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In this study, we explored the diversity of green algal symbionts (photobionts) in sympatric populations of the cosmopolitan lichen-forming fungi Thamnolia and Cetraria . We sequenced with both Sanger and Ion Torrent High-Throughput Sequencing technologies the photobiont ITS-region of 30 lichen thalli from two islands: Iceland and Öland. While Sanger recovered just one photobiont genotype from each thallus, the Ion Torrent data recovered 10–18 OTUs for each pool of 5 lichen thalli, suggesting that individual lichens can contain heterogeneous photobiont populations. Both methods showed evidence for photobiont sharing between Thamnolia and Cetraria on Iceland. In contrast, our data suggest that on Öland the two mycobionts associate with distinct photobiont communities, with few shared OTUs revealed by Ion Torrent sequencing. Furthermore, by comparing our sequences with public data, we identified closely related photobionts from geographically distant localities. Taken together, we suggest that the photobiont composition in Thamnolia and Cetraria results from both photobiont-mycobiont codispersal and local acquisition during mycobiont establishment and/or lichen growth. We hypothesize that this is a successful strategy for lichens to be flexible in the use of the most adapted photobiont for the environment.

The Mediterranean region, comprising the Mediterranean Basin and the Macaronesian Islands, represents a center of diversification for many organisms. The genetic structure and connectivity of mainland and island microbial populations has been poorly explored, in particular in the case of symbiotic fungi. Here we investigated genetic diversity and spatial structure of the obligate outcrossing lichen-forming fungus Parmelina carporrhizans in the Mediterranean region. Using eight microsatellite and mating-type markers we showed that fungal populations are highly diverse but lack spatial structure. This is likely due to high connectivity and long distance dispersal of fungal spores. Consistent with low levels of linkage disequilibrium and lack of clonality, we detected both mating-type idiomorphs in all populations. Furthermore we showed that the Macaronesian Islands are the result of colonization from the Mediterranean Basin. The unidirectional gene flow, though, seemed not to be sufficient to counterbalance the effects of drift, resulting in comparatively allelic poor peripheral populations. Our study is the first to shed light on the high connectivity and lack of population structure in natural populations of a strictly sexual lichen fungus. Our data further support the view of the Macaronesian Islands as the end of the colonization road for this symbiotic ascomycete.

The study investigated the ecophysiological and ultrastructural effects of dust pollution from a cement industry in the lichen species Evernia prunastri and Xanthoria parietina , which were exposed for 30, 90 and 180 days around a cement mill, two quarries, and inhabited and agricultural sites in SW Slovakia. The results showed that dust deposition from quarrying activities and cement works at the cement mill (mainly enriched in Ca, Fe and Ti) significantly affected the photosynthetic apparatus of E. prunastri (sensitive to dust and habitat eutrophication), while X. parietina (tolerant to dust and habitat eutrophication) adapted to the new environment. The length of the exposure strongly affected the vitality of the mycobiont (measured as dehydrogenase activity) in transplanted lichens. Dust deposition led to ultrastructural alterations, including lipid droplets increase, swelling of cellular components, thylakoid degeneration and sometimes plasmolysis, which, on the whole, gave the cells an aged appearance. Photosynthetic parameters deserve further attention as potential indicators for monitoring early biological symptoms of the air pollution caused during cement production.

Anthropic activities such as the emission of pollutants resulting from industrial and agropastoral activities promote several changes in urban and forest areas. Lichens are organisms that are used in air quality evaluations due to their sensitivity to these changes. The aim of this study is to analyze the presence of morphophysiological damages and the metal concentration in samples of the lichen Parmotrema tinctorum, in urban and forest areas, checking for possible parameter variations between these areas, in the different matrices and seasons in the Southern region of Brazil. Six areas were selected (urban and forest), assigned to the rural/urban and urban/industrial matrices belonging to the watershed of the Rio dos Sinos, Brazil. The following parameters were analyzed: Index of photobiont vitality (IPV), photosynthetic pigments (chlorophyll, carotene, and pheophytin) and metals (Cu, Cr, Zn, Pb, and Ni) in the thallus of the lichen. The data were analyzed by an ANOVA one way, Pearson correlation test, and principal component analysis (PCA). Variations in the morphophysiological parameters were recorded in all the areas revealing significant differences. The lowest IPV values and highest concentration of metals were recorded in the urban environment, while low chlorophyll levels were found in the forest areas. The PCA showed a distinction between the areas and the season. The level of urbanization, vehicle traffic, and the weather conditions might have influenced the results. The use of P. tinctorum, the index of photobiont vitality and chlorophyll and pheophytin content, has proved to be an efficient tool to diagnose the air quality in the areas analyzed, allowing its use as a model in air monitoring studies, both in urban and forest areas, as well as in distinct matrices in the subtropical region.

Species recognition in lichen-forming fungi has been a challenge because of unsettled species concepts, few taxonomically relevant traits, and limitations of traditionally used morphological and chemical characters for identifying closely related species. Here we analyze species diversity in the cosmopolitan genus Protoparmelia s.l. The ~25 described species in this group occur across diverse habitats from the boreal-arctic/alpine to the tropics, but their relationship to each other remains unexplored. In this study, we inferred the phylogeny of 18 species currently assigned to this genus based on 160 specimens and six markers: mtSSU, nuLSU, ITS, RPB1, MCM7, and TSR1. We assessed the circumscription of species-level lineages in Protoparmelia s. str. using two coalescent-based species delimitation methods--BP&P and spedeSTEM. Our results suggest the presence of a tropical and an extra-tropical lineage, and eleven previously unrecognized distinct species-level lineages in Protoparmelia s. str. Several cryptic lineages were discovered as compared to phenotype-based species delimitation. Many of the putative species are supported by geographic evidence.

Although classification at supra-specific ranks is inherently arbitrary, comparable taxonomic ranks within clades can facilitate more consistent classifications and objective comparisons among taxa. Different circumscriptions of the hyper-diverse lichen-forming fungal family Parmeliaceae and widely different generic circumscriptions among authors have been proposed. For this study, we use a recently developed temporal approach that uses time-calibrated chronograms to identify temporal bands for specific ranks in Parmeliaceae and allied groups with the overarching goal of establishing a consistent, stable classification. A data set of 330 species, representing 73 genera in the family and 52 species of related families was used to address the circumscription of Parmeliaceae and its genera following the proposed temporal approach. Based on the results of this study, we propose a revised, temporal-based classification for Parmeliaceae, including all clades that share a common ancestor 102.13–112.88 Ma for families and a time window of 29.45–32.55 Ma for genera. Forty-five of the currently accepted genera in Parmeliaceae were supported in their current circumscription. Two subfamilies are accepted within Parmeliaceae: Protoparmelioideae Divakar et al. subfam. nov., including Protoparmelia and the resurrected genus Maronina, and Parmelioideae, including the bulk of genera in the family. The new genus Austromelanelixia Divakar et al. is proposed to accommodate a clade of southern Hemisphere species previously included in Melanelixia. Eumitria and tentatively Dolichousnea are resurrected as genera separate from Usnea . The following genera are reduced to synonymy: Allocetraria, Cetrariella, Usnocetraria , and Vulpicida with Cetraria; Arctocetraria, Cetreliopsis, Flavocetraria, Kaernefeltia, Masonhalea, Tuckermanella , and Tuckermannopsis with Nephromopsis ; and the lichenicolous genera Nesolechia and Raesaenenia with the lichen-forming genera Punctelia and Protousnea, respectively. A total of 47 new combinations and three new names at the species level are proposed.

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.

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 CO2 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.

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.

We evaluate stability of cesium (Cs) and other alkali-metal cation complexes of lichen metabolites in both gas and aqueous phases to discuss why lichens can retain radioactive Cs in the thalli over several years. We focus on oxalic acid, (+)-usnic acid, atranorin, lecanoric acid, and protocetraric acid, which are common metabolite substances in various lichens including, e.g., Flavoparmelia caperata and Parmotrema tinctorum retaining Cs in Fukushima, Japan. By performing quantum chemical calculations, their gas-phase complexation energies and aqueous-solution complexation free energies with alkali-metal cations are computed for their neutral and deprotonated cases. Consequently, all the molecules are found to energetically favor cation complexations and the preference order is Li + > Na + > K + > Rb + > Cs + for all conditions, indicating no specific Cs selectivity but strong binding with all alkali cations. Comparing complexation stabilities among these metabolites, lecanoric and protocetraric acids seen in medullary layer are found to keep higher affinity in their neutral case, while (+)-usnic acid and atranorin in upper cortex exhibit rather strong affinity only in deprotonated cases through forming stable six atoms’ ring containing alkali cation chelated by two oxygens. These results suggest that the medullary layer can catch all alkali cations in a wide pH range around the physiological one, while the upper cortex can effectively block penetration of metal ions when the metal stress grows. Such insights highlight a physiological role of metabolites like blocking of metal-cation migrations into intracellular tissues, and explain long-term retention of alkali cations including Cs in lichens containing enough such metabolites to bind them.