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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's 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-dust 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 included 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 cementing matter and form granules. Organic matter content in cryoconite varied between glaciers, ranging from 1% to >40%. 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 collectors 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.

In both natural and built environments, microbes on occasions manifest in spherical aggregates instead of substratum-affixed biofilms. These microbial aggregates are conventionally referred to as granules. Cryoconites are mineral rich granules that appear on glacier surfaces and are linked with expanding surface darkening, thus decreasing albedo, and enhanced melt. The oxygenic photogranules (OPGs) are organic rich granules that grow in wastewater, which enables wastewater treatment with photosynthetically produced oxygen and which presents potential for net autotrophic wastewater treatment in a compact system. Despite obvious differences inherent in the two, cryoconite and OPG pose striking resemblance. In both, the order Oscillatoriales in Cyanobacteria envelope inner materials and develop dense spheroidal aggregates. We explore the mechanism of photogranulation on account of high similarity between cryoconites and OPGs. We contend that there is no universal external cause for photogranulation. However, cryoconites and OPGs, as well as their intravariations, which are all under different stress fields, are the outcome of universal physiological processes of the Oscillatoriales interfacing with goldilocks interactions of stresses. Finding the rules of photogranulation may enhance engineering of glacier and wastewater systems to manipulate their ecosystem impacts.

To understand the geological origins of minerals in cryoconite and the nutrients sources for microbes on glaciers, we analyzed the Sr–Nd isotopic ratios of the four mineral fractions in cryoconites including saline, carbonate, phosphate, silicate and the organic fraction obtained from Gulkana Glacier in Alaska. The isotopes in the silicate mineral fraction exhibited spatial variation within the glacier (87Sr/86Sr: 0.704533–0.709563, εNd (0): −16.0 to 0.5), which can be explained by the different mixing ratios of the two distinct sources: one of the sources is lateral and terminal moraines or soil, and the other is the medial moraine of the glacier. The minerals in the cryoconite at the lower sites in the glacier are likely derived from the former source, whereas those at the upper sites are from latter sources. The mineralogical and elemental compositions also support mixing of the silicate minerals from the two local sources. The Sr isotopic ratios of the organic fraction also showed spatial variation on the glacier in the middle sites – a trend similar to those of the phosphate fraction. The results suggest that the organic matter is mostly the byproducts of microbes using the phosphate minerals as a nutrient source.

The global cryosphere, Earth's frozen water, is in precipitous decline. The ongoing and predicted impacts of cryosphere loss are diverse, ranging from disappearance of entire biomes to crises of water availability. Covering approximately one-fifth of the planet, mass loss from the terrestrial cryosphere is driven primarily by a warming atmosphere but reductions in albedo (the proportion of reflected light) also contribute by increasing absorption of solar radiation. In addition to dust and other abiotic impurities, biological communities substantially reduce albedo worldwide. In this review, we provide a global synthesis of biological albedo reduction (BAR) in terrestrial snow and ice ecosystems. We first focus on known drivers—algal blooms and cryoconite (granular sediment on the ice that includes both mineral and biological material)—as they account for much of the biological albedo variability in snow and ice habitats. We then consider an array of potential drivers of BAR whose impacts may be overlooked, such as arthropod deposition, resident organisms (e.g., dark-bodied glacier ice worms), and larger vertebrates, including humans, that transiently visit the cryosphere. We consider both primary (e.g., BAR due to the presence of pigmented algal cells) and indirect (e.g., nutrient addition from arthropod deposition) effects, as well as interactions among biological groups (e.g., birds feeding on ice worms). Collectively, we highlight that in many cases, overlooked drivers and interactions among factors have considerable potential to alter BAR, perhaps rivaling the direct effects of algal blooms and cryoconite. We conclude by highlighting knowledge gaps for the field with an emphasis on the underrepresentation of genomic tools, understudied areas (particularly high-elevation glaciers at tropical latitudes), and a dearth of temporal sampling in current efforts. We detail a global framework for long-term BAR monitoring that, if implemented, would yield a tremendous amount of insight for BAR and would be particularly valuable in light of the rapid ecological and physical changes occurring in the contemporary cryosphere.

Cryoconite granules are naturally occurring microbial structures on glacier surfaces worldwide. They play a key role in carbon and nitrogen cycling in glacier ecosystems and can accelerate the melting of snow and ice. However, detailed mechanism of nitrogen cycling in cryoconite granules remains unclear. Here, we demonstrate that redox stratification affects the spatial distribution of N cycling processes in cryoconite granules. Based on microsensor measurements for O2, NH4+, NO2– and NO3–, we identified the presence of fine-scale redox stratification within cryoconite granules. Cyanobacteria at the surface layer of the granules created oxic conditions, whereas the inner core of the granules was anoxic. Metatranscriptomic analyses indicated the active occurrences of nitrification in the inner core, whereas denitrification actively occurred both in the inner core and the surface layer of the granules. Cyanobacteria in the inner core of the granules were inactive, and likely dead and being degraded, providing carbon and nitrogen to support nitrifiers and denitrifiers. Quantities of nitrification genes/transcripts were greater in large cryoconite granules than small ones, most likely because nitrogen substrates were more abundantly present in the inner core of large granules due to distinct redox stratification. Our results suggest that the development of a granular structure of cryoconite granules can largely affect carbon and nitrogen cycling on glaciers.

To understand the microbial composition and diversity patterns, cryoconite granules were collected from two geographical areas, i.e., Nepali Himalaya and Greenland, Arctic. 16S rRNA, ITS and the D1/D2 domain sequencing techniques were used for characterization of microbial communities of the four glaciers. The total 13 species of bacteria such as Bacillus aryabhattai, Bacillus simplex, Brevundimonas vesicularis, Cryobacterium luteum, Cryobacterium psychrotolerans, Dermacoccus nishinomiyaensis, Glaciihabitans tibetensis, Leifsonia kafniensis, Paracoccus limosus, Polaromonas glacialis, Sporosarcina globispora, Staphylococcus saprophyticus, Variovorax ginsengisoli, and 4 species of fungi such as Goffeauzyma gilvescens, Mrakia robertii, Dothideomycetes sp., Helotiales sp. were recorded from Nepali Himalaya. Among these, 12 species of bacteria and 4 species of fungi are new contributions to Himalaya. In contrast to this, six species of bacteria such as Bacillus cereus, Cryobacterium psychrotolerans, Dermacoccus nishinomiyaensis, Enhydrobacter aerosaccus, Glaciihabitans tibetensis, Subtercola frigoramans, and nine species of fungi such as Goffeauzyma gilvescens, Mrakia robertii, Naganishia vaughanmartiniae, Piskurozyma fildesensis, Rhodotorula svalbardensis, Alatospora acuminata, Articulospora sp., Phialophora sp., Thelebolus microspores, and Dothideomycetes sp.), were recorded from Qaanaaq, Isunnguata Sermia and Thule glaciers, Greenland. Among these, five species of bacteria and seven species of fungi are new contributions to Greenland cryoconite. Microbial analyses indicate that the Nepali Himalayan cryoconite colonize higher numbers of microbial species compared to the Greenland cryoconite.

Cryoconite, a sediment found on the surface of glaciers, is known for its ability to accumulate radionuclides. New data on cryoconite from the Morteratsch glacier (Switzerland) are presented to shed light on the mechanisms that control the distribution of radioactivity in cryoconite. Among the radionuclides detected in our samples, we have identified 108mAg, an artificial species which has never been observed in terrestrial environments before. This finding supports that cryoconite has an extraordinary ability to accumulate radioactivity. Our results also show that the radioactivity of cryoconite from a single glacier is far from uniform. Both the absolute amount of radioactivity and the relative contribution of single radionuclides are highly variable in samples from the Morteratsch glacier. To investigate the processes responsible for such variability, we have explored the correlation between radionuclides, organic and inorganic carbon fractions and the morphological features of cryoconite deposits. We have found that the degree of connection between cryoconite and supraglacial hydrology is particularly important, since it strongly influences the accumulation of radionuclides in cryoconite. Cryoconite holes connected with supraglacial channels are rich in cosmogenic 7Be; in contrast, poorly connected deposits are rich in artificial fallout radionuclides and elemental carbon. The very different half-lives of 7Be and artificial radionuclides allowed us to discuss our findings in relation to the age and maturity of cryoconite deposits, highlighting the potential use of radionuclides to investigate hydrological supraglacial processes and material cycling at the surface of glaciers.

Cryoconite holes are water-filled cylindrical holes formed on ablation ice surfaces and commonly observed on glaciers worldwide. Temporal changes of cryoconite holes characteristically −2, 40–55% of the incoming flux). In contrast, holes developed in sunny conditions correspond to high solar radiation (186–278 W m−2, 63–88%). Results suggest that the dimensions of holes drastically changed depending on the weather conditions and that frequent cloudy, warm and windy conditions would cause a decay of holes and weathering crust, inducing an increase in the cryoconite coverage on the ice, consequently darkening the glacier surface.

Cryoconite granules are dark-colored biological aggregates on glaciers. Bacterial community varies with granule size, however, community change in space and their susceptibility to environmental factors has not been described yet. Therefore, we focused on bacterial community from four different granule sizes (30–249 μm, 250–750 μm, 750–1599 μm, more than 1600 μm diameter) in 10 glaciers in northwestern Greenland and their susceptibility to exogenous nutrients in cryoconite hole. A filamentous cyanobacterium Phormidesmis priestleyi, which has been frequently reported from glaciers in Arctic was abundant (10%–26%) across any size of granules on most of glaciers. Bacterial community across glaciers became similar with size increase, and whence smallest size fractions contain more unique genera in each glacier. Multivariate analysis revealed that effect of nutrients to beta diversity is larger in smaller granules (30–249 μm and 250–750 μm diameter), suggesting that bacterial susceptibility to nutrients changes with growth of granule (i.e. P. priestleyi was affected by nitrate in early growth stage).

Cryoconite granules, which are dark-colored biological aggregates on glaciers, effectively accelerate the melting of glacier ice. Bacterial community varies with granule size, however, community change in space and their susceptibility to environmental factors has not been described yet. Therefore, we focused on bacterial community from 4 different granule sizes (30-249 μm, 250- 750 μm, 750-1599 μm, more than 1600 μm diameter) in 10 glaciers in northwestern Greenland and their susceptibility for exogenous nutrients in cryoconite hole. A filamentous cyanobacterium Phormidesmis priestleyi, which has been frequently reported from glaciers in Arctic was abundant (10-26%) across any size of granules on most of glaciers. Bacterial community across glaciers became similar with size increase, and whence smallest size fractions contain more unique genera in each glacier. Multivariate analysis suggests that phosphate, which is significantly higher in one glacier (Scarlet Heart Glacier), is primary associated with bacterial beta diversity. Correlation coefficients between abundance of major genera and nutrients largely changed with granule size, suggesting that nutrients susceptibility to genera changes with growth process of granule (e.g. P. priestleyi was affected by nitrate in early growth stage).

Microbial colonization of glacial ice surfaces incurs feedbacks which affect the melting rate of the ice surface. Ecosystems formed as microbe-mineral aggregates termed cryoconite locally reduce ice surface albedo and represent foci of biodiversity and biogeochemical cycling. Consequently, greater understanding the ecological processes in the formation of functional cryoconite ecosystems upon glacier surfaces is sought. Here, we present the first bacterial biogeography of an ice cap, evaluating the respective roles of dispersal, environmental and biotic filtration occurring at local scales in the assembly of cryoconite microbiota. 16S rRNA gene amplicon semiconductor sequencing of cryoconite colonizing a Svalbard ice cap coupled with digital elevation modelling of physical parameters reveals the bacterial community is dominated by a ubiquitous core of generalist taxa, with evidence for a moderate pairwise distance-decay relationship. While geographic position and melt season duration are prominent among environmental predictors of community structure, the core population of taxa appears highly influential in structuring the bacterial community. Taxon co-occurrence network analysis reveals a highly modular community structured by positive interactions with bottleneck taxa, predominantly Actinobacteria affiliated to isolates from soil humus. In contrast, the filamentous cyanobacterial taxon (assigned to Leptolyngbya/Phormidesmis pristleyi) which dominates the community and binds together granular cryoconite are poorly connected to other taxa. While our study targeted one ice cap, the prominent role of generalist core taxa with close environmental relatives across the global cryosphere indicate discrete roles for cosmopolitan Actinobacteria and Cyanobacteria as respective keystone taxa and ecosystem engineers of cryoconite ecosystems colonizing ice caps.

Glaciers and ice sheets are considered a biome with unique organism assemblages. Tardigrada (water bears) are micrometazoans that play the function of apex consumers on glaciers. Cryoconite samples with the dark-pigmented tardigrade Cryoconicus gen. nov. kaczmareki sp. nov. were collected from four locations on glaciers in China and Kyrgyzstan. The erection of the new genus is based on a unique combination of morphological traits as well as on phylogenetic analyses. The analysis of COI sequences in the new species revealed high genetic differentiation with 9 haplotypes shared among 13 sequenced individuals from three sequenced populations. There was no apparent geographic structure in COI haplotype diversity, which might indicate effective dispersal abilities of the new species. A recovery of numerous live individuals from a sample that was frozen for 11 years suggests high survival rates in the natural environment. The ability to withstand low temperatures, combined with dark pigmentation that is hypothesised to protect from intense UV radiation, could explain how the new taxon is able to dwell in an extreme glacial habitat. We also found that a rare mountain tardigrade Ramazzottius cataphractus (Maucci, 1974) is morphologically similar to the new species, therefore we propose to transfer it to the new genus. Our study indicates that glacier invertebrate fauna is still poorly known and requires intense research.

Cryoconites are microbial aggregates commonly found on glacier surfaces where they tend to take spherical, granular forms. While it has been postulated that the microbes in cryoconite granules play an important role in glacier ecosystems, information on their community structure is still limited, and their functions remain unclear. Here, we present evidence for the occurrence of nitrogen cycling in cryoconite granules on a glacier in Central Asia. We detected marker genes for nitrogen fixation, nitrification and denitrification in cryoconite granules by digital polymerase chain reaction (PCR), while digital reverse transcription PCR analysis revealed that only marker genes for nitrification and denitrification were abundantly transcribed. Analysis of isotope ratios also indicated the occurrence of nitrification; nitrate in the meltwater on the glacier surface was of biological origin, while nitrate in the snow was of atmospheric origin. The predominant nitrifiers on this glacier belonged to the order Nitrosomonadales, as suggested by amoA sequences and 16S ribosomal RNA pyrosequencing analysis. Our results suggest that the intense carbon and nitrogen cycles by nitrifiers, denitrifiers and cyanobacteria support abundant and active microbes on the Asian glacier.

The mineralogical composition of cryoconite on a glacial surface was investigated on six glaciers in northwest Greenland (Qaanaaq Qaqortaq Tugto Bowdoin Sun and Scarlet Heart. The X‑ray diffraction analysis showed that the cryoconites mainly contained seven silicate minerals: hornblende quartz potassium feldspar plagioclase illite kaolinite and chlorite. Semi‑quantitative mineralogical analysis of the silicate mineral composition on the Qaanaaq Glacier showed little variation among the samples collected from five different elevations. This indicates that the minerals on the glacier were probably dominated by dust from a unique source which is recently transported from local sediments including soil and moraine. On the other hand the mineral composition varied significantly among the glaciers. Based on the clay mineral content the glaciers could be classified into three groups. Type A: high clay mineral content composed of illite and kaolinite found on Qaanaaq Qaqortaq and Tugto; Type B: high clay mineral content composed of only kaolinite found on Sun; and Type C: lower clay mineral content composed of only kaolinite found on Bowdoin and Scarlet Heart. The geographical distribution of the types of glaciers did not correspond with the geology in this area indicating that the mineralogical composition is not determined just by the geologi‑ cal conditions around glaciers but also by other factors such as the redistribution of sediments by glacial fluvial or coastal processes.

Spatial variations in impurities (cryoconite) on the glacier surface were investigated on Qaanaaq Ice Cap and Tugto Glacier in the northwest Greenland in the melting season of 2012. Abundance of impurities ranged from 0.36 to 119 g m -2 (dry weight, mean: 18.8 g m -2 ) on bare ice and from 0.01 to 8.7 g m -2

In order to better understand the source of minerals on the dark-colored ice, located in the Greenland ice sheet ablation zone, we analyzed the Sr and Nd isotopic ratios of minerals in cryoconite, which were collected from glaciers in northwest and southwest Greenland. We focused on the following: (i) comparison of the isotopes of minerals in cyroconite with those in sediments from local and distant areas, (ii) regional variations in western Greenland, and (iii) spatial variations across an individual a glacier. The mineral components of the cryoconite showed variable Sr and Nd isotopic ratios (87Sr/86Sr: 0.711335 to 0.742406, εNd (0): −33.1 to −22.9), which corresponded to those of the englacial dust and moraine on and around the glaciers but were significantly different from those of the distant deserts that have been considered to be primary sources of mineral dust on the Greenland Ice Sheet. This suggests that the minerals within the cryoconites were mainly derived from local sediments, rather than from distant areas. The Sr ratios in the northwestern region were significantly higher than those in the southwestern region. This is probably due to geological differences in the source areas, such as the surrounding glaciers in each region. The isotopic ratios further varied spatially within a glacier (Qaanaaq and Kangerlussuaq areas), indicating that the silicate minerals on the glaciers were derived not from a single source but from multiple sources, such as englacial dust and wind-blown minerals from the moraine surrounding the glaciers.

Cryoconite granules are aggregations of microorganisms with mineral particles that form on glacier surfaces. To understand the processes by which the granules develop, this study focused on the altitudinal distribution of the granules and photosynthetic microorganisms on the glacier, bacterial community variation with granules size and environmental factors affecting the growth of the granules. Size-sorted cryoconite granules collected from five different sites on Qaanaaq Glacier were analyzed. C and N contents were significantly higher in large (diameter greater than 250 μm) granules than in smaller (diameter 30–249 μm) granules. Bacterial community structures, based on 16S rRNA gene amplicon sequencing, were different between the smaller and larger granules. The filamentous cyanobacterium Phormidesmis priestleyi was the dominant bacterial species in larger granules. Multivariate analysis suggests that the abundance of mineral particles on the glacier surface is the main factor controlling growth of these cyanobacteria. These results show that the supply of mineral particles on the glacier enhances granule development, that P. priestleyi is likely the key species for primary production and the formation of the granules, and that the bacterial community in the granules changes over the course of the granule development.

Cryoconite is granular sediment found on glacier surfaces comprising both mineral and biological material. Despite long having been recognised as an important glaciological and biological phenomenon cryoconite remains relatively poorly understood. Here, we appraise the literature on cryoconite for the first time, with the aim of synthesising and evaluating current knowledge to direct future investigations. We review the properties of cryoconite, the environments in which it is found, the biology and biogeochemistry of cryoconite, and its interactions with climate and anthropogenic pollutants. We generally focus upon cryoconite in the Arctic in summer, with Antarctic and lower latitude settings examined individually. We then compare the current state-of-the-science with that at the turn of the twentieth century, and suggest directions for future research including specific recommendations for studies at a range of spatial scales and a framework for integrating these into a more holistic understanding of cryoconite and its role in the cryosphere.

Cryoconite granules are dark-colored spherical aggregates of organic and inorganic material on glacier ice, and are commonly observed on glaciers the world over. The structure of cryoconite granules on Ürümqi glacier No. 1, Tien Shan, China, was analyzed. Granules were distributed over the entire ice surface of the ablation area, and ranged in size from 0.26 to 3.5 mm (mean 1.1 mm). The granule surface was densely covered with filamentous cyanobacteria. Microscopy of a thin section revealed various inner structures. Most granules had concentric layers of dense organic matter, which are probably derived from annual growth of the granules by the activity of cyanobacteria. The number of layers averaged 3.5 and ranged up to 7, which is likely to indicate their mean and maximum growth ages, respectively. Some granules contained two or more subgranules, showing that small granules had combined and enlarged. Such structures suggest that granule formation was mainly due to the activity of filamentous cyanobacteria, and that the granules repeatedly grew and disintegrated over a cycle of several years on the glacier.

Spectral reflectances in the visible to near-infrared wavelength range (350–1050 nm) were measured on the surface of Gulkana Glacier, Alaska Range, Alaska, USA, at six different elevations from May to September 2001. The measurements showed variable spectral reflectances on the glacier surface, and the spectra changed over time and elevation. The spectra in May were those of a typical wet snow surface: the reflectance ranged approximately from 0.4 to 0.8, and generally decreased as the wavelength increased. As snowmelt proceeded, the reflectance gradually fell, and specific absorptions appeared in the spectrum at wavelength ranges of 400–600 and 670–680 nm, corresponding to a red snow surface due to snow algal bloom. As the ice surface was exposed, the spectrum finally shifted to a lower (0.1–0.4) and flat reflectance curve, typical of an ice surface contaminated with dust. Analyses of the dust on the glacier surface showed that its quantity and characteristics changed spatially and seasonally and were correlated with spectral albedo. The results showed that the temporal and spatial variations in spectral reflectance on this glacier are due not only to physical properties of the glacier surface, but also to biogenic materials on the surface (e.g red-colored snow algal cells and cryoconite).

Optical characteristics of the cryoconite collected from nine glaciers in the Himalaya, Tibet and the Arctic (Canada and Svalbard) were analyzed. The spectral light reflectance (visible region) of the cryoconite on the six glaciers in the Arctic and the Himalayawas generally low, indicating high light absorbency (dark coloration) of the cryoconite. In contrast, the spectral reflectances of the cryoconite on the three glaciers in Tibet were significantly higher than on the other glaciers. There was no significant difference in the spectral reflectance of mineral particles contained in the cryoconite between the Tibetan and the other glaciers, indicating that the difference in the albedo of the cryoconites is not due to the mineral particles, but due to organic matter contained in the cryoconite. Chemical analysis of the organic matter in the cryoconites revealed that the light absorbency of cryoconites is due to the amount of humic substances, which are dark-colored organic substances, the residue of bacterial decomposition of organic matter. The cryoconite of the three glaciers in Tibet contained significantly smaller amounts of humic substances than that of the other glaciers, probably due to different biological or chemical conditions. Results show that the formation of the humic substances in the cryoconite affects its optical characteristics, and possibly affects the surface albedo of the glaciers.

Glacier environments harbor cold-tolerant bacteria (Amato et al., 2007; Christner et al., 2003; Segawa et al., 2005). However, a variety of bacteria such as soil and enteric bacteria have been detected from these highly oligotrophic and psychrophilic environments (Segawa et al., 2005; Sheridan et al., 2003). The origin of these bacteria remains unknown. These bacteria may have been supplied at least in part from animals by means such as bird droppings (Sjolund et al., 2008). There are other plausible transmissions, such as local and global atmospheric circulation. In the case of the global atmospheric circulation, a glacier environment may receive bacteria originating from hu-man industrial activities, although most glaciers are located at a distance from such sites of human activi-ties. Emergence of antibiotic resistant bacteria has been considered as a human impact on the environment (Cabello, 2006; Chee-Sanford et al., 2001; Goni-Urriza et al., 2000). The prevalence of antibiotic resistant genes in the glacier environment can be used as an indicator of human impact on the global environment. In this experiment, the prevalence of antibiotic resis-tant genes in glacier ice samples taken from various locations in the northern hemisphere (Alaska, Central Asia) and Antarctica were evaluated by real time PCR array approach using a Taqman probe system. We collected samples of the glacial ice (1 cm in depth) or cryoconite hole samples (Table 1) using a sterilized stainless steel scoop or sterilized syringe, re-spectively. The samples were stored in 50-ml sterile plastic conical tubes (Iwaki, Tokyo). All samples were kept frozen and transported to the Tokyo Institute of Technology, Japan or National Institute of Polar Re-search, Japan. Bacterial DNA was extracted as follows within a Class 100 Clean bench (Sanyo, Tokyo, Model MCV-131BNS) at the National Institute of Polar Research (Tokyo). Ice samples were melted and a portion (1‒

Cryoconite is a dark-coloured surface dust deposited on glaciers that consists of wind-blown mineral particles, as well as organic matter derived from microbes living on glaciers. In this paper, we analyse the Sr and Nd isotopic ratios of four mineral fractions (i.e., the saline, carbonate, phosphate, and silicate mineral fractions), as well as the organic fraction, of cryoconite samples obtained from six Asian glaciers (the Altai, Pamir, Tien Shan, Qillian Shan, and Himalayan regions), and discuss their geographical variations in terms of the geological origins of the mineral particles and the biogeochemical processes on the glaciers. The silicate mineral fraction showed lower Sr and higher Nd ratios for the glaciers located to the north (Altai, 87Sr/86Sr: 0.713?490?0.715?284, ?Nd(0): ?6.4 to ?5.6), while higher Sr and lower Nd ratios for the glaciers located to the south (Himalayas, 87Sr/86Sr: 0.740?121?0.742?088, ?Nd(0): ?16.4 to ?15.7); the ratios were similar to those of desert sand, loess, and river sediments in the respective regions of the glaciers. This result suggests that the silicate minerals within the cryoconites were derived from different sources depending on the geographical locations of the glaciers. The isotopic ratios of the saline, carbonate, and phosphate mineral fractions were distinct from those of the silicate fraction, and were similar to those of evaporites and apatite deposits from the Asian deserts, but also varied geographically, indicating that they are likely to reflect their geological origin. The Sr isotopic ratios of the organic fraction were similar to those of the saline and carbonate fractions from glaciers in the central area (Tien Shan and Qillian Shan), but were higher than those of the saline and carbonate fractions, and lower than the phosphate mineral fraction, in the northern and southern areas. The ratios of organic fraction may be determined from the mixing ratio of calcium sources incorporated by microbes on the glaciers.

The worldwide distribution of microinvertebrates on glaciers, the coldest biome, is poorly known. Owing to their tolerance to hostile conditions, small size and dispersal abilities, nematodes, tardigrades and rotifers are considered cosmopolitan and together inhabit various ecosystems. In this study, we investigated their global distribution in cryoconite holes – a type of freshwater reservoir forming directly in the glacial ice that creates biodiversity hotspots on glaciers. We analysed cryoconite samples (using classical microscopic observations and environmental DNA metabarcoding) from 42 glaciers located around the world (the Arctic, Subarctic, Scandinavia, the Alps, the Caucasus, Siberia, Central Asia, Africa, South America and Antarctica), as well as using literature data. Samples from Antarctic, Karakoram and the Alps were analysed using next-generation sequencing (NGS) and classical observations under microscopes, while all other samples were analysed by microscope alone. Three general outcomes were found: (1) tardigrades and rotifers represented the most common invertebrates in cryoconite holes; (2) tardigrades and rotifers often coexisted together, with one or the other dominating, but the dominant taxon varied by region or by glacier; (3) nematodes – the most abundant, hyperdiverse and widespread metazoans on Earth, including in environments surrounding and seeding glacial surfaces – were consistently absent from cryoconite holes. Despite the general similarity of environmental conditions in cryoconite holes, the distribution of tardigrades and rotifers differed among glaciers, but not in any predictable way, suggesting that their distribution mostly depended on the random dispersal, extreme changes of supraglacial zone or competition. Although nematodes have been found in supraglacial habitats, cryoconite hole environments seem not to provide the necessary conditions for their growth and reproduction. Lack of physiological adaptations to permanently low temperatures (~0°C) and competition for different food resources in the cryoconite hole environment may explain the absence of nematodes in cryoconite holes.

Dark-colored material (cryoconite) covering Himalayan glaciers has been reported to greatly accelerate glacier-melting by reducing surface albedo. Structure, formation, and the darkening process of the cryoconite on a Himalayan glacier were analyzed. The cryoconite was revealed to be a stromatolite-like algal mat, a product of microbial activity on the glacier. The granular algal mat contains filamentous blue-green algae (cyanobacteria) and bacteria, and grows on the ice by trapping mineral and organic particles. This structure seems to enable high algal production in nutrient poor glacial meltwater by gathering and keeping nutrient rich particles inside. The dark coloration of the mats promotes melt-hole formation on the ice (cryoconite holes), providing a semistagnant aquatic habitat for various algae and animals in the glacier. Optical and chemical analyses of the cryoconite strongly suggests that their high light- absorbency (dark coloration) is mainly due to dark-colored humic substances, residues from bacterial decomposition of the algal products and other organic matter. Our results strongly suggest that biological activity on the glacier substantially affects the albedo of the glacier surface. The structure of the algal mat seems to be important in the glacier ecosystem and biological process affecting glacier albedo.