Changes in air temperature, but not in precipitation, determine long-term trends in water chemistry of high mountain lakes of the Alps with and without rock glacier influence.

Climate change has strongly affected lakes around the world, but the relative effects of warmer air temperatures and changing precipitation on the water chemistry of alpine systems are not well understood. Here we tested the effect of monthly and seasonal climate on the water chemistry of six high mountain lakes located in the Alps. From 1982 to 2020, water samples were collected annually from different depths during the autumn mixing. We observed a simultaneous increase in electrical conductivity, ionic content, and pH with air temperature. In lakes with rock glacier influence, the increase in conductivity, ionic content, and especially in sulfate was even more pronounced, but accompanied by a strong decrease in pH. These differences are attributed to the direct influence of acidic meltwater from active rock glaciers in catchments with acidic bedrock. We then analyzed changes in lake chemistry, taking into account seasonal trends in air temperature and precipitation, using redundancy analysis. Temperature increase significantly affected water chemistry in five of the six lakes, especially at times of ice breakup. Increasing warming explained 17% to 32% of the changes in electrical conductivity, alkalinity, pH, major ions, and nitrogen. In contrast, precipitation had little effect on the changes of those parameters. Nevertheless, late spring snowfall and high snowfall in early fall, which result in prolonged ice cover, had a dampening effect on the impact of climate warming on lake chemistry. Our results confirm that climate warming remains a major driver of chemical changes in alpine lakes, but provide new evidence that late spring temperatures are the most important triggers.

Factors shaping community patterns of protists and bacteria on a European scale.

Factors shaping community patterns of microorganisms are controversially discussed. Physical and chemical factors certainly limit the survival of individual taxa and maintenance of diversity. In recent years, a contribution of geographic distance and dispersal barriers to distribution patterns of protists and bacteria has been demonstrated. Organismic interactions such as competition, predation and mutualism further modify community structure and maintenance of distinct taxa. Here, we address the relative importance of these different factors in shaping protists and bacterial communities on a European scale using high-throughput sequencing data obtained from lentic freshwater ecosystems. We show that community patterns of protists are similar to those of bacteria. Our results indicate that cross-domain organismic factors are important variables with a higher influence on protists as compared with bacteria. Abiotic physical and chemical factors also contributed significantly to community patterns. The contribution of these latter factors was higher for bacteria, which may reflect a stronger biogeochemical coupling. The contribution of geographical distance was similar for both microbial groups.

The Little Ice Age signature in a 700-year high-resolution chironomid record of summer temperatures in the Central Eastern Alps.

Despite the fact that the Little Ice Age (LIA) is well documented for the European Alps, substantial uncertainties concerning the regional spatio-temporal patterns of temperature changes associated with the LIA still exist, especially for their eastern sector. Here we present a high-resolution (4-10 years) 700-year long mean July air temperature reconstruction based on subfossil chironomid assemblages from a remote lake in the Austrian Eastern Alps to gain further insights into the LIA climatic deterioration in the region. The record provides evidence for a prolonged period of predominantly cooler conditions during AD 1530-1920, broadly equivalent to the climatically defined LIA in Europe. The main LIA phase appears to have consisted of two cold time intervals divided by slightly warmer episodes in the second half of the 1600s. The most severe cooling occurred during the eighteenth century. The LIA temperature minimum about 1.5 °C below the long-term mean recorded in the mid-1780 s coincides with the strongest volcanic signal found in the Greenland ice cores over the past 700 years and may be, at least in part, a manifestation of cooling that followed the long-lasting AD 1783-1784 Laki eruption. A continuous warming trend is evident since ca AD 1890 (1.1 °C in 120 years). The chironomid-inferred temperatures show a clear correlation with the instrumental data and reveal a close agreement with paleotemperature evidence from regional high-elevation tree-ring chronologies. A considerable amount of the variability in the temperature record may be linked to changes in the North Atlantic Oscillation.

Rock glaciers in crystalline catchments: Hidden permafrost-related threats to alpine headwater lakes.

A global warming-induced transition from glacial to periglacial processes has been identified in mountainous regions around the world. Degrading permafrost in pristine periglacial environments can produce acid rock drainage (ARD) and cause severe ecological damage in areas underlain by sulfide-bearing bedrock. Limnological and paleolimnological approaches were used to assess and compare ARDs generated by rock glaciers, a typical landform of the mountain permafrost domain, and their effects on alpine headwater lakes with similar morphometric features and underlying bedrock geology, but characterized by different intensities of frost action in their catchments during the year. We argue that ARD and its effects on lakes are more severe in the alpine periglacial belt with mean annual air temperatures (MAAT) between -2°C and +3°C, where groundwater persists in the liquid phase for most of the year, in contrast to ARD in the periglacial belt where frost action dominates (MAAT < -2°C). The findings clearly suggest that the ambient air temperature is an important factor affecting the ARD production in alpine periglacial environments. Applying the paleoecological analysis of morphological abnormalities in chironomids through the past millennium, we tested and rejected the hypothesis that unfavorable conditions for aquatic life in the ARD-stressed lakes are largely related to the temperature increase over recent decades, responsible for the enhanced release of ARD contaminants. Our results indicate that the ARDs generated in the catchments are of a long-lasting nature and the frequency of chironomid morphological deformities was significantly higher during the Little Ice Age (LIA) than during pre- or post-LIA periods, suggesting that lower water temperatures may increase the adverse impacts of ARD on aquatic invertebrates. This highlights that temperature-mediated modulations of the metabolism and life cycle of aquatic organisms should be considered when reconstructing long-term trends in the ecotoxicological state of lakes.

Trade-off between taxon diversity and functional diversity in European lake ecosystems.

Inferring ecosystem functioning and ecosystem services through inspections of the species inventory is a major aspect of ecological field studies. Ecosystem functions are often stable despite considerable species turnover. Using metatranscriptome analyses, we analyse a thus-far unparalleled freshwater data set which comprises 21 mainland European freshwater lakes from the Sierra Nevada (Spain) to the Carpathian Mountains (Romania) and from northern Germany to the Apennines (Italy) and covers an altitudinal range from 38 m above sea level (a.s.l) to 3110 m a.s.l. The dominant taxa were Chlorophyta and streptophytic algae, Ciliophora, Bacillariophyta and Chrysophyta. Metatranscriptomics provided insights into differences in community composition and into functional diversity via the relative share of taxa to the overall read abundance of distinct functional genes on the ecosystem level. The dominant metabolic pathways in terms of the fraction of expressed sequences in the cDNA libraries were affiliated with primary metabolism, specifically oxidative phosphorylation, photosynthesis and the TCA cycle. Our analyses indicate that community composition is a good first proxy for the analysis of ecosystem functions. However, differential gene regulation modifies the relative importance of taxa in distinct pathways. Whereas taxon composition varies considerably between lakes, the relative importance of distinct metabolic pathways is much more stable, indicating that ecosystem functioning is buffered against shifts in community composition through a functional redundancy of taxa.

Evaluating the performance of water purification in a vegetated groundwater recharge basin maintained by short-term pulsed infiltration events.

Infiltration of surface water constitutes an important pillar in artificial groundwater recharge. However, insufficient transformation of organic carbon and nutrients, as well as clogging of sediments often cause major problems. The attenuation efficiency of dissolved organic carbon (DOC), nutrients and pathogens versus the risk of bioclogging for intermittent recharge were studied in an infiltration basin covered with different kinds of macrovegetation. The quality and concentration of organic carbon, major nutrients, as well as bacterial biomass, activity and diversity in the surface water, the porewater, and the sediment matrix were monitored over one recharge period. Additionally, the numbers of viral particles and Escherichia coli were assessed. Our study showed a fast establishment of high microbial activity. DOC and nutrients have sustainably been reduced within 1.2 m of sediment passage. Numbers of E. coli, which were high in the topmost centimetres of sediment porewater, dropped below the detection limit. Reed cover was found to be advantageous over bushes and trees, since it supported higher microbial activities along with a good infiltration and purification performance. Short-term infiltration periods of several days followed by a break of similar time were found suitable for providing high recharge rates, and good water purification without the risk of bioclogging.

Prokaryotic community structure in deep bedrock aquifers of the Austrian Central Alps.

The bacterial and archaeal diversity of deep groundwater systems was investigated based on 16S rRNA-SSCP (single strand conformation polymorphism) fingerprints. The study site included five boreholes along the projected Brenner Base Tunnel in the central Alps of Tyrol, Austria. To obtain representative samples, packer-sealed fractures were sampled at specific depths between 105 and 780 m below surface. Sequence analysis of SSCP bands obtained from 13 samples showed that between 29 and 62 % of the phylotypes belonged to a variety of Proteobacteria including representatives of typical freshwater bacteria of the genera Acidovorax, Aquabacterium, and Sphingomonas. Bacteroidetes (especially Flavobacterium), Firmicutes (Acetobacterium), and candidate division OP3-related sequences were observed in the majority of the analysed groundwaters. On average, 14 % of the detected prokaryotic phylotypes were affiliated with Archaea, comprising the phyla Euryarchaeota, Crenarchaeota and Thaumarchaeota. Most of the archaeal sequences showed low similarities to known cultivated species, with exception of two sequences having 98 % similarity to Methanosaeta sp. A considerable number of thaumarchaeal sequences belonged to two groups related to Nitrososphaera and Nitrosopumilus phylotypes. An environmental clustering of the groundwater samples, based on the bacterial and archaeal phylogeny, revealed a clear distribution pattern of the samples (sites and depths) reflecting the hydrochemical characteristics and underlying geologies.

Rock glacier outflows may adversely affect lakes: lessons from the past and present of two neighboring water bodies in a crystalline-rock watershed.

Despite the fact that rock glaciers are one of the most common geomorphological expressions of mountain permafrost, the impacts of their solute fluxes on lakes still remain largely obscure. We examined water and sediment chemistry, and biota of two neighboring water bodies with and without a rock glacier in their catchments in the European Alps. Paleolimnological techniques were applied to track long-term temporal trends in the ecotoxicological state of the water bodies and to establish their baseline conditions. We show that the active rock glacier in the mineralized catchment of Lake Rasass (RAS) represents a potent source of acid rock drainage that results in enormous concentrations of metals in water, sediment, and biota of RAS. The incidence of morphological abnormalities in the RAS population of Pseudodiamesa nivosa, a chironomid midge, is as high as that recorded in chironomid populations inhabiting sites heavily contaminated by trace metals of anthropogenic origin. The incidence of morphological deformities in P. nivosa of ∼70% persisted in RAS during the last 2.5 millennia and was ∼40% in the early Holocene. The formation of RAS at the toe of the rock glacier most probably began at the onset of acidic drainage in the freshly deglaciated area. The present adverse conditions are not unprecedented in the lake's history and cannot be associated exclusively with enhanced thawing of the rock glacier in recent years.

Holocene temperature variations at a high-altitude site in the Eastern Alps: a chironomid record from Schwarzsee ob Sölden, Austria.

Few well-dated, quantitative Holocene temperature reconstructions exist from high-altitude sites in the Central Eastern Alps. Here, we present a chironomid-based quantitative reconstruction of mean July air temperatures (T(July)) throughout the Holocene for a remote high-mountain lake, Schwarzsee ob Sölden, situated above the treeline at 2796 m a.s.l. in the Austrian Alps. Applying a chironomid-temperature inference model developed from lakes of the Alpine region to a high-resolution chironomid record from the lake provides evidence for early Holocene (ca 10000-8600 cal yr BP) T(July) of up to 8.5 °C, i.e. >4 °C above the modern (1977-2006) mean July temperature. The reconstruction reveals the so-called '8.2-ka cold event' centered at ca 8250-8000 cal yr BP with temperatures ca 3 °C below the early-Holocene thermal maximum. Rather warm (ca 6 °C) and productive conditions prevailed during ca 7900-4500 cal yr BP. The chironomid record suggests a climate transition between ca 5200 and 4500 cal yr BP to cooler T(July). A distinct cooling trend is evident from ca 4500 until ca 2500 cal yr BP. Thereafter, the study site experienced its coldest conditions (around 4 °C or less) throughout the rest of the Holocene, with the exception of the warming trend during the late 20th century. Beside other factors, the Northern Hemisphere summer insolation seems to be the major driving force for the long-term trends in T(July) at high altitudes in the Eastern Alps. Due to the extreme location of the lake and the limited temperature range represented by the applied calibration data set, the chironomid-based temperature reconstruction fails to track phases of the late-Holocene climatic history with T(July) cooler than 4 °C. Further chironomid-based palaeoclimate model and down-core studies are required to address this problem, provide more realistic T(July) estimates from undisturbed high-altitude lakes in the Alps, and extract a reliable regional temperature signal.

Microorganisms in the atmosphere over Antarctica.

Antarctic microbial biodiversity is the result of a balance between evolution, extinction and colonization, and so it is not possible to gain a full understanding of the microbial biodiversity of a location, its biogeography, stability or evolutionary relationships without some understanding of the input of new biodiversity from the aerial environment. In addition, it is important to know whether the microorganisms already present are transient or resident - this is particularly true for the Antarctic environment, as selective pressures for survival in the air are similar to those that make microorganisms suitable for Antarctic colonization. The source of potential airborne colonists is widespread, as they may originate from plant surfaces, animals, water surfaces or soils and even from bacteria replicating within the clouds. On a global scale, transport of air masses from the well-mixed boundary layer to high-altitude sites has frequently been observed, particularly in the warm season, and these air masses contain microorganisms. Indeed, it has become evident that much of the microbial life within remote environments is transported by air currents. In this review, we examine the behaviour of microorganisms in the Antarctic aerial environment and the extent to which these microorganisms might influence Antarctic microbial biodiversity.

Distribution and diversity of autotrophic bacteria in groundwater systems based on the analysis of RubisCO genotypes.

A molecular approach, based on the detection of ribulose-1,5-bisphosphate carboxylase/oxygenase (RubisCO) large subunit genes, was applied to investigate the distribution and diversity of autotrophic bacteria in groundwater systems. DNA extracts from 48 sampling stations, including a variety of pristine and polluted, shallow and deep-subsurface groundwater samples obtained from Germany and Austria, served as a template for the PCR amplification of form I (cbbL) and form II (cbbM) large subunit RubisCO genes. The majority of the samples (>80%) contained two different forms of RubisCO. In 17 samples, all three forms of RubisCO were identified. PCR products from four selected groundwater habitats containing all three forms of RubisCO were used to construct clone libraries. Based on restriction fragment length polymorphism (RFLP) analysis, 109 RubisCO-clone-inserts were subjected to sequencing and phylogenetic analysis. With the exception of a form IA RubisCO sequence cluster obtained from deep subsurface samples, which was identical to the RubisCO genes described for Ralstonia metallidurans CH34, most sequences were distantly related to a variety of RubisCO species in chemolithoautotrophic Proteobacteria. Several sequences occurred in isolated lineages. These findings suggest that autotrophic bacteria with the capability to assimilate CO2 via the Calvin Cycle pathway are widespread inhabitants of groundwater systems.

Spatio-temporal niche separation of planktonic Betaproteobacteria in an oligo-mesotrophic lake.

We investigated the diversity of planktonic Betaproteobacteria and the seasonal population changes of betaproteobacterial taxa in an oligo-mesotrophic lake (Piburger See, Austria). Focus was put on the vertical distribution of the investigated populations and on differences between their respective cell fractions with apparent amino acid incorporation. On average, 66% of betaproteobacterial cells and 73% of their diversity could be attributed to four clades within three lineages that were further analysed by fluorescence in situ hybridization. The numbers of bacteria from the R-BT subclade of the beta I lineage and from the PnecB subgroup of the beta II lineage were rather constant throughout the water column. In contrast, members of another subgroup of beta II (PnecC) and bacteria related to Methylophilus (beta IV) were particularly numerous in the oxygen-depleted zone. In general, only moderate seasonal changes in abundance were observed in the upper water layers, whereas there was a clear relationship between decreasing oxygen levels and the rise of bacteria from the PnecC and beta IV clades in deeper strata. On average, almost 80% of beta I bacteria, but < 15% of cells from the beta IV clade, showed amino acid incorporation. Our results suggest that the studied populations occupy distinct vertical and ecophysiological niches in Piburger See.

Unexpected response of high Alpine Lake waters to climate warming.

Over the past two decades, we have observed a substantial rise in solute concentration at two remote high mountain lakes in catchments of metamorphic rocks in the European Alps. At Rasass See, the electrical conductivity increased 18-fold. Unexpectedly high nickel concentrations at Rasass See, which exceeded the limit in drinking water by more than 1 order of magnitude, cannot be related to catchment geology. We attribute these changes in lake water quality to solute release from the ice of an active rock glacier in the catchment as a response to climate warming. Similar processes occurred at the higher elevation lake Schwarzsee ob Sölden, where electrical conductivity has risen 3-fold during the past two decades.

Factors governing the atmospheric deposition of polycyclic aromatic hydrocarbons to remote areas.

Polycyclic aromatic hydrocarbons (PAH) were measured in bulk atmospheric deposition collected in three remote areas of Europe during 1997-1998. Mean total PAH fluxes over a period of 18 months were 1560 +/- 750 and 1150 +/- 630 ng m(-2) mo(-1) in the Pyrenees and the Alps, respectively. In the Caledonian mountains (Scandinavia) the observed mean fluxes were 1900 +/- 940 ng m(-2) mo(-1) (6 month collection). Similar qualitative PAH compositions (p values <0.05) in the bulk atmospheric deposition have been observed between sites, which are dominated by the more volatile parent compounds. The main differences between lakes are related to the high molecular weight compounds. Atmospheric deposition of PAH to these remote sites appears to be independent of their concentrations in the atmosphere, which are similar between sites (in the range of 1.8-3.0 ng x m(-3)), being controlled mainly by particle deposition, followed by precipitation and air temperature. A multilevel regression model including these three variables accounted for 74% of the total variability in total PAH bulk deposition; however, the contribution of each variable in the model is compound and site-dependent. The deposition of high molecular weight PAH depends more on particle deposition and precipitation, whereas air temperature is the main factor controlling the deposition fluxes of the low molecular weight PAH.

Atmospherc deposition of organochlorine compounds to remote high mountain lakes of Europe.

Bulk deposition samples were taken near three mountain lakes located in the Pyrenees (Estany Redó), Alps (Gossenköllesee), and Caledonian Mountains (Ovre Neådalsvatn) for evaluation of the atmospheric deposition load of organochlorine compounds (OC), namely, polychlorobiphenyls (PCBs), hexachlorocyclohexanes (HCHs), hexachlorobenzene (HCB), and endosulfans, in the remote European high mountain areas. The compounds of present use in agriculture, namely, endosulfans and gamma-HCH, exhibit large differences in mean deposition fluxes between the three sites. They occur in large amounts in Estany Redó (340 and 430 ng m(-2) month(-1) for endosulfans and gamma-HCH, respectively), reflecting the impact of agricultural activities in southern Europe. This lake showed also the highest proportion of the more labile endosulfan isomers (alpha and beta = 82%) whereas only the most recalcitrant species, endosulfan sulfate, was found in Ovre Neådalsvatn. In contrast, the OC whose use is now banned exhibit a more uniform geographic distribution with deposition fluxes of 31-40, 30-100, and 1.4-15 ng m(-2) month(-1) for alpha-HCH, PCBs, and HCB. Both compounds of present and past use exhibit a clear seasonal pattern, with higher deposition in the warm periods, which is consistent with enhanced volatilization at higher temperatures. In the case of the agricultural pesticides it may also reflect higher use during application periods. The OC distributions in the atmospheric deposition of the three sites are rather uniform and highly enriched in compounds with volatilities larger than 0.0032 Pa. However, more than 90% of these compounds are not retained in the lake waters or sediments. Comparison of OC composition in atmospheric and sedimentary deposition evidences a selective trapping of the less volatile compounds. Trapping efficiencies increase at decreasing air temperatures of the lacustrine systems.

Improvement of ciliate identification and quantification: a new protocol for fluorescence in situ hybridization (FISH) in combination with silver stain techniques.

A new protocol for taxon specific probe based fluorescent in situ hybridization was developed for the identification and quantification of ciliates in microbial communities. Various fixatives and experimental parameters were evaluated and optimized with respect to cell permeability and morphological preservation. Optimum results were adaption by obatined of a modified fixation method using Bouin's solution. Furthermore, conventional staining procedures such as different Protargol stain techniques and a silver nitrate impregnation method were modified and can now be applied in combination with fluorescence in situ hybridization. The new protocol allows a rapid and reliable identification as well as quantification of ciliates based upon classical morphological aspects and rRNA based phylogenetic relationships performed in one experiment. Furthermore, a set of specific probes targeting different regions of the 18S rRNA was designed for Glaucoma scintillans Ehrenberg, 1830 and tested by applying this new approach of combining in situ cell hybridization with conventional staining techniques.