It is fundamental to understand the development of Zygnematophycean (Streptophyte) micro-algal blooms within Greenland Ice Sheet (GrIS) supraglacial environments, given their potential to significantly impact both physical (melt) and chemical (carbon and nutrient cycling) surface characteristics. Here, we report on a space-for-time assessment of a GrIS ice algal bloom, achieved by sampling an ∼85 km transect spanning the south-western GrIS bare ice zone during the 2016 ablation season. Cell abundances ranged from 0 to 1.6 × 104 cells ml−1, with algal biomass demonstrated to increase in surface ice with time since snow line retreat (R2 = 0.73, P < 0.05). A suite of light harvesting and photo-protective pigments were quantified across transects (chlorophylls, carotenoids and phenols) and shown to increase in concert with algal biomass. Ice algal communities drove net autotrophy of surface ice, with maximal rates of net production averaging 0.52 ± 0.04 mg C l−1 d−1, and a total accumulation of 1.306 Gg C (15.82 ± 8.14 kg C km−2) predicted for the 2016 ablation season across an 8.24 × 104 km2 region of the GrIS. By advancing our understanding of ice algal bloom development, this study marks an important step toward projecting bloom occurrence and impacts into the future.

Blooms of Zygnematophycean "glacier algae" lower the bare ice albedo of the Greenland Ice Sheet (GrIS), amplifying summer energy absorption at the ice surface and enhancing meltwater runoff from the largest cryospheric contributor to contemporary sea-level rise. Here, we provide a step change in current understanding of algal-driven ice sheet darkening through quantification of the photophysiological mechanisms that allow glacier algae to thrive on and darken the bare ice surface. Significant secondary phenolic pigmentation (11 times the cellular content of chlorophyll a) enables glacier algae to tolerate extreme irradiance (up to ∼4,000 μmol photons·m −2 ·s −1 ) while simultaneously repurposing captured ultraviolet and short-wave radiation for melt generation. Total cellular energy absorption is increased 50-fold by phenolic pigmentation, while glacier algal chloroplasts positioned beneath shading pigments remain low-light-adapted (E k ∼46 μmol photons·m −2 ·s −1 ) and dependent upon typical nonphotochemical quenching mechanisms for photoregulation. On the GrIS, glacier algae direct only ∼1 to 2.4% of incident energy to photochemistry versus 48 to 65% to ice surface melting, contributing an additional ∼1.86 cm water equivalent surface melt per day in patches of high algal abundance (∼10 4 cells·mL −1 ). At the regional scale, surface darkening is driven by the direct and indirect impacts of glacier algae on ice albedo, with a significant negative relationship between broadband albedo (Moderate Resolution Imaging Spectroradiometer [MODIS]) and glacier algal biomass (R 2 = 0.75, n = 149), indicating that up to 75% of the variability in albedo across the southwestern GrIS may be attributable to the presence of glacier algae.Greenland Ice Sheet | glacier algae | photophysiology | melt | cryosphere

Abstract. Glaciers and ice sheets host abundant and dynamic communities of microorganisms on the ice surface (supraglacial environments). Recently, it has been shown that Streptophyte glacier algae blooming on the surface ice of the south-western coast of the Greenland Ice Sheet are a significant contributor to the 15-year marked decrease in albedo. Currently, little is known about the constraints, such as nutrient availability, on this large-scale algal bloom. In this study, we investigate the relative abundances of dissolved inorganic and dissolved organic macronutrients (N and P) in these darkening surface ice environments. Three distinct ice surfaces, with low, medium and high visible impurity loadings, supraglacial stream water and cryoconite hole water, were sampled. Our results show a clear dominance of the organic phase in all ice surface samples containing low, medium and high visible impurity loadings, with 93 % of the total dissolved nitrogen and 67 % of the total dissolved phosphorus in the organic phase. Mean concentrations in low, medium and high visible impurity surface ice environments are 0.91, 0.62 and 1.0 µM for dissolved inorganic nitrogen (DIN), 5.1, 11 and 14 µM for dissolved organic nitrogen (DON), 0.03, 0.07 and 0.05 µM for dissolved inorganic phosphorus (DIP) and 0.10, 0.15 and 0.12 µM for dissolved organic phosphorus (DOP), respectively. DON concentrations in all three surface ice samples are significantly higher than DON concentrations in supraglacial streams and cryoconite hole water (0 and 0.7 µM, respectively). DOP concentrations are higher in all three surface ice samples compared to supraglacial streams and cryoconite hole water (0.07 µM for both). Dissolved organic carbon (DOC) concentrations increase with the amount of visible impurities present (low: 83 µM, medium: 173 µM and high: 242 µM) and are elevated compared to supraglacial streams and cryoconite hole water (30 and 50 µM, respectively). We speculate that the architecture of the weathering crust, which impacts on water flow paths and storage in the melting surface ice and/or the production of extracellular polymeric substances (EPS), containing both N and P in conjunction with C, is responsible for the temporary retention of DON and DOP in the melting surface ice. The unusual presence of measurable DIP and DIN, principally as NH4+, in the melting surface ice environments suggests that factors other than macronutrient limitation are controlling the extent and magnitude of the glacier algae.

Current research into bacterial dynamics on the Greenland Ice Sheet (GrIS) is biased toward cryoconite holes, despite this habitat covering less than 8% of the ablation (melt) zone surface. In contrast, the expansive surface ice, which supports wide-spread Streptophyte micro-algal blooms thought to enhance surface melt, has been relatively neglected. This study aims to understand variability in bacterial abundance and production across an ablation season on the GrIS, in relation to micro-algal bloom dynamics. Bacterial abundance reached 3.3 ± 0.3 × 10 5 cells ml −1 in surface ice and was significantly linearly related to algal abundances during the middle and late ablation periods ( R 2 = 0.62, p < 0.05; R 2 = 0.78, p < 0.001). Bacterial production (BP) of 0.03–0.6 μg C L −1 h −1 was observed in surface ice and increased in concert with glacier algal abundances, indicating that heterotrophic bacteria consume algal-derived dissolved organic carbon. However, BP remained at least 28 times lower than net primary production, indicating inefficient carbon cycling by heterotrophic bacteria and net accumulation of carbon in surface ice throughout the ablation season. Across the supraglacial environment, cryoconite sediment BP was at least four times greater than surface ice, confirming that cryoconite holes are the true “hot spots” of heterotrophic bacterial activity.

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.