Abstract: Lichen specimen collections for bryophytes, primarily from Washington and Oregon. Only Darwin Core 2 fields are accessible through this interface. Localities and geographic coordinates for rare taxa have been withheld.

Abstract: The Mars Desert Research Station is a Mars analog research site located in the desert outside of Hanksville, Utah, U.S.A. Here we present a preliminary checklist of the vascular plant and lichen flora for the station, based on collections made primarily during a two-week simulated Mars mission in November, 2014. Additionally, we present notes on the endolithic chlorophytes and cyanobacteria, and the identification of a fungal genus also based on these collections. Altogether, we recorded 38 vascular plant species from 14 families, 13 lichen species from seven families, six algae taxa including both chlorophytes and cyanobacteria, and one fungal genus from the station and surrounding area. We discuss this floristic diversity in the context of the ecology of the nearby San Rafael Swell and the desert areas of Wayne and Emery counties in southeastern Utah.

Abstract: Lichens are commonly recognized as a symbiotic association of a fungus and chlorophyll containing partner, either green algae or cyanobacteria, or both. The fungus provides a suitable habitat for the partner, which provides photosynthetically fixed carbon as energy source for the system. The evolutionary result of the self-sustaining partnership is a unique joint structure, the lichen thallus, which is indispensable for fungal sexual reproduction. The classical view of a dual symbiosis has been challenged by recent microbiome research, which revealed host-specific bacterial microbiomes. The recent results about bacterial associations with lichens symbioses corroborate their notion as a multi-species symbiosis. Multi-omics approaches have provided evidence for functional contribution by the bacterial microbiome to the entire lichen meta-organism while various abiotic and biotic factors can additionally influence the bacterial community structure. Results of current research also suggest that neighboring ecological niches influence the composition of the lichen bacterial microbiome. Specificity and functions are here reviewed based on these recent findings, converging to a holistic view of bacterial roles in lichens. Finally we propose that the lichen thallus has also evolved to function as a smart harvester of bacterial symbionts. We suggest that lichens represent an ideal model to study multi-species symbiosis, using the recentily available omics tools and other cutting edge methods.

Abstract: Previous studies of the impacts of climate change on lichens and fungi have focused largely on alpine and subalpine habitats, and have not investigated the potential impact on narrowly endemic species. Here, we estimate the impacts of climate change on high-elevation, endemic lichens in the southern Appalachians, a global diversity hotspot for many groups of organisms, including lichens. We conducted extensive field surveys in the high elevations of the region to accurately document the current distributions of eight narrowly endemic lichen species. Species distribution modeling was used to predict how much climatically suitable area will remain within, and north of, the current range of the target species under multiple climate change scenarios at two time points in the future. Our field work showed that target species ranged from extreme rarity to locally abundant. Models predicted over 93 % distributional loss for all species investigated and very little potentially suitable area north of their current distribution in the coming century. Our results indicate that climate change poses a significant threat to high-elevation lichens, and provide a case study in the application of current modeling techniques for rare, montane species.

Abstract: Original data comes from a project which takes or took place as part of the DFG priority program "Exploratories for large-scale and long-term functional biodiversity research". The data is stored together with descriptive metadata, in combination called a dataset, in the project repository (https://www.bexis.uni-jena.de). Species information was extracted from that original dataset. The second paragraph is part of the metadata of the original dataset. species list with abundance per plot

Abstract: A wide variety of terrestrial ecosystems in tundra have a ground vegetation cover composed of reindeer lichens (genera Cladonia and Cetraria). The microbial communities of two lichen-dominated ecosystems typical of the sub-arctic zone of northwestern Siberia, that is a forested tundra soil and a shallow acidic peatland, were examined in our study. As revealed by molecular analyses, soil and peat layers just beneath the lichen cover were abundantly colonized by bacteria from the phylum Planctomycetes. Highest abundance of planctomycetes detected by fluorescence in situ hybridization was in the range 2.2-2.7 × 107 cells per gram of wet weight. 16S rRNA gene fragments from the Planctomycetes comprised 8-13% of total 16S rRNA gene reads retrieved using Illumina pair-end sequencing from the soil and peat samples. Lichen-associated assemblages of planctomycetes displayed unexpectedly high diversity, with a total of 89,662 reads representing 1723 operational taxonomic units determined at 97% sequence identity. The soil of forested tundra was dominated by uncultivated members of the family Planctomycetaceae (53-71% of total Planctomycetes-like reads), while sequences affiliated with the Phycisphaera-related group WD2101 (recently assigned to the order Tepidisphaerales) were most abundant in peat (28-51% of total reads). Representatives of the Isosphaera-Singulisphaera group (14-28% of total reads) and the lineages defined by the genera Gemmata (1-4%) and Planctopirus-Rubinisphaera (1-3%) were present in both habitats. Two strains of Singulisphaera-like bacteria were isolated from studied soil and peat samples. These planctomycetes displayed good tolerance of low temperatures (4-15°C) and were capable of growth on a number of polysaccharides, including lichenan, a characteristic component of lichen-derived phytomass.

Abstract: The fungal genus Collemopsidium comprises species that develop so-called borderline lichen symbioses with algae or cyanobacteria. Together with morphologically similar pyrenocarpous fungi it has been assigned to the family Xanthopyreniaceae. The adscription of this family to higher taxonomic ranks remain uncertain. Using sequence data of five nuclear genomic regions (nuLSU, nuSSU, tef1-α, rpb1 and rpb2) and one mitochondrial locus (mtSSU) we found that the studied representatives of this family are placed in the Dothideomyceta, yet relationships with the classes Dothideomycetes and Arthoniomycetes remain uncertain. We describe the new order Collemopsidiales to accommodate the genus Collemopsidium (paraphyletic as currently understood) and the lichenicolous genus Zwackhiomyces. Using five fungal fossils as calibrations points, we infer an age of c. 230 Mya for the crown of Collemopsidiales. Based on two molecular markers, we also provide insight into the global diversity of marine species of the genus Collemopsidium. According to the species delimitation algorithm GMYC, c. 26 putative species exist, far more than the six species recognized hitherto. We have confirmed this result by comparing the two alternative species models by means of Bayes factors, using path sampling and stepping-stone sampling algorithms to estimate the marginal likelihood of each model. Finally, our observations suggest rock-boring ability evolved in parallel in the different lineages within this group of fungi.

Abstract: Lichens, self-supporting mutualistic associations between a fungal partner and one or more photosynthetic partners, also harbor non-photosynthetic bacteria. The diversity and contribution of these bacteria to the functioning of lichen symbiosis have recently begun to be studied, often by culture-independent techniques due to difficulties in their isolation and culture. However, culturing as yet unculturable lichenic bacteria is critical to unravel their potential functional roles in lichen symbiogenesis, to explore and exploit their biotechnological potential and for the description of new taxa. Our objective was to improve the recovery of lichen associated bacteria by developing novel isolation and culture approaches, initially using the lichen Pseudevernia furfuracea. We evaluated the effect of newly developed media enriched with novel lichen extracts, as well as the influence of thalli washing time and different disinfection and processing protocols of thalli. The developed methodology included: i) the use of lichen enriched media to mimic lichen nutrients, supplemented with the fungicide natamycin; ii) an extended washing of thalli to increase the recovery of ectolichenic bacteria, thus allowing the disinfection of thalli to be discarded, hence enhancing endolichenic bacteria recovery; and iii) the use of an antioxidant buffer to prevent or reduce oxidative stress during thalli disruption. The optimized methodology allowed significant increases in the number and diversity of culturable bacteria associated with P. furfuracea, and it was also successfully applied to the lichens Ramalina farinacea and Parmotrema pseudotinctorum. Furthermore, we provide, for the first time, data on the abundance of culturable ecto- and endolichenic bacteria that naturally colonize P. furfuracea, R. farinacea and P. pseudotinctorum, some of which were only able to grow on lichen enriched media. This innovative methodology is also applicable to other microorganisms inhabiting these and other lichen species.

Abstract: In this chapter we examine the structure, composition, and function of soil biocrust (i.e., biological soil crust) communities dominated by lichens. Lichens are composite organisms resulting from a symbiotic relationship between a fungus and an alga or cyanobacterium. Biocrust lichens can tolerate a wide range of abiotic stresses such as desiccation, extreme temperature, and high light intensity, making them well adapted to life in arid, semiarid, and polar deserts where resources are limited and competition from vascular plants is low. The ability of some lichens to fix atmospheric nitrogen also gives them a competitive advantage over other organisms in environments that are recovering from disturbance. Biocrust lichens perform many important ecological functions such as providing habitat for microfauna, stabilizing soils, fixing nitrogen and carbon, and enhancing water flow through the soil. However, as with other biocrust taxa, they are highly susceptible to physical disturbances such as trampling by livestock, disturbance by vehicular traffic, and fire. Biocrust lichen ecology is hampered by a lack of information on the distribution of many taxa, difficulties of identifying small organisms, and the high cost of sampling environments where biocrust lichens are likely to form a major component of the surface biota. The adoption of morphological groups for classifying biocrust lichens has helped to simplify the study of biocrust community ecology, but more sophisticated DNA sequencing techniques are needed in order to better understand their ecology, distribution, and how they interact with other soil crust organisms.

Abstract: Investigations were done on two former open-cast lignite mining sites under reclamation, an artificial sand dune in Welzow Sud, and a forest plantation in Schlabendorf Sud (Brandenburg, Germany). The aim was to associate the topsoil hydrological characteristics of green algae dominated as well as moss and soil lichen dominated biological soil crusts during crustal succession with their water retention and the repellency index on sandy soils under temperate cli-mate and different reliefs. The investigation of the repellency index showed on the one hand an increase due to the cross-linking of sand parti-cles by green algae which resulted in clogging of pores. On the other hand, the occurrence of moss plants led to a de-crease of the repellency index due to absorption caused by bryophytes. The determination of the water retention curves showed an increase of the water holding capacity, especially in conjunction with the growth of green algae layer. The pore-related van Genuchten parameter indicate a clay-like behaviour of the developed soil crusts. Because of the inho-mogeneous distribution of lichens and mosses as well as the varying thickness of green algae layers, the water retention differed between the study sites and between samples of similar developmental stages. However, similar tendencies of water retention and water repellency related to the soil crust formation were observed. Biological soil crusts should be considered after disturbances in the context of reclamation measures, because the ini-tial development of green algae biocrusts lead to an increasing repellency index, while the occurrence of mosses and a gain in organic matter enhance the water holding capacity. Thus, the succession of biocrusts and their small-scale succes-sion promote the development of soil and ecosystem.

Abstract: Biological soil crusts (biocrusts) are fundamental components of drylands worldwide, and are of great importance for the regulation of ecosystem functioning. However, little is known on the role of species identify of biocrust-forming lichens in mediating the response of nitrogen (N) cycling to concurring global environmental change. Here, we conducted a microcosm study to evaluate how the species identity of biocrust-forming lichens (Diploschistes thunbergianus, Psora crystallifera and Xanthoparmelia reptans) regulate key processes of N cycling in response to simulated changes in rainfall frequency and N addition. We explicitly considered both direct and indirect effects (i.e. driven via microbial diversity and abundance) of global changes on N availability and losses using structural equation models. Our results showed that species of biocrust-forming lichens differentially mediated effects of N amendment and altered rainfall frequencies on belowground nitrate availability and N2O flux rate. For instance, soils under P. crystallifera species showed the highest increase in nitrate content in response to N amendment under low rainfall frequency. Moreover, soils under D. thunbergianus showed the highest N2O flux under high rainfall frequency without N addition. Interestingly, soils under X. reptans showed lowest and highest resistance in nitrate availability and N2O flux, respectively, in response to N addition regardless of different rainfall frequencies. Strikingly, we only found an indirect impact of either rainfall frequency or N amendment on the nitrate availability (but not N2O flux) driven via the ammonia-oxidizing community under X. reptans. Our results provide evidence that the species identity of biocrust-forming lichens modulates the response of N cycling to global change drivers. These findings have implications for predicting the potential consequence of altered rainfall patterns and environmental N inputs in dryland ecosystems.

Abstract: We studied the relative importance of several environmental factors for tundra plant communities in five locations across Svalbard (High Arctic) that differed in geographical location, oceanographic and climatic influence, and soil characteristics. The amount of marine-derived nitrogen in the soil supplied by seabirds was locally the most important of the studied environmental factors influencing the tundra plant community. We found a strong positive correlation between δ15N isotopic values and total N content in the soil, confirming the fundamental role of marine-derived matter to the generally nutrient-poor Arctic tundra ecosystem. We also recorded a strong correlation between the δ15N values of soil and of the tissues of vascular plants and mosses, but not of lichens. The relationship between soil δ15N values and vascular plant cover was linear. In the case of mosses, the percentage ground cover reached maximum around a soil δ 15N value of 8‰, as did plant community diversity. This soil δ15N value clearly separated the occurrence of plants with low nitrogen tolerance (e.g. Salix polaris) from those predominating on high N content soils (e.g. Cerastium arcticum, Poa alpina). Large colonies of planktivorous little auks have a great influence on Arctic tundra vegetation, either through enhancing plant abundance or in shaping plant community composition at a local scale.

Abstract: Climate warming leads to more intensive evaporation from the Arctic sea resulting in increased precipitation in the low Arctic, e.g., higher snowfall during winter. Deeper snow keeps the arctic soils warmer and alters soil attributes and vegetation, e.g., increase in nitrogen availability, expansion of shrubs and decline in shade-intolerant lichens and bryophytes. Changes in soil properties and vegetation are expected to influence on saprotrophic and plant-symbiotic fungi, but how increased snow depth affects their community composition remain unknown. In the present work, we used DNA metabarcoding to study the effects of long-term experimental manipulations of snow depth on soil fungal communities in dry heath and moist tussock tundra in Arctic Alaska. We report strong changes in fungal community compositions in the two tundra types, with pronounced declines observed in the majority of fungal functional guilds, including ectomycorrhizal, lichenized, plant pathogenic, saprotrophic and bryophyte-associated species. The observed changes in lichenized and bryophyte-associated fungi are in agreement with previously published above-ground changes, i.e. decrease of lichen and bryophyte cover and diversity. However, the majority of observed trends, including the decline of ectomycorrhizal fungi (that were anticipated to benefit from the expansion of their host plants), suggest that changes in fungal communities do not entirely correspond to and are not primarily driven by shifts in vegetation. Instead, arctic fungal communities appear to exhibit faster turnover that may be influenced by dynamic interactions with numerous biotic and abiotic factors, e.g., soil nutrient cycling and community dynamics in other groups of soil microorganisms. We highlight the importance of “below-ground studies” in assessing ecosystem responses to climatic changes, because faster turnover of microbial communities may be applicable for monitoring early-stage alterations caused by climatic changes.

Abstract: Seabirds in the Arctic are known to link marine and terrestrial ecosystems. They feed in the sea and breed on land, where they deposit enormous amounts of guano. Soil in the vicinity of seabird colonies is much more enriched with nutrients as compared to areas beyond their impact, which positively affects primary and secondary production. Water bears (Tardigrada) are microinvertebrates which constitute a permanent and ubiquitous faunal component of polar regions. Here, we tested the influence of seabird guano on Tardigrada communities; we established two transects in Hornsund (SW Spitsbergen): (1) SEABIRD where little auks (Alle alle) nested, and (2) CONTROL, an area without a little auk colony. In total, we collected 160 moss, lichen and mixed (moss/lichen) samples from those areas. In total, we found 1990 specimens belonging to 32 taxa (25 identified to species level). The average density of water bears was higher in the SEABIRD transect (9.31 ind g−1), where mosses predominated over lichens, in comparison with the CONTROL transect (5.83 ind g−1), where more lichens occurred. Thus, ornithogenic enrichment of soil and locally facilitated development of mosses over lichens might be important factors responsible for the increase in invertebrate abundance. According to canonical correspondence analysis, the little auk colony effect explained 13.2 % of the tardigrade species composition, and this factor appeared to be more important than the vegetation type itself. Four taxa, i.e. Isohypsibius cf. reticulatus, Microhypsibius bertolanii, Minibiotus cf. formosus and Ramazzottius cf. rupeus, have been recorded in the Svalbard archipelago for the first time.

Abstract: Nitrogen (N) fixation is the main source of ‘new’ N for N-limited ecosystems like subarctic and arctic tundra. This crucial ecosystem function is performed by a wide range of N2 fixer (diazotroph) associations that could differ fundamentally in their timing and amount of N release to the soil. To assess the importance of different associative N2 fixers for ecosystem N cycling, we tracked 15N-N2 into four N2-fixer associations (with a legume, lichen, free-living, moss) and into soil, microbial biomass and non-diazotroph-associated plants 3 days and 5 weeks after in situ labelling. In addition, we tracked 13C from 13CO2 labelling to assess if N and C fixation are linked. Three days after labelling, half of the fixed 15N was recovered in the legume soils, indicating a fast release of fixed N2. Within 5 weeks, the free-living N2 fixers released two-thirds of the fixed 15N into the soil, whereas the lichen and moss retained the fixed 15N. Carbon and N2 fixation were linked in the lichen shortly after labelling, in free-living N2 fixers 5 weeks after labelling, and in the moss at both sampling times. The four investigated N2-fixer associations released fixed N2 at different rates into the soil, and non-diazotroph-associated plants have no access to ‘new’ N within several weeks after N2 fixation. Although legumes and free-living N2 fixers are immediate sources of ‘new’ N for N-limited tundra ecosystems, lichens and especially mosses, do not contribute to increase the N pool via N2 fixation in the short term.

Abstract: Large, architecturally complex lichen symbioses arose only a few times in evolution, increasing thallus size by orders of magnitude over those from which they evolved. The innovations that enabled symbiotic assemblages to acquire and maintain large sizes are unknown. We mapped morphometric data against an eight-locus fungal phylogeny across one of the best-sampled thallus size transition events, the origins of the Placopsis lichen symbiosis, and used a phylogenetic comparative framework to explore the role of nitrogen-fixing cyanobacteria in size differences. Thallus thickness increased by >150% and fruiting body core volume increased ninefold on average after acquisition of cyanobacteria. Volume of cyanobacteria-containing structures (cephalodia), once acquired, correlates with thallus thickness in both phylogenetic generalized least squares and phylogenetic generalized linear mixed-effects analyses. Our results suggest that the availability of nitrogen is an important factor in the formation of large thalli. Cyanobacterial symbiosis appears to have enabled lichens to overcome size constraints in oligotrophic environments such as acidic, rain-washed rock surfaces. In the case of the Placopsis fungal symbiont, this has led to an adaptive radiation of more than 60 recognized species from related crustose members of the genus Trapelia. Our data suggest that precyanobacterial symbiotic lineages were constrained to forming a narrow range of phenotypes, so-called cryptic species, leading systematists until now to recognize only six of the 13 species clusters we identified in Trapelia.

Abstract: Lichens are obligate symbioses between fungi and green algae or cyanobacteria. Most lichens resynthesize their symbiotic thalli from propagules, but some develop within the structures of already existing lichen symbioses. Diploschistes muscorum starts as a parasite infecting the lichen Cladonia symphycarpa and gradually develops an independent Diploschistes lichen thallus. Here we studied how this process influences lichen-associated microbiomes and photobionts by sampling four transitional stages, at sites in Sweden and Germany, and characterizing their microbial communities using high-throughput 16S rRNA gene and photobiont-specific ITS rDNA sequencing, and fluorescence in situ hybridization. A gradual microbiome shift occurred during the transition, but fractions of Cladonia-associated bacteria were retained during the process of symbiotic reorganization. Consistent changes observed across sites included a notable decrease in the relative abundance of Alphaproteobacteria with a concomitant increase in Betaproteobacteria. Armatimonadia, Spartobacteria and Acidobacteria also decreased during the infection of Cladonia by Diploschistes. The lichens differed in photobiont specificity. Cladonia symphycarpa was associated with the same algal species at all sites, but Diploschistes muscorum had a flexible strategy with different photobiont combinations at each site. This symbiotic invasion system suggests that partners can be reorganized and selected for maintaining potential roles rather than depending on particular species.

Abstract: Biological soil crusts (biocrusts) are found in a broad range of climatic zones as components of open vegetation. In addition to photosynthetically active and N-fixing Cyanobacteria, heterotrophic bacteria and fungi are integral components of biocrusts. In this chapter, we present an overview of the diversity and interactions of fungi and bacteria. Both can occur “free-living” or in tight associations. Lichen symbioses, which frequently form a dominating aspect of biocrusts, tend to be a particular hotspot of complex bacterial and fungal interactions. Biocrust lichens can be colonized by other fungi or by other lichens, and they represent a particularly rich habitat for bacterial communities as shown by recent research results. Nevertheless, metabolic details and the regulatory principle of fungal and bacterial dynamics remain largely unclear and are in need of refined analysis. According to present data, biocrust bacteria and fungi seem to be widespread, diverse, and capable of withstanding the harsh and fluctuating conditions of biocrust habitats as long as stability of soil is warranted.

Abstract: Fourteen species of lichens, fifteen lichenicolous fungi and one saprobic fungus are reported for the first time for St. Petersburg, Western or Eastern Leningrad Region. The lichen Lecidella meiococca and the lichenicolous fungus Tremella phaeophysciae are reported as new to Russia, the lichen Lecania sambucina and the lichenicolous fungus Endococcus tricolorans are new for the European Russia, the lichens Buellia arborea, Chaenotheca cinerea, Bellemerea sanguinea , resinicolous calicioid fungus Chaenothecopsis mediarossica and lichenicolous fungi Arthonia molendoi, Lichenochora obscuroides, Pronectria leptaleae, Sphaerellothecium cladoniae are new for the North-Western European Russia. The most interesting records are briefly discussed.