Growing interest in understanding the relevance of marine fungi to food webs, biogeochemical cycling, and biological patterns necessitates establishing a context for interpreting future findings. To help establish this context, we summarize the diversity of cultured and observed marine planktonic fungi from across the world. While exploring this diversity, we discovered that only half of the known marine fungal species have a publicly available DNA locus, which we hypothesize will likely hinder accurate highthroughput sequencing classification in the future, as it does currently. Still, we reprocessed >600 high-throughput datasets and analyzed 4.9 × 10 9 sequences (4.8 × 10 9 shotgun metagenomic reads and 1.0 × 10 8 amplicon sequences) and found that every fungal phylum is represented in the global marine planktonic mycobiome; however, this mycobiome is generally predominated by three phyla: the Ascomycota, Basidiomycota, and Chytridiomycota. We hypothesize that these three clades are the most abundant due to a combination of evolutionary histories, as well as physical processes that aid in their dispersal. We found that environments with atypical salinity regimes (>5 standard deviations from the global mean: Red Sea, Baltic Sea, sea ice) hosted higher proportions of the Chytridiomycota, relative to open oceans that are dominated by Dikarya. The Baltic Sea and Mediterranean Sea had the highest fungal richness of all areas explored. An analysis of similarity identified significant differences between oceanographic regions. There were no latitudinal gradients of marine fungal richness and diversity observed. As more high-throughput sequencing data become available, expanding the collection of reference loci and genomes will be essential to understanding the ecology of marine fungi.

Recent molecular evidence suggests a global-distribution of marine fungi; however, the ecological relevance and corresponding biological contributions of fungi to marine ecosystems remains largely unknown. We assessed fungal biomass from the open Arctic Ocean by applying novel biomass conversion factors from cultured-isolates to environmental sterol and CARD-FISH data. We found an average of 16.54 nmol m -3 of ergosterol in sea ice and seawater, which corresponds to 1.74 mg C m -3 (444.56 mg C m -2 in seawater). Using Chytridiomycota-specific probes, we observed freeliving and particulate-attached cells that averaged 34.07 µg C m -3 in sea ice and seawater (11.66 mg C m -2 in seawater). Summed CARD-FISH and ergosterol values approximate 1.77 mg C m -3 in sea ice and seawater (456.23 mg C m -2 in seawater), which is similar to biomass estimates of other marine taxa generally considered integral to marine food webs and ecosystem processes. Using the GeoChip microarray, we detected evidence for fungal viruses within the Partitiviridae in sediment, as well as fungal genes involved in the degradation of biomass and the assimilation of nitrate. To bridge our observations of fungi on particulate and the detection of degradative genes, we germinated fungal conidia in zooplankton fecal pellets and grew fungal conidia after eight months incubation in sterile seawater. Ultimately, these data suggest that fungi could be as important in oceanic ecosystems as they are in freshwater environments. Significance Statement:The oceans cover 71% of the world's surface; however, the contributions of marine fungi to these environments remain a gap to understanding marine ecosystem processes and associated biogeochemical cycling. Here we provide the first biomass estimates of fungi from the Arctic Ocean, supplemented with a functional gene inventory, and experiment-based ecological data demonstrating putative niches occupied by fungi in the marine environment. We provide evidence that marine fungi comprise a comparable quantity of biomass relative to other taxonomic groups that are generally considered to be essential ecosystem components.

Future supplies of rare minerals for global industries with high-tech products may depend on deep-sea mining. However, environmental standards for seafloor integrity and recovery from environmental impacts are missing. We revisited the only midsize deep-sea disturbance and recolonization experiment carried out in 1989 in the Peru Basin nodule field to compare habitat integrity, remineralization rates, and carbon flow with undisturbed sites. Plough tracks were still visible, indicating sites where sediment was either removed or compacted. Locally, microbial activity was reduced up to fourfold in the affected areas. Microbial cell numbers were reduced by ~50% in fresh “tracks” and by <30% in the old tracks. Growth estimates suggest that microbially mediated biogeochemical functions need over 50 years to return to undisturbed levels. This study contributes to developing environmental standards for deep-sea mining while addressing limits to maintaining and recovering ecological integrity during large-scale nodule mining.

Sedimentary biofilms comprising microbial communities mediating the anaerobic oxidation of methane are rare. Here, we describe two biofilm communities discovered in sediment cores recovered from Arctic cold seep sites (gas hydrate pingos) in the north-western Barents Sea, characterized by steady methane fluxes. We found macroscopically visible biofilms in pockets in the sediment matrix at the depth of the sulphate-methane-transition zone. 16S rRNA gene surveys revealed that the microbial community in one of the two biofilms comprised exclusively of putative anaerobic methanotrophic archaea of which ANME-1 was the sole archaeal taxon. The bacterial community consisted of relatives of sulphate-reducing bacteria (SRB) belonging to uncultured Desulfobacteraceae clustering into SEEP-SRB1 (i.e. the typical SRB associated to ANME-1), and members of the atribacterial JS1 clade. Confocal laser scanning microscopy demonstrates that this biofilm is composed of multicellular strands and patches of ANME-1 that are loosely associated with SRB cells, but not tightly connected in aggregates. Our discovery of methanotrophic biofilms in sediment pockets closely associated with methane seeps constitutes a hitherto overlooked and potentially widespread sink for methane and sulphate in marine sediments.

Abstract. Glaciers are known to harbor surprisingly complex ecosystems. On their surface, distinct cylindrical holes filled with meltwater and sediments are considered as hot spots for microbial life. The present paper addresses possible biological interactions within the community of prokaryotic cyanobacteria and eukaryotic microalgae (microalgae) and relations to their potential grazers, additional to their environmental controls. Svalbard glaciers with substantial allochthonous input of material from local sources reveal high microalgal densities. Small valley glaciers with high sediment coverages and high impact of birds show high biomasses and support a high biological diversity. Invertebrate grazer densities do not show any significant negative correlation with microalgal abundances, but a positive correlation with eukaryotic microalgae. Most microalgae found in this study form large colonies (< 10 cells, or > 25 μm), which may protect them against invertebrate grazing. This finding rather indicates grazing as a positive control on eukaryotic microalgae by nutrient recycling. Density differences between the eukaryotic microalgae and prokaryotic cyanobacteria and their high distinction in RDA and PCA analyses indicate that these two groups are in strong contrast. Eukaryotic microalgae occurred mainly in unstable cryoconite holes with high sediment loads, high N : P ratios, and a high impact of bird guano, as a proxy for nutrients. In these environments autochthonous nitrogen fixation appears to be negligible. Selective wind transport of Oscillatoriales via soil and dust particles is proposed to explain their dominance in cryoconites further away from the glacier margins. We propose that, for the studied glaciers, nutrient levels related to recycling of limiting nutrients is the main factor driving variation in the community structure of microalgae and grazers.