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Abstract 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 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. [ABSTRACT FROM AUTHOR]

This study investigates the phylogeny and symbiotic relationships of the foliose lichen genus Remototrachyna, R. crenata and R. rhabdiformis from the Western Ghats through morphological, chemical and molecular phylogenetic tools using concatenated ITS and LSU data analyses. The study further identified the photobiont Trebouxia species in R. crenata and R. rhabdiformis based on ITS sequence data and phylogeny. The Trebouxia species of R. crenata and R. rhabdiformis were delineated as a major clade closely related to Trebouxia sp. voucher SF31 sequenced from Usnea ghattensis belonging to Trebouxia Clade I. This study represents a pioneering effort to unravel the enigmatic lichen symbiosis that exists in the genus Remototrachyna from the Western Ghats of India. [ABSTRACT FROM AUTHOR]

Cyanobacteria require liquid water for photosynthesis, whereas green algae can photosynthesise with water vapour alone. We discovered that several Lobaria spp. which normally have cyanobacteria as the sole photobiont, in some regions of the trans-Himalayas also harboured green algae. We tested whether green algal acquisition was: limited to high elevations; obtained from neighbouring chloro- Lobaria species; enabled photosynthesis at low humidity. Lobaria spp. were collected from 2000 to 4000 m elevation. Spectrophotometry quantified green algal abundance by measuring chlorophyll b (absent in cyanobacteria). Thalli cross-sections visually confirmed green algal presence. We sequenced gene regions: Lobaria (ITS-EF-1α-RPB2), green algae (18S-RBC-L) and Nostoc (16S). Phylogenetic analysis determined myco-photobiont associations. We used a custom closed-circuit gas exchange system with an infrared gas analyser to measure CO2 exchange rates for desiccated specimens at 33%, 76%, 86% and 98% humidity. Cross-sections revealed that the photobiont layers in putative cyano -Lobaria contained both cyanobacteria and green algae, indicating that they should be considered chloro-cyanolichens. Chloro- Lobaria had no visible cephalodia nor cyanobacteria in the photobiont layer. Chloro- Lobaria and chloro-cyano- Lobaria had comparable levels of chlorophyll b. Chloro- Lobaria usually contained Symbiochloris. Chloro-cyano- Lobaria mainly associated with Parachloroidium and Nostoc ; infrequently with Symbiochloris , Apatococcus, Chloroidium , Pseudochlorella, Trebouxia. Sequences from two green algal genera were obtained from within some thalli. Desiccated specimens of every Lobaria species could attain net photosynthesis with light exposure and 33% humidity. CO2 exchange dynamics over a five-day period differed between species. At all elevations, chloro-cyano- Lobaria spp. had abundant green algae in the photobiont layer, but green algal strains mostly differed to those of chloro- Lobaria spp. Both chloro- Lobaria and chloro-cyano- Lobaria were capable of conducting photosynthesis without liquid water. The data strongly suggest that they attained positive net photosynthesis. [ABSTRACT FROM AUTHOR]

The reproduction and dispersal strategies of lichens play a major role in shaping their population structure and photobiont diversity. Sexual reproduction, which is common, leads to high lichen genetic diversity and low photobiont selectivity. However, the lichen genus Endocarpon adopts a special co-dispersal model in which algal cells from the photobiont and ascospores from the mycobiont are released together into the environment. To explore the dispersal strategy impact on population structures, a total of 62 Endocarpon individuals and 12 related Verrucariaceae genera individuals, representing co-dispersal strategy and conventional independent dispersal mode were studied. Phylogenetic analysis revealed that Endocarpon, with a large-scale geographical distribution, showed an extremely high specificity of symbiotic associations with their photobiont. Furthermore, three types of group I intron at 1769 site have been found in most Endocarpon mycobionts, which showed a high variety of group I intron in the same insertion site even in the same species collected from one location. This study suggested that the ascospore-alga co-dispersal mode of Endocarpon resulted in this unusual mycobiont-photobiont relationship; also provided an evidence for the horizontal transfer of group I intron that may suggest the origin of the complexity and diversity of lichen symbiotic associations. [ABSTRACT FROM AUTHOR]

This paper, with Italy as a case-study, provides a general overview on the ecology of lichenicolous lichens, i.e. those which start their life-cycle on the thallus of other lichens. It aims at testing whether some ecological factors do exert a positive selective pressure on the lichenicolous lifestyle. The incidence of some biological traits (photobionts, growth-forms and reproductive strategies) in lichenicolous and non-lichenicolous lichens was compared, on a set of 3005 infrageneric taxa potentially occurring in Italy, 189 of which are lichenicolous. Lichenicolous lichens have a much higher incidence of coccoid (non-trentepohlioid) green algae, crustose growth-forms and sexual reproduction. A matrix of the 2762 species with phycobionts and some main ecological descriptors was subjected to ordination. Lichenicolous lichens occupy a well-defined portion of the ecological space, tending to grow on rocks in dry, well-lit habitats where a germinating spore is likely to have a short life-span, at all altitudes. This corroborates the hypothesis that at least some of them are not true "parasites", as they are often called, but gather the photobionts - which have already adapted to local ecological conditions - from their hosts, eventually developing an independent thallus. [ABSTRACT FROM AUTHOR]

The article discusses the current proposals to redefine lichens and highlights the disadvantages of these proposals. Lichens are currently defined as a symbiotic relationship between a fungus (mycobiont) and an alga or cyanobacterium (photobiont). However, some lichen biologists have suggested expanding this definition to include other microorganisms that colonize the lichen thallus. The article argues that these proposals are not justified because the roles and significance of these additional microorganisms are still poorly understood and their inclusion would blur the distinction between lichens and other ecosystems. The article concludes that the current definition of lichens as a partnership between mycobiont and photobiont remains meaningful and should not be amended. [Extracted from the article]

Carbon‐concentrating mechanisms (CCMs) are a widespread phenomenon in photosynthetic organisms. In vascular plants, the evolution of CCMs ([C44‐carbon compound] and crassulacean acid metabolism [CAM]) is associated with significant shifts, most often to hot, dry and bright, or aquatic environments. If and how CCMs drive distributions of other terrestrial photosynthetic organisms, remains little studied. Lichens are ecologically important obligate symbioses between fungi and photosynthetic organisms. The primary photosynthetic partner in these symbioses can include CCM‐presenting cyanobacteria (as carboxysomes), CCM‐presenting green algae (as pyrenoids) or green algae lacking any CCM. We use an extensive dataset of lichen communities from eastern North America, spanning a wide climatic range, to test the importance of CCMs as predictors of lichen ecology and distribution. We show that the presence or absence of CCMs leads to opposite responses to temperature and precipitation in green algal lichens, and different responses in cyanobacterial lichens. These responses contrast with our understanding of lichen physiology, whereby CCMs mitigate carbon limitation by water saturation at the cost of efficient use of vapor hydration. This study demonstrates that CCM status is a key functional trait in obligate lichen symbioses, equivalent in importance to its role in vascular plants, and central for studying present and future climate responses. [ABSTRACT FROM AUTHOR]

Lichenized fungi as a multicomponent biological system are an interesting but difficult research object. In view of the complex interaction of the myco- and photobiont, the slow growth of natural thalli, their exceptional exactingness to environmental factors, the study of the biological characteristics of lichens is significantly difficult. This article discusses the problem of studying the anatomical and morphological structure of the myco- and photobiont of lichenized fungi using the tissue culture method on the example of rare species (Lobaria pulmonaria) and species with pharmaceutical potential (Usnea dasopoga, Cetraria islandica). The authors proposed a method for cultivating of myco- and photobionts of these species on synthetic nutrient media for the purpose of further research on the characteristics of lichenized fungi. Fragments of thalli, apothecia (for C. islandica), and soredia (for U. dasopoga and L. pulmonaria) were used as donor material. For the introduction of the photobiont under sterile conditions, a homogenate of the thallus region was prepared. Pure cultures of the myco- and photobiont of L. pulmonaria, U. dasopoga, and C. islandica were obtained on three types of hormone-free nutrient media, namely MS nutrient medium, modified MS medium with a reduced nitrogen content, Czapek medium. The verification of the research results was carried out taking into account the microscopy of the obtained cultures of myco- and photobionts. Using microscopy, the dimensional characteristics of the obtained mycobionts were determined. The diameters of the hyphae of L. pulmonaria, U. dasopoga, and C. islandica were 4.3–4.4 µm, 4.6 µm, and 4.1–4.3 µm, respectively, which corresponds to the size of the hyphae in natural samples. The use of the tissue culture method makes it possible to study and analyze the biological characteristics of lichenized fungi as a multicomponent biological system, as well as to contribute to the preservation of rare species and their components in the Red Data Book; while optimizing the method of cultivating myco- and photobiont on synthetic nutrient media, to reduce the anthropogenic load on natural populations of lichens when using them as medicinal raw materials.

Lichens have long been considered as composite organisms composed of algae and/or cyanobacteria hosted by a fungus in a mutualistic relationship. Other organisms have been gradually discovered within the lichen thalli, such as multiple algal species, yeasts, or even viruses. Of pivotal relevance is the existence of the lichen microbiome, which is a community of microorganisms that can be found living together on the lichen surface. This community performs a growing number of functions. In this entry, we explore the journey of lichens being considered from a dual partnership to a multi-species symbiotic relationship.

Gyalectidium setiferum Vězda & Sérus., a foliicolous lichen characterized by having some vertically oriented whitish translucent cilia surrounding a swollen greenish diahyphal mass, is reported as new to East Asia. It was collected from two localities on the lowlands of central Honshu in Japan where it grew on leaves of Aucuba japonica and Maesa japonica. The description with illustration based on the Japanese material is given. Gyalectidium setiferum has been included in this genus due to its morphological features even though no fertile specimen has been found so far. The monophyly of this species and its position in this genus were newly confirmed by the molecular phylogenetic analysis based on mtSSU and nrLSU. [ABSTRACT FROM AUTHOR]

The heavy metal Cd accumulates in trophic chains, constituting a toxic element for photosynthesizing organisms, including the algal photobionts of lichen. Thus, as lichens respond differently to heavy metal toxicity, we hypothesized that the species Parmotrema tinctorum and Usnea barbata, commonly sampled in the Cerrado ecoregion, could be sensitive to Cd and, therefore, be used to biomonitor the dispersion of this metal. We also aimed to indicate the responsiveness of biological markers to Cd in these species by exposing the thalli to simulated rainfall with increasing metal concentrations. We observed that both lichen species are responsive to Cd stress; however, different pathways are accessed. The synthesis of carotenoids by P. tinctorum and the production of antioxidant enzymes by U. barbata seem to constitute relevant response strategies to Cd-induced stress. The lichen morphoanatomy, cell viability, photobiont vitality index, chlorophyll a fluorescence, and chlorophyll a synthesis were efficient biomarkers for the effects of increasing Cd exposure in P. tinctorum, being the variables primarily associated with damage to the photobiont. For U. barbata, the lichen morphoanatomy, photochemistry, and antioxidant enzyme activity (catalase, superoxide dismutase and ascorbate peroxidase) were essential to reflect Cd toxicity. However, the species P. tinctorum was characterized as the most sensitive to Cd toxicity, constituting a good bioindicator for the presence of this metal. It can be used in the diagnosis of air quality in urban and industrial areas or even in forest areas influenced by Cd in phosphate fertilizers. [ABSTRACT FROM AUTHOR]

Definition: Lichens have long been considered as composite organisms composed of algae and/or cyanobacteria hosted by a fungus in a mutualistic relationship. Other organisms have been gradually discovered within the lichen thalli, such as multiple algal species, yeasts, or even viruses. Of pivotal relevance is the existence of the lichen microbiome, which is a community of microorganisms that can be found living together on the lichen surface. This community performs a growing number of functions. In this entry, we explore the journey of lichens being considered from a dual partnership to a multi-species symbiotic relationship. [ABSTRACT FROM AUTHOR]

We propose that a qualitative trait approach based on more detailed nuanced traits may reveal previously overlooked patterns, especially when combined with phylogenetic perspectives. By sampling epiphytic lichens and using a functional approach based on nuanced qualitative traits, such as a much greater resolution over photobiont identity, type of cortex, and chemical compounds, we evaluated the effects of environmental filtering and phylogenetic constraints on community assembly along a natural succession of Atlantic rainforest. We found changes in taxonomic, functional, and phylogenetic composition, structure, and diversity. Functional traits such as photobiont genera, type of cortex, reproductive structures, propagule size, and protection strategies showed strong responses to succession. Mature forests with a closed canopy impose strong environment filtering that is reflected in lichen species turnover, limiting diversity, but also holding different functional and phylogenetic composition. The use of a nuanced qualitative trait approach may overcome some of the limitations of using this type of traits and shows the importance of often‐overlooked key lichen functional traits, including the presence of carbon‐concentrating mechanisms in photobionts and cortex properties. Furthermore, this is the first study showing how patterns of phylogenetic assembly along forest succession structure lichen communities. [ABSTRACT FROM AUTHOR]

Abstract We propose that a qualitative trait approach based on more detailed nuanced traits may reveal previously overlooked patterns, especially when combined with phylogenetic perspectives. By sampling epiphytic lichens and using a functional approach based on nuanced qualitative traits, such as a much greater resolution over photobiont identity, type of cortex, and chemical compounds, we evaluated the effects of environmental filtering and phylogenetic constraints on community assembly along a natural succession of Atlantic rainforest. We found changes in taxonomic, functional, and phylogenetic composition, structure, and diversity. Functional traits such as photobiont genera, type of cortex, reproductive structures, propagule size, and protection strategies showed strong responses to succession. Mature forests with a closed canopy impose strong environment filtering that is reflected in lichen species turnover, limiting diversity, but also holding different functional and phylogenetic composition. The use of a nuanced qualitative trait approach may overcome some of the limitations of using this type of traits and shows the importance of often‐overlooked key lichen functional traits, including the presence of carbon‐concentrating mechanisms in photobionts and cortex properties. Furthermore, this is the first study showing how patterns of phylogenetic assembly along forest succession structure lichen communities.

Lichen associations are overwhelmingly supported by carbon produced by photosynthetic algal symbionts. These algae have diversified to occupy nearly all climates and continents; however, we have a limited understanding of how their climatic niches have evolved through time. Here we extend previous work and ask whether phylogenetic signal in, and the evolution of, climatic niche, varies across climatic variables, phylogenetic scales, and among algal lineages in Trebouxia— the most common genus of lichen-forming algae. Our analyses reveal heterogeneous levels of phylogenetic signal across variables, and that contrasting models of evolution underlie the evolution of climatic niche divergence. Together these analyses demonstrate the variable processes responsible for shaping climatic tolerance in Trebouxia , and provide a framework within which to better understand potential responses to climate change-associated perturbations. Such predictions reveal a disturbing trend in which the pace at which modern climate change is proceeding will vastly exceed the rate at which Trebouxia climatic niches have previously evolved. [ABSTRACT FROM AUTHOR]

Lichenized algae and cyanobacteria are known to be shared and selected by unrelated lichen-forming fungi coexisting in so-called photobiont-mediated guilds. Life in such a guild could be crucial for the survival of a large group of lichen fungi dependent on horizontal transmission of photobionts. Here, we investigate frequent lichen phycobionts of the genus Trebouxia in rock-dwelling lichen communities. We found intensive and repeated sharing of specific Trebouxia assemblages by co-occurring lichens across distant localities. Rock chemistry, expressed as pH, determined the composition of photobiont pools and separated three saxicolous lichen guilds, sharing environmentally specific photobiont groups. Moreover, unlike the majority of lichen fungi, many Trebouxia photobionts represented opportunists in the choice of general substrate form (soil-rock-tree bark/wood), maintaining their pH preferences. Thus, saxicolous communities form just a part of a complex guild system that is in principle mediated by environmentally conditioned groups of naturally co-occurring photobionts. The complexity of the system is influenced by diverse photobiont life strategies, including also dispersal style. The findings of photobionts strictly or predominantly associated with sexually reproducing fungi stimulated us to emphasize the role of free-dispersing photobionts in the establishment and maintenance of lichen guilds. [ABSTRACT FROM AUTHOR]

Climatic factors, soil chemistry and geography are considered as major factors affecting lichen distribution and diversity. To determine how these factors limit or support the associations between the symbiotic partners, we revise the lichen symbiosis as a network of relationships here. More than one thousand thalli of terricolous Cladonia lichens were collected at sites with a wide range of soil chemical properties from seven biogeographical regions of Europe. A total of 18 OTUs of the algal genus Asterochloris and 181 OTUs of Cladonia mycobiont were identified. We displayed all realized pairwise mycobiont–photobiont relationships and performed modularity analysis. It revealed four virtually separated modules of cooperating OTUs. The modules differed in mean annual temperature, isothermality, precipitation, evapotranspiration, soil pH, nitrogen, and carbon contents. Photobiont switching was strictly limited to algae from one module, i.e., algae of similar ecological preferences, and only few mycobionts were able to cooperate with photobionts from different modules. Thus, Cladonia mycobionts generally cannot widen their ecological niches through photobiont switching. The modules also differed in the functional traits of the mycobionts, e.g., sexual reproduction rate, presence of soredia, and thallus type. These traits may represent adaptations to the environmental conditions that drive the differentiation of the modules. In conclusion, the promiscuity in Cladonia mycobionts is strictly limited by climatic factors and soil chemistry. [ABSTRACT FROM AUTHOR]

Lichen associations are overwhelmingly supported by carbon produced by photosynthetic algal symbionts. These algae have diversified to occupy nearly all climates and continents; however, we have a limited understanding of how their climatic niches have evolved through time. Here we extend previous work and ask whether phylogenetic signal in, and the evolution of, climatic niche, varies across climatic variables, phylogenetic scales, and among algal lineages in Trebouxia—the most common genus of lichen-forming algae. Our analyses reveal heterogeneous levels of phylogenetic signal across variables, and that contrasting models of evolution underlie the evolution of climatic niche divergence. Together these analyses demonstrate the variable processes responsible for shaping climatic tolerance in Trebouxia, and provide a framework within which to better understand potential responses to climate change-associated perturbations. Such predictions reveal a disturbing trend in which the pace at which modern climate change is proceeding will vastly exceed the rate at which Trebouxia climatic niches have previously evolved.

Climatic factors, soil chemistry and geography are considered as major factors affecting lichen distribution and diversity. To determine how these factors limit or support the associations between the symbiotic partners, we revise the lichen symbiosis as a network of relationships here. More than one thousand thalli of terricolous Cladonia lichens were collected at sites with a wide range of soil chemical properties from seven biogeographical regions of Europe. A total of 18 OTUs of the algal genus Asterochloris and 181 OTUs of Cladonia mycobiont were identified. We displayed all realized pairwise mycobiont–photobiont relationships and performed modularity analysis. It revealed four virtually separated modules of cooperating OTUs. The modules differed in mean annual temperature, isothermality, precipitation, evapotranspiration, soil pH, nitrogen, and carbon contents. Photobiont switching was strictly limited to algae from one module, i.e., algae of similar ecological preferences, and only few mycobionts were able to cooperate with photobionts from different modules. Thus, Cladonia mycobionts generally cannot widen their ecological niches through photobiont switching. The modules also differed in the functional traits of the mycobionts, e.g., sexual reproduction rate, presence of soredia, and thallus type. These traits may represent adaptations to the environmental conditions that drive the differentiation of the modules. In conclusion, the promiscuity in Cladonia mycobionts is strictly limited by climatic factors and soil chemistry.

The heavy metal Cd accumulates in trophic chains, constituting a toxic element for photosynthesizing organisms, including the algal photobionts of lichen. Thus, as lichens respond differently to heavy metal toxicity, we hypothesized that the species Parmotrema tinctorum and Usnea barbata, commonly sampled in the Cerrado ecoregion, could be sensitive to Cd and, therefore, be used to biomonitor the dispersion of this metal. We also aimed to indicate the responsiveness of biological markers to Cd in these species by exposing the thalli to simulated rainfall with increasing metal concentrations. We observed that both lichen species are responsive to Cd stress; however, different pathways are accessed. The synthesis of carotenoids by P. tinctorum and the production of antioxidant enzymes by U. barbata seem to constitute relevant response strategies to Cd-induced stress. The lichen morphoanatomy, cell viability, photobiont vitality index, chlorophyll a fluorescence, and chlorophyll a synthesis were efficient biomarkers for the effects of increasing Cd exposure in P. tinctorum, being the variables primarily associated with damage to the photobiont. For U. barbata, the lichen morphoanatomy, photochemistry, and antioxidant enzyme activity (catalase, superoxide dismutase and ascorbate peroxidase) were essential to reflect Cd toxicity. However, the species P. tinctorum was characterized as the most sensitive to Cd toxicity, constituting a good bioindicator for the presence of this metal. It can be used in the diagnosis of air quality in urban and industrial areas or even in forest areas influenced by Cd in phosphate fertilizers.

Interactions within lichen communities include, in addition to close mutualistic associations between the main partners of specific lichen symbioses, also more elusive relationships between members of a wider symbiotic community. Here, we analyze association patterns of cyanolichen symbionts in the tropical montane forests of Taita Hills, southern Kenya, which is part of the Eastern Afromontane biodiversity hotspot. The cyanolichen specimens analyzed represent 74 mycobiont taxa within the order Peltigerales (Ascomycota), associating with 115 different variants of the photobionts genus Nostoc (Cyanobacteria). Our analysis demonstrates wide sharing of photobionts and reveals the presence of several photobiont-mediated lichen guilds. Over half of all mycobionts share photobionts with other fungal species, often from different genera or even families, while some others are strict specialists and exclusively associate with a single photobiont variant. The most extensive symbiont network involves 24 different fungal species from five genera associating with 38 Nostoc photobionts. The Nostoc photobionts belong to two main groups, the Nephroma-type Nostoc and the Collema/Peltigera-type Nostoc, and nearly all mycobionts associate only with variants of one group. Among the mycobionts, species that produce cephalodia and those without symbiotic propagules tend to be most promiscuous in photobiont choice. The extent of photobiont sharing and the structure of interaction networks differ dramatically between the two major photobiont-mediated guilds, being both more prevalent and nested among Nephroma guild fungi and more compartmentalized among Peltigera guild fungi. This presumably reflects differences in the ecological characteristics and/or requirements of the two main groups of photobionts. The same two groups of Nostoc have previously been identified from many lichens in various lichen-rich ecosystems in different parts of the world, indicating that photobiont sharing between fungal species is an integral part of lichen ecology globally. In many cases, symbiotically dispersing lichens can facilitate the dispersal of sexually reproducing species, promoting establishment and adaptation into new and marginal habitats and thus driving evolutionary diversification.

Interactions within lichen communities include, in addition to close mutualistic associations between the main partners of specific lichen symbioses, also more elusive relationships between members of a wider symbiotic community. Here, we analyze association patterns of cyanolichen symbionts in the tropical montane forests of Taita Hills, southern Kenya, which is part of the Eastern Afromontane biodiversity hotspot. The cyanolichen specimens analyzed represent 74 mycobiont taxa within the order Peltigerales (Ascomycota), associating with 115 different variants of the photobionts genus Nostoc (Cyanobacteria). Our analysis demonstrates wide sharing of photobionts and reveals the presence of several photobiont-mediated lichen guilds. Over half of all mycobionts share photobionts with other fungal species, often from different genera or even families, while some others are strict specialists and exclusively associate with a single photobiont variant. The most extensive symbiont network involves 24 different fungal species from five genera associating with 38 Nostoc photobionts. The Nostoc photobionts belong to two main groups, the Nephroma -type Nostoc and the Collema / Peltigera -type Nostoc , and nearly all mycobionts associate only with variants of one group. Among the mycobionts, species that produce cephalodia and those without symbiotic propagules tend to be most promiscuous in photobiont choice. The extent of photobiont sharing and the structure of interaction networks differ dramatically between the two major photobiont-mediated guilds, being both more prevalent and nested among Nephroma guild fungi and more compartmentalized among Peltigera guild fungi. This presumably reflects differences in the ecological characteristics and/or requirements of the two main groups of photobionts. The same two groups of Nostoc have previously been identified from many lichens in various lichen-rich ecosystems in different parts of the world, indicating that photobiont sharing between fungal species is an integral part of lichen ecology globally. In many cases, symbiotically dispersing lichens can facilitate the dispersal of sexually reproducing species, promoting establishment and adaptation into new and marginal habitats and thus driving evolutionary diversification. [ABSTRACT FROM AUTHOR]

Lichens are classic models of symbiosis, and one of the most frequent nutritional modes among fungi. The ecologically and geographically widespread lichen-forming algal (LFA) genus Trebouxia is one of the best-studied groups of LFA and associates with over 7000 fungal species. Despite its importance, little is known about its diversification. We synthesized twenty years of publicly available data by characterizing the ecological preferences of this group and testing for time-variant shifts in climatic regimes over a distribution of trees. We found evidence for limited shifts among regimes, but that disparate lineages convergently evolved similar ecological tolerances. Early Trebouxia lineages were largely forest specialists or habitat generalists that occupied a regime whose extant members occur in moderate climates. Trebouxia then convergently diversified in non-forested habitats and expanded into regimes whose modern representatives occupy wet-warm and cool-dry climates. We rejected models in which climatic diversification slowed through time, suggesting climatic diversification is inconsistent with that expected under an adaptive radiation. In addition, we found that climatic and vegetative regime shifts broadly coincided with the evolution of biomes and associated or similar taxa. Together, our work illustrates how this keystone symbiont from an iconic symbiosis evolved to occupy diverse habitats across the globe. [ABSTRACT FROM AUTHOR]

This article is a Commentary on Spribille et al. (2022), 234: 1566–1582. [ABSTRACT FROM AUTHOR]

Lichens are remarkable and classic examples of symbiotic organisms that have fascinated scientists for centuries. Yet, it has only been for a couple of decades that significant advances have focused on the diversity of their green algal and/or cyanobacterial photobionts. Cyanolichens, which contain cyanobacteria as their photosynthetic partner, include up to 10% of all known lichens and, as such, studies on their cyanobionts are much rarer compared to their green algal counterparts. For the unicellular cyanobionts, i.e. cyanobacteria that do not form filaments, these studies are even scarcer. Nonetheless, these currently include at least 10 different genera in the cosmopolitan lichen order Lichinales. An international consortium (International Network of CyanoBionts; INCb) will tackle this lack of knowledge. In this article, we discuss the status of current unicellular cyanobiont research, compare the taxonomic resolution of photobionts from cyanolichens with those of green algal lichens (chlorolichens), and give a roadmap of research on how to recondition the underestimated fraction of symbiotic unicellular cyanobacteria in lichens., Competing Interests: None declared., (© The Author(s) 2024. Published by Oxford University Press on behalf of the International Society for Microbial Ecology.)

The reproduction and dispersal strategies of lichens play a major role in shaping their population structure and photobiont diversity. Sexual reproduction, which is common, leads to high lichen genetic diversity and low photobiont selectivity. However, the lichen genus Endocarpon adopts a special co-dispersal model in which algal cells from the photobiont and ascospores from the mycobiont are released together into the environment. To explore the dispersal strategy impact on population structures, a total of 62 Endocarpon individuals and 12 related Verrucariaceae genera individuals, representing co-dispersal strategy and conventional independent dispersal mode were studied. Phylogenetic analysis revealed that Endocarpon , with a large-scale geographical distribution, showed an extremely high specificity of symbiotic associations with their photobiont. Furthermore, three types of group I intron at 1769 site have been found in most Endocarpon mycobionts, which showed a high variety of group I intron in the same insertion site even in the same species collected from one location. This study suggested that the ascospore-alga co-dispersal mode of Endocarpon resulted in this unusual mycobiont-photobiont relationship; also provided an evidence for the horizontal transfer of group I intron that may suggest the origin of the complexity and diversity of lichen symbiotic associations., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (2024, by The Mycological Society of Japan.)

Lichens are fungi (mycobionts) that form symbiotic associations with photoautotrophic prokaryotes or eukaryotes (photobionts); however, some species can exchange photosynthetic partners during their lifecycles. This phenomenon modifies the morphology of lichens and consequently influences the taxonomy of lichenized fungi. Here, a few such cases in which the photobionts influenced the taxonomy and systematics of lichenized fungi are reviewed. Two different morphotypes of the same species – known as photomorphs – were classified as different species and sometimes different genera. Moreover, different types of photobionts and the absence or presence (optional lichenization) of an alga in the thallus were believed to be diagnostic characters for discriminating genera. However, the taxonomy and systematics of lichens are based always, according to Article F.1.1. of the International Code of Nomenclature for Algae, Fungi, and Plants, on the fungal partner and only one name is applied.

Lichens are fungi (mycobionts) that form symbiotic associations with photoautotrophic prokaryotes or eukaryotes (photobionts); however, some species can exchange photosynthetic partners during their lifecycles. This phenomenon modifies the morphology of lichens and consequently influences the taxonomy of lichenized fungi. Here, a few such cases in which the photobionts influenced the taxonomy and systematics of lichenized fungi are reviewed. Two different morphotypes of the same species - known as photomorphs - were classified as different species and sometimes different genera. Moreover, different types of photobionts and the absence or presence (optional lichenization) of an alga in the thallus were believed to be diagnostic characters for discriminating genera. However, the taxonomy and systematics of lichens are based always, according to Article F.1.1. of the International Code of Nomenclature for Algae, Fungi, and Plants, on the fungal partner and only one name is applied. [ABSTRACT FROM AUTHOR]

Background and Aims Lichens represent a symbiotic relationship between at least one fungal and one photosynthetic partner. The association between the lichen-forming fungus Mastodia tessellata (Verrucariaceae) and different species of Prasiola (Trebouxiophyceae) has an amphipolar distribution and represents a unique case study for the understanding of lichen symbiosis because of the macroalgal nature of the photobiont, the flexibility of the symbiotic interaction and the co-existence of free-living and lichenized forms in the same microenvironment. In this context, we aimed to (1) characterize the photosynthetic performance of co-occurring populations of free-living and lichenized Prasiola and (2) assess the effect of the symbiosis on water relations in Prasiola , including its tolerance of desiccation and its survival and performance under sub-zero temperatures. Methods Photochemical responses to irradiance, desiccation and freezing temperature and pressure–volume curves of co-existing free-living and lichenized Prasiola thalli were measured in situ in Livingston Island (Maritime Antarctica). Analyses of photosynthetic pigment, glass transition and ice nucleation temperatures, surface hydrophobicity extent and molecular analyses were conducted in the laboratory. Key Results Free-living and lichenized forms of Prasiola were identified as two different species: P. crispa and Prasiola sp. respectively. While lichenization appears to have no effect on the photochemical performance of the alga or its tolerance of desiccation (in the short term), the symbiotic lifestyle involves (1) changes in water relations, (2) a considerable decrease in the net carbon balance and (3) enhanced freezing tolerance. Conclusions Our results support improved tolerance of sub-zero temperature as the main benefit of lichenization for the photobiont, but highlight that lichenization represents a delicate equilibrium between a mutualistic and a less reciprocal relationship. In a warmer climate scenario, the spread of the free-living Prasiola to the detriment of the lichen form would be likely, with unknown consequences for Maritime Antarctic ecosystems. [ABSTRACT FROM AUTHOR]

To evaluate the importance of morphological and chemical characters used in the recognition of species within the Parmelia omphalodes group, we performed phylogenetic, morphological and chemical analyses of 335 specimens, of which 34 were used for molecular analyses. Phylogenetic analyses, based on ITS rDNA sequences, show that P. pinnatifida is distinct from P. omphalodes and the most important difference between those species is the development of pseudocyphellae. In P. pinnatifida, they are mostly marginal and form white rims along lobes margins, but laminal pseudocyphellae can develop in older parts of thalli and are predominantly connected with marginal pseudocyphellae. In contrast, in P. omphalodes laminal pseudocyphellae are common and are predominantly not connected to marginal pseudocyphellae. Chemical composition of secondary lichen metabolites in both analysed species is identical and therefore this feature is not diagnostic in species recognition. Few samples of P. discordans, species morphologically similar to P. omphalodes and P. pinnatifida, were also included in the analyses and they are nested within the clade of P. omphalodes, despite the different chemistry (protocetraric acid present versus salazinic acid in P. omphalodes). All taxa of the P. omphalodes group occupy similar niches, but their potential distributions are wider than those currently known. The absence of specimens in some localities may be limited by the photobiont availability. Parmelia omphalodes and P. pinnatifida are moderately selective in photobiont choice as they form associations with at least two or three lineages of Trebouxia clade S. Parmelia pinnatifida, as well as P. discordans are associated with Trebouxia OTU S02 which seems to have a broad ecological amplitude. Other lineages of Trebouxia seem to be rarer, especially Trebouxia sp. OTU S04, which is sometimes present in P. pinnatifida. This study indicates the importance of extensive research including morphology, chemistry and analysis of molecular markers of both bionts in taxonomical studies of lichens. [ABSTRACT FROM AUTHOR]

The ‘green’ ciliate Paramecium bursaria lives in mutualistic symbiosis with green algae belonging to the species Chlorella variabilis or Micractinium conductrix. We analysed the diversity of algal endosymbionts and their P. bursaria hosts in nine strains from geographically diverse origins. Therefore, their phylogenies using different molecular markers were inferred. The green paramecia belong to different syngens of P. bursaria. The intracellular algae were assigned to Chl. variabilis, M. conductrix or, surprisingly, Choricystis parasitica. This usually free-living alga co-occurs with M. conductrix in the host’s cytoplasm. Addressing the potential status of Chor. parasitica as second additional endosymbiont, we determined if it is capable of symbiosis establishment and replication within a host cell. Symbiont-free P. bursaria were generated by cycloheximid treatment. Those aposymbiotic P. bursaria were used for experimental infections to investigate the symbiosis specificity not only between P. bursaria and Chor. parasitica but including also Chl. variabilis and M. conductrix. For each algae we observed the uptake and incorporation in individual perialgal vacuoles. These host-symbiont associations are stable since more than five months. Thus, Chor. parasitica and P. bursaria can form an intimate and long-term interaction. This study provides new insights into the diversity of P. bursaria algal symbionts.

This article is a Commentary on Mark et al. (2020), 227: 1362–1375. [ABSTRACT FROM AUTHOR]

The anatomical (thickness of layers of fungal and algal symbionts) and ecophysiological parameters (the content of photosynthetic pigments) of two lichen species (Hypogymnia physodes and Umbilicaria hyperborea) from rocky forest communities of northernmost boreal zone (North-West of Russia) were studied. H. physodes is a lichen with foliose thallus morphology and circumpolar distribution which colonizes almost every substrate. U. hyperborea is strictly epilithic lichen with umbilicate type of thallus morphology and arcto-alpine distribution. Despite the fact that the photobiont in two species are closely related taxa (unicellular green algae of the genus Trebouxia) a high variability of photosynthetic pigments content in H. physodes thalli has been revealed, whereas in U. hyperborea, the mycobiotic showed high structural variability. It can be concluded that both mycobiont and photobiont contribute to their adaptation to environmental factors. In the present study it was obtained that the proportion of variation in anatomical structures and the content of photosynthetic pigments probably less depends on taxonomic position of photobiont and more determined by the plasticity of the individual components of the symbiotrophic organism. [ABSTRACT FROM AUTHOR]

Anthropogenic disturbances can have strong impacts on lichen communities, as well as on individual species of lichenized fungi. Traditionally, lichen monitoring studies are based on the presence and abundance of fungal morphospecies. However, the photobionts, as well photobiont mycobiont interactions also contribute to the structure, composition, and resilience of lichen communities. Here we assess the genetic diversity and interaction patterns of algal and fungal partners in lichen communities along an anthropogenic disturbance gradient in Białowieża Forest (Poland). We sampled a total of 224 lichen thalli in a protected, a managed, and a disturbed area of the forest, and sequenced internal transcribed spacer (ITS) ribosomal DNA (rDNA) of both, fungal and algal partners. Sequence clustering using a 97% similarity threshold resulted in 46 fungal and 23 green algal operational taxonomic units (OTUs). Most of the recovered photobiont OTUs (14 out of 23) had no similar hit in the NCBI-BLAST search, suggesting that even in well studied regions, such as central Europe, a lot of photobiont diversity is yet undiscovered. If a mycobiont was present at more than one site, it was typically associated with the same photobiont OTU(s). Generalist species, i.e., taxa that associate with multiple symbiont partners, occurred in all three disturbance regimes, suggesting that such taxa have few limitations in colonizing or persisting in disturbed areas. Trebouxia jamesii associated with 53% of the fungal OTUs, and was generally the most common photobiont OTU in all areas, implying that lichens that associate with this symbiont are not limited by the availability of compatible photobionts in Central European forests, regardless of land use intensity.

Lichens are symbiotic organisms that are very sensitive to heavy metal pollution. However, there is little evidence of how heavy metal pollution affects the physiological status, ultrastructural changes and distribution of elements in the layers of lichen thalli. For this purpose we simulated metal pollution to lichens and studied its impact on Xanthoria parietina . Thalli were treated with the heavy metals Cu, Zn, Ni, Cd in the form of sulfates at concentrations of 100 µM and 500 µM during 24, 48 and 72 h. Untreated lichens served as controls. We assessed the status of physiological parameters (fluorescence and integrity of chlorophyll a , content of soluble proteins and ergosterol), ultrastructural changes, especially to the photobiont, and the distribution of elements in the layers of thalli in relation to treatment with heavy metals. We found positive correlations between the content of all tested heavy metals and the physiological response. We assessed the toxicity of the selected metals as follows: Cd >= Cu >= Ni > Zn, based on the effects on the photobiont layer in the lichen thallus and physiological measurements. [ABSTRACT FROM AUTHOR]

Secondary metabolites of lichens can be involved in production of chelates with heavy metals. We hypothesized that parietin plays important role in protection of photobiont cells in Xanthoria parietina from an excess of cadmium ions. Two types of X. parietina lichen thalli, natural with presence of secondary metabolite parietin (p+) as well as without parietin (p−) were exposed to different doses of cadmium (up to 300 μmol g −1 dw). Based on determination of the total and intracellular Cd-accumulation, ergosterol and thiobarbituric acid reactive substances (TBARS) content did not show statistically significant differences in the response of both types of thalli (p+ and p−). However, a stronger toxic effect of the highest Cd-dose on photosynthetic pigment content and chlorophyll a fluorescence was observed in the parietin-depleted thalli. The protective role of parietin against Cd excess was better supported and concluded from the differences observed in the production of non-protein thiol compounds (cysteine, glutathione and phytochelatins) involved in Cd detoxification. In the p+ thalli Cys content was stable but GSH content slightly decreased in the studied Cd range, while in the p− thalli these compounds were completely absent at high Cd doses. At Cd doses higher than 37.5 μmol Cd g −1 dw, toxic to both types of X. parietina thalli, Cys and GSH contents were significantly higher in p+ than in p− thalli. Also, the photobiont partner in the p+ thalli was better protected of the metal exposition, and able to produce phytochelatins (PCs) over the whole range of metal, while in the p− thalli the production was completely inhibited at 75 μmol Cd g −1 dw and higher concentrations, together with the inhibition of cysteine (Cys) and reduced glutathione (GSH) production. The obtained results indicate that the parietin layer is a natural barrier decreasing Cd access to algal cells in X. parietina . Comparison of PCs production appeared to be the most sensitive marker for estimation of Cd availability to photobiont in the symbiotic system. [ABSTRACT FROM AUTHOR]

Background: The diversity of lichen fungal components and their photosynthetic partners reflects both ecological and evolutionary factors. In present study, molecular investigations of the internal transcribed spacer of the nuclear ribosomal DNA (ITS nrDNA) region were conducted to analyze the genetic diversity of Umbilicaria esculenta and U. muehlenbergii together with their associated green algae. Result: It was here demonstrated that the reproductive strategy is a principal reason for fungal selectivity to algae. U. muehlenbergii, which disperses via sexual spores, exhibits lower selectivity to its photosynthetic partners than U. esculenta, which has a vegetative reproductive strategy. The difference of genotypic diversity (both fungal and algal) between these two Umbilicaria species is low, although their nucleotide diversity can vary greatly. Conclusions: The present study illustrates that lichen-forming fungi with sexual reproductive strategies are less selective with respect to their photobionts; and reveals that both sexual and vegetative reproduction allow lichens to generate similar amounts of diversity to adapt to the environments. The current study will be helpful for elucidating how lichens with different reproductive strategies adapt to changing environments. [ABSTRACT FROM AUTHOR]

* Premise of the study: The life histories of lichen fungi are not well known and cannot be readily studied in laboratory culture. This work documents in situ the complete life cycle of the widespread crustose lichen Calopadia puiggarii, which reproduces sexually and asexually on the surfaces of leaves. * Methods: Plastic cover slips held in a mesh frame were placed over leaves in the field and successively removed for microphotography of colonizing lichens. * Key results: Macroconidia produced within campylidia encircled photobiont cells and codispersed with them, a feature not reported previously for C. puiggarii. Dispersed macroconidia readily germinated and lichenized the photobionts. Algal cells were often dislodged from the encircling macroconidia, providing a likely source for the free-living populations observed. Aposymbiotically dispersed ascospores germinated and lichenized nearby algal cells soon after dispersal. Thallus areolae merged readily in early development, although adjacent mature thalli were often separated by growth inhibition zones. Pycnidia are reported for the first time in Calopadia-, their pyriform microconidia probably function as male gametes (spermatia). Pycnidia, apothecia, and campylidia began development similarly as darkly pigmented primordia on the fungal prothallus. * Conclusions: Abundant dispersal of ascospores, conidia, and photobionts allows C. puiggarii to quickly colonize leaves with the dual advantages of sexual and asexual reproduction, and with the added convenience of having its algal partner on hand. Fusions and prothallic capture of additional algae provide many opportunities for multiple mycobiont and photobiont genotypes to be combined in a single thallus, but the outcomes of such events remain to be explored. [ABSTRACT FROM AUTHOR]

Background and Aims The integrity and evolution of lichen symbioses depend on a fine-tuned combination of algal and fungal genotypes. Geographically widespread species complexes of lichenized fungi can occur in habitats with slightly varying ecological conditions, and it remains unclear how this variation correlates with symbiont selectivity patterns in lichens. In an attempt to address this question, >300 samples were taken of the globally distributed and ecologically variable lichen-forming species complex Tephromela atra, together with closely allied species, in order to study genetic diversity and the selectivity patterns of their photobionts. Methods Lichen thalli of T. atra and of closely related species T. grumosa, T. nashii and T. atrocaesia were collected from six continents, across 24 countries and 62 localities representing a wide range of habitats. Analyses of genetic diversity and phylogenetic relationships were carried out both for photobionts amplified directly from the lichen thalli and from those isolated in axenic cultures. Morphological and anatomical traits were studied with light and transmission electron microscopy in the isolated algal strains. Key Results Tephromela fungal species were found to associate with 12 lineages of Trebouxia. Five new clades demonstrate the still-unrecognized genetic diversity of lichen algae. Culturable, undescribed lineages were also characterized by phenotypic traits. Strong selectivity of the mycobionts for the photobionts was observed in six monophyletic Tephromela clades. Seven Trebouxia lineages were detected in the poorly resolved lineage T. atra sensu lato, where co-occurrence of multiple photobiont lineages in single thalli was repeatedly observed. Conclusions Low selectivity apparently allows widespread lichen-forming fungi to establish successful symbioses with locally adapted photobionts in a broader range of habitats. This flexibility might correlate with both lower phylogenetic resolution and evolutionary divergence in species complexes of crustose lichen-forming fungi. [ABSTRACT FROM AUTHOR]

* Premise of the study: Many lichens exhibit extensive ranges spanning several ecoregions. It has been hypothesized that this wide ecological amplitude is facilitated by fungal association with locally adapted photobiont strains. * Methods: We studied the identity and geographic distribution of photobionts of the widely distributed North American lichen Ramalina menziesii based on rbcL (chloroplast DNA) and nuclear ribosomal ITS DNA sequences. To test for ecological specialization, we associate photobiont genotypes with local climate and phorophyte. * Key results: Of the photobiont lineages of R. menziesii, 94% belong to a clade including Trebouxia decolorans. The remaining are related to T. jamesii. The photobionts showed (1) significant structure according to ecoregion and phorophyte species and (2) genetic associations with phorophyte species and climate. * Conclusions: Geography, climate, and ecological specialization shape genetic differentiation of lichen photobionts. One great advantage of independent dispersal of the fungus is symbiotic association with locally adapted photobiont strains. [ABSTRACT FROM AUTHOR]