Ice-lidded cryoconite holes on glaciers in the Taylor Valley, Antarctica, provide a unique system of natural mesocosms for studying community structure and assembly. We used high-throughput DNA sequencing to characterize both microbial eukaryotic communities and bacterial communities within cryoconite holes across three glaciers to study similarities in their spatial patterns. We expected that the alpha (phylogenetic diversity) and beta (pairwise community dissimilarity) diversity patterns of eukaryotes in cryoconite holes would be related to those of bacteria, and that they would be related to the biogeochemical gradient within the Taylor Valley. We found that eukaryotic alpha and beta diversity were strongly related to those of bacteria across scales ranging from 140 m to 41 km apart. Alpha diversity of both was significantly related to position in the valley and surface area of the cryoconite hole, with pH also significantly correlated with the eukaryotic diversity. Beta diversity for both bacteria and eukaryotes was significantly related to position in the valley, with bacterial beta diversity also related to nitrate. These results are consistent with transport of sediments onto glaciers occurring primarily at local scales relative to the size of the valley, thus creating feedbacks in local chemistry and diversity.

Although cryoconite holes, sediment-filled melt holes on glacier surfaces, appear small and homogenous, their microbial inhabitants may be spatially partitioned. This partitioning could be particularly important for maintaining biodiversity in holes that remain isolated for many years, such as in Antarctica. We hypothesized that cryoconite holes with greater species richness and biomass should exhibit greater partitioning between the sediments and water, promoting greater biodiversity through spatial niche partitioning. We tested this hypothesis by sampling frozen cryoconite holes along a gradient of biomass and biodiversity in the Taylor Valley, Antarctica, where ice-lidded cryoconite holes are a ubiquitous feature of glaciers. We extracted DNA and chlorophyll a from the sediments and water of these samples to describe biodiversity and quantify proxies for biomass. Contrary to our expectation, we found that cryoconite holes with greater richness and biomass showed less partitioning of phylotypes by the sediments versus the water, perhaps indicating that the probability of sediment microbes being mixed into the water is higher from richer sediments. Another explanation may be that organisms from the water were compressed by freezing down to the sediment layer, leaving primarily relic DNA of dead cells to be detected higher in the frozen water. Further evidence of this explanation is that the dominant sequences unique to water closely matched organisms that do not live in cryoconite holes or the Dry Valleys (e.g., vertebrates); so this cryptic biodiversity could represent unknown microbial animals or DNA from atmospheric deposition of dead biomass in the otherwise low-biomass water. Although we cannot rule out spatial niche partitioning occurring at finer scales or in melted cryoconite holes, we found no evidence of partitioning between the sediments and water in frozen holes. Future work should include more sampling of cryoconite holes at a finer spatial scale, and characterizing the communities of the sediments and water when cryoconite holes are melted and active.

Here we describe recent breakthroughs in our understanding of microbial life in dry volcanic tephra ("soil") that covers much of the surface area of the highest elevation volcanoes on Earth. Dry tephra above 6000 m.a.s.l. is perhaps the best Earth analog for the surface of Mars because these "soils" are acidic, extremely oligotrophic, exposed to a thin atmosphere, high UV fluxes, and extreme temperature fluctuations across the freezing point. The simple microbial communities found in these extreme sites have among the lowest alpha diversity of any known earthly ecosystem and contain bacteria and eukaryotes that are uniquely adapted to these extreme conditions. The most abundant eukaryotic organism across the highest elevation sites is a Naganishia species that is metabolically versatile, can withstand high levels of UV radiation and can grow at sub-zero temperatures, and during extreme diurnal freeze-thaw cycles (e.g. - 10 to + 30 °C). The most abundant bacterial phylotype at the highest dry sites sampled (6330 m.a.s.l. on Volcán Llullaillaco) belongs to the enigmatic B12-WMSP1 clade which is related to the Ktedonobacter/Thermosporothrix clade that includes versatile organisms with the largest known bacterial genomes. Close relatives of B12-WMSP1 are also found in fumarolic soils on Volcán Socompa and in oligotrophic, fumarolic caves on Mt. Erebus in Antarctica. In contrast to the extremely low diversity of dry tephra, fumaroles found at over 6000 m.a.s.l. on Volcán Socompa support very diverse microbial communities with alpha diversity levels rivalling those of low elevation temperate soils. Overall, the high-elevation biome of the Atacama region provides perhaps the best "natural experiment" in which to study microbial life in both its most extreme setting (dry tephra) and in one of its least extreme settings (fumarolic soils).

Viral metagenomics has expanded our knowledge of the ecology of uncultured viruses, within both environmental (e.g., terrestrial and aquatic) and host-associated (e.g., plants and animals, including humans) contexts. Here, we emphasize the implementation of an ecological framework in viral metagenomic studies to address questions in virology rarely considered ecological, which can change our perception of viruses and how they interact with their surroundings. An ecological framework explicitly considers diverse variants of viruses in populations that make up communities of interacting viruses, with ecosystem-level effects. It provides a structure for the study of the diversity, distributions, dynamics, and interactions of viruses with one another, hosts, and the ecosystem, including interactions with abiotic factors. An ecological framework in viral metagenomics stands poised to broadly expand our knowledge in basic and applied virology. We highlight specific fundamental research needs to capitalize on its potential and advance the field. Expected final online publication date for the Annual Review of Virology, Volume 8 is September 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Predator avoidance behavior, in which prey limit foraging activities in the presence of predation threats, affects the dynamics of many ecological communities. Despite the growing theoretical appreciation of the role predation plays in coexistence, predator avoidance behavior has yet to be incorporated into the theory in a general way. We introduce adaptive avoidance behavior to a consumerresource model with three trophic levels to ask whether the ability of prey-the middle trophic level-to avoid predators alters their ability to coexist. We determine the characteristics of cases in which predator avoidance behavior changes prey coexistence or the order of competitive dominance. The mechanism underlying such changes is the weakening of apparent competition relative to resource competition in determining niche overlap, even with resource intake costs. Avoidance behavior thus generally promotes coexistence if prey partition resources but not predators, whereas it undermines coexistence if prey partition predators but not resources. For any given case, the changes in the average fitness difference between two species resulting from avoidance behavior interact with changes in niche overlap to determine coexistence. These results connect the substantial body of theoretical work on avoidance behavior and population dynamics with the body of theory on competitive coexistence.

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.

Cryoconite holes are holes in a glacier's surface caused by sediment melting into the glacier. These holes are self-contained ecosystems that include abundant bacterial life within their sediment and liquid water, and have recently gained the attention of microbial ecologists looking to use cryoconite holes as "natural microcosms" to study microbial community assembly. Here, we explore the idea that cryoconite holes can be viewed as "islands," in the same sense that an island in the ocean is an area of habitat surrounded by a barrier to entry. In the case of a classic oceanic island, the ocean is a barrier between islands, but in the case of cryoconite holes, the ocean is comprised of impermeable solid ice. We test two hypotheses, born out of island biogeographic theory, that can be readily applied to cryoconite hole bacteria. First, we ask to what extent the size of a cryoconite hole is related to the amount of bacterial diversity found within it. Second, we ask to what extent cryoconite holes exhibit distance decay of similarity, meaning that geographically close holes are expected to harbor similar bacterial communities, and distant holes are expected to harbor more different bacterial communities. To test the island size hypothesis, we measured the sizes of cryoconite holes on three glaciers in Antarctica's Taylor Valley and used DNA sequencing to measure diversity of bacterial communities within them. We found that for two of these glaciers, there is a strong relationship between hole size and bacterial phylogenetic diversity, supporting the idea that cryoconite holes on those glaciers are "islands." The high biomass dispersing to the third glacier we measured could explain the lack of size-diversity relationship, remaining consistent with island biogeography. To test the distance decay of similarity hypothesis, we used DNA sequence data from several previous studies of cryoconite hole bacteria from across the world. Combined with our Taylor Valley data, those data showed that cryoconite holes have strong spatial structuring at scales of one to several hundred kilometers, also supporting the idea that these dirty holes on glaciers are really islands in the cryosphere.

Antarctic cryoconite holes, or small melt-holes in the surfaces of glaciers, create habitable oases for isolated microbial communities with tightly linked microbial population structures. Viruses may influence the dynamics of polar microbial communities, but the viromes of the Antarctic cryoconite holes have yet to be characterized. We characterize single-stranded DNA (ssDNA) viruses from three cryoconite holes in the Taylor Valley, Antarctica, using metagenomics. Half of the assembled metagenomes cluster with those in the viral family Microviridae (n = 7), and the rest with unclassified circular replication associated protein (Rep)-encoding single-stranded (CRESS) DNA viruses (n = 7). An additional 18 virus-like circular molecules encoding either a Rep, a capsid protein gene, or other unidentified but viral-like open reading frames were identified. The samples from which the genomes were identified show a strong gradient in microbial diversity and abundances, and the number of viral genomes detected in each sample mirror that gradient. Additionally, one of the CRESS genomes assembled here shares ~90% genome-wide pairwise identity with a virus identified from a freshwater pond on the McMurdo Ice Shelf (Antarctica). Otherwise, the similarity of these viruses to those previously identified is relatively low. Together, these patterns are consistent with the presence of a unique regional virome present in fresh water host populations of the McMurdo Dry Valley region.