsis, and D. rivas-martinezii occurs physically close to D. diacapsis. Microalgal diversity was studied by Sanger sequencing and 454-pyrosequencing of the nrITS region, and the microalgae were characterized ultrastructurally. Mycobionts were studied by performing phylogenetic analyses. Mineralogical and macro- and micro-element patterns were analysed to evaluate their influence on the microalgal pool available in the substrate. The intrathalline coexistence of various microalgal lineages was confirmed in all mycobionts. D. diacapsis was confirmed as an algal donor, and the associated lichenicolous lichens acquired their phycobionts in two ways: maintenance of the hosts' microalgae and algal switching. Fe and Sr were the most abundant microelements in the substrates but no significant relationship was found with the microalgal diversity. The range of associated phycobionts are influenced by thallus morphology.

Lichens are symbiotic organisms that are well adapted to desiccation/rehydration cycles. Over the last decades, the physiological background of their photosynthetic response-specifically activation of the protective mechanism during desiccation-has been studied at the level of photosystem II of the lichen photobiont by means of several biophysical methods. In our study, the effects of desiccation and low temperatures on chlorophyll fluorescence and spectral reflectance parameters were investigated in Antarctic chlorolichen Dermatocarpon polyphyllizum. Lichen thalli were collected from James Ross Island, Antarctica, and following transfer to a laboratory, samples were fully hydrated and exposed to desiccation at temperatures of 18, 10, and 4 °C. During the desiccation process, the relative water content (RWC) was measured gravimetrically and photosynthetic parameters related to the fast transient of chlorophyll fluorescence (OJIP) were measured repeatedly. Similarly, the change in spectral reflectance parameters (e.g., NDVI, PRI, G, NPCI) was monitored during thallus dehydration. The dehydration-response curves showed a decrease in a majority of the OJIP-derived parameters (e.g., maximum quantum yield of photosystem II photochemistry: F V /F M , and performance index: PI in D. polyphyllizum, which were more apparent at RWCs below 20%. The activation of protective mechanisms in severely dehydrated thalli was documented by increased thermal dissipation (DI 0 /RC) and its quantum yield (Phi&#95;D 0 ). Low temperature accelerated these processes. An analysis of the OJIP shape reveals the presence of K-bands (300 μs), and L-bands (80 μs), which can be attributed to dehydration-induced stress. Spectral reflectance indices decreased in a majority of cases with an RWC decrease and were positively related to the OJIP-derived parameters: F V /F M (capacity of photosynthetic processes in PSII), Phi&#95;E 0 (effectiveness of electron transport), and PI&#95;tot (total performance index), which was more apparent in NDVI. A negative relation was found for NPCI. These indices could be used in follow-up ecophysiological photosynthetic studies of lichens that are undergoing rehydration/dehydration cycles.

The objective of this study is to compare the physiological response (content and degradation of photosynthetic pigments, membrane oxidation products and soluble proteins) and multi-element content of Ramalina celastri (lichenized fungi) growing on agricultural fences with no-tillage (associated with transgenic crops and agrochemical application), organic cropping and a non-cultivated area. We found that R. celastri did not differ in its physiological response to agricultural practices, except for the contents of chlorophyll b and phaeophytin a which were high in both cultivated areas. Lichens growing in organic cropping fields have higher arsenic, chromium, uranium and internal transition elements common in the earth's crust, possibly due to the greater resuspension of the material during soil tillage. Lichens that grow on posts close to no-tillage field had higher bromine contents (present in numerous pesticides). We found evidence that R. celastri behaves as a tolerant species to air pollution in agricultural environments.

Lithobionts (rock-dwelling organisms) have been recognized as agents of aesthetic and physico-chemical deterioration of stonework. In consequence, their removal from cultural heritage stone surfaces (CHSS) is widely considered a necessary step in conservation interventions. On the other hand, lithobiontic communities, including microbial biofilms ('biological patinas'), can help integrate CHSS with their environmental setting and enhance biodiversity. Moreover, in some cases bioprotective effects have been reported and even interpreted as potential biotechnological solutions for conservation. This paper reviews the plethora of traditional and innovative methodologies to characterize lithobionts on CHSS in terms of biodiversity, interaction with the stone substrate and impacts on durability. In order to develop the best management and conservation strategies for CHSS, such diagnosis should be acquired on a case-by-case basis, as generalized approaches are unlikely to be suitable for all lithobionts, lithologies, environmental and cultural contexts or types of stonework. Strategies to control biodeteriogenic lithobionts on CHSS should similarly be based on experimental evaluation of their efficacy, including long-term monitoring of the effects on bioreceptivity, and of their environmental safety. This review examines what is known about the efficacy of control methods based on traditional-commercial biocides, as well as those based on innovative application of substances of plant and microbial origin, and physical techniques. A framework for providing a balanced scientific assessment of the role of lithobionts on CHSS and integrating this knowledge into management and conservation decision-making is presented.

As part of the Biology and Mars Experiment (BIOMEX; ILSRA 2009-0834), samples of the lichen Circinaria gyrosa were placed on the exposure platform EXPOSE-R2, on the International Space Station (ISS) and exposed to space and to a Mars-simulated environment for 18 months (2014-2016) to study: (1) resistance to space and Mars-like conditions and (2) biomarkers for use in future space missions (Exo-Mars). When the experiment returned (June 2016), initial analysis showed rapid recovery of photosystem II activity in the samples exposed exclusively to space vacuum and a Mars-like atmosphere. Significantly reduced recovery levels were observed in Sun-exposed samples, and electron and fluorescence microscopy (transmission electron microscope and field emission scanning electron microscope) data indicated that this was attributable to the combined effects of space radiation and space vacuum, as unirradiated samples exhibited less marked morphological changes compared with Sun-exposed samples. Polymerase chain reaction analyses confirmed that there was DNA damage in lichen exposed to harsh space and Mars-like environmental conditions, with ultraviolet radiation combined with space vacuum causing the most damage. These findings contribute to the characterization of space- and Mars-resistant organisms that are relevant to Mars habitability.

Mountain pine beetle (MPB) has become an invasive forest pest of mature pine in western North America as it spreads beyond its former endemic range. Management actions such as timber harvest can reduce the spread of MPB but may affect species of conservation concern like woodland caribou. Our goal was to inform MPB management within caribou ranges by exploring the impacts of MPB on caribou habitat-focusing on terrestrial lichens, an important winter food for caribou. We evaluated differences in lichen cover among four MPB management actions: timber harvest, wildfires, leaving MPB killed trees as-is, and single-tree cut-and-burn control. We found little evidence that leaving MPB killed trees as-is or controlling MPB using single-tree cut-and-burn impacted lichen cover. However, we found that lichen cover was lower in timber harvested and burned areas compared to intact undisturbed forest but only 10 to 20 years post-disturbance, respectively. Our results suggest that despite short-term reductions in lichen cover, using timber harvesting and prescribed burns to control MPB may balance management needs for MPB while maintaining lichen cover over time. However, using timber harvesting and prescribed burns to manage MPB is likely to have detrimental population-level effects on caribou by increasing the proportion of disturbed habitat and thus predators within caribou ranges. Among the four management actions that we evaluated, the cut-and-burn control program balances the need to limit the spread of MPB while also limiting negative impacts on caribou food. Our work addresses some of the challenges of managing competing forest and ecosystem values by evaluating the consequence of forest pest management actions on an important food resource for a species-at-risk., Competing Interests: The full list of fRI Research partners and funders are: Alberta Agriculture and Forestry, Alberta Biodiversity Monitoring Institute, Alberta Conservation Association, Alberta Energy Regulator, Alberta Environment and Parks, Alberta Fish and Game Association, Alberta Institute of Agrologists, Alberta Innovates, Alberta Newsprint Company, Alberta-Pacific Forest Industries Inc., Alberta Professional Planners Institute, Alberta Riparian Habitat Management Society, Apache Canada Ltd., Arctos Ecological Consulting, Aseniwuche Winewak Nation of Canada, Athabasca Watershed Council, Bandaloop Landscape-Ecosystem Services, Battle River Watershed Alliance, BC Oil and Gas Commission, BC Oil and Gas Research and Innovation Society, Beaver River Watershed Alliance, Borealis Ecology Wildlife Research, Bow River Basin Council, Canadian Association of Petroleum Producers, Canadian Boreal Forest Agreement, Canadian Institute of Forestry, Canadian International Oil Corp., Canadian Natural Resources Limited, Canadian Wildlife Health Cooperative, Canfor Corporation, Cenovus Energy Inc., City of Grande Prairie, ConocoPhillips Canada, Cumulative Environmental Management Association, Daishowa-Marubeni International Ltd., Devon Energy Corporation, Ducks Unlimited Canada, Encana Corporation, Environment and Climate Change Canada, Explorers and Producers Association of Canada, Fisheries and Oceans Canada, Followit Sweden, Foothills Forest Products, FORCORP, Forest Resource Improvement Association of Alberta (FRIAA), Forest Stewardship Council, Forsite Consultants Ltd., Fuse Consulting Ltd., Golder Associates, Government of British Columbia (Ministry of Environment; Ministry of Forests, Lands, and Natural Resource Operations), Government of Northwest Territories (Ministry of Environment and Natural Resources), Government of Saskatchewan (Ministry of Environment), Grande Cache Tourism and Interpretative Centre, Hinton and District Chamber of Commerce, Husky Energy Inc., Inside Education, Integrated Ecological Research, Jasper-Yellowhead Museum & Archives, Joss Wind Power Inc., Lesser Slave Watershed Council, Louisiana-Pacific Corporation, Mighty Peace Watershed Alliance, Millar Western Forest Products Ltd., Mistik Management Ltd., Mitacs, Natural Sciences and Engineering Research Council of Canada (NSERC), Natural Resources Canada, Canadian Forest Service, Norbord Inc., North Saskatchewan Watershed Alliance, Northern Rockies Museum of Culture and Heritage, Norwegian University of Life Sciences, Norwegian Institute of Bioeconomy Research, Oldman Watershed Council, Paramount Resources Ltd., Parks Canada, Pembina Pipeline Corporation, Peregrine Helicopters, Peter J. Murphy Forest Consulting Ltd., Petroleum Technology Alliance Canada, Red Deer River Watershed Alliance, Repsol Oil and Gas Inc., Scandinavian Brown Bear Research Project, Seven Generations Energy Ltd., Shell Canada Ltd., South East Alberta Watershed Alliance, Spray Lake Sawmills, St’at’imc Government Services, Suncor Energy Inc., Sustainable Forestry Initiative, TAQA North Ltd., Teck Resources Ltd., TerrainWorks, Timberworks Inc., Tolko Industries Ltd., Tom Peterson, Toronto Zoo, Tourmaline Oil Corp., Town of Hinton, TransCanada Corporation, Trout Unlimited Canada, United States Department of Agriculture (United States Forest Service), University of Alberta, University of British Columbia, University of Calgary, Université Laval, University of Oslo, University of Saskatchewan, University of Victoria, Washington State University, Vanderwell Contractors (1971) Ltd., West Fraser Mills Ltd., Westmoreland Coal Company, Weyerhaeuser Company, Wild Year Productions Ltd., Woodlands Operations Learning Foundation, XTO Energy Inc., Yellowhead County, Yellowstone to Yukon Conservation Initiative. Authors TL and LF are employed by fRI Research – a non-profit. fRI Research did not play a role in the study design, data collection and analysis, decision to publish, or preparation of the manuscript and only provided financial support in the form of authors’ salaries and/or research materials. Author BN was employed by fRI Research at the time of data analysis and preparation of the draft manuscript and is currently employed by Alberta Environment and Parks. Alberta Environment and Parks and fRI Research did not play a role in the study design, data collection and analysis, decision to publish, or preparation of the manuscript. Author KEP was employed by fRI Research at the time of data collection, analysis, and preparation of the draft manuscript and is currently employed by Carelton University. Carelton University and fRI Research did not play a role in the study design, data collection and analysis, decision to publish, or preparation of the manuscript. The authors past and current employers did not alter our adherence to PLOS ONE policies on sharing data and materials.

Stable, long-term interactions between fungi and algae or cyanobacteria, collectively known as lichens, have repeatedly evolved complex architectures with little resemblance to their component parts. Lacking any central scaffold, the shapes they assume are casts of secreted polymers that cement cells into place, determine the angle of phototropic exposure and regulate water relations. A growing body of evidence suggests that many lichen extracellular polymer matrices harbor unicellular, non-photosynthesizing organisms (UNPOs) not traditionally recognized as lichen symbionts. Understanding organismal input and uptake in this layer is key to interpreting the role UNPOs play in lichen biology. Here, we review both polysaccharide composition determined from whole, pulverized lichens and UNPOs reported from lichens to date. Most reported polysaccharides are thought to be structural cell wall components. The composition of the extracellular matrix is not definitively known. Several lines of evidence suggest some acidic polysaccharides have evaded detection in routine analysis of neutral sugars and may be involved in the extracellular matrix. UNPOs reported from lichens include diverse bacteria and yeasts for which secreted polysaccharides play important biological roles. We conclude by proposing testable hypotheses on the role that symbiont give-and-take in this layer could play in determining or modifying lichen symbiotic outcomes., (© FEMS 2020.)

The tendency for species to retain their ancestral biological properties has been widely demonstrated, but the effect of phylogenetic constraints when progressing from species to ensemble-level properties requires further assessment. Here we test whether community-level patterns (environmental shifts in local species richness and turnover) are phylogenetically conserved, assessing whether their similarity across different families of lichens, insects, and birds is dictated by the relatedness of these families. We show a significant phylogenetic signal in the shape of the species richness-elevation curve and the decay of community similarity with elevation: closely related families share community patterns within the three major taxa. Phylogenetic influences are partly explained by similarities among families in conserved traits defining body plan and interactions, implying a scaling of phylogenetic effects from the organismal to the community level. Consequently, the phylogenetic signal in community-level patterns informs about how the historical legacy of a taxon and shared responses among related taxa to similar environments contribute to community assembly and diversity patterns.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur., (© 2019 The Authors New Phytologist © 2019 New Phytologist Trust.)

In dryland regions, biological soil crusts (BSCs) have numerous important ecosystem functions. Crust species and functions are, however, highly spatially heterogeneous and remain poorly understood at a range of scales. In this study, chlorophyll fluorescence imaging was used to quantify millimeter-scale patterns in the distribution and activity of photosynthetic organisms in BSCs of different successional stages (including cyanobacterial, lichen, moss three main successional stages and three intermixed transitional stages) from the Tengger Desert, China. Chlorophyll fluorescence images derived from the Imaging PAM (Pulse Amplitude Modulation) showed that with the succession from cyanobacterial to lichen and to moss crusts, crust photosynthetic efficiency (including the maximum and effective photosynthetic efficiency, respectively) and fluorescence coverage increased significantly (P < 0.05), and that increasing photosynthetically active radiation (PAR) reduced the effective photosynthetic efficiency (Yield). The distribution of photosynthetic organisms in crusts determined Fv/Fm (ratio of variable fluorescence to maximum fluorescence) frequency pattern, although the photosynthetic heterogeneity (SHI index) was not significantly different (P > 0.05) between cyanobacterial and moss crusts, and showed a unimodal pattern of Fv/Fm values. In contrast, photosynthetic heterogeneity was significantly higher in lichen, cyanobacteria-moss and lichen-moss crusts (P < 0.05), with a bimodal pattern of Fv/Fm values. Point pattern analysis showed that the distribution pattern of chlorophyll fluorescence varied at different spatial scales and also among the different crust types. These new results provide a detailed (millimeter-scale) insight into crust photosynthetic mechanisms and spatial distribution patterns associated with their community types. Collectively, this information provides an improved theoretical basis for crust maintenance and management in dryland regions.

The view of lichens as a symbiosis only between a mycobiont and a photobiont has been challenged by discoveries of diverse associated organisms. Specific basidiomycete yeasts in the cortex of a range of macrolichens were hypothesized to influence the lichens' phenotype. The present study explores the occurrence and diversity of cystobasidiomycete yeasts in the lichen genus Cladonia. We obtained seven cultures and 56 additional sequences using specific primers from 27 Cladonia species from all over Europe and performed phylogenetic analyses based on ITS, LSU and SSU rDNA loci. We revealed yeast diversity distinct from any previously reported. Representatives of Cyphobasidiales, Microsporomycetaceae and of an unknown group related to Symmetrospora have been found. We present evidence that the Microsporomycetaceae contains mainly lichen-associated yeasts. Lichenozyma pisutiana is circumscribed here as a new genus and species. We report the first known associations between cystobasidiomycete yeasts and Cladonia (both corticate and ecorticate), and find that the association is geographically widespread in various habitats. Our results also suggest that a great diversity of lichen associated yeasts remains to be discovered., (Copyright © 2019 British Mycological Society. Published by Elsevier Ltd. All rights reserved.)