Strong Dispersal Limitation of Microbial Communities at Shackleton Glacier, Antarctica.

Microbial communities can be structured by both deterministic and stochastic processes, but the relative importance of these processes remains unknown. The ambiguity partly arises from an inability to disentangle soil microbial processes from confounding factors, such as aboveground plant communities or anthropogenic disturbance. In this study, we characterized the relative contributions of determinism and stochasticity to assembly processes of soil bacterial communities across a large environmental gradient of undisturbed Antarctic soils. We hypothesized that harsh soils would impose a strong environmental selection on microbial communities, whereas communities in benign soils would be structured largely by dispersal. Contrary to our expectations, dispersal was the dominant assembly mechanism across the entire soil environmental gradient, including benign environments. The microbial community composition reflects slowly changing soil conditions and dispersal limitation of isolated sites. Thus, stochastic processes, as opposed to deterministic, are primary drivers of soil ecosystem assembly across space at our study site. This is especially surprising given the strong environmental constraints on soil microorganisms in one of the harshest environments on the planet, suggesting that dispersal could be a driving force in microbial community assembly in soils worldwide. IMPORTANCE Because of their diversity and ubiquity, microbes provide an excellent means to tease apart how natural communities are structured. In general, ecologists believe that stochastic assembly processes, like random drift and dispersal, should dominate in benign environments while deterministic processes, like environmental filtering, should be prevalent in harsh environments. To help resolve this debate, we analyzed microbial community composition in pristine Antarctic soils devoid of human influence or plant communities for eons. Our results demonstrate that dispersal limitation is a surprisingly potent force of community limitation throughout all soil conditions. Thus, dispersal appears to be a driving force of microbial community assembly, even in the harshest of conditions.

Carbonate weathering, phosphate fertilizer, and hydrologic controls on dissolved uranium in rivers in the US Corn Belt: Disentangling seasonal geogenic- and fertilizer-derived sources.

Soil and bedrock weathering and phosphate (P) fertilizers may both contribute to the uranium (U) load of rivers in agricultural regions, but controls over their relative influence are not well known. This study investigates the U sources to rivers in Ohio, United States, part of the Eastern Corn Belt in the Mississippi River watershed. We present a regional picture of seasonal U sources to rivers based on four analyses: 1) a spatial analysis of legacy soil and water data, 2) new measurements of U and carbonate weathering products from rivers at 50 locations across the state collected seasonally over two years, 3) a weekly time series with additional 234U/238U (n = 5) and 87Sr/86Sr (n = 5) measurements from an agricultural river, and 4) a mass-balance approach to U addition to the landscape based on reported P fertilizer use. Uranium concentrations in surface waters collected statewide ranged 0.1-21 nM (n = 132), with significantly higher concentrations in the glaciated agricultural portion of the state (mean = 7.3 nM; n = 105) than the non-glaciated portion (mean = 2.0 nM; n = 24). Concentrations in the glaciated region were highest during the spring and summer and decreased during baseflow. In the time-series, concentrations were ~7 nM during baseflow and ~14 nM during intermediate seasonal discharge conditions, indicating a second more surficial endmember source of U in addition to bedrock weathering that is well correlated with other carbonate weathering products. Systematic increases in 87Sr/86Sr and decreases in 234U/238U with increasing discharge confirm a changing source of carbonate and U weathering and a third surficial endmember during high discharge events. Our mass balance approach and geochemical analysis suggest that elevated U concentrations are the result of carbonate weathering deep in the soil column during elevated seasonal flow. Further work on U dynamics in agricultural rivers is required to understand mechanism controlling seasonal changes in U concentrations and 234U/238U in downstream rivers and U flux.

Elevational Constraints on the Composition and Genomic Attributes of Microbial Communities in Antarctic Soils.

The inland soils found on the Antarctic continent represent one of the more challenging environments for microbial life on Earth. Nevertheless, Antarctic soils harbor unique bacterial and archaeal (prokaryotic) communities able to cope with extremely cold and dry conditions. These communities are not homogeneous, and the taxonomic composition and functional capabilities (genomic attributes) of these communities across environmental gradients remain largely undetermined. We analyzed the prokaryotic communities in soil samples collected from across the Shackleton Glacier region of Antarctica by coupling quantitative PCR, marker gene amplicon sequencing, and shotgun metagenomic sequencing. We found that elevation was the dominant factor explaining differences in the structures of the soil prokaryotic communities, with the drier and saltier soils found at higher elevations harboring less diverse communities and unique assemblages of cooccurring taxa. The higher-elevation soil communities also had lower maximum potential growth rates (as inferred from metagenome-based estimates of codon usage bias) and an overrepresentation of genes associated with trace gas metabolism. Together, these results highlight the utility of assessing community shifts across pronounced environmental gradients to improve our understanding of the microbial diversity found in Antarctic soils and the strategies used by soil microbes to persist at the limits of habitability. IMPORTANCE Antarctic soils represent an ideal system to study how environmental properties shape the taxonomic and functional diversity of microbial communities given the relatively low diversity of Antarctic soil microbial communities and the pronounced environmental gradients that occur across soils located in reasonable proximity to one another. Moreover, the challenging environmental conditions typical of most Antarctic soils present an opportunity to investigate the traits that allow soil microbes to persist in some of the most inhospitable habitats on Earth. We used cultivation-independent methods to study the bacterial and archaeal communities found in soil samples collected from across the Shackleton Glacier region of the Transantarctic Mountains. We show that those environmental characteristics associated with elevation have the greatest impact on the structure of these microbial communities, with the colder, drier, and saltier soils found at higher elevations sustaining less diverse communities that were distinct from those in more hospitable soils with respect to their composition, genomic attributes, and overall life-history strategies. Notably, the harsher conditions found in higher-elevation soils likely select for taxa with lower maximum potential growth rates and an increased reliance on trace gas metabolism to support growth.

Response of Antarctic soil fauna to climate-driven changes since the Last Glacial Maximum.

Understanding how terrestrial biotic communities have responded to glacial recession since the Last Glacial Maximum (LGM) can inform present and future responses of biota to climate change. In Antarctica, the Transantarctic Mountains (TAM) have experienced massive environmental changes associated with glacial retreat since the LGM, yet we have few clues as to how its soil invertebrate-dominated animal communities have responded. Here, we surveyed soil invertebrate fauna from above and below proposed LGM elevations along transects located at 12 features across the Shackleton Glacier region. Our transects captured gradients of surface ages possibly up to 4.5 million years and the soils have been free from human disturbance for their entire history. Our data support the hypothesis that soils exposed during the LGM are now less suitable habitats for invertebrates than those that have been exposed by deglaciation following the LGM. Our results show that faunal abundance, community composition, and diversity were all strongly affected by climate-driven changes since the LGM. Soils more recently exposed by the glacial recession (as indicated by distances from present ice surfaces) had higher faunal abundances and species richness than older exposed soils. Higher abundances of the dominant nematode Scottnema were found in older exposed soils, while Eudorylaimus, Plectus, tardigrades, and rotifers preferentially occurred in more recently exposed soils. Approximately 30% of the soils from which invertebrates could be extracted had only Scottnema, and these single-taxon communities occurred more frequently in soils exposed for longer periods of time. Our structural equation modeling of abiotic drivers highlighted soil salinity as a key mediator of Scottnema responses to soil exposure age. These changes in soil habitat suitability and biotic communities since the LGM indicate that Antarctic terrestrial biodiversity throughout the TAM will be highly altered by climate warming.

Enhanced trace element mobilization by Earth's ice sheets.

Trace elements sustain biological productivity, yet the significance of trace element mobilization and export in subglacial runoff from ice sheets is poorly constrained at present. Here, we present size-fractionated (0.02, 0.22, and 0.45 µm) concentrations of trace elements in subglacial waters from the Greenland Ice Sheet (GrIS) and the Antarctic Ice Sheet (AIS). Concentrations of immobile trace elements (e.g., Al, Fe, Ti) far exceed global riverine and open ocean mean values and highlight the importance of subglacial aluminosilicate mineral weathering and lack of retention of these species in sediments. Concentrations are higher from the AIS than the GrIS, highlighting the geochemical consequences of prolonged water residence times and hydrological isolation that characterize the former. The enrichment of trace elements (e.g., Co, Fe, Mn, and Zn) in subglacial meltwaters compared with seawater and typical riverine systems, together with the likely sensitivity to future ice sheet melting, suggests that their export in glacial runoff is likely to be important for biological productivity. For example, our dissolved Fe concentration (20,900 nM) and associated flux values (1.4 Gmol y-1) from AIS to the Fe-deplete Southern Ocean exceed most previous estimates by an order of magnitude. The ultimate fate of these micronutrients will depend on the reactivity of the dominant colloidal size fraction (likely controlled by nanoparticulate Al and Fe oxyhydroxide minerals) and estuarine processing. We contend that ice sheets create highly geochemically reactive particulates in subglacial environments, which play a key role in trace elemental cycles, with potentially important consequences for global carbon cycling.

Genetic diversity of soil invertebrates corroborates timing estimates for past collapses of the West Antarctic Ice Sheet.

During austral summer field seasons between 1999 and 2018, we sampled at 91 locations throughout southern Victoria Land and along the Transantarctic Mountains for six species of endemic microarthropods (Collembola), covering a latitudinal range from 76.0°S to 87.3°S. We assembled individual mitochondrial cytochrome c oxidase subunit 1 (COI) sequences (n = 866) and found high levels of sequence divergence at both small (<10 km) and large (>600 km) spatial scales for four of the six Collembola species. We applied molecular clock estimates and assessed genetic divergences relative to the timing of past glacial cycles, including collapses of the West Antarctic Ice Sheet (WAIS). We found that genetically distinct lineages within three species have likely been isolated for at least 5.54 My to 3.52 My, while the other three species diverged more recently (<2 My). We suggest that Collembola had greater dispersal opportunities under past warmer climates, via flotation along coastal margins. Similarly increased opportunities for dispersal may occur under contemporary climate warming scenarios, which could influence the genetic structure of extant populations. As Collembola are a living record of past landscape evolution within Antarctica, these findings provide biological evidence to support geological and glaciological estimates of historical WAIS dynamics over the last ca 5 My.

Uranium in Ohio, USA Surface Waters: Implications for a Fertilizer Source in Waters Draining Agricultural lands.

Synthetic fertilizer is a potential source of uranium to natural waters, yet evidence is lacking. We analyzed dissolved uranium concentrations in lakes, reservoirs, and rivers in Ohio, USA during the summer of 2017. All water bodies drain areas of extensive agriculture where phosphate-rich fertilizer is applied. Uranium concentrations ranged from 0.3 to 3.9 µg L-1, with the lowest concentrations observed in the most offshore Lake Erie samples. These results, especially when placed in the context of previous work on both surface and groundwater, suggest that dissolved uranium concentrations in this water emanating from agricultural lands are higher than background, and uranium should be categorized similarly to nitrate and phosphate in that it originates in part from fertilizer application.

Microbial diversity of an Antarctic subglacial community and high-resolution replicate sampling inform hydrological connectivity in a polar desert.

Antarctic subglacial environments host microbial ecosystems and are proving to be geochemically and biologically diverse. The Taylor Glacier, Antarctica, periodically expels iron-rich brine through a conduit sourced from a deep subglacial aquifer, creating a dramatic red surface feature known as Blood Falls. We used Illumina MiSeq sequencing to describe the core microbiome of this subglacial brine and identified previously undetected but abundant groups including the candidate bacterial phylum Atribacteria and archaeal phylum Pacearchaeota. Our work represents the first microbial characterization of samples collected from within a glacier using a melt probe, and the only Antarctic subglacial aquatic environment that, to date, has been sampled twice. A comparative analysis showed the brine community to be stable at the operational taxonomic unit level of 99% identity over a decade. Higher resolution sequencing enabled deconvolution of the microbiome of subglacial brine from mixtures of materials collected at the glacier surface. Diversity patterns between this brine and samples from the surrounding landscape provide insight into the hydrological connectivity of subglacial fluids to the surface polar desert environment. Understanding subice brines collected on the surfaces of thick ice covers has implications for analyses of expelled materials that may be sampled on icy extraterrestrial worlds.

Variations in Dissolved Nitrate, Chloride, and Sulfate in Precipitation, Reservoir, and Tap Waters, Columbus, Ohio.

Urban potable waters can be very susceptible to human activities that can impact water quality and, hence, public health. Columbus, Ohio, is currently the 14th largest city in the United States with an estimated population of ~860,000. Much of the urban population receives its water supply from a series of reservoirs located north of the city proper. These reservoirs are fed by river systems that drain either large agricultural lands, or rapidly growing suburban areas, or both. The agricultural activities introduce dissolved nitrate, and increased usage of de-icing salts on roads and highways within the drainage area introduce chloride into the river/reservoir systems. High nitrate in drinking water poses a potential health risk, particularly to infants, while high chloride, applied as halite, in drinking water can aid in the development of cardiovascular disease. In this work, we present a 19-month time series measuring nitrate, chloride, and sulfate in local precipitation, reservoir and household tap waters in order to better understand the relationship of the hydrologic residence time on the tap water chemistry, as well as to evaluate the anion concentrations. The highest chloride tap water concentration, 6.9 mM, occurred in early February 2011, while increases in nitrate occurred in both early summer and the middle of winter. In general, the anion concentrations in the precipitation are all equal to or lower than the reservoir waters. Similarly, the tap water had concentrations of chloride and sulfate higher than reservoir water, while nitrate was similar to reservoir water. Tap water had higher fluoride and sulfate concentrations, suggesting that they are added during the treatment of the reservoir water prior to residential distribution. These data clearly demonstrate the importance of watershed lands on the quality of water in the human distribution system.

Decadal ecosystem response to an anomalous melt season in a polar desert in Antarctica.

Amplified climate change in polar regions is significantly altering regional ecosystems, yet there are few long-term records documenting these responses. The McMurdo Dry Valleys (MDV) cold desert ecosystem is the largest ice-free area of Antarctica, comprising soils, glaciers, meltwater streams and permanently ice-covered lakes. Multi-decadal records indicate that the MDV exhibited a distinct ecosystem response to an uncharacteristic austral summer and ensuing climatic shift. A decadal summer cooling phase ended in 2002 with intense glacial melt ('flood year')-a step-change in water availability triggering distinct changes in the ecosystem. Before 2002, the ecosystem exhibited synchronous behaviour: declining stream flow, decreasing lake levels, thickening lake ice cover, decreasing primary production in lakes and streams, and diminishing soil secondary production. Since 2002, summer air temperatures and solar flux have been relatively consistent, leading to lake level rise, lake ice thinning and elevated stream flow. Biological responses varied; one stream cyanobacterial mat type immediately increased production, but another stream mat type, soil invertebrates and lake primary productivity responded asynchronously a few years after 2002. This ecosystem response to a climatic anomaly demonstrates differential biological community responses to substantial perturbations, and the mediation of biological responses to climate change by changes in physical ecosystem properties.

Hydrological Controls on Ecosystem Dynamics in Lake Fryxell, Antarctica.

The McMurdo Dry Valleys constitute the largest ice free area of Antarctica. The area is a polar desert with an annual precipitation of ∼ 3 cm water equivalent, but contains several lakes fed by glacial melt water streams that flow from four to twelve weeks of the year. Over the past ∼20 years, data have been collected on the lakes located in Taylor Valley, Antarctica as part of the McMurdo Dry Valley Long-Term Ecological Research program (MCM-LTER). This work aims to understand the impact of climate variations on the biological processes in all the ecosystem types within Taylor Valley, including the lakes. These lakes are stratified, closed-basin systems and are perennially covered with ice. Each lake contains a variety of planktonic and benthic algae that require nutrients for photosynthesis and growth. The work presented here focuses on Lake Fryxell, one of the three main lakes of Taylor Valley; it is fed by thirteen melt-water streams. We use a functional regression approach to link the physical, chemical, and biological processes within the stream-lake system to evaluate the input of water and nutrients on the biological processes in the lakes. The technique has been shown previously to provide important insights into these Antarctic lacustrine systems where data acquisition is not temporally coherent. We use data on primary production (PPR) and chlorophyll-A (CHL)from Lake Fryxell as well as discharge observations from two streams flowing into the lake. Our findings show an association between both PPR, CHL and stream input.

Antarctic subglacial lake exploration: first results and future plans.

After more than a decade of planning, three attempts were made in 2012-2013 to access, measure in situ properties and directly sample subglacial Antarctic lake environments. First, Russian scientists drilled into the top of Lake Vostok, allowing lake water to infiltrate, and freeze within, the lower part of the ice-core borehole, from which further coring would recover a frozen sample of surface lake water. Second, UK engineers tried unsuccessfully to deploy a clean-access hot-water drill, to sample the water column and sediments of subglacial Lake Ellsworth. Third, a US mission successfully drilled cleanly into subglacial Lake Whillans, a shallow hydraulically active lake at the coastal margin of West Antarctica, obtaining samples that would later be used to prove the existence of microbial life and active biogeochemical cycling beneath the ice sheet. This article summarizes the results of these programmes in terms of the scientific results obtained, the operational knowledge gained and the engineering challenges revealed, to collate what is known about Antarctic subglacial environments and how to explore them in future. While results from Lake Whillans testify to subglacial lakes as being viable biological habitats, the engineering challenges to explore deeper more isolated lakes where unique microorganisms and climate records may be found, as exemplified in the Lake Ellsworth and Vostok missions, are considerable. Through international cooperation, and by using equipment and knowledge of the existing subglacial lake exploration programmes, it is possible that such environments could be explored thoroughly, and at numerous sites, in the near future.

The fate of minor alkali elements in the chemical evolution of salt lakes.

Alkaline earth elements and alkali metals (Mg, Ca, Na and K) play an important role in the geochemical evolution of saline lakes as the final brine type is defined by the abundance of these elements. The role of major ions in brine evolution has been studied in great detail, but little has been done to investigate the behaviour of minor alkali elements in these systems despite their similar chemical affinities to the major cations. We have examined three major anionic brine types, chloride, sulphate, and bicarbonate-carbonate, in fifteen lakes in North America and Antarctica to determine the geochemical behaviour of lithium, rubidium, strontium, and barium. Lithium and rubidium are largely conservative in all water types, and their concentrations are the result of long-term solute input and concentration through evaporation and/or sublimation. Strontium and barium behaviours vary with anionic brine type. Strontium can be removed in sulphate and carbonate-rich lakes by the precipitation of carbonate minerals. Barium may be removed in chloride and sulphate brines by either the precipitation of barite and perhaps biological uptake.

Mercury deposition in a polar desert ecosystem.

Trace metals have received considerable attention in the recent decades due to their potential toxic nature. Glacial snow and ice have been used extensively to elucidate historical changes in the atmospheric composition of trace metals and other compounds. Mercury concentrations in Antarctic ice have described changes in atmospheric mercury deposition during the transition from the Last Glacial Maximum to the Holocene, however the record of modern mercury deposition in Antarctica is limited. Here we present a record of net mercury deposition to Antarctic snow over the past two decades. Over decadal periods, mercury is conserved in the snowpack and is dependent on a regional oceanic source. Annual to subannual mercury concentrations in snow are to some extent preserved and show covariance with marine aerosols as evidenced by calcium concentrations. Aeolian inputs from exposed rock and soil also play a critical role in depositing mercury to Antarctic snow. Such identifications along with previous data illustrate that mercury transport directly from the glaciers may account for 25-65% of the total mercury concentration in proglacial streams and the surface waters of perennially ice-covered lakes.

Effect of watershed parameters on mercury distribution in different environmental compartments in the Mobile Alabama River Basin, USA.

Total mercury (THg) and mono-methylmercury (MeHg) levels in water, sediment, and largemouth bass (LMB) (Micropterus salmoides) were investigated at 52 sites draining contrasting land use/land cover and habitat types within the Mobile Alabama River Basin (MARB). Aqueous THg was positively associated with iron-rich suspended particles and highest in catchments impacted by agriculture. Sediment THg was positively associated with sediment organic mater and iron content, with the highest levels observed in smaller catchments influenced by wetlands, followed by those impacted by agriculture or mixed forest, agriculture, and wetlands. The lowest sediment THg levels were observed in main river channels, except for reaches impacted by coal mining. Sediment MeHg levels were a positive function of sediment THg and organic matter and aqueous nutrient levels. The highest levels occurred in agricultural catchments and those impacted by elevated sulfate levels associated with coal mining. Aqueous MeHg concentrations in main river channels were as high as those in smaller catchments impacted by agriculture or wetlands, suggesting these areas were sources to rivers. Elevated Hg levels in some LMB were observed across all types of land use and land cover, but factors such as shallow water depth, larger wetland catchment surface area, low aqueous potassium levels, and higher Chl a concentrations were associated with higher Hg burdens, particularly in the Coastal Plain province. It is suggested that the observed large variability in LMB Hg burdens is linked to fish displacement by anglers, differences in food web structure, and sediment biogeochemistry, with surficial sediment iron oxides buffering the flux of MeHg from sediments to deeper water pelagic food webs.

Impact of land use and physicochemical settings on aqueous methylmercury levels in the Mobile-Alabama River System.

Mercury (Hg) concentrations above levels that could pose health risks have been measured recently in predatory fish from many aquatic systems in the southeastern region of the United States. Based on hypotheses derived from published experimental data on the aqueous geochemistry of Hg, we investigated the effect of certain natural and human-imposed conditions on in situ levels of methylmercury (MeHg) in the Mobile-Alabama River System (MARS). Water samples were collected from different types of environments, hypothesized to have contrasting levels of MeHg in the aqueous phase, and were analyzed for total-Hg (THg) and MeHg concentrations, as well as some key geochemical parameters. The results showed the following. i) Overall, total Hg concentrations in waters of the MARS are quite uniformly distributed and vary from 0.2 to 6 ng L(-1), suggesting that besides geological sources, atmospheric deposition is certainly the main source of Hg inputs in the studied system. ii) In locations with comparable THg levels, the Hg fraction present as MeHg was consistently higher in samples collected from the Coastal Plain portion of the MARS as compared to those from other geological provinces. iii) Our in situ observations confirmed conclusions derived from laboratory experiments, in that, MeHg abundance in aquatic systems correlates with sulfate (but only within a narrow range of concentrations); decreasing pH; and has no direct relationships with either nitrate or phosphate. iv) The investigation of Hg accumulation in biota at a single site showed that an aquatic system with low THg concentrations but a high MeHg:THg ratios, could have organisms with Hg content above safe levels. Therefore, potential health risks to fish eating populations can exist even when the aqueous phase does not show signs of significant Hg enrichment.

Antarctic climate cooling and terrestrial ecosystem response.

The average air temperature at the Earth's surface has increased by 0.06 degrees C per decade during the 20th century, and by 0.19 degrees C per decade from 1979 to 1998. Climate models generally predict amplified warming in polar regions, as observed in Antarctica's peninsula region over the second half of the 20th century. Although previous reports suggest slight recent continental warming, our spatial analysis of Antarctic meteorological data demonstrates a net cooling on the Antarctic continent between 1966 and 2000, particularly during summer and autumn. The McMurdo Dry Valleys have cooled by 0.7 degrees C per decade between 1986 and 2000, with similar pronounced seasonal trends. Summer cooling is particularly important to Antarctic terrestrial ecosystems that are poised at the interface of ice and water. Here we present data from the dry valleys representing evidence of rapid terrestrial ecosystem response to climate cooling in Antarctica, including decreased primary productivity of lakes (6-9% per year) and declining numbers of soil invertebrates (more than 10% per year). Continental Antarctic cooling, especially the seasonality of cooling, poses challenges to models of climate and ecosystem change.