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A sample of a sediment record contains diatom species that have grown in disparate habitats and eventually accumulated in a deep part of the lake. The original habitats may differ in substrate, depth location, and availability of resources. Identifying the species characteristic of each habitat should improve our ecological and environmental interpretation of the sediment record by distinguishing habitat specific responses. With this aim, we studied the benthic diatom communities of a deep oligotrophic lake across several habitats. The main source of variation in the diatom composition was the substrate type; particularly, sediment biofilms. Depth was the second factor. The thermocline defined a shift in diatom communities that also included changes in the dominant lifeforms. A third factor was the mesoscale heterogeneity (i.e., rock sides). Although most species were present in many habitats, characteristic species were identified for all the main habitats and used for an improved interpretation of the deep sediment record. Appropriate standardization showed increasing species richness and diversity from epilimnetic epilithic samples to hypolimnetic sediment samples. We estimate that more than 5000 valve counts are required for appropriate comparisons. Consequently, in sediment records with lower counts per sample, one has to amalgamate samples—losing temporal resolution—to achieve reliable analyses of diversity changes over time. Deep sediment samples are representative of the gamma-diversity of the lake diatom metacommunity, which result from the local alpha diversity of the habitats and the beta-diversity of the variability in composition among them. This double source of diversity has to be taken into account when using the sediment record for estimating lake biodiversity changes. On the other hand, we show that an estimation of the spatial (habitat) heterogeneity of a reconstructed environmental variable can be achieved using subsets of species characteristic of each habitat. We demonstrate the procedure by reconstructing the pH fluctuations during the last 200 years in several habitats from a single sediment record. The results are coherent with the expected differences between predominantly trophogenic or tropholithic habitats.

Este artículo contiene 9 páginas, 2 tablas, 2 figuras., During the last decades, atmospheric nitrogen loading in mountain ranges of the Northern Hemisphere has increased substantially, resulting in high nitrate concentrations in many lakes. Yet, how increased nitrogen has affected denitrification, a key process for nitrogen removal, is poorly understood. We measured actual and potential (nitrate and carbon amended) denitrification rates in sediments of several lake types and habitats in the Pyrenees during the ice-free season. Actual denitrification rates ranged from 0 to 9 μmol N2O m−2 h−1 (mean, 1.5 ± 1.6 SD), whereas potential rates were about 10-times higher. The highest actual rates occurred in warmer sediments with more nitrate available in the overlying water. Consequently, littoral habitats showed, on average, 3-fold higher rates than the deep zone. The highest denitrification potentials were found in more productive lakes located at relatively low altitude and small catchments, with warmer sediments, high relative abundance of denitrification nitrite reductase genes, and sulphate-rich waters. We conclude that increased nitrogen deposition has resulted in elevated denitrification rates, but not sufficiently to compensate for the atmospheric nitrogen loading in most of the highly oligotrophic lakes. However, there is potential for high rates, especially in the more productive lakes and landscape features largely govern this., The Spanish Government provided funds through the research grants: NitroPir (CGL2010-19737), Lacus (CGL2013-45348-P) and Transfer (CGL2016-80124-C2-1-P) and a predoctoral fellowship to CP-L (FPU12-00644).

The ratio between chrysophycean cysts and diatom valves (CD ratio) in lake sediments has been suggested as a useful indicator of changing trophic state conditions in oligotrophic lakes. Other environmental factors, however, may influence the CD ratio because chrysophycean cysts usually reflect conditions in the planktonic environment and diatoms reflect benthic conditions. We investigated the CD ratio in 76 mountain lakes in the Pyrenees to determine the environmental drivers that influence the ratio and assess its value for paleoenvironmental inference. The lakes surveyed included a broad range with respect to bedrock type, altitude and surface area, characteristics that cover much of the variability that can be found in cold, oligotrophic mountain lakes. Lake depth and Ca2+ concentration explain most of the variation in the CD ratio. Trophic state factors (e.g. total phosphorus, TP) play a secondary role. As a predictor, CD ratio performs primarily as a lake depth indicator. The predictive models can be improved if trophic state (i.e. TP) and chemical conditions (Ca2+) are known or can be estimated independently. Use of the CD ratio for inferring Ca2+ oscillations only makes sense in lakes with Ca2+

10 páginas, 3 figuras., Rate limitation due to encounters is fundamental to many ecological interactions. Since encounter rate governs reaction rates, and thus, dynamics of systems, it deserves systematic study. In classical population biology, ecological dynamics rely on the assumption of perfectly mixed interacting entities (e.g., individuals, populations, etc.) in a spaceless world. The so-called mean field assumption assumes that encounter rates are driven exclusively by changes in the density of the interacting entities and not on how they are distributed or move in space. Therefore, the mean field assumption does not give any insight into relevant spatiotemporal statistical properties produced by the trajectories of moving entities through space. In the present study, we develop spatially explicit simulations of random walking particles (i.e., L´evy walkers) to evaluate encounter rate constraints beyond the mean field assumption. We show that encounter rate fluctuations are driven not only by physical aspects such as the size or the velocity of the interacting particles, but also by different motion patterns. In particular, superdiffusion phenomena might be relevant at low densities and/or low spatial dimensionality. Finally, we discuss potential adaptive responses of living organisms that may allow individuals to control how they diffuse through space and/or the spatial dimensions employed in the exploration process., This study was supported by funds provided by DARPA (HR0011-05-1-0057) and the Andrew W. Mellon Foundation. FB’s work was also supported by a postdoctoral fellowhip from the Spanish Government (EX-2005- 1011) and the National Science Foundation (DEB-0083566).

Chrysophyte cysts were identified from the surface sediment of 105 mountain lakes in the Pyrenees (NE Spain), and their statistical relationship to water chemistry was examined using canonical correspondence analysis (CCA). The chemical parameters that explained significant and independent amounts of variability were alkalinity, pH, potassium, nitrate and magnesium. In a CCA using these parameters, the first canonical axis was related to a gradient of alkalinity and pH, which reflected the varying nature of the watershed bedrock in the Pyrenees, while the second axis was correlated with potassium (negatively) and nitrate (positively). The potential for environmental reconstructions of the five chemical parameters was further studied by: (i) analyzing the distribution of optima and tolerances calculated by weighted-averaging (WA); (ii) carrying out detrended canonical correspondence analysis (DCCA) with a single environmental variable; and (iii) examining the performance of WA-PLS transfer functions. Acceptable transfer functions were obtained for alkalinity, pH and nitrate. However, for potassium and magnesium the tolerance of cysts was too broad and the distribution of optima too skewed, respectively. The possibility of reconstructing nitrogen-related issues using chrysophyte cysts is particularly interesting because of the lack of direct chemical records of nitrogen compounds in sediments. Nitrate reconstructions using transfer functions may be complemented by a holistic reconstruction using partial CCA, where, after subtracting the effects of other chemicals, samples are ordered on a plain defined by potassium and nitrate. This ordination could show down-core trends in lake productivity and renewal time.

The ecosystem response of Lake Redo (Central Pyrenees) to fluctuations in seasonal air temperature during the last two centuries was investigated by comparison of reconstructed air temperatures with the sediment record. Fine slicing allowed a resolution of 3–6 years according to the 210Pb dating, although it was still difficult to easily investigate the response to air temperature forcing, since extreme fluctuations in temperature occur on interannual time-scales. However, the resolution was sufficient to show responses on decadal and century scales. An overall tendency to warming in mean annual temperature in the Central Pyrenees has been caused by summer and in particular by autumn increases. Many of the measured sediment variables apparently responded to these long term trends, but the significance of the relationships was highly conditioned by the structure of the data. The variables responding most on the finer time scales were the microfossils. For diatoms, chironomids and chrysophytes the main variability correlated to summer and to autumn temperatures. For two planktonic species, Fragilaria nanana and Cyclotella pseudostelligera, we found a link of their variability with temperature fluctuations in their growing months (September and October, respectively). This relationship appeared at a certain point during a general warming trend, indicating a threshold in the response. On the other hand, no significant changes in the dominant species could be linked to temperature, nor in any significant subgroup of the 180 diatom species present in the core. In contrast, for most chironomids (particularly Paratanytarsus austriacus, Heterotrissocladius marcidus and Micropsectra radialis) a negative relationship with summer temperature extended throughout the studied period. This response of the whole group gives chironomids a more robust role as indicators for recording temperature changes on long time-scales (e.g., through the Holocene) and for lake signal inter-comparison. Finally, our results indicated that, in all cases, there was a significant resilience to high frequency changes and hysteresis despite extreme fluctuations. Although we were dealing with organisms with one or many generations per year, their populations seemed to follow the decadal trends in air temperature.

Weather variation and climate fluctuations are the main sources of ecosystem variability in remote mountain lakes. Here we describe the main patterns of seasonal variability in the ecosystems of nine lakes in Europe, and discuss the implications for recording climatic features in their sediments. Despite the diversity in latitude and size, the lakes showed a number of common features. They were ice-covered between 5–9 months, and all but one were dimictic. This particular lake was long and shallow, and wind action episodically mixed the water column throughout the ice-free period. All lakes showed characteristic oxygen depletion during the ice-covered-period, which was greater in the most productive lakes. Two types of lakes were distinguished according to the number of production peaks during the ice-free season. Lakes with longer summer stratification tended to have two productive periods: one at the onset of stratification, and the other during the autumn overturn. Lakes with shorter stratification had a single peak during the ice-free period. All lakes presented deep chlorophyll maxima during summer stratification, and subsurface chlorophyll maxima beneath the ice. Phosphorus limitation was common to all lakes, since nitrogen compounds were significantly more abundant than the requirements for the primary production observed. The major chemical components present in the lakes showed a short but extreme dilution during thawing. Certain lake features may favour the recording of particular climatic fluctuations, for instance: lakes with two distinct productive periods, climatic fluctuations in spring or autumn (e.g., through chrysophycean cysts); lakes with higher oxygen consumption, climatic factors affecting the duration of the ice-cover (e.g., through low-oxygen tolerant chironomids); lakes with higher water retention time; changes in atmospheric deposition (e.g., through carbon or pigment burial); lakes with longer stratification, air temperature changes during summer and autumn (e.g., through all epilimnetic species).

This paper compares the palaeolimnological evidence for climate change over the last 200 years with instrumental climate data for the same period at seven European remote mountain lakes. The sites are Ovre Neadalsvatn (Norway), Saanajarvi (Finland), Gossenkollesee (Austria), Hagelseewli (Switzerland), Jezero v Ledvici (Slovenia), Estany Redo (Spain, Pyrenees), and NiOne Terianske Pleso (Slovakia). We used multiple regression analysis to transfer homogenised lowland air temperature records to each of the sites, and these reconstructions were validated using data from on-site automatic weather stations. These data showed that mean annual temperature has varied over the last 200 years at each site by between 1 and 2 °C, typical of the high frequency variability found throughout the Holocene, and appropriate, therefore, to test the sensitivity of the various proxy methods used. Sediment cores from each site were radiometrically dated using 210 Pb, 137 Cs and 241 Am and analysed for loss-on-ignition, C, N, S, pigments, diatoms, chrysophytes, Cladocera and chironomids. Comparisons between the proxy data and the instrumental data were based on linear regression analysis with the proxy data treated as response variables and the instrumental data (after smoothing using LOESS regressions) as predictor variables. The results showed few clear or consistent patterns with generally low or very low r 2 values. Highest values were found when the data were compared after smoothing using a broad span, indicating that some of the proxy data were capturing climate variability but only at a relatively coarse time resolution. Probable reasons for the weak performance of the methods used include inaccurate dating, especially for earlier time periods, the influence of confounding forcing factors at This is the eleventh of 11 papers published in this special issue on the palaeolimnology of remote mountain lakes in Europe resulting from the MOLAR project funded by the European Union. The guest editor was Richard W. Battarbee.

Here we present the chemical trends of seven high altitude lakes, analysed within the AL:PE and MOLAR Projects of the EU (1999) and selected on the basis of the availability of complete and reliable data for the period 1984–1999. The lakes are representative of the Scandinavian Alps, the Cairngorm Mountains in Scotland, the Alps and the Pyrenees. Significant trends were identified for some indicators of acidification, for instance pH and alkalinity, but not all lakes reacted similarly to decreasing depositions of sulphate and base cations. Differences in lake response are discussed in relation to recent variations of atmospheric deposition chemistry and associated changes in climatic conditions. Beside individual variations of the studied lakes, depending, among other things, on altitude and morphology, catchment characteristics and climate trends play a major role for the reaction of high altitude lakes on changes in atmospheric depositions.

This paper introduces the results of an integrated project designed to compare high resolution analysis of proxy records of climate change in the sediments of seven mountain lakes across Europe with reconstructed instrumental records of climate change over the last 200 years. Palaeolimnological methods used include radiometric dating (210Pb, 137Cs), mineral magnetics, dry weight, loss-on-ignition, carbon, nitrogen, sulphur, pigments, diatoms, chrysophyte cysts, cladocera and chironomids. Changes in fossil assemblages were summarised using principal components analysis. The stratigraphic data were compared with the instrumental record using linear regression techniques. The dated sediment records for each proxy from each site were treated as the response variables and the various attributes of the instrumental climate record as the predictor variables. The predictor variables were generated for each site for the period 1781 to 1997 using temperature reconstructions based on meteorological records. To harmonise the climatic predictors and the response variables, the climatic variables were smoothed along time with a LOESS regression. The results of the various analyses at the seven sites are presented in the following papers. A synthesis of the project and the relative performance of the different proxy methods are discussed in the final paper.

A modern diatom-pH calibration data-set consisting of surface-sediment diatom assemblages from 118 lakes and 530 taxa is presented. The AL:PE data-set is from high-altitude or high-latitude lakes in the Alps, Norway, Svalbard, Kola Peninsula, UK, Slovenia, Slovakia, Poland, Portugal, and Spain (pH range = 4.5-8.0; DOC range = 0.2-3.2 mg l-1). In addition, 92 epilithon samples from 22 high-altitude or high-latitude lakes comprise an AL:PE epilithon diatom-pH data-set. Weighted averaging partial least squares regression is used to develop pH-inference models. The AL:PE data-set has a root-mean-square-error of prediction (RMSEP) of 0.33 and a maximum bias of 0.36 pH units and r2 of 0.82, as assessed by leave-one-out cross-validation. The epilithon data-set has, after data-screening and the deletion of one very obvious outlier, a RMSEP of 0.23 and a maximum bias of 0.18 pH units and r2 of 0.88. The 167 sample SWAP diatom-pH data-set from lowland or upland lakes in the UK, Norway, and Sweden has a RMSEP of 0.29 and a maximum bias of 0.23 pH units and r2 of 0.86. The pH optima, as estimated by weighted averaging and Gaussian regression, are compared for the three data-sets (AL:PE, SWAP, AL:PE epilithon). There is a good correspondence between the AL:PE and the AL:PE epilithon optima, but a consistent bias between the AL:PE and SWAP optima, with the SWAP optima being lower than the AL:PE estimates. The predictive performances of the AL:PE and SWAP calibration data-sets are compared using independent test samples and six core sequences, all from high-altitude lakes, one in south-east Siberia and five in eastern Scotland. The results show the importance of using the AL:PE data-set for inferring lake-water pH from diatom assemblages in high-altitude or high latitude lakes with low DOC concentrations.

In the framework of the AL:PE 2 project, studies on acidification and organic pollution in mountain lakes have been conducted in several ranges in the Iberian peninsula: Pyrenees (Northeastern Spain), Sierra de Gredos (Central Spain), Sierra Nevada (Southern Spain) and Serra da Estrela (Central Portugal). These studies focused on water and sediment chemistry and organisms (benthic diatoms, Zooplankton, aquatic macroinvertebrates, and fish) as indicators of acidification. Organic micropollutants (PAH, PCB, DDE, hexachlorobenzene and others) in lake sediments and fish have been studied as tracers of atmospheric pollution. The Iberian peninsula lakes do not show severe anthropogenic acidification. pH values are in the range of sensitive lakes, but the levels of acidic pollutants are low. The status of the organisms surveyed agreed with this diagnosis. Pyrenean lakes showed the highest fluxes of organic pollutants related to fossil fuel combustion., higher pollutioninduced versus natural acidity ratios, and modeled alkalinity and pH declines.

13 páginas, 10 figuras, 1 tabla., Baseline and historical environmental data are sparse in high-altitude regions. Diatom assemblages preserved in lake sediment records can, however, provide proxy data of past environmental and biological conditions. Sediment cores were retrieved from 209 high altitude lakes from 11 countries in Europe. Diatoms were extracted and counted from surface-sediment samples and sediments representing pre-industrial conditions. Regional changes in diatom assemblages covering at least the last c. 150 years are discussed and quantified using ordination techniques and dissimilarity indices. Distinct changes in diatom composition are identified and regional patterns highlighted, with two lake regions showing particularly large changes in diatom composition (Central Swiss Alps and the Pyrenees). Several driving Aquat. Sci. 67 (2005) 224–236 1015-1621/05/030224-13 DOI 10.1007/s00027-004-0745-2 © EAWAG, Dübendorf, 2005 Aquatic Sciences mechanisms that might have caused the changes in the diatom community are discussed. Pre-industrial and present-day lake-water pH are inferred from the diatom assemblages and their prediction accuracy compared to contemporary water-chemistry measurements for each lake. Regional pH changes are minimal with no lake region exceeding 0.27 pH units of change in the period covering the last c. 150 years. The majority of lakes studied show an increase in planktonic diatom species over the period covering the past c. 150 years. We hypothesise that changes in the ratio of planktonic to non-planktonic diatoms within the study lake-regions are related to changes in climate and to associated changes in nutrients, ice-cover and erosion caused by climate warming., This research is part of the EMERGE Framework 5 project, supported by the European Union.