The polar regions in a 2°C warmer world.

Over the past decade, the Arctic has warmed by 0.75°C, far outpacing the global average, while Antarctic temperatures have remained comparatively stable. As Earth approaches 2°C warming, the Arctic and Antarctic may reach 4°C and 2°C mean annual warming, and 7°C and 3°C winter warming, respectively. Expected consequences of increased Arctic warming include ongoing loss of land and sea ice, threats to wildlife and traditional human livelihoods, increased methane emissions, and extreme weather at lower latitudes. With low biodiversity, Antarctic ecosystems may be vulnerable to state shifts and species invasions. Land ice loss in both regions will contribute substantially to global sea level rise, with up to 3 m rise possible if certain thresholds are crossed. Mitigation efforts can slow or reduce warming, but without them northern high latitude warming may accelerate in the next two to four decades. International cooperation will be crucial to foreseeing and adapting to expected changes.

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.

Primary productivity as a control over soil microbial diversity along environmental gradients in a polar desert ecosystem.

Primary production is the fundamental source of energy to foodwebs and ecosystems, and is thus an important constraint on soil communities. This coupling is particularly evident in polar terrestrial ecosystems where biological diversity and activity is tightly constrained by edaphic gradients of productivity (e.g., soil moisture, organic carbon availability) and geochemical severity (e.g., pH, electrical conductivity). In the McMurdo Dry Valleys of Antarctica, environmental gradients determine numerous properties of soil communities and yet relatively few estimates of gross or net primary productivity (GPP, NPP) exist for this region. Here we describe a survey utilizing pulse amplitude modulation (PAM) fluorometry to estimate rates of GPP across a broad environmental gradient along with belowground microbial diversity and decomposition. PAM estimates of GPP ranged from an average of 0.27 µmol O2/m2/s in the most arid soils to an average of 6.97 µmol O2/m2/s in the most productive soils, the latter equivalent to 217 g C/m2/y in annual NPP assuming a 60 day growing season. A diversity index of four carbon-acquiring enzyme activities also increased with soil productivity, suggesting that the diversity of organic substrates in mesic environments may be an additional driver of microbial diversity. Overall, soil productivity was a stronger predictor of microbial diversity and enzymatic activity than any estimate of geochemical severity. These results highlight the fundamental role of environmental gradients to control community diversity and the dynamics of ecosystem-scale carbon pools in arid systems.

Microbial Community Responses to Increased Water and Organic Matter in the Arid Soils of the McMurdo Dry Valleys, Antarctica.

The soils of the McMurdo Dry Valleys, Antarctica are an extreme polar desert, inhabited exclusively by microscopic taxa. This region is on the threshold of anticipated climate change, with glacial melt, permafrost thaw, and the melting of massive buried ice increasing liquid water availability and mobilizing soil nutrients. Experimental water and organic matter (OM) amendments were applied to investigate how these climate change effects may impact the soil communities. To identify active taxa and their functions, total community RNA transcripts were sequenced and annotated, and amended soils were compared with unamended control soils using differential abundance and expression analyses. Overall, taxonomic diversity declined with amendments of water and OM. The domain Bacteria increased with both amendments while Eukaryota declined from 38% of all taxa in control soils to 8 and 11% in water and OM amended soils, respectively. Among bacterial phyla, Actinobacteria (59%) dominated water-amended soils and Firmicutes (45%) dominated OM amended soils. Three bacterial phyla (Actinobacteria, Proteobacteria, and Firmicutes) were primarily responsible for the observed positive functional responses, while eukaryotic taxa experienced the majority (27 of 34) of significant transcript losses. These results indicated that as climate changes in this region, a replacement of endemic taxa adapted to dry, oligotrophic conditions by generalist, copiotrophic taxa is likely.

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.

Niche and metabolic principles explain patterns of diversity and distribution: theory and a case study with soil bacterial communities.

The causes of biodiversity patterns are controversial and elusive due to complex environmental variation, covarying changes in communities, and lack of baseline and null theories to differentiate straightforward causes from more complex mechanisms. To address these limitations, we developed general diversity theory integrating metabolic principles with niche-based community assembly. We evaluated this theory by investigating patterns in the diversity and distribution of soil bacteria taxa across four orders of magnitude variation in spatial scale on an Antarctic mountainside in low complexity, highly oligotrophic soils. Our theory predicts that lower temperatures should reduce taxon niche widths along environmental gradients due to decreasing growth rates, and the changing niche widths should lead to contrasting α- and ß-diversity patterns. In accord with the predictions, α-diversity, niche widths and occupancies decreased while ß-diversity increased with increasing elevation and decreasing temperature. The theory also successfully predicts a hump-shaped relationship between α-diversity and pH and a negative relationship between α-diversity and salinity. Thus, a few simple principles explained systematic microbial diversity variation along multiple gradients. Such general theory can be used to disentangle baseline effects from more complex effects of temperature and other variables on biodiversity patterns in a variety of ecosystems and organisms.

Characterization of growing bacterial populations in McMurdo Dry Valley soils through stable isotope probing with (18) O-water.

Soil microbial communities of the McMurdo Dry Valleys, Antarctica (MDV) contain representatives from at least fourteen bacterial phyla. However, given low rates of microbial activity, it is unclear whether this richness represents functioning rather than dormant members of the community. We used stable isotope probing (SIP) with (18) O-water to determine if microbial populations grow in MDV soils. Changes in the microbial community were characterized in soils amended with H2 (18) O and H2 (18) O-organic matter. Sequencing the 16S rRNA genes of the heavy and light fractions of the bacterial community DNA shows that DNA of microbial populations was labeled with (18) O-water, indicating these micro-organisms grew in the MDV soils. Significant differences existed in the community composition of the heavy and light fractions of the H2 (18) O and H2 (18) O-organic matter amended samples (Anosim P < 0.05 of weighted Unifrac distance). Control samples and the light DNA fraction of the H2 (18) O amended samples were dominated by representatives of the phyla Deinococcus-Thermus, Proteobacteria, Planctomyces, Gemmatimonadetes, Actinobacteria and Acidobacteria, whereas Proteobacteria were more prevalent in the heavy DNA fractions from the H2 (18) O-water and the H2 (18) O-water-organic matter treatments. Our results indicate that SIP with H2 (18) O can be used to distinguish active bacterial populations even in this low organic matter environment.

Soil microbial responses to increased moisture and organic resources along a salinity gradient in a polar desert.

Microbial communities in extreme environments often have low diversity and specialized physiologies suggesting a limited resistance to change. The McMurdo Dry Valleys (MDV) are a microbially dominated, extreme ecosystem currently undergoing climate change-induced disturbances, including the melting of massive buried ice, cutting through of permafrost by streams, and warming events. These processes are increasing moisture across the landscape, altering conditions for soil communities by mobilizing nutrients and salts and stimulating autotrophic carbon inputs to soils. The goal of this study was to determine the effects of resource addition (water/organic matter) on the composition and function of microbial communities in the MDV along a natural salinity gradient representing an additional gradient of stress in an already extreme environment. Soil respiration and the activity of carbon-acquiring extracellular enzymes increased significantly (P < 0.05) with the addition of resources at the low- and moderate-salinity sites but not the high-salinity site. The bacterial community composition was altered, with an increase in Proteobacteria and Firmicutes with water and organic matter additions at the low- and moderate-salinity sites and a near dominance of Firmicutes at the high-salinity site. Principal coordinate analyses of all samples using a phylogenetically informed distance matrix (UniFrac) demonstrated discrete clustering among sites (analysis of similarity [ANOSIM], P < 0.05 and R > 0.40) and among most treatments within sites. The results from this experimental work suggest that microbial communities in this environment will undergo rapid change in response to the altered resources resulting from climate change impacts occurring in this region.

Bacterial community composition of divergent soil habitats in a polar desert.

Edaphic factors such as pH, organic matter, and salinity are often the most significant drivers of diversity patterns in soil bacterial communities. Desert ecosystems in particular are model locations for examining such relationships as food web complexity is low and the soil environment is biogeochemically heterogeneous. Here, we present the findings from a 16S rRNA gene sequencing approach used to observe the differences in diversity and community composition among three divergent soil habitats of the McMurdo Dry Valleys, Antarctica. Results show that alpha diversity is significantly lowered in high pH soils, which contain higher proportions of the phyla Acidobacteria and Actinobacteria, while mesic soils with higher soil organic carbon (and ammonium) content contain high proportions of Nitrospira, a nitrite-oxidizing bacteria. Taxonomic community resolution also had a significant impact on our conclusions, as pH was the primary predictor of phylum-level diversity, while moisture was the most significant predictor of diversity at the genus level. Predictive power also increased with increasing taxonomic resolution, suggesting a potential increase in niche-based drivers of bacterial community composition at such levels.

Factors Controlling Soil Microbial Biomass and Bacterial Diversity and Community Composition in a Cold Desert Ecosystem: Role of Geographic Scale.

Understanding controls over the distribution of soil bacteria is a fundamental step toward describing soil ecosystems, understanding their functional capabilities, and predicting their responses to environmental change. This study investigated the controls on the biomass, species richness, and community structure and composition of soil bacterial communities in the McMurdo Dry Valleys, Antarctica, at local and regional scales. The goals of the study were to describe the relationships between abiotic characteristics and soil bacteria in this unique, microbially dominated environment, and to test the scale dependence of these relationships in a low complexity ecosystem. Samples were collected from dry mineral soils associated with snow patches, which are a significant source of water in this desert environment, at six sites located in the major basins of the Taylor and Wright Valleys. Samples were analyzed for a suite of characteristics including soil moisture, pH, electrical conductivity, soil organic matter, major nutrients and ions, microbial biomass, 16 S rRNA gene richness, and bacterial community structure and composition. Snow patches created local biogeochemical gradients while inter-basin comparisons encompassed landscape scale gradients enabling comparisons of microbial controls at two distinct spatial scales. At the organic carbon rich, mesic, low elevation sites Acidobacteria and Actinobacteria were prevalent, while Firmicutes and Proteobacteria were dominant at the high elevation, low moisture and biomass sites. Microbial parameters were significantly related with soil water content and edaphic characteristics including soil pH, organic matter, and sulfate. However, the magnitude and even the direction of these relationships varied across basins and the application of mixed effects models revealed evidence of significant contextual effects at local and regional scales. The results highlight the importance of the geographic scale of sampling when determining the controls on soil microbial community characteristics.

How does subsurface characterization affect simulations of hyporheic exchange?

We investigated the role of increasingly well-constrained geologic structures in the subsurface (i.e., subsurface architecture) in predicting streambed flux and hyporheic residence time distribution (RTD) for a headwater stream. Five subsurface realizations with increasingly resolved lithological boundaries were simulated in which model geometries were based on increasing information about flow and transport using soil and geologic maps, surface observations, probing to depth to refusal, seismic refraction, electrical resistivity (ER) imaging of subsurface architecture, and time-lapse ER imaging during a solute tracer study. Particle tracking was used to generate RTDs for each model run. We demonstrate how improved characterization of complex lithological boundaries and calibration of porosity and hydraulic conductivity affect model prediction of hyporheic flow and transport. Models using hydraulic conductivity calibrated using transient ER data yield estimates of streambed flux that are three orders of magnitude larger than uncalibrated models using estimated values for hydraulic conductivity based on values published for nearby hillslopes (10(-4) vs. 10(-7) m(2)/s, respectively). Median residence times for uncalibrated and calibrated models are 10(3) and 10(0) h, respectively. Increasingly well-resolved subsurface architectures yield wider hyporheic RTDs, indicative of more complex hyporheic flowpath networks and potentially important to biogeochemical cycling. The use of ER imaging to monitor solute tracers informs subsurface structure not apparent from other techniques, and helps to define transport properties of the subsurface (i.e., hydraulic conductivity). Results of this study demonstrate the value of geophysical measurements to more realistically simulate flow and transport along hyporheic flowpaths.

Bacterial community structure along moisture gradients in the parafluvial sediments of two ephemeral desert streams.

Microorganisms inhabiting stream sediments mediate biogeochemical processes of importance to both aquatic and terrestrial ecosystems. In deserts, the lateral margins of ephemeral stream channels (parafluvial sediments) are dried and rewetted, creating periodically wet conditions that typically enhance microbial activity. However, the influence of water content on microbial community composition and diversity in desert stream sediments is unclear. We sampled stream margins along gradients of wet to dry sediments, measuring geochemistry and bacterial 16S rRNA gene composition, at streams in both a cold (McMurdo Dry Valleys, Antarctica) and hot (Chihuahuan Desert, New Mexico, USA) desert. Across the gradients, sediment water content spanned a wide range (1.6-37.9% w/w), and conductivity was highly variable (12.3-1,380 µS cm(-2)). Bacterial diversity (at 97% sequence similarity) was high and variable, but did not differ significantly between the hot and cold desert and was not correlated with sediment water content. Instead, conductivity was most strongly related to diversity. Water content was strongly related to bacterial 16S rRNA gene community composition, though samples were distributed in wet and dry clusters rather than as assemblages shifting along a gradient. Phylogenetic analyses showed that many taxa from wet sediments at the hot and cold desert site were related to, respectively, halotolerant Gammaproteobacteria, and one family within the Sphingobacteriales (Bacteroidetes), while dry sediments at both sites contained a high proportion of taxa related to the Acidobacteria. These results suggest that bacterial diversity and composition in desert stream sediments is more strongly affected by hydrology and conductivity than temperature.

Moving beyond the banks: hyporheic restoration is fundamental to restoring ecological services and functions of streams.

Stream restoration needs to consider the hyporheic zone just as much as the surface and benthic regions.