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• Bacterial communities form the basis of biogeochemical processes and determine plant growth and health. Mosses harbour diverse bacterial communities that are involved in nitrogen fixation and carbon cycling. Global climate change is causing changes in aboveground plant biomass and shifting species composition in the Arctic, but little is known about the response of moss microbiomes in these environments. • Here, we studied the total and potentially active bacterial communities associated with Racomitrium lanuginosum in response to a 20-yr in situ warming in an Icelandic heathland. We evaluated the effect of warming and warming-induced shrub expansion on the moss bacterial community composition and diversity, and nifH gene abundance. • Warming changed both the total and the potentially active bacterial community structure, while litter abundance only affected the total bacterial community structure. The abundance of nifH genes was negatively affected by litter abundance. We also found shifts in the potentially nitrogen-fixing community, with Nostoc decreasing and noncyanobacterial diazotrophs increasing in relative abundance. • Our data suggest that the moss microbial community and potentially nitrogen fixing taxa will be sensitive to future warming, partly via changes in litter and shrub abundance.

Primary producers shape terrestrial biodiversity, but most research has focused on vascular plants, while the role of cryptogams (mosses, lichens and algae) remains under‐explored. Cryptogams dominate Antarctic vegetation and support diverse microarthropod communities. However, how cryptogam traits influence these communities remains poorly understood. We therefore investigated the role of 28 cryptogam species and one vascular plant, via their functional traits, in shaping microarthropod communities across three contrasting sites (Signy Island, Byers Peninsula and Rothera) in the maritime Antarctic. We hypothesized that vegetation nitrogen and moisture content, major microarthropod taxa, and abiotic drivers interact to influence community patterns. Vegetation type effects on total microarthropod abundance were context‐dependent. Mosses hosted more microarthropods than lichens at Signy Island, but these differences diminished further south. Microarthropod richness and springtail abundance were consistently higher in mosses than lichens across all sites, whereas mite abundance did not differ between vegetation types. Cryptogam nitrogen and moisture content strongly predicted microarthropod community patterns, although their influence varied with vegetation type and location. Among mosses, moisture increased springtail abundance but reduced diversity due to the dominance of Cryptopygus antarcticus. In lichens, nitrogen had a stronger influence than in mosses, particularly on mite abundance and Shannon diversity. As hypothesized, moisture was more important at the harshest southern site, while nitrogen had stronger effects at more productive northern locations. These findings emphasize that the influence of cryptogam traits in structuring Antarctic terrestrial biodiversity is modulated by the environmental context. With future shifts predicted in vegetation composition, the functional traits of emerging dominant species may restructure microarthropod communities and their ecological functions.

Mosses are among the first colonizing organisms after glacier retreat and can develop into thick moss mats during later successional stages. They are key players in N2 fixation through their microbiome, which is an important process for nutrient buildup during primary succession. How these moss–microbe interactions develop during succession is not well studied and is relevant in the light of climate change and increased glacier retreat. We examined how the bacterial communities associated with two moss species of the genus Racomitrium and the underlying soil, as well as moss traits and nitrogen fixation, develop along a successional gradient in the glacier forefield of Fláajökull in southeast Iceland. In addition, we tested whether moss functional traits, such as total carbon (TC) and total nitrogen (TN) contents, moss moisture content, and moss shoot length are drivers of moss and underlying soil bacterial communities. Although time since deglaciation did not affect TN and moss moisture contents, TC and shoot length increased with time since deglaciation. Moss and underlying soil bacterial communities were distinct. While the soil bacterial community structure was driven by moss C/N ratios, the moss bacterial community structure was linked to time since deglaciation, moss C/N ratio, and moss moisture content. Moss N2‐fixation rates were linked to bacterial community composition and nifH gene abundance rather than moss TN or time since deglaciation. This was accompanied by a shift from autotrophic to heterotrophic diazotrophs. Overall, our results suggest that there is little lateral transfer between moss and soil bacterial communities and that moss traits affect moss and soil bacterial community structure. Only moss bacterial community changed with time since deglaciation. In addition, moss N2‐fixation rates are determined by bacterial community structure, rather than moss traits or time since deglaciation. This study on the interplay between succession, mosses, soils, and their bacterial communities will inform future work on the fate of newly exposed areas as a result of glacier retreat.

Lava tubes on Mars hold exciting potential for the preservation of biosignatures, which may survive on geological timescales in these isolated, stable environments. To support the development of future astrobiological mission concepts, we turn to terrestrial lava tubes, host to a variety of microbial communities and secondary minerals. Following a multidisciplinary sampling protocol, we retrieved biological, molecular, and mineralogical data from several lava tubes in Iceland. We report on blue‐colored copper‐rich secondary minerals and their associated bacterial communities using a multi‐method approach, and an amalgam of 16S rRNA gene sequencing, Raman spectroscopy, scanning electron microscopy, and energy‐dispersive X‐ray spectroscopy data sets. We found numerous bacterial genera known for their high metal resistance and ability to survive in low‐nutrient environments. Both are characteristics to be expected for any potential life in Martian lava tubes, and should be considered when checking for contaminants in Mars mission preparations. Associated with the microbial mats, we identified several types of copper‐rich secondary minerals, indicating localized copper enrichments in the groundwater, possibly stemming from overlying ash deposits and nearby hyaloclastite formations. Molecular analysis revealed carotenoid signals preserved within the copper speleothems. If found in Martian lava tubes, blue copper‐rich mineral precipitates would be deserving of astrobiological investigation, as they have potential to preserve biosignatures and harbor life. Plain Language Summary Subterranean lava tubes on Mars are exciting locations to study in the potential discovery of signs of life outside of Earth, as the surface of Mars does not have conditions conducive to the preservation of life as we know it. In order to better study these Martian environments we look first to comparable lava tubes on Earth. Within Icelandic lava tubes we found blue‐colored copper minerals, host to microbial life. The microbes that thrive in these caves are able to withstand extreme conditions, and leave behind detectable molecular traces indicative of life, a type of biosignature. Using a variety of tools and techniques, we describe the nature of the blue minerals and their provenance, the role of the microbial populations within them, and the value of the molecular traces as biosignatures. We discuss the potential for such minerals and microbes in Martian lava tubes, and how we might successfully sample them in future missions to Mars. Key Points Icelandic lava tubes are appropriate analog test sites for multi‐disciplinary sampling protocols for future missions to Martian lava tubes Blue copper‐rich speleothems serve as biotopes for metal‐resistant organisms in lava tubes Copper‐enriched groundwater filters into lava tubes and precipitates as chrysocolla, a mechanism that may be relevant to lava tubes on Mars

Mosses are among the first colonizing organisms after glacier retreat and can develop into thick moss mats during later successional stages. They are key players in N2 fixation through their microbiome, which is an important process for nutrient build-up during primary succession. How these moss-microbe interactions develop during succession is not well-studied and is relevant in the light of climate change and increased glacier retreat. We examined how the bacterial communities associated with two moss species of the genus Racomitrium and the underlying substrate, as well as moss traits and nitrogen fixation, develop along a successional gradient in the glacier forefield of Fláajökull in southeast Iceland. In addition, tested whether moss functional traits, such as total carbon (TC) and nitrogen contents (TN) are drivers of moss and underlying soil bacterial communities. Although time since deglaciation did not affect TN and moisture content, TC and shoot length increased with time since deglaciation. Moss and underlying soil bacterial communities were distinct. While the soil bacterial community structure was driven by the time since deglaciation and moss C/N ratios, the moss bacterial community structure was linked to time since deglaciation and moss moisture content. Moss N2-fixation rates were linked to bacterial community composition and nifH gene abundance rather than moss TN or time since deglaciation. This was accompanied by a shift from autotrophic to heterotrophic diazotrophs. Overall, our results suggest that there is little lateral transfer between moss and soil bacterial communities and that moss traits and time since deglaciation affect moss and soil bacterial community structure. In addition, moss N2-fixation rates are determined by bacterial community structure, rather than moss traits or time since deglaciation. Competing Interest Statement The authors have declared no competing interest.

Lichens are traditionally defined as a symbiosis between a fungus and a green alga and or a cyanobacterium. This idea has been challenged by the discovery of bacterial communities inhabiting the lichen thalli. These bacteria are thought to contribute to the survival of lichens under extreme and changing environmental conditions. How these changing environmental conditions affect the lichen-associated bacterial community composition remains unclear. We describe the total (rDNA-based) and potentially metabolically active (rRNA-based) bacterial community of the lichen Cetaria islandica and its response to long-term warming using a 20-year warming experiment in an Icelandic sub-Arctic tundra. 16S rRNA and rDNA amplicon sequencing showed that the orders Acetobacterales (of the class Alphaproteobacteria) and Acidobacteriales (of the phylum Acidobacteria) dominated the bacterial community. Numerous ASVs (amplicon sequence variants) taxa could only be detected in the potentially active community but not in the total community. Long-term warming led to increases in relative abundance on class, order and ASV level. Warming altered the relative abundance of ASVs of the most common bacterial genera, such as Granulicella and Endobacter. The potentially metabolically active bacterial community was also more responsive to warming than the total community. Our results suggest that the bacterial community of the lichen C. islandica is dominated by acidophilic taxa and harbors disproportionally active rare taxa. We also show for the first time that climate warming can lead to shifts in lichen-associated bacterial community composition.

Abstract 1. Bacterial communities form the basis of biogeochemical processes and determine plant growth and health. Mosses, an abundant plant group in Arctic ecosystems, harbour diverse bacterial communities that are involved in nitrogen fixation and carbon cycling. Global climate change is causing changes in aboveground plant biomass and shifting species composition in the Arctic, but little is known about the response of moss microbiomes. 2. Here, we studied the total and potentially active bacterial community associated with Racomitrium lanuginosum, in response to 20-year in situ warming in an Icelandic heathland. We evaluated the effect of warming and warming-induced shrub expansion on the moss bacterial community composition and diversity, nifH gene abundance and nitrogen-fixation rates. 3. Warming changed both the total and the potentially active bacterial community structure, while litter abundance only affected the total bacterial community structure. The relative abundance of Proteobacteria increased, while the relative abundance of Cyanobacteria and Acidobacteria decreased. NifH gene abundance and nitrogen-fixation rates were negatively affected by litter and Betula nana abundance, respectively. We also found shifts in the potentially nitrogen-fixing community, with Nostoc decreasing and non-cyanobacterial diazotrophs increasing in relative abundance. Our data suggests that the moss microbial community including the potentially nitrogen-fixing taxa is sensitive to future warming. 4. Synthesis. Long-term warming led to a shift in moss-associated bacterial community composition, while the abundance of nitrogen-fixing bacteria and nitrogen-fixation rates were negatively affected by increased litter and Betula nana abundance respectively. Warming and increased shrub abundance as a result of warming can affect moss-associated bacterial communities and nitrogen fixation rates in tundra ecosystems. Competing Interest Statement The authors have declared no competing interest.