Abstract: Context: Lichens are composite organisms consisting of a symbiotic association of a fungus (the mycobiont) with a photosynthetic partner (the phytobiont), usually either a green alga or cyanobacter...

Abstract: Lichens are symbiotic associations between fungi and photosynthetic algae or cyanobacteria. Microcystins are potent toxins that are responsible for the poisoning of both humans and animals. These toxins are mainly associated with aquatic cyanobacterial blooms, but here we show that the cyanobacterial symbionts of terrestrial lichens from all over the world commonly produce microcystins. We screened 803 lichen specimens from five different continents for cyanobacterial toxins by amplifying a part of the gene cluster encoding the enzyme complex responsible for microcystin production and detecting toxins directly from lichen thalli. We found either the biosynthetic genes for making microcystins or the toxin itself in 12% of all analyzed lichen specimens. A plethora of different microcystins was found with over 50 chemical variants, and many of the variants detected have only rarely been reported from free-living cyanobacteria. In addition, high amounts of nodularin, up to 60 μg g−1, were detected from some lichen thalli. This microcystin analog and potent hepatotoxin has previously been known only from the aquatic bloom-forming genus Nodularia. Our results demonstrate that the production of cyanobacterial hepatotoxins in lichen symbiosis is a global phenomenon and occurs in many different lichen lineages. The very high genetic diversity of the mcyE gene and the chemical diversity of microcystins suggest that lichen symbioses may have been an important environment for diversification of these cyanobacteria.

Abstract: In order to investigate succession of biological soil crusts (BSCs) and their microstructure variability, we conducted this work in Shapotou revegetation region at the southeast edge of Tengger Deser. The results showed that BSCs generally succeeded as a pathway of “Algae crusts, algae–lichen crusts, lichen crusts, lichen–moss crusts and moss crusts”. Occasionally mosses directly occurred on algae crusts, and BSCs succeeded from algae crusts to moss crusts. Crust vertical stratification was a common phenomenon, from top to bottom an inorganic layer, algae-dense layer and algae-sparse layer were divided in algae crusts; a thallus layer, rhizoid layer and sub-rhizoid layer in lichen crusts; a “stem-leaf” layer, rhizoid layer and sub-rhizoid layer in moss crusts, respectively. The main crust binding organisms varied from filamental cyanobacteria (dominated by Microcoleus) in algae crusts to lichen rhizoids, free-living cyanobacterial filaments and fungal hyphaes in lichen crusts, and to moss rhizoids and fungal hyphaes in moss crusts. The dominant phototrophic organisms varied from Microcoleus (algae) in algae crusts to Collema (lichens) in lichen crusts, and to Bryum (or Didymodon and Tortula; mosses) in moss crusts. Total phototrophic biomass increased while the free-living algal biomass decreased with the succession of BSCs. In addition, exopolysaccharides and fine particles accumulated in the course of development and succession of BSCs, all of which lead to a gradual increase in crust thickness and porosity, while decrease in the bulk density.

Abstract: Little is known about the impact of changing temperature regimes on composition and diversity of cryptogam communities in the Arctic and Subarctic, despite the well-known importance of lichens and bryophytes to the functioning and climate feedbacks of northern ecosystems. We investigated changes in diversity and abundance of lichens and bryophytes within long-term (9–16 years) warming experiments and along natural climatic gradients, ranging from Swedish subarctic birch forest and subarctic/subalpine tundra to Alaskan arctic tussock tundra. In both Sweden and Alaska, lichen diversity responded negatively to experimental warming (with the exception of a birch forest) and to higher temperatures along climatic gradients. Bryophytes were less sensitive to experimental warming than lichens, but depending on the length of the gradient, bryophyte diversity decreased both with increasing temperatures and at extremely low temperatures. Among bryophytes, Sphagnum mosses were particularly resistant to experimental warming in terms of both abundance and diversity. Temperature, on both continents, was the main driver of species composition within experiments and along gradients, with the exception of the Swedish subarctic birch forest where amount of litter constituted the best explanatory variable. In a warming experiment in moist acidic tussock tundra in Alaska, temperature together with soil ammonium availability were the most important factors influencing species composition. Overall, dwarf shrub abundance (deciduous and evergreen) was positively related to warming but so were the bryophytes Sphagnum girgensohnii, Hylocomium splendens and Pleurozium schreberi; the majority of other cryptogams showed a negative relationship to warming. This unique combination of intercontinental comparison, natural gradient studies and experimental studies shows that cryptogam diversity and abundance, especially within lichens, is likely to decrease under arctic climate warming. Given the many ecosystem processes affected by cryptogams in high latitudes (e.g. carbon sequestration, N2-fixation, trophic interactions), these changes will have important feedback consequences for ecosystem functions and climate.

Abstract: Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.

Abstract: Lichens are widespread symbioses and play important roles in many terrestrial ecosystems. The genetic structure of lichens is the result of the association between fungal and algal populations constituting the lichen thallus. Using eight fungus- and seven alga-specific highly variable microsatellite markers on within-population spatial genetic data from 62 replicate populations across Europe, North America, Asia and Africa, we investigated the contributions of vertical and horizontal transmission of the photobiont to the genetic structure of the epiphytic lichen Lobaria pulmonaria. Based on pairwise comparisons of multilocus genotypes defined separately for the mycobiont and for the photobiont, we inferred the transmission mode of the photobiont and the relative contribution of somatic mutation and recombination. After constraining the analysis of one symbiont to pairs of individuals with genetically identical symbiotic partners, we found that 77% of fungal and 70% of algal pairs were represented by clones. Thus, the predominant dispersal mode was by means of symbiotic vegetative propagules (vertical transmission), which dispersed fungal and algal clones co-dependently over a short distance, thus shaping the spatial genetic structure up to distances of 20m. Evidence for somatic mutation generating genetic diversity was found in both symbionts, accounting for 30% of pairwise comparisons in the alga and 15% in the fungus. While the alga did not show statistically significant evidence of recombination, recombination accounted for 7.7% of fungal pairs with identical algae. This implies that, even in a mostly vegetatively reproducing species, horizontal transmission plays a role in shaping the symbiotic association, as shown in many coral and other symbioses in nature.

Abstract: Summary Bacterial communities colonize the surfaces of lichens in a biofilm-like manner. The overall structure of the bacterial communities harboured by the lichens shows similarities, in particular the dominance of not yet cultured Alphaproteobacteria. Parameters causing variation in abundance, composition and spatial organization of the lichen-associated bacterial communities are so far poorly understood. As a first step, we used a microscopic approach to test the significance of both lichen-intrinsic and extrinsic environmental factors on the bacterial communities associated with 11 lichen samples, belonging to six species. Some of these species have thalli with a distinct age gradient. A statistically significant effect can be attributed to the age of the thallus parts, which is an intrinsic factor: growing parts of the lichens host bacterial communities that significantly differ from those of the ageing portions of the thalli. The substrate type (rock, tree, understory) and (at a lower extent) the exposition to the sun also affected the bacterial communities. Interestingly, the abundance of bacterial cells in the lichens was also influenced by the same structure-triggering factors. No effect on the composition with main bacterial groups was attributed to different lichen species, differentiated thallus parts or thallus growth type. Our results are important for the experimental designs in lichen-bacterial ecology.

Abstract: Due to climatic and orographic reasons, the occurrence of vascular plants in high alpine regions is limited. At locations that are not suitable for the establishment of higher plants because of exposure, substrate or other abiotic factors, cryptogams can be the dominant life forms. Mosses, lichens and algae particularly thrive on places such as bare rocks, permafrost soils or, exceptionally, even in melting snow and permanent ice. Since these lower plants are poikilohydric and lack complex morphological tissues like the cormophytes, unfavourable conditions (like drought) can be overcome by physiological inactivity, and structural damage is not the critical issue for these poikilohydric organisms. The vegetation period of cryptogams can be very short (from days to a few weeks per year), and growth and reproduction have to be adapted to limiting factors such as low temperatures, limited water-availability or irradiation stress.

Abstract: In lichen symbiosis, fungal and algal partners form close associations, often codispersed by vegetative propagules. Due to the particular interdependence, processes such as colonization, dispersal or genetic drift are expected to result in congruent patterns of genetic structure in the symbionts. To study the population structure of an obligate symbiotic system in Europe, we genotyped the fungal and algal symbionts of the epiphytic lichen Lobaria pulmonaria at eight and seven microsatellite loci, respectively, and analysed about 4300 L. pulmonaria thalli from 142 populations from the species’ European distribution range. Based on a centroid approach, which localizes centres of genetic differentiation with a high frequency of geographically restricted alleles, we identified the South Italy–Balkan region as the primary glacial refugial area of the lichen symbiosis. Procrustean rotation analysis and a distance congruence test between the fungal and algal population graphs indicated general concordance between the phylogeographies of the symbionts. The incongruent patterns found in areas of postglacial recolonization may show the presence of an additional refugial area for the fungal symbiont, and the impact that horizontal photobiont transmission and different mutation rates of the symbionts have on their genotypic associations at a continental scale.

Abstract: Nitrogen (N) deposition has increased globally over the last 150 years and further increases are predicted. Epiphytic lichens decline in abundance and diversity in areas with high N loads, and the ...

Abstract: The vegetation of the Antarctic tundra is dominated by mosses and lichens. Deschampsia antarctica, the Antarctic hairgrass, is one of two vascular plant species which grow along the west coast of the Antarctic Peninsula. However, little is known about its recruitment and interaction with non-vascular tundra plants. Although several authors propose that tolerance and/or competition should be the main forms of interaction between moss carpets and D. antarctica, no relevant studies exist so far. We investigated whether positive interactions are predominant at the Shetland Islands and the west coast of the Antarctic Peninsula and focussed on the role that moss carpets play in the recruitment of D. antarctica. Across the studied zone, D. antarctica showed a significant association with moss carpets, with higher frequencies as well as more and larger individuals than on bare ground. At one site, we conducted moss removal and seedlings transplant experiments to assess the relevance of the moss carpets for different life stages of hairgrass. All experimental individuals survived until the following summer whether the moss carpet was removed or not, but growth rate was significantly lower in tussocks with moss carpets removed. Likewise, tiller size was higher in plants growing in moss carpets than on bare ground. The detected positive interactions with mosses seem to be important for the expansion of D. antarctica, raising the question about their importance under future climate change scenarios.

Abstract: As part of a comprehensive study on lecideoid lichens in Antarctica, we investigated the photobiont diversity and abundance in 119 specimens of lecideoid lichens from 11 localities in the continental and maritime Antarctic. A phylogeny of these photobiont ITS sequences, including samples from arctic, alpine and temperate lowland regions, reveals the presence of five major Trebouxia clades in Antarctic lecideoid lichens. Two clades are formed by members of the T. jamesii and T. impressa aggregates but for all other clades no close match to any known Trebouxia species could be found in sequence databases. One genetically uniform and well-supported Trebouxia clade was found only in the climatically unique cold desert regions of the Antarctic (preliminarily called Trebouxia sp.URa1), where it is preferentially associated with the highly adapted Antarctic endemic lichen Lecidea cancriformis. Levels of genetic photobiont diversity differ slightly, but insignificantly among ecological regions of the Antarctic and do not decrease towards regions with more unfavourable ecological conditions. The genetic diversity of photobionts varies among mycobiont species. Most pairwise comparisons reveal that these differences are insignificant, probably due to the small sample size for most species. The Antarctic lichens studied here are predominantly not specific for a single photobiont species or lineage, except for Lecidella greenii and L. siplei. These two species are preferably associated with Trebouxia sp. URa2, although in the sampling areas of both species, a pool of several other photobionts is available. Lecidea cancriformis associates with the highest diversity of photobionts followed by L. andersonii.

Abstract: As defined in the broadest sense, algae are oxygen-generating, photosynthetic organisms other than embryophyte land plants and lichens (Cavalier-Smith, T. 2007. Evolution and relationships of algae: major branches of the tree of life. In Unravelling the algae: the past, present and future of algal systematics, eds. Brodie, J. and Lewis J, pp. 21–55. CRC Press. The Systematics Association Special Volume 75). They are an artificial and highly heterogeneous aggregation of organisms belonging to many different evolutionary lineages, and therefore highly diverse from a genetic point of view. This genetic diversity is reflected in the huge diversity exhibited by algae in terms of morphological, ultrastructural, ecological, biochemical, and physiological traits.

Abstract: The McMurdo Dry Valleys of Antarctica present extreme environmental challenges. Life is restric- ted to patchy occurrence of lichens, mosses and inverte- brates, plus soil microbial communities. Fungi have been described in lichen symbioses but relatively little is known about the occurrence of free-living soil fungi in the Dry Valleys. A challenge in estimating fungal species richness has been the extent to which estimates based on either cultivation or environmental DNA reflect the active assemblage in cold-arid soils. Here, we describe analysis for inland Dry Valleys soil of environmental DNA and RNA (cDNA) to infer total and putative metabolically active assemblages, respectively, plus cultivation approa- ches using a variety of laboratory growth conditions. Environmental sequences indicated a highly restricted assemblage of just seven phylotypes that affiliated phylo- genetically within two known genera, Helicodendron and Zalerion, plus previously unidentified fungal phylotypes. None of the commonly encountered molds and mitosporic genera recorded from maritime Antarctic locations were encountered. A striking difference was observed in the frequency of recovery for phylotypes between libraries. This suggests that both species richness and beta diversity estimates based on DNA libraries have the potential to misinform putatively active assemblages. Cultivation yielded a cold-tolerant Zalerion strain that affiliated with DNA and RNA library clones, and a psychrotrophic yeast (Debaryomyces hansenii), which was not detected using either culture-independent approach.

Abstract: Elevated temperatures and diminished precipitation amounts accompanying climate warming in arid ecosystems are expected to have adverse effects on the photosynthesis of lichen species sensitive to elevated temperature and/or water limitation. This premise was tested by artificially elevating temperatures (increase 2.1–3.8°C) and reducing the amounts of fog and dew precipitation (decrease 30.1–31.9%), in an approximation of future climate warming scenarios, using transparent hexagonal open-top warming chambers placed around natural populations of four lichen species (Xanthoparmelia austroafricana, X. hyporhytida , Xanthoparmelia. sp., Xanthomaculina hottentotta) at a dry inland site and two lichen species (Teloschistes capensis and Ramalina sp.) at a humid coastal site in the arid South African Succulent Karoo Biome. Effective photosynthetic quantum yields (\( \Updelta F/F_{\text{m}}^{\prime } \)) were measured hourly throughout the day at monthly intervals in pre-hydrated lichens present in the open-top warming chambers and in controls which comprised demarcated plots of equivalent open-top warming chamber dimensions constructed from 5-cm-diameter mesh steel fencing. The cumulative effects of the elevated temperatures and diminished precipitation amounts in the open-top warming chambers resulted in significant decreases in lichen \( \Updelta F/F_{\text{m}}^{\prime } \). The decreases were more pronounced in lichens from the dry inland site (decline 34.1–46.1%) than in those from the humid coastal site (decline 11.3–13.7%), most frequent and prominent in lichens at both sites during the dry summer season, and generally of greatest magnitude at or after the solar noon in all seasons. Based on these results, we conclude that climate warming interacting with reduced precipitation will negatively affect carbon balances in endemic lichens by increasing desiccation damage and reducing photosynthetic activity time, leading to increased incidences of mortality.

Abstract: The production and regulation of secondary metabolites in non-lichenized fungi, mainly ascomycetes, has been reviewed by a number of authors with an emerging understanding of the biosynthesis and the pathways involved in regulation (Keller et al, 2005; Yu & Keller 2005; and others) However, lichenized fungi make up almost half of all known ascomycetes (Kirk et al, 2001) and are known to produce over 800 secondary metabolites, most of which are unique to lichenized fungi Many of these compounds have bioactive properties (Huneck, 1999) and some studies have shown or suggested that secondary metabolite production is influenced by changes in culture conditions, which might be regarded as environmental changes Intense investigation of the changes in production of these unique bioactive secondary metabolites from lichen fungi have been hampered by problems associated with isolating and growing cultures of lichen fungi Lichens have been studied for more than two centuries as morphological entities but experimental lichenology has remained a nearly unexplored scientific field for many decades because of the slow growing nature of lichens Thomas (1939 in Stocker-Worgotter, 2001) reported the first successful resynthesis of Cladonia pyxidata Since the 1970’s, one major goal of experimental lichenology has been the improvement and optimization of culture conditions of lichen fungi Culture techniques for lichen fungi have improved in recent years allowing for further research on these challenging organisms Therefore, with greater access to cultures of lichen symbionts and progression of knowledge of non-lichenized fungi, studies are just beginning to accumulate on genes involved in production of secondary metabolites from lichen fungi; and the effects of the environment on the expression of these genes by observations in ecological studies, and through experimentation by manipulating culture conditions Fungal secondary metabolism is covered by extensive body of literature (see Bennett & Ciegler, 1983) Secondary metabolism is not required for survival and its products are dispensible whereas primary metabolism is essential for survival with anabolic and catabolic activities to maintain life Secondary metabolites are chemically diverse but are produced from a few key intermediates of primary metabolism, and are generally categorized by the intermediates from which they are produced Bennett and Ciegler (1983) summarize six categories of secondary metabolites derived from different primary intermediates Although fungal secondary metabolites are extensive, they are generally

Abstract: Five new monophyletic groups of xanthorioid lichens have been identified by phylogenetic analyses using nuclear ITS and 18S and mitochondrial 12 mt and 23 mt sequences. As a consequence, the following new genera are described: Gallowayella, Jesmurraya, Honeggeria, Massjukiella and Martinjahnsia. New combinations for 26 species are proposed: Gallowayella aphrodites, G. borealis, G. concinna, G. coppinsii, G. fulva, G. hasseana, G. hermonii, G. montana, G. galericulata, G. gallowayi, G. oregana, G. poeltii, G. sogdiana, G. tibellii, G. weberi, G. wetmorei, Honeggeria rosmarieae, Jesmurraya novozelandica, Massjukiella alaskana, M. candelaria, M. kaernefeltii, M. nowakii, M. polycarpa, M. ucrainica, M. ucrainica ssp. marginata and Martinjahnsia resendei.

Abstract: Overstory Populus has been shown to facilitate establishment by cyano- and cephalolichens on conifer branches within its dripzone; though the mechanisms behind this “dripzone effect” have yet to be...

Abstract: Colonization studies may function as natural experiments and have the potential of addressing important questions about community assembly. We studied colonization for a guild of epiphytic lichens in a former treeless heathland area of 170 km2 in southwest Norway. We investigated if epiphytic lichen species richness and composition on aspen (Populus tremula) trees corresponded to a random draw of lichen individuals from the regional species pool. We compared lichen communities of isolated young (55-120 yr) and old (140-200 yr) forest patches in the heathland area to those of aspen forest in an adjacent reference area that has been forested for a long time. All thalli (lichen bodies) of 32 selected lichen species on trunks of aspen were recorded in 35 aspen sites. When data for each site category (young, old, and reference) were pooled, we found the species richness by rarefaction to be similar for reference sites and old sites, but significantly lower for young sites. The depauperated species richness of young sites was accompanied by a skew in species composition and absence of several species that were common in the reference sites. In contrast, genetic variation screened with neutral microsatellite markers in the lichen species Lobaria pulmonaria showed no significant differences between site categories. Our null hypothesis of a neutral species assembly in young sites corresponding to a random draw from the regional species pool was rejected, whereas an alternative hypothesis based on differences in colonization capacity among species was supported. The results indicate that for the habitat configuration in the heathland area (isolated patches constituting < 0.4% of the area) lichen communities may need a colonization time of 100-150 yr for species richness to level off, but given enough time, isolation will not affect species richness. We suggest that this contradiction to expectations from classical island equilibrium theory results from low extinction rates.

Abstract: A biomonitoring study, using transplanted lichens Flavoparmelia caperata, was conducted to assess the indoor air quality in primary schools in urban (Lisbon) and rural (Ponte de Sor) Portuguese sites. The lichens exposure period occurred between April and June 2010 and two types of environments of the primary schools were studied: classrooms and outdoor/courtyard. Afterwards, the lichen samples were processed and analyzed by instru- mental neutron activation analysis (INAA) to assess a total of 20 chemical elements. Accumulated elements in the exposed lichens were assessed and enrichment factors (EF) were determined. Indoor and outdoor biomonitoring results were compared to evaluate how biomonitors (as lichens) react at indoor environments and to assess the type of pollutants that are prevalent in those environments.