Empirical testing of cryoconite granulation: Role of cyanobacteria in the formation of key biogenic structure darkening glaciers in polar regions.

Cryoconite, the dark sediment on the surface of glaciers, often aggregates into oval or irregular granules serving as biogeochemical factories. They reduce a glacier's albedo, act as biodiversity hotspots by supporting aerobic and anaerobic microbial communities, constitute one of the organic matter (OM) sources on glaciers, and are a feeder for micrometazoans. Although cryoconite granules have multiple roles on glaciers, their formation is poorly understood. Cyanobacteria are ubiquitous and abundant engineers of cryoconite hole ecosystems. This study tested whether cyanobacteria may be responsible for cryoconite granulation as a sole biotic element. Incubation of Greenlandic, Svalbard, and Scandinavian cyanobacteria in different nutrient availabilities and substrata for growth (distilled water alone and water with quartz powder, furnaced cryoconite without OM, or powdered rocks from glacial catchment) revealed that cyanobacteria bind mineral particles into granules. The structures formed in the experiment resembled those commonly observed in natural cryoconite holes: they contained numerous cyanobacterial filaments protruding from aggregated mineral particles. Moreover, all examined strains were confirmed to produce extracellular polymeric substances (EPS), which suggests that cryoconite granulation is most likely due to EPS secretion by gliding cyanobacteria. In the presence of water as the only substrate for growth, cyanobacteria formed mostly carpet-like mats. Our data empirically prove that EPS-producing oscillatorialean cyanobacteria isolated from the diverse community of cryoconite microorganisms can form granules from mineral substrate and that the presence of the mineral substrate increases the probability of the formation of these important and complex biogeochemical microstructures on glaciers.

Functional and Taxonomic Diversity of Anaerobes in Supraglacial Microbial Communities.

Cryoconite holes are small ponds present on the surface of most glaciers filled with meltwater and sediment at the bottom. Although they are characterized by extreme conditions, they host bacterial communities with high taxonomic and functional biodiversity. Despite that evidence for a potential niche for anaerobic microorganisms and anaerobic processes has recently emerged, the composition of the microbial communities of the cryoconite reported so far has not shown the relevant presence of anaerobic taxa. We hypothesize that this is due to the lower growth yield of anaerobes compared to aerobic microorganisms. In this work, we aim at evaluating whether the anaerobic bacterial community represents a relevant fraction of the biodiversity of the cryoconite and at describing its structure and functions. We collected sediment samples from cryoconite holes on the Forni Glacier (Italy) and sequenced both 16S rRNA amplicon genes and 16S rRNA amplicon transcripts at different times of the day along a clear summer day. Results showed that a relevant fraction of taxa has been detected only by 16S rRNA transcripts and was undetectable in 16S rRNA gene amplicons. Furthermore, in the transcript approach, anaerobic taxa were overrepresented compared with DNA sequencing. The metatranscriptomics approach was used also to investigate the expression of the main metabolic functions. Results showed the occurrence of syntrophic and commensalism relationships among fermentative bacteria, hydrogenothrophs, and consumers of fermentation end products, which have never been reported so far in cryoconite. IMPORTANCE Recent evidence disclosed the presence of a potential niche for anaerobic microorganisms and anaerobic processes in supraglacial sediments (cryoconite), but a detailed description of the structure and functions of the anaerobic population is still lacking. This work used rRNA and mRNA sequencing and demonstrated that anaerobes are very active in these environments and represent a relevant albeit neglected part of the ecosystem functions in these environments.

Impact of anthropogenic contamination on glacier surface biota.

Glaciers are ecosystems and they host active biological communities. Despite their remoteness, glaciers act as cold condensers where high precipitation rates and cold temperatures favor the deposition of pollutants. These contaminants include a broad range of substances, including legacy pollutants, but also compounds still largely used. Some of these compounds are monitored in the environment and their effects on the ecosystems are known, in contrast others can be defined as emerging pollutants since their presence and their impact on the environment are still poorly understood (e.g. microplastics, radionuclides). This review aim to provide an overview of the studies that have investigated the effects of pollutants on the supraglacial ecosystem so far. Despite the distribution of the pollutants in glacier environments has been discussed in several studies, no review paper has summarized the current knowledge on the effects of these substances on the ecological communities living in glacier ecosystems.

Trophic and symbiotic links between obligate-glacier water bears (Tardigrada) and cryoconite microorganisms.

Insights into biodiversity and trophic webs are important for understanding ecosystem functions. Although the surfaces of glaciers are one of the most productive and biologically diverse parts of the cryosphere, the links between top consumers, their diet and microbial communities are poorly understood. In this study, for the first time we investigated the relationships between bacteria, fungi and other microeukaryotes as they relate to tardigrades, microscopic metazoans that are top consumers in cryoconite, a biologically rich and productive biogenic sediment found on glacier surfaces. Using metabarcoding (16S rDNA for bacteria, ITS1 for fungi, and 18S rDNA for other microeukaryotes), we analyzed the microbial community structures of cryoconite and compared them with the community found in both fully fed and starved tardigrades. The community structure of each microbial group (bacteria, fungi, microeukaryotes) were similar within each host group (cryoconite, fully fed tardigrades and starved tardigrades), and differed significantly between groups, as indicated by redundancy analyses. The relative number of operational taxonomic units (ZOTUs, OTUs) and the Shannon index differed significantly between cryoconite and tardigrades. Species indicator analysis highlighted a group of microbial taxa typical of both fully fed and starved tardigrades (potential commensals), like the bacteria of the genera Staphylococcus and Stenotrophomonas, as well as a group of taxa typical of both cryoconite and fully fed tardigrades (likely part of the tardigrade diet; bacteria Flavobacterium sp., fungi Preussia sp., algae Trebouxiophyceae sp.). Tardigrades are consumers of bacteria, fungi and other microeukaryotes in cryoconite and, being hosts for diverse microbes, their presence can enrich the microbiome of glaciers.

Unveiling the extreme environmental radioactivity of cryoconite from a Norwegian glacier.

This study is a first survey of the occurrence of artificial (137Cs, 241Am, 207Bi, Pu isotopes) and natural (210Pb, 228Ac, 214Bi, 40K) radionuclides in Norwegian cryoconite. Cryoconite samples were collected before (12 samples) and after (5 samples) a rainfall event, after which 7 cryoconite holes dissapeared. The concentrations of radionuclides in cryoconite samples from the Blåisen Glacier are compared with data from the Arctic and Alpine glaciers. Cryoconite samples from the studied glacier had extremely high activity concentrations of 137Cs, 241Am, 207Bi and 239+240Pu (up to 25,000 Bq/kg, 58 Bq/kg, 13 Bq/kg and 131 Bq/kg, respectively) and also high concentrations of organic matter (OM), comparing to other Scandinavian and Arctic glaciers, reaching up to ~40% of total mass. The outstandingly high concentrations of 137Cs, 241Am, Pu isotopes, and 207Bi on the Blåisen Glacier are primarily related to bioaccumulation of radionuclides in organic-rich cryoconite and might be enhanced by additional transfers of contamination from the tundra by lemmings during their population peaks. The presumed influence of intense rainfall on radionuclide concentrations in the cryoconite was not confirmed.

Cryoconite - From minerals and organic matter to bioengineered sediments on glacier's surfaces.

Cryoconite is a mixture of mineral and organic material covering glacial ice, playing important roles in biogeochemical cycles and lowering the albedo of a glacier surface. Understanding the differences in structure of cryoconite across the globe can be important in recognizing past and future changes in supraglacial environments and ice-organisms-minerals interactions. Despite the worldwide distribution and over a century of studies, the basic characteristics of cryoconite, including its forms and geochemistry, remain poorly studied. The major purpose of our study is the presentation and description of morphological diversity, chemical and photoautotrophs composition, and organic matter content of cryoconite sampled from 33 polar and mountain glaciers around the globe. Observations revealed that cryoconite is represented by various morphologies including loose and granular forms. Granular cryoconite includes smooth, rounded, or irregularly shaped forms; with some having their surfaces covered by cyanobacteria filaments. The occurrence of granules increased with the organic matter content in cryoconite. Moreover, a major driver of cryoconite colouring was the concentration of organic matter and its interplay with minerals. The structure of cyanobacteria and algae communities in cryoconite differs between glaciers, but representatives of cyanobacteria families Pseudanabaenaceae and Phormidiaceae, and algae families Mesotaeniaceae and Ulotrichaceae were the most common. The most of detected cyanobacterial taxa are known to produce polymeric substances (EPS) that may cement granules. Organic matter content in cryoconite varied between glaciers, ranging from 1% to 38%. The geochemistry of all the investigated samples reflected local sediment sources, except of highly concentrated Pb and Hg in cryoconite collected from European glaciers near industrialized regions, corroborating cryoconite as element-specific collector and potential environmental indicator of anthropogenic activity. Our work supports a notion that cryoconite may be more than just simple sediment and instead exhibits complex structure with relevance for biodiversity and the functioning of glacial ecosystems.

What do orange spots reveal about male (and female) guppies? A test using correlated responses to selection.

Female preferences for male ornamental traits can arise from indirect benefits, such as increased attractiveness or better viability of progeny, but empirical evidence for such benefits is inconsistent. Artificial selection offers a powerful way to investigate indirect effects of male ornaments. Here, we selected for the area of orange spots on male guppies, a trait subject to female preferences in our population, in replicated up- and down-selected lines. We found a significant direct response to selection, and a correlated response in female preferences, with females from down-selected lines showing less interest in more orange males. Nevertheless, up-selected males sired more offspring in direct competition with low-selected males, irrespective of female origin. We did not find a significantly correlated response to selection among any other fitness correlates we measured. Our results imply that female preferences for orange spots can lead to increased reproductive success of their sons, with no effect on general viability of progeny. Furthermore, although we demonstrate that female preferences may evolve via linkage disequilibrium with the preferred trait, the potential for runaway selection by positive feedback may be constrained by the lack of corresponding linkage with male reproductive competitiveness.

Biotope and biocenosis of cryoconite hole ecosystems on Ecology Glacier in the maritime Antarctic.

Despite recent great interest in glacier ecosystems in the continental Antarctic, little is known about their maritime counterparts. Our study presents descriptive data on cryoconite sediments and cryoconite holes on Ecology Glacier (King George Island) to accomplish three main objectives: (a) to identify main eukaryotic (algae, invertebrates) and prokaryotic (cyanobacteria) components of microbial communities; (b) to provide a "baseline" of community composition, organic matter and artificial contamination; and (c) to identify key abiotic factors that might be important in community assembly. Cryoconite holes were sampled along an altitudinal gradient of Ecology Glacier in January, mid Austral Summer 2017. Cryoconite holes located in lower altitude were deeper than those located in the middle and the highest altitude. Seventeen species of algae and cyanobacteria with biomass of 0.79 to 5.37 µg/cm3 have been found in sediments. Dominant species were cyanobacterial Pseudanabaena frigida and Bacillariophyceae Microcostaus sp. Biomass of Bacillariophyceae was significantly higher than that of Chlorophyta and Cyanobacteria. We found three species of rotifers (potentially two new to science) and for the first time a glacier dwelling Acari (suspension feeder, Nanorchestes nivalis). Organic matter content ranged from 5.4% to 7.6%. Investigated artificial radionuclides included 137Cs, 238Pu, 239+240Pu and 241Am. 210Pb seems to be related to organic matter content. Overall, cryoconite holes on Ecology Glacier present unique habitats that serve as biodiversity hotspots of psychrophiles, source of organic matter, matrices for radioactivity tracking and model for observing changes in supraglacial ecosystems in the maritime Antarctic.

An integrative redescription of the nominal taxon for the Mesobiotus harmsworthi group (Tardigrada: Macrobiotidae) leads to descriptions of two new Mesobiotus species from Arctic.

The Mesobiotus harmsworthi group has a global distribution, with localities in polar, temperate and tropical zones. Since the first species of the harmsworthi group was described in the beginning of the 20th century, tens of new species within the group were found and named. However, the diagnosis of the nominal Mesobiotus harmsworthi is insufficient and enigmatic, thus it can be is a serious obstacle in solving the taxonomy of this group. Here, we integratively redescribe the nominal species for the genus Mesobiotus, i.e., Mesobiotus harmsworthi and clarify taxonomic statuses of the two subspecies: M. harmsworthi harmsworthi and M. harmsworthi obscurus that have been recognised as distinct taxa for more than three decades. Traditionally, egg chorion in M. harmsworthi was considered almost smooth and without any traces of areolation, however here we report many misunderstandings that accumulated across decades and we show that, in fact, the chorion in this species exhibits a partially developed areolation. We present an integrative (morphological, morphometric and molecular) diagnosis of the nominal taxon and we confirm that it differs from other species of the harmsworthi group by morphological characters of both animals and eggs. Additionally, we describe two new species of the genus Mesobiotus: M. skorackii sp. nov. from the Kyrgyz Republic (using classical morphological description) and M. occultatus sp. nov. from Svalbard Archipelago (by means of integrative taxonomy). Finally, we also provide the first genetic phylogeny of the genus Mesobiotus based on COI sequences which, together with molecular species delimitation, independently confirms the validity of the analysed taxa.

Tardigrada and Rotifera from moss microhabitats on a disappearing Ugandan glacier, with the description of a new species of water bear.

Glaciers and ice sheets are a peculiar biome with characteristic abiotic and biotic components. Mountain glaciers are predicted to decrease their volume and even to melt away within a few decades. Despite the threat of a disappearing biome, the diversity and the role of microscopic animals as consumers at higher trophic levels in the glacial biome still remain largely unknown. In this study, we report data on tardigrades and rotifers found in glacial mosses on Mount Stanley, Uganda, and describe a new tardigrade species. Adropion afroglacialis sp. nov. differs from the most similar species by having granulation on the cuticle, absence of cuticular bars under the claws, and a different macroplacoid length sequence. We also provide a morphological diagnosis for another unknown tardigrade species of the genus Hypsibius. The rotifers belonged to the families Philodinidae and Habrotrochidae. In addition, we discuss the diversity of microinvertebrates and potential role of tardigrades and rotifers on mountain glaciers as top consumers. As for any organism living apparently exclusively in glacial habitats on tropical glaciers, their extinction in the near future is inevitable, possibly before we can even discover their existence.