In four samples of mosses and mosses mixed with lichens collected in the Peruvian region of Cusco, 344 tardigrades, 78 free-laid eggs and six simplexes were found. In total, nine species were identified: Cornechiniscus lobatus, Echiniscus dariae, E. ollantaytamboensis, Isohypsibius condorcanquii sp. nov., Macrobiotus pisacensis sp. nov., Milnesium krzysztofi, Minibiotus intermedius, Paramacrobiotus intii sp. nov. and Pseudechiniscus ramazzottii ramazzottii. Isohypsibius condorcanquii sp. nov. is most similar to I. baldii, but differs mainly by the absence of ventral sculpture, the presence of the oral cavity armature, a different macroplacoid length sequence and a different shape of macroplacoids. The new species also differs from other congeners by a different dorsal sculpture, the absence of cuticular bars under the claws and the absence of eyes. Macrobiotus pisacensis sp. nov. differs from the most similar M. ariekammensis and M. kirghizicus by a different oral cavity armature, the presence of cuticular pores, details of egg morphology and some morphometric characters of both animals and eggs. Paramacrobiotus intii sp. nov. differs from most similar species of the genus by a different type of the oral cavity armature, details of egg morphology and some morphometric characters of both animals and eggs. In addition, we briefly discuss the tardigrade fauna of Peru, and propose a simple and economic system of describing relative lengths of pharyngeal macroplacoids. The system is especially useful in interspecific comparisons and differential diagnoses.

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.

Despite the large number of studies on glaciers, knowledge regarding biota in cryoconite holes is limited. Cryophilic animals are often neglected in ecological studies on glacial habitats, but are important for the functioning of these environments. Owing to climate change and the melting of polar ice, cryophilic fauna could be threatened in the near future. We provide the first comprehensive survey of invertebrates inhabiting the cryoconite holes of Alpine, Antarctic, Arctic, Himalayan and Patagonian glaciers. At present, the list of taxa is rather short and includes five phyla (Rotifera, Annelida, Tardigrada, Nematoda and Arthropoda). Owing to generally poor knowledge of the fauna of cryoconite holes, there could be more than the 25 currently known species. In addition, we present the geographical distributions of the taxa and discuss the diversity of invertebrates living in cryoconite holes and make predictions regarding the faunal diversity of these habitats.

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.

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 ...

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.