Bacterial diversity across individual lichens

Symbioses are unique habitats for bacteria. We surveyed the spatial diversity of bacterial communities across multiple individuals of closely related lichens using terminal restriction fragment length polymorphism (T-RFLP) and pyrosequencing. Centers of lichens house richer, more consistent assemblages than species-poor and compositionally disparate lichen edges, suggesting that ecological succession plays a role in structuring these communities.     

The weight of the crust: Biomass of crustose lichens in tropical dry forest represents more than half of foliar biomass

In recent years, our ecological knowledge of tropical dry forests has increased dramatically. However, the functional contributions of whole ecosystem components, such as lichens, remain mostly unknown. In these forests, the abundance of epiphyte crustose lichens is responsible for the characteristic white bark on most woody plants, conspicuous during the dry season, but the amount of resources that the lichen component represents remains unexplored. We estimated lichen biomass in a Mexican tropical dry forest using the bark area of trees, the dry mass of lichens per unit area and the percentage of bark covered by lichens, together with previously known tree densities. The lowest 2.5 m of the forests main trunks contained 188 kg/ha of lichen biomass, with lichens covering 85% of the available bark for trees <12 cm DBH and 38% for trees >12 cm. Total epiphytic lichen biomass was 1.34–1.99 Mg/ha. Lichen biomass represented 61% of the foliar biomass in the forest. To our knowledge, this is the first time that a lichen biomass estimate is provided for an ecosystem in which crustose lichens are the dominant lichen growth form. Crustose lichens are typically considered to contribute little to the total lichen biomass and to be difficult to include in ecological analyses. The high lichen biomass in this ecosystem implies a significant ecological role which so far is unexplored. We suggest the crustose lichen component should not be underestimated a priori in ecological studies, especially in ecosystems with abundant lichen cover.     

Spatial patterns of photobiont diversity in some Nostoc-containing lichens

Patterns of photobiont diversity were examined in some Nostoc -containing lichens using the nucleotide sequence of the cyanobacterial tRNA^Leu (UAA) intron. Lichen specimens collected in northwestern USA were analysed and the sequence data were compared with tRNA^Leu(UAA) intron sequences previously obtained from lichens in northern Europe. Generally, it is the species identity of a lichen rather than the geographical origin of the specimen that determines the identity of the cyanobiont. Identical intron sequences were found in Peltigera membranacea specimens collected in Oregon (USA) and in Sweden, and very similar sequences were also found in Nephroma resupinatum thalli collected in Oregon and Finland. Furthermore, in mixed assemblages where two Peltigera species grew in physical contact with each other, the different lichen species housed different photobiont strains. There is however not a one-to-one relation between mycobiont and photobiont as some intron sequences were found in more than one lichen species, and different intron sequences were found in different samples of some lichen taxa. Peltigera venosa exhibited a higher level of photobiont diversity than any other lichen species studied, and several intron sequences could for the first time be obtained from a single thallus. It is not clear whether this is evidence of lower cyanobiont specificity, or reflects an ability to exhibit different degrees of lichenization with different Nostoc strains. In one specimen of P. venosa , which contained bipartite cyanosymbiodemes and tripartite, cephalodiate thalli, both thallus types contained the same intron sequence.     

肉疣衣属中国新记录种——乌普萨拉肉疣衣

采用形态解剖、化学、生态类型及地理成分分析等传统分类方法对新疆天山南部巴音布鲁克山区地衣外部形态和内部构造、地衣附属结构、地衣繁殖器官进行观察。利用显色反应法(CT)、微量化学结晶法(MCT)、薄层层析法(TLC)等生物化学方法,鉴定地衣次生代谢产物,发现了1个中国新记录种:乌普萨拉肉疣衣[Ochrolechia upsaliensis(L.)A.Massal]。文中描述了该新记录种地衣的形态解剖、化学特征和生境,并提供了地衣体、子囊及子囊孢子的彩色图片。     

Spatial structure, rock type, and local environmental conditions drive moss and lichen distribution on calcareous boulders

Although mosses and lichens are a relevant component of the biota of rock habitats targeted for biodiversity conservation in Europe, the ecological factors driving their distribution are still poorly known. In this work, we examined the epilithic moss and lichen assemblages colonizing boulders of different types of calcareous rocks co-occurring in the same area in the Italian Alps. The goals were: (1) to evaluate if and to what extent different calcareous rocks host different assemblages; (2) to identify species associated to each rock type; (3) to quantify the relative importance of rock type, local environmental factors, and habitat spatial structure in explaining species distribution. Our results demonstrated that different calcareous rocks host different moss and lichen assemblages with some typical species, indicating that each rock type contributes to the total diversity of both mosses and lichens. Local environmental conditions influenced mosses and not lichens whose distribution is mainly associated to rock type. The patterns of both organism groups were also significantly related to habitat spatial structure, species assemblages tending to have a patchy distribution, which may reflect dispersal dynamics. Our results have implications for conservation: (1) each rock type may play a relevant role in maintaining the overall diversity contributing with unique assemblages and typical species; (2) the patchy distribution of both moss and lichen assemblages should warn from considering rock patches as a monotonous repetition of the same habitat across space.     

ITS1 metabarcoding highlights low specificity of lichen mycobiomes at a local scale

As self‐supporting and long‐living symbiotic structures, lichens provide a habitat for many other organisms beside the traditionally considered lichen symbionts—the myco‐ and the photobionts. The lichen‐inhabiting fungi either develop diagnostic phenotypes or occur asymptomatically. Because the degree of specificity towards the lichen host is poorly known, we studied the diversity of these fungi among neighbouring lichens on rocks in an alpine habitat. Using a sequencing metabarcoding approach, we show that lichen mycobiomes clearly reflect the overlap of multiple ecological sets of taxa, which differ in their trophic association with lichen thalli. The lack of specificity to the lichen mycobiome is further supported by the lack of community structure observed using clustering and ordination methods. The communities encountered across samples largely result from the subsampling of a shared species pool, in which we identify three major ecological components: (i) a generalist environmental pool, (ii) a lichenicolous/endolichenic pool and (iii) a pool of transient species. These taxa majorly belong to the fungal classes Dothideomycetes, Eurotiomycetes and Tremellomycetes with close relatives in adjacent ecological niches. We found no significant evidence that the phenotypically recognized lichenicolous fungi influence the occurrence of the other asymptomatic fungi in the host thalli. We claim that lichens work as suboptimal habitats or as a complex spore and mycelium bank, which modulate and allow the regeneration of local fungal communities. By performing an approach that minimizes ambiguities in the taxonomic assignments of fungi, we present how lichen mycobiomes are also suitable targets for improving bioinformatic analyses of fungal metabarcoding.     

Lichen diversity on dolmen and menhir in the Megalithic complex of Sa Coveccada (Mores,Sardinia)

This work describes the lichen diversity found on the megalithic Dolmen of Sa Coveccada (Mores, Sardinia) until 2010. After that year, a restoration with chemical removal of lichen crusts took place, which destroyed a great part of the lichen communities. These were studied again after removal and lichen communities occurring on rock outcrops in the surroundings of the Dolmen and on a contiguous menhir were investigated as well for comparison. Before the restoration, 33 species had been recorded on the Dolmen, most being crustose, followed by foliose and fruticose forms. Among these, eight species are regarded as rare in Sardinia and five rare at lower elevations. Most of the recorded species are typical for eutrophic substrates and for meso- to xerophytic conditions. Studies on lichen diversity on archaeological monuments in Sardinia are limited. This is the first report on the lichens of a Sardinian dolmen. This paper questions whether the lichen diversity of such monuments should be preserved as lichens have been an important part of the monument ecosystem and of the landscape for many centuries. This work also aims to improve collaboration among lichen and monument experts, in order to avoid hasty restoration decisions.     

The impact of nitrogen deposition on photobiont‐mycobiont balance of epiphytic lichens in subtropical forests of central China

Excessive nitrogen (N) deposition can impact lichen diversity in forest ecosystems, and this is a particular situation in China. Here, we examined the N uptake, assimilation, and the impact of excessive N deposition on the symbiotic balance of dominant epiphytic lichens in the subtropical forests in the Mts. Shennongjia of central China. The results show that lichen species took up, assimilated and utilized more ammonium than nitrate in a species‐specific way, following the increase of N availability. The photobiont of the lichens decreased with the increase of N concentration following an initial increase, while the mycobiont response to the N addition was not apparent. Considerable variation in response to excessive N deposition exists among the lichen species. Usnea longissima could regulate its N uptake, resulting in a stable photobiont‐mycobiont ratio among N treatments. In contrast, the photobiont‐mycobiont ratio of other four lichens increased initially but decreased when N concentration exceeded a certain level, and N stress may have broken the balance between photobiont and mycobiont of these lichens. Our results suggest that most epiphytic lichens in subtropical forest of central China could uptake and assimilate more ammonium than nitrate and that the balance between photobiont and mycobiont of many epiphytic lichens might change with the increasing N deposition load, which could impact the lichen diversity of this forest ecosystem.     

Exploring functional contexts of symbiotic sustain within lichen-associated bacteria by comparative omics

Symbioses represent a frequent and successful lifestyle on earth and lichens are one of their classic examples. Recently, bacterial communities were identified as stable, specific and structurally integrated partners of the lichen symbiosis, but their role has remained largely elusive in comparison to the well-known functions of the fungal and algal partners. We have explored the metabolic potentials of the microbiome using the lung lichen Lobaria pulmonaria as the model. Metagenomic and proteomic data were comparatively assessed and visualized by Voronoi treemaps. The study was complemented with molecular, microscopic and physiological assays. We have found that more than 800 bacterial species have the ability to contribute multiple aspects to the symbiotic system, including essential functions such as (i) nutrient supply, especially nitrogen, phosphorous and sulfur, (ii) resistance against biotic stress factors (that is, pathogen defense), (iii) resistance against abiotic factors, (iv) support of photosynthesis by provision of vitamin B₁₂, (v) fungal and algal growth support by provision of hormones, (vi) detoxification of metabolites, and (vii) degradation of older parts of the lichen thallus. Our findings showed the potential of lichen-associated bacteria to interact with the fungal as well as algal partner to support health, growth and fitness of their hosts. We developed a model of the symbiosis depicting the functional multi-player network of the participants, and argue that the strategy of functional diversification in lichens supports the longevity and persistence of lichens under extreme and changing ecological conditions.     

Topographic and forest-stand variables determining epiphytic lichen diversity in the primeval beech forest in the Ukrainian Carpathians

The Uholka-Shyrokyi Luh area of the Carpathian Biosphere Reserve is considered the largest and the most valuable primeval beech forest in Europe for biodiversity conservation. To study the impact of different topographic and forest-stand variables on epiphytic lichen diversity a total of 294 systematically distributed sampling plots were surveyed and 198 epiphytic lichen species recorded in this forest landscape, which has an uneven-aged structure. The obtained data were analysed using a non-metric multidimensional ordination and a generalized linear model. The epiphytic lichen species density at the plot level was mainly influenced by altitude and forest-stand variables. These variables are related to both the light availability i.e. canopy closure, and the habitat diversity, i.e. the developmental stage of the forest stands and the mean stem diameter. We found that lichen species density on plots with a relatively open canopy was significantly higher than on plots with a fairly loose or closed canopy structure. The late developmental stage of forest stands, which is characterized by a large number of old trees with rough and creviced bark, had a strong positive effect on lichen species density. In the Uholka-Shyrokyi Luh primeval forest the mean stem diameter of beech trees significantly correlated with lichen species density per plot. Similar trends in the species diversity of nationally red-listed lichens were revealed. Epiphytic lichens with a high conservation value nationally and internationally were found to be rather abundant in the Uholka-Shyrokyi Luh area, which shows its international importance for the conservation of forest-bound lichens.     

Bole epiphytic lichens as potential indicators of environmental change in subtropical forest ecosystems in southwest China

Lichen epiphytes are applied as excellent environmental indicators worldwide. However, very little is known about epiphytic lichen communities and their response to forest dynamics in subtropical China. This paper proposes the applications of the cover, diversity, and functional traits of epiphytic lichens to assess environmental changes associated with succession in subtropical forests of southwest China. Bole lichens were sampled from 120 plots of eight representative forest types in the Ailao Mountains. Total cover, species richness, diversity and community structure of bole lichens differed significantly among forest types, and the highest cover and diversity occurred in the Populus bonatii secondary forest (PBSF). Sixty-one indicator species were associated with particular forest types and more than 50% occurred in the PBSF. Both cover and diversity of most lichen functional groups varied regularly during forest succession. Lichen pioneer species were not displaced by competitively superior species as succession proceeds and cyanolichens were more prevalent in secondary forests. The results also highlight the importance of habitat variables such as canopy openness, host diversity, forest age, tree size, the size of the largest tree, tree density, and basal area on the lichen community. Consequently, our findings support the notion that epiphytic lichens, in terms of cover, diversity, species composition and functional traits can be used as effective indicators for large-scale and long-term forest monitoring. More importantly, the narrowly lobed foliose group was the best candidate indicator of environmental conditions in this region. The combined application of lichen indicator species and functional groups seemed to be a more reliable and more powerful method for monitoring forest dynamics in subtropical montane ecosystems.     

FREQUENCY,DIVERSITY AND ECOLOGICAL STRATEGIES OF EPIPHYTIC LICHENS IN THE SWISS CENTRAL PLATEAU AND THE PRE-ALPS

To identify representative quantitative criteria for the creation of a future Red List of epiphytic lichens, 849 trees in 132 long-term ecological observation plots in the Swiss Central Plateau and the Pre-Alps were surveyed by standard sampling. Based on the trees, frequency data of the lichen taxa observed are described by the log series model, indicating the controlling effect of few ecological factors. Based on the plots, four classes of scarcity, each comprising 25% of the species, were established. As a contribution to the development of a national, representative survey of lichens, α-diversity (species richness, species density) andβ-diversity (dissimilarity) were calculated in terms of region, vegetation formation, vegetation belt and for their combinations. Differences in lichen diversity between the Central Plateau and the Pre-Alps were caused by the bigger elevational range in the Pre-Alps, which resulted in a higher species richness. α-Diversity of forest and non-forest were similar, whereas each vegetation formation showed one third of its species restricted to it. The contributions to the total lichen diversity of crustose, foliose and fruticose as well as of generative and vegetative species was calculated. Specific features along the altitudinal gradient of vegetation belts emerged: the percentage of crustose and generative lichens declined with every altitudinal step, increased in fruticose and vegetative lichens, and was the same in foliose species.     

Significant decrease in epiphytic lichen diversity in a remote area in the European Alps,Austria

We present long-term (1993–2010) monitoring results of lichens from a remote site in Austria. Whereas gaseous pollution is negligible in this area, levels of long-range air pollution of S and N via rain and fog are high. Lichen cover on tree trunks have decreased significantly. Accordingly, we found that the population of all species had declined. The analyses demonstrated significant changes in lichen community composition. The lichen flora suffered a significant decrease in diversity. Our hypothesis is that the breakdown of epiphytic lichens is mainly due to eutrophication through long-term N deposition, and more particularly to ammonia in precipitation and in fog. Since many areas in the European Alps receive high wet deposition loads, a decrease in the abundance and diversity of epiphytic lichens is most probably a widespread phenomenon.     

Case study of the implications of climate change for lichen diversity and distributions

There is ample evidence for species distributional changes in response to recent climate change, but most studies are biased toward better known taxa. Thus, an integrated approach is needed that includes the “cryptic diversity” represented partly by lichens, which are among the most sensitive organisms to environmental change due to their physiological characteristics. The use of functional traits and ecological attributes may improve the interpretation of how species respond to climate change. Thus, we quantified the future climate change impacts on 41 lichen species distributed in the Iberian Peninsula using ensemble climatic suitability maps (derived from generalized linear and generalized additive models, and classification and regression tree analysis) and different metrics. We also determined the lichen traits/attributes that might be related to a shared response to climate change. The results indicated a loss of bioclimatic space for 75% of the species studied and an increase for 10 species, especially in Mediterranean ones. Most of the species that will lose more than 70% of their current modeled distribution area comprised big macrolichens with cyanobacteria as the photobiont, thereby indicating a great biomass loss in forests, which might affect nutrient cycles. We also found that the predicted distributions were trait-related. Smaller species, green-algae lichens, and saxicolous and epiphyte species will respond better to future climate change. The results of this type of study may help to identify the species that are most vulnerable to climate change and facilitate the development of conservation measures to avoid their decline.     

Differential sharing and distinct co‐occurrence networks among spatially close bacterial microbiota of bark,mosses and lichens‬‬

Knowledge of bacterial community host‐specificity has increased greatly in recent years. However, the intermicrobiome relationships of unrelated but spatially close organisms remain little understood. Trunks of trees covered by epiphytes represent complex habitats with a mosaic of ecological niches. In this context, we investigated the structure, diversity and interactions of microbiota associated with lichens, mosses and the bare tree bark. Comparative analysis revealed significant differences in the habitat‐associated community structures. Corresponding co‐occurrence analysis indicated that the lichen microbial network is less complex and less densely interconnected than the moss‐ and bark‐associated networks. Several potential generalists and specialists were identified for the selected habitats. Generalists belonged mainly to Proteobacteria, with Sphingomonas as the most abundant genus. The generalists comprise microorganisms with generally beneficial features, such as nitrogen fixation or other supporting functions, according to a metagenomic analysis. We argue that beneficial strains shared among hosts contribute to ecological stability of the host biocoenoses.     

Edge and land-use effects on epiphytic lichen diversity in the forest-steppe ecotone of the Mongolian Altai

Epiphytic lichen diversity was studied in forests of Siberian larch (Larix sibirica) in the forest-steppe ecotone of the Mongolian Altai. These forests are utilized for livestock grazing, fuelwood collection and occasional logging by pastoral nomads. The density of nomad households in the proximity of the forests influences epiphytic lichen diversity more strongly than the position of sample trees in the forest interior or at the forest line to the steppe. This suggests that land use exerts a stronger effect on lichen diversity than the distinct gradient in microclimate between the forest interior and the forest edge. The co-occurrence of nitrophytes with anitrophytic acidophytes on a small spatial scale as well as higher N and Ca concentrations and pH values in the bark of larch trees at the forest edge than in the forest interior indicate that moderate livestock grazing increases the epiphytic lichen diversity due to an increase of the diversity of chemically different microhabitats. Preference of many lichen species (and of rare species in particular) for overmature and decaying trees suggests that logging and fuelwood collection has adverse effects on epiphytic lichen diversity. This adverse effect is likely to be more crucial for lichen diversity than the putatively positive effect of livestock grazing, since more lichens with a preference for old and decaying trees than for nitrogen-enriched bark were found. The present study is the first one investigating the impact of pastoral nomadism in Central Asia on epiphyte diversity.     

云南哀牢山原生林及次生林群落附生地衣物种多样性与分布

附生地衣是森林附生植物的重要类群之一, 在维护森林生态系统的物种多样性以及水分和养分循环等方面发挥着重要作用。作者于2005年12月至2006年5月利用树干取样法调查了云南哀牢山徐家坝地区原生山地常绿阔叶林及其次生群落栎类萌生林、滇山杨(Populus bonatii)林和花椒(Zanthoxylum bungeanum)人工林中525株不同种类和径级树木距地面 0–2.0 m处附生地衣的组成和分布, 并收集了各个群落地面上凋落的地衣, 分析了林冠层附生地衣的物种组成。研究结果表明, 该区森林中附生地衣物种比较丰富。共收集到附生地衣61种, 分属17科29属, 其中原生林、栎类萌生林、滇山杨林和花椒人工林分别有51、53、46和23种。在树干距地面 0–2.0 m位置, 各群落中的附生地衣组成明显不同;但在林冠层中, 各群落内的附生地衣基本相似。原生林中附生地衣种类较多, 但分布不均匀。树干附生地衣的Shannon-Wiener和Simpson多样性指数以栎类萌生林最高, 分别为2.71和0.89;花椒林和滇山杨林次之, 分别为2.43–2.45和0.88–0.89;原生林最低, 为1.25和0.67。树干方位、宿主种类和宿主径级等都对附生地衣的物种组成和多样性有着重要影响, 附生地衣更多地出现于树干南向方位, 云南越桔(Vaccinium duclouxii)的附生地衣最为丰富, 胸径5.0–25.0 cm的树木上附生地衣较多。哀牢山山地森林群落中丰富的附生地衣种类及物种多样性在维系本区山地森林生态系统生物多样性格局方面具有重要的作用。     

Bacterial communities associated with the lichen symbiosis

Lichens are commonly described as a mutualistic symbiosis between fungi and "algae" (Chlorophyta or Cyanobacteria); however, they also have internal bacterial communities. Recent research suggests that lichen-associated microbes are an integral component of lichen thalli and that the classical view of this symbiotic relationship should be expanded to include bacteria. However, we still have a limited understanding of the phylogenetic structure of these communities and their variability across lichen species. To address these knowledge gaps, we used bar-coded pyrosequencing to survey the bacterial communities associated with lichens. Bacterial sequences obtained from four lichen species at multiple locations on rock outcrops suggested that each lichen species harbored a distinct community and that all communities were dominated by Alphaproteobacteria. Across all samples, we recovered numerous bacterial phylotypes that were closely related to sequences isolated from lichens in prior investigations, including those from a lichen-associated Rhizobiales lineage (LAR1; putative N(2) fixers). LAR1-related phylotypes were relatively abundant and were found in all four lichen species, and many sequences closely related to other known N(2) fixers (e.g., Azospirillum, Bradyrhizobium, and Frankia) were recovered. Our findings confirm the presence of highly structured bacterial communities within lichens and provide additional evidence that these bacteria may serve distinct functional roles within lichen symbioses.     

Community Analysis Reveals Close Affinities Between Endophytic and Endolichenic Fungi in Mosses and Lichens

Endolichenic fungi live in close association with algal photobionts inside asymptomatic lichen thalli and resemble fungal endophytes of plants in terms of taxonomy, diversity, transmission mode, and evolutionary history. This similarity has led to uncertainty regarding the distinctiveness of endolichenic fungi compared with endophytes. Here, we evaluate whether these fungi represent distinct ecological guilds or a single guild of flexible symbiotrophs capable of colonizing plants or lichens indiscriminately. Culturable fungi were sampled exhaustively from replicate sets of phylogenetically diverse plants and lichens in three microsites in a montane forest in southeastern Arizona (USA). Intensive sampling combined with a small spatial scale permitted us to decouple spatial heterogeneity from host association and to sample communities from living leaves, dead leaves, and lichen thalli to statistical completion. Characterization using data from the nuclear ribosomal internal transcribed spacer and partial large subunit (ITS-LSU rDNA) provided a first estimation of host and substrate use for 960 isolates representing five classes and approximately 16 orders, 32 families, and 65 genera of Pezizomycotina. We found that fungal communities differ at a broad taxonomic level as a function of the phylogenetic placement of their plant or lichen hosts. Endolichenic fungal assemblages differed as a function of lichen taxonomy, rather than substrate, growth form, or photobiont. In plants, fungal communities were structured more by plant lineage than by the living vs. senescent status of the leaf. We found no evidence that endolichenic fungi are saprotrophic fungi that have been “entrapped” by lichen thalli. Instead, our study reveals the distinctiveness of endolichenic communities relative to those in living and dead plant tissues, with one notable exception: we identify, for the first time, an ecologically flexible group of symbionts that occurs both as endolichenic fungi and as endophytes of mosses.     

Commercial and ethnic use of lichens in India

The diversity of Indian lichen flora has undergone a considerable decline in the last five decades. Among the various anthropogenic activities, overexploitation and selective removal of economically important lichens have become major threats to the lichen flora of the country. Commercial samples of lichens collected from different states of India were studied for their source and commercial use. Among samples offered for sale, 38 different lichen species were identified. The western Himalayas proved to be the main area for lichen collection, while some lichen species also come from the central and Western Ghats. Local traders coordinate the lichen collection and then sell the material to traders in the foothills, and from there it is processed and distributed to other parts of the country and abroad. Because lichens belonging to the families Parmeliaceae and Physciaceae are the ones most exploited commercially, they are recommended for inclusion in the CITES list.