Trends in Antarctic soil fungal research in the context of environmental changes.

Antarctic soils represent one of the most pristine environments on Earth, where highly adapted and often endemic microbial species withstand multiple extremes. Specifically, fungal diversity is extremely low in Antarctic soils and species distribution and diversity are still not fully characterized in the continent. Despite the unique features of this environment and the international interest in its preservation, several factors pose severe threats to the conservation of inhabiting ecosystems. In this light, we aimed to provide an overview of the effects on fungal communities of the main changes endangering the soils of the continent. Among these, the increasing human presence, both for touristic and scientific purposes, has led to increased use of fuels for transport and energy supply, which has been linked to an increase in unintentional environmental contamination. It has been reported that several fungal species have evolved cellular processes in response to these soil contamination episodes, which may be exploited for restoring contaminated areas at low temperatures. Additionally, the effects of climate change are another significant threat to Antarctic ecosystems, with the expected merging of previously isolated ecosystems and their homogenization. A possible reduction of biodiversity due to the disappearance of well-adapted, often endemic species, as well as an increase of biodiversity, due to the spreading of non-native, more competitive species have been suggested. Despite some studies describing the specialization of fungal communities and their correlation with environmental parameters, our comprehension of how soil communities may respond to these changes remains limited. The majority of studies attempting to precisely define the effects of climate change, including in situ and laboratory simulations, have mainly focused on the bacterial components of these soils, and further studies are necessary, including the other biotic components.

The hidden network of biocrust successional stages in the High Arctic: Revealing abiotic and biotic factors shaping microbial and metazoan communities.

Despite the important role that biocrust communities play in maintaining ecosystem structure and functioning in deglaciated barren soil, few studies have been conducted on the dynamics of biotic communities and the impact of physicochemical characteristics in shaping the different successional stages. In this study an integrated approach encompassing physicochemical parameters and molecular taxonomy was used for identifying the indicator taxa and the presence of intra- and inter-kingdom interactions in five different crust/biocrust successional stages: i) physical crust, ii) cyanobacteria-dominated biocrust, iii) cyanobacteria/moss-dominated biocrust, iv) moss-dominated biocrust and v) bryophyte carpet. The phylum Gemmatimonadota was the bacterial indicator taxon in the early stage, promoting both inter- and intra-kingdom interactions, while Cyanobacteria and Nematoda phyla played a pivotal role in formation and dynamics of cyanobacteria-dominated biocrusts. A multitrophic community, characterized by a shift from oligotrophic to copiotrophic bacteria and the presence of saproxylic arthropod and herbivore insects was found in the cyanobacteria/moss-dominated biocrust, while a more complex biota, characterized by an increased fungal abundance (classes Sordariomycetes, Leotiomycetes, and Dothideomycetes, phylum Ascomycota), associated with highly trophic consumer invertebrates (phyla Arthropoda, Rotifera, Tardigrada), was observed in moss-dominated biocrusts. The class Bdelloidea and the family Hypsibiidae (phyla Rotifera and Tardigrada, respectively) were metazoan indicator taxon in bryophyte carpet, suggesting their potential role in shaping structure and function of this late successional stage. Nitrogen and phosphorus were the main physicochemical limiting factors driving the shift among different crust/biocrust successional stages. Identification and characterization of indicator taxa, biological intra- and inter-kingdom interactions and abiotic factors driving the shift among different crust/biocrust successional stages provide a detailed picture on crust/biocrust dynamics, revealing a strong interconnection among micro- and macrobiota systems. These findings enhance our understanding of biocrust ecosystems in High Arctic, providing valuable insights for their conservation and management in response to environmental shifts due to climate change.

Wide divergence of fungal communities inhabiting rocks and soils in a hyper-arid Antarctic desert.

Highly simplified microbial communities colonise rocks and soils of continental Antarctica ice-free deserts. These two habitats impose different selection pressures on organisms, yet the possible filtering effects on the diversity and composition of microbial communities have not hitherto been fully characterised. We hence compared fungal communities in rocks and soils in three localities of inner Victoria Land. We found low fungal diversity in both substrates, with a mean species richness of 28 across all samples, and significantly lower diversity in rocks than in soils. Rock and soil communities were strongly differentiated, with a multinomial species classification method identifying just three out of 328 taxa as generalists with no affinity for either substrate. Rocks were characterised by a higher abundance of lichen-forming fungi (typically Buellia, Carbonea, Pleopsidium, Lecanora, and Lecidea), possibly owing to the more protected environment and the porosity of rocks permitting photosynthetic activity. In contrast, soils were dominated by obligate yeasts (typically Naganishia and Meyerozyma), the abundances of which were correlated with edaphic factors, and the black yeast Cryomyces. Our study suggests that strong differences in selection pressures may account for the wide divergences of fungal communities in rocks and soils of inner Victoria Land.

Effects of 5-year experimental warming in the Alpine belt on soil Archaea: Multi-omics approaches and prospects.

We currently lack a predictive understanding of how soil archaeal communities may respond to climate change, particularly in Alpine areas where warming is far exceeding the global average. Here, we characterized the abundance, structure, and function of total (by metagenomics) and active soil archaea (by metatranscriptomics) after 5-year experimental field warming (+1°C) in Italian Alpine grasslands and snowbeds. Our multi-omics approach unveiled an increasing abundance of Archaea during warming in snowbeds, which was negatively correlated with the abundance of fungi (by qPCR) and micronutrients (Ca and Mg), but positively correlated with soil water content. In the snowbeds transcripts, warming resulted in the enrichment of abundances of transcription and nucleotide biosynthesis. Our study provides novel insights into possible changes in soil Archaea composition and function in the climate change scenario.

Knufia obscura sp. nov. and Knufia victoriae sp. nov., two new species from extreme environments.

Six strains of black meristematic fungi were isolated from Antarctic soils, gasoline car tanks and from the marine alga Flabellia petiolata. These fungi were characterized by morphological, physiological and phylogenetic analyses. According to the maximum-likelihood analysis reconstructed with ITS and LSU sequences, these strains belonged to the genus Knufia. Knufia obscura sp. nov. (holotype CBS 148926) and Knufia victoriae sp. nov. (holotype CBS 149015) are proposed as two novel species and descriptions of their morphological, physiological and phylogenetic features are presented. Based on the maximum-likelihood analyses, K. obscura was closely related to Knufia hypolithi (99 % bootstrap support), while K. victoriae clustered in the clade of Knufia cryptophialidica and Knufia perfecta (93 % bootstrap support). Knufia victoriae, recorded in Antarctic soil samples, had a psychrophilic behaviour, with optimal growth between 10 and 15 °C and no growth recorded at 20 °C. Knufia obscura, from a gasoline car tank and algae, displayed optimal growth between 20 and 25 °C and was more tolerant to salinity than K. victoriae.

Highly differentiated soil bacterial communities in Victoria Land macro-areas (Antarctica).

Ice-free areas of Victoria Land, in Antarctica, are characterized by different terrestrial ecosystems, that are dominated by microorganisms supporting highly adapted communities. Despite the unique conditions of these ecosystems, reports on their bacterial diversity are still fragmentary. From this perspective, 60 samples from 14 localities were analyzed. These localities were distributed in coastal sites with differently developed biological soil crusts, inner sites in the McMurdo Dry Valleys with soils lacking of plant coverage, and a site called Icarus Camp, with a crust developed on a thin locally weathered substrate of the underlying parent granitic-rock. Bacterial diversity was studied through 16S rRNA metabarcoding sequencing. Communities diversity, composition and the abundance and composition of different taxonomic groups were correlated to soil physicochemical characteristics. Firmicutes, Bacteroidetes, Cyanobacteria and Proteobacteria dominated these communities. Most phyla were mainly driven by soil granulometry, an often disregarded parameter and other abiotic parameters. Bacterial composition differed greatly among the three macrohabitats, each having a distinct bacterial profile. Communities within the two main habitats (coastal and inner ones) were well differentiated from each other as well, therefore depending on site-specific physicochemical characteristics. A core community of the whole samples was observed, mainly represented by Firmicutes and Bacteroidetes.

Growth Forms and Functional Guilds Distribution of Soil Fungi in Coastal Versus Inland Sites of Victoria Land, Antarctica.

In Victoria Land, Antarctica, ice-free areas are restricted to coastal regions and dominate the landscape of the McMurdo Dry Valleys. These two environments are subjected to different pressures that determine the establishment of highly adapted fungal communities. Within the kingdom of fungi, filamentous, yeasts and meristematic/microcolonial growth forms on one side and different lifestyles on the other side may be considered adaptive strategies of particular interest in the frame of Antarctic constraints. In this optic, soil fungal communities from both coastal and Dry Valleys sites, already characterized thorough ITS1 metabarcoding sequencing, have been compared to determine the different distribution of phyla, growth forms, and lifestyles. Though we did not find significant differences in the richness between the two environments, the communities were highly differentiated and Dry Valleys sites had a higher evenness compared to coastal ones. Additionally, the distribution of different growth forms and lifestyles were well differentiated, and their diversity and composition were likely influenced by soil abiotic parameters, among which soil granulometry, pH, P, and C contents were the potential main determinants.

Extracellular Enzymes and Bioactive Compounds from Antarctic Terrestrial Fungi for Bioprospecting.

Antarctica, one of the harshest environments in the world, has been successfully colonized by extremophilic, psychrophilic, and psychrotolerant microorganisms, facing a range of extreme conditions. Fungi are the most diverse taxon in the Antarctic ecosystems, including soils. Genetic adaptation to this environment results in the synthesis of a range of metabolites, with different functional roles in relation to the biotic and abiotic environmental factors, some of which with new biological properties of potential biotechnological interest. An overview on the production of cold-adapted enzymes and other bioactive secondary metabolites from filamentous fungi and yeasts isolated from Antarctic soils is here provided and considerations on their ecological significance are reported. A great number of researches have been carried out to date, based on cultural approaches. More recently, metagenomics approaches are expected to increase our knowledge on metabolic potential of these organisms, leading to the characterization of unculturable taxa. The search on fungi in Antarctica deserves to be improved, since it may represent a useful strategy for finding new metabolic pathways and, consequently, new bioactive compounds.

Expansion of shrubs could result in local loss of soil bacterial richness in Western Greenland.

Climate warming in Greenland is facilitating the expansion of shrubs across wide areas of tundra. Given the close association between plants and soil microorganisms and the important role of soil bacteria in ecosystem functioning, it is of utmost importance to characterize microbial communities of arctic soil habitats and assess the influence of plant edaphic factors on their composition. We used 16S rRNA gene amplicons to explore the bacterial assemblages of three different soil habitats representative of a plant coverage gradient: bare ground, biological soil crusts dominated by mosses and lichens and vascular vegetation dominated by shrubs. We investigated how bacterial richness and community composition were affected by the vegetation coverage, and soil pH, moisture and carbon (C), nitrogen (N) and phosphorus (P) contents. Bacterial richness did not correlate with plant coverage complexity, while community structure varied between habitats. Edaphic variables affected both the taxonomic richness and community composition. The high number of Amplicon Sequence Variants (ASVs) indicators of bare ground plots suggests a risk of local bacterial diversity loss due to expansion of vascular vegetation.

Vegetation, pH and Water Content as Main Factors for Shaping Fungal Richness, Community Composition and Functional Guilds Distribution in Soils of Western Greenland.

Fungi are the most abundant and one of the most diverse components of arctic soil ecosystems, where they are fundamental drivers of plant nutrient acquisition and recycling. Nevertheless, few studies have focused on the factors driving the diversity and functionality of fungal communities associated with these ecosystems, especially in the scope of global warming that is particularly affecting Greenland and is leading to shrub expansion, with expected profound changes of soil microbial communities. We used soil DNA metabarcoding to compare taxonomic and functional composition of fungal communities in three habitats [bare ground (BG), biological soil crusts (BSC), and vascular vegetation (VV) coverage] in Western Greenland. Fungal richness increased with the increasing complexity of the coverage, but BGs and BSCs samples showed the highest number of unique OTUs. Differences in both fungal community composition and distribution of functional guilds identified were correlated with edaphic factors (mainly pH and water content), in turn connected with the different type of coverage. These results suggest also possible losses of diversity connected to the expansion of VV and possible interactions among the members of different functional guilds, likely due to the nutrient limitation, with potential effects on elements recycling.

Altitude and fungal diversity influence the structure of Antarctic cryptoendolithic Bacteria communities.

Endolithic growth within rocks is a critical adaptation of microbes living in harsh environments where exposure to extreme temperature, radiation, and desiccation limits the predominant life forms, such as in the ice-free regions of Continental Antarctica. The microbial diversity of the endolithic communities in these areas has been sparsely examined. In this work, diversity and composition of bacterial assemblages in the cryptoendolithic lichen-dominated communities of Victoria Land (Continental Antarctica) were explored using a high-throughput metabarcoding approach, targeting the V4 region of 16S rDNA. Rocks were collected in 12 different localities (from 14 different sites), along a gradient ranging from 1000 to 3300 m a.s.l. and at a sea distance ranging from 29 to 96 km. The results indicate Actinobacteria and Proteobacteria are the dominant taxa in all samples and defined a 'core' group of bacterial taxa across all sites. The structure of bacteria communities is correlated with the fungal counterpart and among the environmental parameters considered, altitude was found to influence bacterial biodiversity, while distance from sea had no evident influence.

Effect of environmental parameters on biodiversity of the fungal component in lithic Antarctic communities.

A wide sampling of rocks, colonized by microbial epi-endolithic communities, was performed along an altitudinal gradient from sea level to 3600 m asl and sea distance from the coast to 100 km inland along the Victoria Land Coast, Antarctica. Seventy-two rock samples of different typology, representative of the entire survey, were selected and studied using denaturing gradient gel electrophoresis to compare variation in fungal diversity according to environmental conditions along this altitudinal and sea distance transect. Lichenized fungi were largely predominant in all the samples studied and the biodiversity was heavily influenced even by minimal local variations. The n-MDS analysis showed that altitude and sea distance affect fungal biodiversity, while sandstone allows the communities to maintain high biodiversity indices. The Pareto-Lorenz curves indicate that all the communities analyzed are highly adapted to extreme conditions but scarcely resilient, so any external perturbation may have irreversible effects on these fragile ecosystems.