Abstract: Lichens are symbiotic associations between fungi and a photosynthetic alga and/or cyanobacteria. Lichenized fungi have been found to produce a wide array of secondary metabolites, most of which are unique to the lichenized condition. These secondary metabolites have shown an impressive range of biological activities including antibiotics, antifungal, anti-HIV, anticancer, anti-protozoan, etc. This review focuses primarily on the antibiotic and anticancer properties of lichen secondary chemicals. We have reviewed various publications related to antibiotic and anticancer drug therapies emphasizing results about specific lichens and/or lichen compounds, which microbes or cancer cells were involved and the main findings of each study. We found that crude lichen extracts and various isolated lichen compounds often demonstrate significant inhibitory activity against various pathogenic bacteria and cancer cell lines at very low concentrations. There were no studies examining the specific mechanism of action against pathogenic bacteria; however, we did find a limited number of studies where the mechanism of action against cancer cell lines had been explored. The molecular mechanism of cell death by lichen compounds includes cell cycle arrest, apoptosis, necrosis, and inhibition of angiogenesis. Although lichens are a reservoir for various biologically active compounds, only a limited number have been tested for their biological significance. There is clearly an urgent need for expanding research in this area of study, including in depth studies of those compounds which have shown promising results as well as a strong focus on identifying specific mechanisms of action and extensive clinical trials using the most promising lichen based drug therapies followed by large scale production of the best of those compounds.

Abstract: The effect of management related factors on species richness of epiphytic bryophytes and lichens was studied in managed deciduous-coniferous mixed forests in Western-Hungary. At the stand level, the potential explanatory variables were tree species composition, stand structure, microclimate and light conditions, landscape and historical variables; while at tree level host tree species, tree size and light were studied. Species richness of the two epiphyte groups was positively correlated. Both for lichen and bryophyte plot level richness, the composition and diversity of tree species and the abundance of shrub layer were the most influential positive factors. Besides, for bryophytes the presence of large trees, while for lichens amount and heterogeneity of light were important. Tree level richness was mainly determined by host tree species for both groups. For bryophytes oaks, while for lichens oaks and hornbeam turned out the most favourable hosts. Tree size generally increased tree level species richness, except on pine for bryophytes and on hornbeam for lichens. The key variables for epiphytic diversity of the region were directly influenced by recent forest management; historical and landscape variables were not influential. Forest management oriented to the conservation of epiphytes should focus on: (i) the maintenance of tree species diversity in mixed stands; (ii) increment the proportion of deciduous trees (mainly oaks); (iii) conserving large trees within the stands; (iv) providing the presence of shrub and regeneration layer; (v) creating heterogeneous light conditions. For these purposes tree selection and selective cutting management seem more appropriate than shelterwood system.

Abstract: Lichens and bryophytes are abundant globally and they may even form the dominant autotrophs in (sub)polar ecosystems, in deserts and at high altitudes. Moreover, they can be found in large amounts as epiphytes in old-growth forests. Here, we present the first process-based model which estimates the net carbon uptake by these organisms at the global scale, thus assessing their significance for biogeochemical cycles. The model uses gridded climate data and key properties of the habitat (e.g. disturbance intervals) to predict processes which control net carbon uptake, namely photosynthesis, respiration, water uptake and evaporation. It relies on equations used in many dynamical vegetation models, which are combined with concepts specific to lichens and bryophytes, such as poikilohydry or the effect of water content on CO 2 diffusivity. To incorporate the great functional variation of lichens and bryophytes at the global scale, the model parameters are characterised by broad ranges of possible values instead of a single, globally uniform value. The predicted terrestrial net uptake of 0.34 to 3.3 Gt yr −1 of carbon and global patterns of productivity are in accordance with empirically-derived estimates. Considering that the assimilated carbon can be invested in processes such as weathering or nitrogen fixation, lichens and bryophytes may play a significant role in biogeochemical cycles.

Abstract: Summary Lichenization is assumed to be a very ancient mode of fungal nutrition, but fossil records are rare. Here we describe two fragments of exceptionally preserved, probably charred, lichen thalli with internal stratification. Cyanolichenomycites devonicus has a cyanobacterial and Chlorolichenomycites salopensis a unicellular, presumably green algal photobiont. Fruiting bodies are missing. Cyanolichenomycites devonicus forms asexual spores in a pycnidium. All specimens were examined with scanning electron microscopy techniques. The fossils were extracted by maceration. Extant lichens and free-living cyanobacteria were either experimentally charcoalified for comparison or conventionally prepared. Based on their septate hyphal structure, both specimens are tentatively interpreted as representatives of the Pezizomycotina (Ascomycota). Their presence in 415 million yr (Myr) old rocks from the Welsh Borderland predates existing Late Cretaceous records of pycnidial conidiomata by some 325 Myr and Triassic records of lichens with broadly similar organization by some 195 Myr. These fossils represent the oldest known record of lichens with symbionts and anatomy as typically found in morphologically advanced taxa today. The latter does not apply to Winfrenatia reticulata, the enigmatic crustose lichen fossil from the Lower Devonian, nor to presumed lichen-like organisms such as the Cambrian Farghera robusta or to the Lower Devonian Spongiophyton minutissimum.

Abstract: Epiphytic bryophytes and lichens are an important component of the endangered forest biota in temperate forests, their diversity and composition patterns being regulated by tree, stand and landscape scale factors. The aim of this study is to improve ecological understanding of such factors in managed coniferous–deciduous mixed forests of Hungary in the context of forest management. In particular, this study investigate the effect of tree species composition, stand structure (tree size distribution, shrub layer and dead wood), microclimate (light, temperature and air humidity), landscape and historical factors on the stand level and tree level composition of epiphytic bryophytes and lichens. The relationships were explored by multivariate methods (redundancy analysis, canonical correspondence analysis and variation partitioning) and indicator species analysis. Tree species is among the most important driver of species composition in both organism groups. For bryophytes, the continuity of forest microclimate and the presence of shrub layer are also important, while lichen assemblages are influenced by light availability. Landscape and historical variables were less influential than stand scale factors. On the basis of our results, the main strategy of management focusing on epiphyte diversity conservation should include: (1) the maintenance of tree species diversity in mixed stands; (2) increasing the proportion of deciduous trees (mainly oaks and hornbeam); (3) the maintenance of large trees within the stands; (4) the presence of shrub and regeneration layer; (5) the creation of heterogeneous light conditions.

Abstract: Many boreal and polar lichens occupy bipolar distributional ranges that frequently extend into high mountains at lower latitudes. Although such disjunctions are more common among lichens than in other groups of organisms, the geographic origin of bipolar lichen taxa, and the way and time frame in which they colonized their ranges have not been studied in detail. We used the predominantly vegetative, widespread lichen Cetraria aculeata as a model species. We surveyed the origin and history of its bipolar pattern using population genetics, phylogenetic and genealogical reconstruction methods. Cetraria aculeata originated in the Northern Hemisphere and dispersed southwards during the Pleistocene. The genetic signal suggests a Pleistocene dispersive burst in which a population size expansion concurred with the acquisition of a South-American range that culminated in the colonization of the Antarctic.

Abstract: Biological soil crusts of arid and semiarid regions are one of the least explored habitats with respect to the diversity of their fungal communities and the Arabian deserts, in particular, remains mycologically poorly investigated. Here, we investigate the diversity of free-living and lichen-forming fungal communities associated with crusts at two locations in Oman, using intensive cultivation and pyrosequencing, and their role in improving soil stability and hydrology. A total of 226 fungal isolates were recovered and phylogenetic analysis placed 98% of the isolates within the Ascomycota phylum, most of which belonged to Dothideomycetes class and Pleosporales order. The isolates were phylogenetically affiliated to 101 different species within 44 different genera with >55% of the total isolates belonging to Alternaria, Aspergillus, Cladosporium, Cochliobolus, Fusarium, Myrothecium, Phoma and Ulocladium. Using pyrosequencing, a total of 26,998 sequence reads were obtained with Ascomycota, Basidiomycota and Chytridiomycota encompassing >96% of the total sequences. In cyanobacterial crusts, between 67.2 and 70.6% of the total fungal sequences belonged to the classes Dothideomycetes and Eurotiomycetes with the dominance of yeast-like fungi of the genera Sarcinomyces and Aureobasidium. On the other hand, the sequences obtained from the lichen crusts mainly belonged to the classes Lichinomycetes, Lecanoromycetes and Eurotiomycetes. Among the identified lichens were Placidium lacinulatum, Psora decipiens, Peccania fontqueriana, Stromatella bermudana, Verrucaria chiloensis, Pecania arizonica, Lempholemma polyanthes and Lichinella cribellifera. Although detected fungi confirmed earlier trends in fungal diversity in other deserts, quite a number of isolates and sequences representing novel taxa were recovered. The presence of lichen in crusts improved their resistance to erosion and increased their water holding capacity. We conclude that desert crusts of Oman harbor a large diversity of fungal communities that varies with crust type and desert ecoregions, and the presence of lichens in these crusts improves soil characteristics.

Abstract: Sixteen epilithic lichen samples (13 species), collected from seven locations in Northern and Southern Victoria Land in Antarctica, were investigated for the presence of black fungi. Thirteen fungal strains isolated were studied by both morphological and molecular methods. Nuclear ribosomal 18S gene sequences were used together with the most similar published and unpublished sequences of fungi from other sources, to reconstruct an ML tree. Most of the studied fungi could be grouped together with described or still unnamed rock-inhabiting species in lichen dominated Antarctic cryptoendolithic communities. At the edge of life, epilithic lichens withdraw inside the airspaces of rocks to find conditions still compatible with life; this study provides evidence, for the first time, that the same microbes associated to epilithic thalli also have the same fate and chose endolithic life. These results support the concept of lichens being complex symbiotic systems, which offer attractive and sheltered habitats for other microbes.

Abstract: I was looking forward to reviewing this book because I hoped it would be a complete treatment of the lichens of the Great Smoky Mountains National Park (GSMNP), but sadly I found it fell short of that goal. The title is a misnomer as the book is really a preliminary checklist and a vehicle to publish some new taxa. Readers expecting to find a thorough presentation of a flora will be disappointed. The table of contents lists a foreword by Keith Langdon of the park staff, an author preface, acknowledgements, a summary of taxonomic changes, introduction, methodological summary, discussion, results (eight sections), literature cited, and an index of scientific names. At over 500,000 acres, GSMNP is the largest national park in the eastern U. S. and contains the highest vascular plant diversity. It is reasonable to expect the lichen diversity to be high as well. Given the area of the park and its’ rugged terrain one would expect more than ‘‘...three years of limited field work...’’ would be needed to inventory the park, yet that is what the authors admit to doing. In addition, no dates are given for collecting periods (other than in the selected specimen lists), and there is no list or map of the localities. There is a map at the very end of the book, probably added as an afterthought, with no collecting localities marked. It is customary to include a collecting locality map near the front as part of the methods. There is no presentation of the substrates in the park, which are critical to understanding what lichens to expect. For example, from the species section dolomite is mentioned a few times, yet there are none of the typical limestone loving species listed (e.g., Lichinella sp., Lecanora muralis, Xanthoria elegans). The coverage of the park is not presented so the reader has no idea how well-studied the park is, even though there is a section on how well-known it is (and this question is left unanswered). The authors admit in the species list that they have verified 84% of the 804 taxa they report, saying they have been unable to study the high elevation areas of the park or collect taxonomic groups that are not easy to find, giving the Caliciales as the example. The question of how many species are in the park remains unclear: is it 672 or 804? Identification methods are not described at all. Knowing two of the authors, I don’t doubt their identifications but what about other readers in the future? How will they know how the identifications were made? The species list is strangely limited to the authors’ work and the inclusion of some previous lists, which is good for a start, but I did a search in the Consortium of North American Lichen Herbaria (CNALH) database which produced a list of 251 taxa in the park, including 28 that are not in the book. Leaving these out seems short-sighted. Other species I expected to find but didn’t include Julella fallaciosa (collected by the first author north and south of the park), Teloschistes chrysophthalmus (expected in the coves and lowlands), and other species of Acarospora on acidic rocks, for examples. An example of a premature declaration is where the authors claim they have found the first confirmed

Abstract: Recently deglaciated areas are ideal environments to study soil formation and primary microbial succession where phototrophic microorganisms may play a role as primary producers. The aim of our study was to investigate the cyanobacterial and green algal community composition in three different successional stages of the Damma glacier forefield in the Swiss Alps using 16S rDNA and ITS rDNA clone libraries. Cyanobacterial target sequences varied along the glacier forefield, with the highest cyanobacterial 16S rRNA gene copies found in sparsely vegetated soils. Sequence analysis revealed that the phototrophic communities were distinct in each of the three soil environments. The majority of the cyanobacterial sequences retrieved from barren soils were related to the Oscillatoriales. The diversity in sparsely vegetated soils was low, and sequences closely related to Nostoc sp. dominated. The majority of the algal phylotypes are related to members of the Trebouxiophyceae known to live as symbiotic partners in lichens. We conclude that the community composition appears to shift markedly along the chronosequence, indicating that each soil environment selects for its phototrophic community. When cyanobacteria occur together with eukaryotic microalgae, they form a rich source of organic matter and may be important contributors of carbon in nutrient-deficient deglaciated soils.

Abstract: Many black meristematic fungi persist on rock surfaces—hostile and exposed habitats where high doses of radiation and periods of desiccation alternate with rain and temperature extremes. To cope with these extremes, rock-inhabiting black fungi show phenotypic plasticity and produce melanin as cell wall pigments. The rather slow growth rate seems to be an additional prerequisite to oligotrophic conditions. At least some of these fungi can undergo facultative, lichen-like associations with photoautotrophs. Certain genera presenting different lifestyles are phylogenetic related among the superclass Dothideomyceta. In this paper, we focus on the genus Lichenothelia, which includes border-line lichens, that is, associations of melanised fungi with algae without forming proper lichen thalli. We provide a first phylogenetic hypothesis to show that Lichenothelia belongs to the superclass Dothideomyceta. Further, culture experiments revealed the presence of co-occurring fungi in Lichenothelia thalli. These fungi are related to plant pathogenic fungi (Mycosphaerellaceae) and to other rock-inhabiting lineages (Teratosphaeriaceae). The Lichenothelia thallus-forming fungi represent therefore consortia of different black fungal strains. Our results suggest a common link between rock-inhabiting meristematic and lichen-forming lifestyles of ascomycetous fungi.

Abstract: We present ataxonomic revision of the ‘manna lichens’ based on morphological, chemical, ecological and molecular data. A large number of herbarium specimens and fresh collections were examined. Phylogenetic analyses were performed using nuclear ribosomal (nrITS, nrLSU) and mitochondrial small subunit (mtSSU) sequences. Some notable phenotypic characters were plotted on the phylogenetic tree, and the analysis reveals that some of these characters are useful for genus and species level distinction of certain ‘manna lichens.’ Phylogeny of the Megasporaceae was revised using a combined data set of nrLSU and mtSSU and performing parsimony and Bayesian analyses. Five genera (Aspicilia, Circinaria, Lobothallia, Megaspora and Sagedia) are recognized. Further, the relationships of five presumably closely related genera of ‘manna lichens’, namely Agrestia (vagrant), Aspicilia (crustose) Circinaria (crustose), Chlorangium (vagrant) and Sphaerothallia (vagrant) with different growth forms were analysed. The analyses revealed that ‘manna lichens’ do not form a monophyletic group but occur in different clades within the genus Circinaria. The genera Agrestia, Chlorangium and Sphaerothallia are assigned as new synonyms under the genus Circinaria and no vagrant or erratic species remain in the genus Aspicilia. The analyses also show that five new erratic, vagrant and crustose species can be recognized. In this study two ‘manna lichens’, viz. Circinaria rostamii sp. nov. (Azerbaijan, Iran and Turkey), and Circinaria gyrosa sp. nov. (Armenia, Azerbaijan, Iran, Turkey, Turkmenistan and Spain) are described as new to science. Three potentially new species with crustose and erratic forms need additional study. Aspicilia fruticolosofoliacea is reduced to synonymy under C. alpicola, and a lectotype is designated for C. aspera. Thirteen new combinations in Circinaria are presented. The phenomenon of vagrancy is briefly discussed, and the biogeography of the ‘manna lichens’ is outlined. Illustrations, distribution maps, and an identification key to the species are provided.

Abstract: The vertical gradient of the community structure of epiphytic lichens in forest canopies was studied in southern Estonian coniferous forests. All lichen species on 15 Norway spruces and 15 Scots pines were recorded; age of trees ≥100 years. Species were sampled in height ranges on entire trees from the bottom to the top. The number of lichen species on pine trunks decreased with height from the ground, whereas the number of species on branches increased. There was no significant vertical change in the number of lichen species on spruce trunks; number of species on spruce branches was highest in mid canopy. The mean number of lichen species on a tree was 41 in spruces and 34 in pines; the mean number of species on the first 2 m was 14 in spruces and 12 in pines. According to the results, about two thirds of a tree's lichen species remain unrecorded if only the first 2 m near the ground are surveyed. Many lichen species were found only higher than 2 m, including several common species (e.g. Buellia griseovirens, Lecanora pulicaris, and Melanohalea exasperatula), but also some locally rather rare (Fellhanera subtilis, Micarea nitschkeana, Rinodina efflorescens) or red-listed ones (Evernia mesomorpha, Usnea barbata, U. fulvoreagens, U. substerilis, U. wasmuthii) and one new species for Estonia, Lecanora farinaria. There were also some species, such as Chaenotheca stemonea, Cladonia cenotea and C. norvegica, that were restricted to the lowest 2 m. The results indicate that canopy lichens form a significant part of lichen diversity in coniferous forests, and could add valuable information when estimating forest lichen diversity for conservation and other purposes.

Abstract: This study investigated the influence of angular exposure and distance from vehicular traffic on the diversity of epiphytic lichens and the bioaccumulation of traffic-related elements in a town of central Italy. An Index of Lichen Diversity (ILD) was calculated on the street-facing and the opposite side of road-lining trees and in a urban park 250 m away, and the content of selected trace elements (Al, Ba, Ce, Cd, Cr, Cu, Fe, Mn, Ni, Pb, Sb, V, and Zn) was determined in samples of the lichen Punctelia borreri (Sm.) Krog growing on tree bark, both on the exposed and opposite sides. ILD increases with distance from traffic emissions. However, at the site with vehicle traffic, non-nitrophilous lichens decreased while nitrophilous ones increased. The concentration of the traffic-related elements Ba, Cr, Cu, Mn, Sb, and Zn accumulated in thalli of P. borreri was higher on roadside trees than in trees from the urban park. ILD was not affected by the angular exposure to the road and the bioaccumulation of traffic-related elements was similar in lichens from the side of the bole exposed to traffic emissions and particulate resuspension and from the opposite side. The angular exposure in respect to the traffic source does not influence trace element accumulation. These results are important when using lichens for biomonitoring purposes, both for planning future studies and for the reliability of the interpretation of past surveys that do not report information about the angular exposure of the collected lichen material.

Abstract: An annotated checklist comprised of 458 taxa of orchids known from Nepal is provided, including 104 genera, 437 species, 16 varieties, 3 subspecies and 2 forma and 18 endemic species. In Nepal, orchid species are distributed from 60–5200 m a.s.l. In the checklist, notes on altitudinal ranges, habit, habitat, global distribution, phenology, etc. are presented. Nepal, a small Himalayan country, lies between 26°12′ to 30°27′N and 80°04′E to 88°12′E covering an area of 147 181 km 2 . The diverse physiography, climate, edaphic factors, etc. within an altitude range of 60 – 8848 m a.s.l. has favored vegetation within six bioclimatic zones from tropical luxuriant forests in the south to the alpine scrub or barren high mountain in the north (Chaudhary 1998). The tropical zone is dominated by sal (Shorea robusta), tropical deciduous riverine and tropical evergreen forests. The sal forest is associated with Adina cardifolia, Albizia procera, Butea monosperma, Acacia catechu, Bombax ceiba, Dalbergia sisoo, Syzygium cumini, Terminalia bellirica, T. chebula, Zizyphus mauritiana, etc. In the subtropical zone, Schima–Castanopsis, pine and alder forest types are dominant. The temperate zone is represented by lower temperate mixed broad-leaved, temperate mixed evergreen and upper temperate mixed broad-leaved forests. The subalpine zone is characterized by silver fir (Abies spectabilis) and birch– Rhododendron forest. This zone also includes a pure forest belt of Betula utilis, in association with several species of Rhododendron, Junipererus, etc. In this region, tree trunks and branches are festooned with parasitic orchids, filaments of mosses and various lichens. The alpine zone covers areas above 4100 m a.s.l. and is associated with Juniperus, Rhododendron, Caragana–Lonicera and alpine meadows. In the nival zone, above 5000 m a.s.l., some mosses and lichens are found up to 7400 m a.s.l. (Baniya et al. 2010) and they are known as the ‘Arctic or cold deserts.’

Abstract: The increasing demand for forest-derived bio-fuel may decrease the amount of dead wood and hence also the amount of available substrate for saproxylic ( = dead-wood dependent) organisms. Cut stumps constitute a large portion of dead wood in managed boreal forests. The lichen flora of such stumps has received little interest. Therefore, we investigated which lichens that occur on stumps in young (4–19 years), managed forests and analyzed how species richness and occurrence of individual species were related to stump and stand characteristics. We performed lichen inventories of 576 Norway spruce stumps in 48 forest stands in two study areas in Central Sweden, recording in total 77 lichen species. Of these, 14 were obligately lignicolous, while the remaining were generalists that also grow on bark, soil or rocks. We tested the effect of characteristics reflecting successional stage, microclimate, substrate patch size, and the species pool in the surrounding area on (1) total lichen species richness, (2) species richness of obligately lignicolous lichens and (3) the occurrence of four obligately lignicolous lichen species. The most important variables were stump age, with more species on old stumps, and study area, with similar total species richness but differences in occupancy for individual species. Responses for total lichen species richness and species richness of obligately lignicolous lichens were overall similar, indicating similar ecological requirements of these two groups. Our results indicate that species richness measurements serve as poor proxies for the responses of individual, obligately lignicolous lichen species.

Abstract: Premise of the study Colonists of even the most inhospitable environments, lichens are present in all terrestrial ecosystems. Because of their ecological versatility and ubiquity, they have been considered excellent candidates for early colonizers of terrestrial environments. Despite such predictions, good preservation potential, and the extant diversity of lichenized fungi, the fossil record of lichen associations is sparse. Unequivocal lichen fossils are rare due, in part, to difficulties in ascertaining the presence of both symbionts and in characterizing their interactions. This study describes an exceptionally well-preserved heteromerous lichen from the Lower Cretaceous of Vancouver Island. Methods The fossil occurs in a marine carbonate concretion collected from the Apple Bay locality on Vancouver Island, British Columbia, and was prepared for light microscopy and SEM using the cellulose acetate peel technique. Key results The lichen, Honeggeriella complexa gen. et sp. nov., is formed by an ascomycete mycobiont and a chlorophyte photobiont, and exhibits heteromerous thallus organization. This is paired with a mycobiont-photobiont interface characterized by intracellular haustoria, previously not documented in the fossil record. Conclusions Honeggeriella adds a lichen component to one of the richest and best characterized Early Cretaceous floras and provides a significant addition to the sparse fossil record of lichens. As a heteromerous chlorolichen, it bridges the >350 million-year gap between previously documented Early Devonian and Eocene occurrences.