Ice phenology interactions with water and air temperatures in high mountain lakes.

Ice phenology is of great importance for the thermal structure of lakes and ponds and the biology of lake species. Under the current climate change conditions, ice-cover duration has been reduced by an advance in ice-off, and a delay in ice-on, and future projections foresee this trend as continuing. Here, we describe the current ice phenology of Pyrenean high mountain lakes and ponds, including ice-cover duration and ice-on and ice-off dates. We used mixed models to identify the variables that explained the observed patterns, extrapolated them across all water bodies in the mountain range, and related the seasonality of air and water temperatures with ice phenology using structural equation models. Ice phenology was obtained from the temperature series of 85 lakes and ponds for fourteen years, including 2001 to 2004 and 2009 to 2019. We discovered that high autumn precipitation was related to earlier ice-on dates, and that earlier ice-off dates were associated with higher following-summer water temperatures. We found a greater predictability of ice-off dates and ice-cover duration than ice-on dates. Altitude was the most important variable explaining the variation in ice phenology, followed by latitude, which was related to climatic differences among the northern and southern slopes of the mountain range. The lake area was significant for ice-on dates and ice-cover duration. The interannual variability in air temperature and radiation was remarkable for the ice-off date and ice-cover duration but not for the ice-on date. In contrast, wind speed was related to an earlier ice-off date and shorter ice-cover duration. All the measured lakes and ponds froze in winter during the studied period, a feature maintained in the extrapolation to the whole set of water bodies.

Reduced precipitation can induce ecosystem regime shifts in lakes by increasing internal nutrient recycling.

Eutrophication is a main threat to continental aquatic ecosystems. Prevention and amelioration actions have been taken under the assumption of a stable climate, which needs reconsideration. Here, we show that reduced precipitation can bring a lake ecosystem to a more productive regime even with a decline in nutrient external load. By analyzing time series of several decades in the largest lake of the Iberian Peninsula, we found autocorrelated changes in the variance of state variables (i.e., chlorophyll and oxygen) indicative of a transient situation towards a new ecosystem regime. Indeed, exceptional planktonic diatom blooms have occurred during the last few years, and the sediment record shows a shift in phytoplankton composition and an increase in nutrient retention. Reduced precipitation almost doubled the water residence time in the lake, enhancing the relevance of internal processes. This study demonstrates that ecological quality targets for aquatic ecosystems must be tailored to the changing climatic conditions for appropriate stewardship.

Female proportion has a stronger influence on dispersal than body size in nematodes of mountain lakes.

Nematodes disperse passively and are amongst the smallest invertebrates on Earth. Free-living nematodes in mountain lakes are highly tolerant of environmental variations and are thus excellent model organisms in dispersal studies, since species-environment relationships are unlikely to interfere. In this study, we investigated how population or organism traits influence the stochastic physical nature of passive dispersal in a topologically complex environment. Specifically, we analyzed the influence of female proportion and body size on the geographical distribution of nematode species in the mountain lakes of the Pyrenees. We hypothesized that dispersal is facilitated by (i) a smaller body size, which would increase the rate of wind transport, and (ii) a higher female proportion within a population, which could increase colonization success because many nematode species are capable of parthenogenetic reproduction. The results showed that nematode species with a low proportion of females tend to have clustered spatial distributions that are not associated with patchy environmental conditions, suggesting greater barriers to dispersal. When all species were pooled, the overall proportion of females tended to increase at the highest elevations, where dispersal between lakes is arguably more difficult. The influence of body size was barely relevant for nematode distributions. Our study highlights the relevance of female proportion as a mechanism that enhances the dispersal success of parthenogenetic species, and that female sex is a determining factor in metacommunity connectivity.

Self-filling enclosures to experimentally assess plankton response to pulse nutrient enrichments.

Experimental nutrient additions are a fundamental approach to investigating plankton ecology. Possibilities range from whole-lake fertilization to flask assays encompassing a trade-off between closeness to the "real world" and feasibility and replication. Here we describe an enclosure type that minimizes the manipulation of planktonic communities during the enclosure filling. The enclosure (typically ~100 L volume) consists of a narrow translucent cylinder that can comprise the entire photic zone (or a large part of it in clear deep lakes, e.g. 20-m long) and holds a sediment trap at the bottom for recovering the sinking material. The enclosures are inexpensive and straightforward to build. Thus, many can be used in an experiment, favoring the diversity of treatments and the number of replicates. They also are lightweight with easy transport and use in lakes that cannot be reached by road. The enclosures are fundamentally aimed at investigating the short-term response of the planktonic community, integrated across the photic zone, to pulse perturbations using before and after comparisons and multiple replication and treatments. The pros and cons of the enclosure design are evaluated based on experience gained in Lake Redon, a high mountain ultraoligotrophic deep lake in the Pyrenees.

Bacterioplankton seasonality in deep high-mountain lakes.

Due to global warming, shorter ice cover duration might drastically affect the ecology of lakes currently undergoing seasonal surface freezing. High-mountain lakes show snow-rich ice covers that determine contrasting conditions between ice-off and ice-on periods. We characterized the bacterioplankton seasonality in a deep high-mountain lake ice-covered for half a year. The lake shows a rich core bacterioplankton community consisting of three components: (i) an assemblage stable throughout the year, dominated by Actinobacteria, resistant to all environmental conditions; (ii) an ice-on-resilient assemblage dominating during the ice-covered period, which is more diverse than the other components and includes a high abundance of Verrucomicrobia; the deep hypolimnion constitutes a refuge for many of the typical under-ice taxa, many of which recover quickly during autumn mixing; and (iii) an ice-off-resilient assemblage, which members peak in summer in epilimnetic waters when the rest decline, characterized by a dominance of Flavobacterium, and Limnohabitans. The rich core community and low random elements compared to other relatively small cold lakes can be attributed to its simple hydrological network in a poorly-vegetated catchment, the long water-residence time (ca. 4 years), and the long ice-cover duration; features common to many headwater deep high-mountain lakes.

Scientists' warning of threats to mountains.

Mountains are an essential component of the global life-support system. They are characterized by a rugged, heterogenous landscape with rapidly changing environmental conditions providing myriad ecological niches over relatively small spatial scales. Although montane species are well adapted to life at extremes, they are highly vulnerable to human derived ecosystem threats. Here we build on the manifesto 'World Scientists' Warning to Humanity', issued by the Alliance of World Scientists, to outline the major threats to mountain ecosystems. We highlight climate change as the greatest threat to mountain ecosystems, which are more impacted than their lowland counterparts. We further discuss the cascade of "knock-on" effects of climate change such as increased UV radiation, altered hydrological cycles, and altered pollution profiles; highlighting the biological and socio-economic consequences. Finally, we present how intensified use of mountains leads to overexploitation and abstraction of water, driving changes in carbon stock, reducing biodiversity, and impacting ecosystem functioning. These perturbations can provide opportunities for invasive species, parasites and pathogens to colonize these fragile habitats, driving further changes and losses of micro- and macro-biodiversity, as well further impacting ecosystem services. Ultimately, imbalances in the normal functioning of mountain ecosystems will lead to changes in vital biological, biochemical, and chemical processes, critically reducing ecosystem health with widespread repercussions for animal and human wellbeing. Developing tools in species/habitat conservation and future restoration is therefore essential if we are to effectively mitigate against the declining health of mountains.

Individual and population-scale carbon and nitrogen isotopic values of Procambarusclarkii in invaded freshwater ecosystems.

Background: Freshwater ecosystems are amongst the most threatened habitats on Earth; nevertheless, they support about 9.5% of the known global biodiversity while covering less than 1% of the globe's surface. A number of anthropogenic pressures are impacting species diversity in inland waters and, amongst them, the spread of invasive alien species is considered one of the main drivers of biodiversity loss and homogenisation in freshwater habitats.Crayfish species are widely distributed freshwater invaders and, while alien species introductions occur mostly accidentally, alien crayfish are often released deliberately into new areas for commercial purposes. After their initial introduction, crayfish species can rapidly establish and reach high-density populations as a result of their adaptive functional traits, such as their generalist diet.The Louisiana crayfish Procambarusclarkii (Girard, 1852) is globally considered one of the worst invaders and its impact on recipient freshwater communities can vary from predation and competition with native species, to modification of food webs and habitat structure and introduction of pathogens. Native to the south United States and north Mexico, P.clarkii has been introduced in Europe, Asia and Africa, determining negative ecological and economic impacts in the majority of invaded habitats where it became dominant within the receiving benthic food webs. Due to its flexible feeding strategy, P.clarkii exerts adverse effects at different trophic levels, ultimately affecting the structure and dynamics of invaded food webs. It is, therefore, paramount to evaluate the ecological consequences of P.clarkii invasion and to quantify its impact in a spatially explicit context. New information: In the past decades, the analysis of stable isotopes of carbon, nitrogen and other elements has become a popular methodology in food web ecology. Notably, stable isotope analysis has emerged as a primary tool for addressing applied issues in biodiversity conservation and management, such as the assessment of the trophic ecology of non-indigenous species in invaded habitats. Here, we built two geo-referenced datasets, resolved respectively at the population and individual scale, by collating information on δ13C and δ15N values of P.clarkii within invaded inland waters. The population-scale dataset consists of 160 carbon and nitrogen isotopic values of the Louisiana crayfish and its potential prey, including living and non-living primary producers and benthic invertebrates. The dataset resolved at individual scale consists of 1,168 isotopic records of P.clarkii. The isotopic values included within the two datasets were gathered from 10 countries located in Europe, Asia, Africa and North America, for a total of 41 studies published between 2005 and 2021. To the best of the authors' knowledge, this effort represents the first attempt to collate in standardised datasets the sparse isotopic information of P.clarkii available in literature. The datasets lend themselves to being used for providing a spatially explicit resolution of the trophic ecology of P.clarkii and to address a variety of ecological questions concerning its ecological impact on recipient aquatic food webs.

Factors of surface thermal variation in high-mountain lakes of the Pyrenees.

Thermal variables are crucial drivers of biological processes in lakes and ponds. In the current context of climate change, determining which factors better constrain their variation within lake districts become of paramount importance for understanding species distribution and their conservation. In this study, we describe the regional and short-term interannual variability in surface water temperature of high mountain lakes and ponds of the Pyrenees. And, we use mixed regression models to identify key environmental factors and to infer mean and maximum summer temperature, accumulated degree-days, diel temperature ranges and three-days' oscillation. The study is based on 59 lake-temperature series measured from 2001 to 2014. We found that altitude was the primary explicative factor for accumulated degree-days and mean and maximum temperature. In contrast, lake area showed the most relevant effect on the diel temperature range and temperature oscillations, although diel temperature range was also found to decline with altitude. Furthermore, the morphology of the catchment significantly affected accumulated degree-days and maximum and mean water temperatures. The statistical models developed here were applied to upscale spatially the current thermic conditions across the whole set of lakes and ponds of the Pyrenees.

Phragmites australis as a dual indicator (air and sediment) of trace metal pollution in wetlands - the key case of Flix reservoir (Ebro River).

Evaluation of trace metal pollution in an environmentally complex context may require the use of a suite of indicators. Common reed, Phragmites australis, is a well-known biomonitor of sediment pollution. Here, we show its potential for also assessing air pollution. The plant panicles, holding silky hairs with high surface to volume ratio, are appropriate collectors of atmospheric contaminants, which perform independently from root bioconcentration. We applied the dual value of common reed as an indicator of trace metal pollution to the case of a chlor-alkali plant in the Ebro river bank (Spain). This factory had historically damped waste to the shallow Flix reservoir. Extensive common reed meadows are growing on the top of the waste, in a nearby nature reserve across the reservoir and a meander immediately downriver. Three replicated individuals from a total of 11 sites were sampled, and the trace metal content measured in the main plant compartments (roots, rhizomes, stems, leaves, and panicles). Panicles and roots showed a much larger concentration of trace metals than the other plant compartments. Levels of Hg, Cu, and Ni were markedly higher in panicles at the factory and nearby points of the reserve and lowered at the meander. In contrast, Cd, Zn, and Mn in roots increased from the factory to the meander downriver. We conclude that panicles show recent (less than a year) airborne pollution, whereas roots indicate the long-term transport of pollutants from the waste in the shoreline of the factory to downriver sedimentation hotspots, where they become more bioavailable than in the factory waste. The Hg spatial pattern in panicles agree with air measurements in later years, therefore, confirming the panicles suitability for assessing airborne pollution and, consequently, Phragmites as a potential dual biomonitor of air and sediments.

Denitrification rates in lake sediments of mountains affected by high atmospheric nitrogen deposition.

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.

Diatoms as indicators of the multivariate environment of mountain lakes.

Diatom assemblages are used widely as indicators of environmental conditions. They have been particularly useful in the assessment of some acute environmental problems such as water acidification, eutrophication and salinisation. High mountain lakes are currently a research focus as sentinels of global change, predominantly of long-distance atmospheric pollution and climate warming. The diatom assemblages in these lakes are extremely rich, and the sediment record provides short and long historical perspectives of the changes. We investigated the relative sensitivity of diatoms to the main environmental gradients found in mountain lakes and evaluated the strength and uncertainties for predicting simultaneously several environmental variables associated with these gradients based on a survey of 83 lakes in the Pyrenees. Variables related to the ionic composition and acid-base balance (calcium*, acid neutralizing capacity*, magnesium, sodium, and sulphates), trophic conditions (total phosphorus* and dissolved organic carbon) and physical factors (water temperature*, irradiance at the lake bottom*, and macrophyte cover) explained independently the variation in the diatom assemblages. However, the assemblage predictive capacity of these variables- tested by developing transfer functions (Weighted Averaging- Partial Least Squares) - was only acceptable for a subset of the variables (*). The spatial autocorrelation of the environmental variables had no influence on the performance of the transfer functions except for water temperature, which is highly dependent on altitude. Our results indicate that diatom assemblages have great potential for assessments of multiple environmental variables in mountain lakes and, consequently, in applications of global change surveillance.

Taxonomy and functional interactions in upper and bottom waters of an oligotrophic high-mountain deep lake (Redon, Pyrenees) unveiled by microbial metagenomics.

High mountain lakes are, in general, highly sensitive systems to external forcing and good sentinels of global environmental changes. For a better understanding of internal lake processes, we examined microbial biodiversity and potential biogeochemical interactions in the oligotrophic deep high-mountain Lake Redon (Pyrenees, 2240 m altitude) using shotgun metagenomics. We analyzed the two ends of the range of environmental conditions found in Lake Redon, at 2 and 60 m depths. Bacteria were the most abundant component of the metagenomic reads (>90%) and the diversity indices of both taxonomic (16S and 18S rRNA) and functional (carbon-, nitrogen-, sulfur-, and phosphorous-cycling) related genes were higher in the bottom dark layer than in the upper compartment. A marked segregation was observed both in biodiversity and in the dominant energy and biomass generating pathways between the extremes. The aerobic respiration was mainly dominated by heterotrophic Burkholderiales at the top and Actinobacteria and Burkholderiales at the lake bottom. The potential for an active nitrogen cycle (nitrogen fixation, nitrification, nitrite oxidation, and nitrate reduction) was mainly found at 60 m, and potential for methanogenesis, anaerobic ammonia oxidation and dissimilatory sulfur pathways were only observed there. Some unexpected and mostly unseen energy and biomass pathways were found relevant for the biogeochemical cycling in lake Redon, i.e., those related to carbon monoxide oxidation and phosphonates processing. We provide a general scheme of the main biogeochemical processes that may operate in the sentinel deep Lake Redon. This framework may help for a better understanding of the whole lake metabolism.

The DNRA-Denitrification Dichotomy Differentiates Nitrogen Transformation Pathways in Mountain Lake Benthic Habitats.

Effects of nitrogen (N) deposition on microbially-driven processes in oligotrophic freshwater ecosystems are poorly understood. We quantified guilds in the main N-transformation pathways in benthic habitats of 11 mountain lakes along a dissolved inorganic nitrogen gradient. The genes involved in denitrification (nirS, nirK, nosZ), nitrification (archaeal and bacterial amoA), dissimilatory nitrate reduction to ammonium (DNRA, nrfA) and anaerobic ammonium oxidation (anammox, hdh) were quantified, and the bacterial 16S rRNA gene was sequenced. The dominant pathways and associated bacterial communities defined four main N-transforming clusters that differed across habitat types. DNRA dominated in the sediments, except in the upper layers of more productive lakes where nirS denitrifiers prevailed with potential N2O release. Loss as N2 was more likely in lithic biofilms, as indicated by the higher hdh and nosZ abundances. Archaeal ammonia oxidisers predominated in the isoetid rhizosphere and rocky littoral sediments, suggesting nitrifying hotspots. Overall, we observed a change in potential for reactive N recycling via DNRA to N losses via denitrification as lake productivity increases in oligotrophic mountain lakes. Thus, N deposition results in a shift in genetic potential from an internal N accumulation to an atmospheric release in the respective lake systems, with increased risk for N2O emissions from productive lakes.

Estimating Sediment Denitrification Rates Using Cores and N2O Microsensors.

Denitrification is the primary biogeochemical process removing reactive nitrogen from the biosphere. The quantitative evaluation of this process has become particularly relevant for assessing the anthropogenic-altered global nitrogen cycle and the emission of greenhouse gases (i.e., N2O). Several methods are available for measuring denitrification, but none of them are completely satisfactory. Problems with existing methods include their insufficient sensitivity, and the need to modify the substrate levels or alter the physical configuration of the process using disturbed samples.This work describes a method for estimating sediment denitrification rates that combines coring, acetylene inhibition, and microsensor measurements of the accumulated N2O. The main advantages of this method are a low disturbance of the sediment structure and the collection of a continuous record of N2O accumulation; these enable estimates of reliable denitrification rates with minimum values up to 0.4-1 µmol N2O m-2 h-1. The ability to manipulate key factors is an additional advantage for obtaining experimental insights. The protocol describes procedures for collecting the cores, calibrating the sensors, performing the acetylene inhibition, measuring the N2O accumulation, and calculating the denitrification rate. The method is appropriate for estimating denitrification rates in any aquatic system with retrievable sediment cores. If the N2O concentration is above the detection limit of the sensor, the acetylene inhibition step can be omitted to estimate the N2O emission instead of denitrification. We show how to estimate both actual and potential denitrification rates by increasing nitrate availability as well as the temperature dependence of the process. We illustrate the procedure using mountain lake sediments and discuss the advantages and weaknesses of the technique compared to other methods. This method can be modified for particular purposes; for instance, it can be combined with 15N tracers to assess nitrification and denitrification or field in situ measurements of denitrification rates.

Experimental evidence of the quantitative relationship between the prokaryote ingestion rate and the food vacuole content in mixotrophic phytoflagellates.

The verification that many phytoflagellates ingest prokaryotes has changed the view of the microbial loop in aquatic ecosystems. Still, progress is limited because the phagotrophic activity is difficult to quantify in natural assemblages. Linking the food vacuole content in protist with the ingestion rate of prokaryotes would provide a crucial step forward. In this study, using the catalysed reporter deposition - fluorescence in situ hybridization protocol (CARD-FISH), which allows the visualization of labelled prokaryotes inside protists without relying on incubation procedures, we experimentally relate the food vacuole content of prokaryotes (Vc ) to the population-averaged ingestion rates (Ir ) estimated using bacteria-size fluorescent microspheres. The two variables relate according to the equation Ir = 7.52 Vc 0.9 , which indicates a prokaryote half-life of about 6 min in the protist vacuole. Five mixotrophic flagellate species from natural and culture populations were evaluated seven times during 24 h; they provided a broad range of average vacuole content (0.01 to 2.02 prokaryote protist-1 ) and ingestion rates (0.18 to 23 prokaryote protist-1 h-1 ). Consequently, the relationship found can be applied to quantify the mixotrophy activity in a large variety of field and experimental studies.

High planktonic diversity in mountain lakes contains similar contributions of autotrophic, heterotrophic and parasitic eukaryotic life forms.

A rich eukaryotic planktonic community exists in high-mountain lakes despite the diluted, oligotrophic and cold, harsh prevailing conditions. Attempts of an overarching appraisal have been traditionally hampered by observational limitations of small, colorless, and soft eukaryotes. We aimed to uncover the regional eukaryotic biodiversity of a mountain lakes district to obtain general conclusions on diversity patterns, dominance, geographic diversification, and food-web players common to oligotrophic worldwide distributed freshwater systems. An unprecedented survey of 227 high-altitude lakes comprising large environmental gradients was carried out using Illumina massive tag sequencing of the 18S rRNA gene. We observed a large Chrysophyceae dominance in richness, abundance and novelty, and unveiled an unexpected richness in heterotrophic phagotrophs and parasites. In particular, Cercozoa and Chytridiomycota showed diversity features similar to the dominant autotrophic groups. The prominent beta-dispersion shown by parasites suggests highly specific interactions and a relevant role in food webs. Interestingly, the freshwater Pyrenean metacommunity contained more diverse specific populations than its closest marine oligotrophic equivalent, with consistently higher beta-diversity. The relevance of unseen groups opens new perspectives for the better understanding of planktonic food webs. Mountain lakes, with remarkable environmental idiosyncrasies, may be suitable environments for the genetic diversification of microscopic eukaryotic life forms.

Nematode distributions as spatial null models for macroinvertebrate species richness across environmental gradients: A case from mountain lakes.

Nematode species are widely tolerant of environmental conditions and disperse passively. Therefore, the species richness distribution in this group might largely depend on the topological distribution of the habitats and main aerial and aquatic dispersal pathways connecting them. If so, the nematode species richness distributions may serve as null models for evaluating that of other groups more affected by environmental gradients. We investigated this hypothesis in lakes across an altitudinal gradient in the Pyrenees. We compared the altitudinal distribution, environmental tolerance, and species richness, of nematodes with that of three other invertebrate groups collected during the same sampling: oligochaetes, chironomids, and nonchironomid insects. We tested the altitudinal bias in distributions with t-tests and the significance of narrow-ranging altitudinal distributions with randomizations. We compared results between groups with Fisher's exact tests. We then explored the influence of environmental factors on species assemblages in all groups with redundancy analysis (RDA), using 28 environmental variables. And, finally, we analyzed species richness patterns across altitude with simple linear and quadratic regressions. Nematode species were rarely biased from random distributions (5% of species) in contrast with other groups (35%, 47%, and 50%, respectively). The altitudinal bias most often shifted toward low altitudes (85% of biased species). Nematodes showed a lower portion of narrow-ranging species than any other group, and differed significantly from nonchironomid insects (10% and 43%, respectively). Environmental variables barely explained nematode assemblages (RDA adjusted R2 = 0.02), in contrast with other groups (0.13, 0.19 and 0.24). Despite these substantial differences in the response to environmental factors, species richness across altitude was unimodal, peaking at mid elevations, in all groups. This similarity indicates that the spatial distribution of lakes across altitude is a primary driver of invertebrate richness. Provided that nematodes are ubiquitous, their distribution offers potential null models to investigate species richness across environmental gradients in other ecosystem types and biogeographic regions.

Some Mixotrophic Flagellate Species Selectively Graze on Archaea.

Many phototrophic flagellates ingest prokaryotes. This mixotrophic trait becomes a critical aspect of the microbial loop in planktonic food webs because of the typical high abundance of these flagellates. Our knowledge of their selective feeding upon different groups of prokaryotes, particularly under field conditions, is still quite limited. In this study, we investigated the feeding behavior of three species (Rhodomonas sp., Cryptomonas ovata, and Dinobryon cylindricum) via their food vacuole content in field populations of a high mountain lake. We used the catalyzed reporter deposition-fluorescence in situ hybridization (CARD-FISH) protocol with probes specific for the domain Archaea and three groups of Eubacteria: Betaproteobacteria, Actinobacteria, and Cytophaga-Flavobacteria of Bacteroidetes Our results provide field evidence that contrasting selective feeding exists between coexisting mixotrophic flagellates under the same environmental conditions and that some prokaryotic groups may be preferentially impacted by phagotrophic pressure in aquatic microbial food webs. In our study, Archaea were the preferred prey, chiefly in the case of Rhodomonas sp., which rarely fed on any other prokaryotic group. In general, prey selection did not relate to prey size among the grazed groups. However, Actinobacteria, which were clearly avoided, mostly showed a size of <0.5 µm, markedly smaller than cells from the other groups. IMPORTANCE: That mixotrophic flagellates are not randomly feeding in the main prokaryotic groups under field conditions is a pioneer finding in species-specific behavior that paves the way for future studies according to this new paradigm. The particular case that Archaea were preferentially affected in the situation studied shows that phagotrophic pressure cannot be disregarded when considering the distribution of this group in freshwater oligotrophic systems.

Ecology under lake ice.

Winter conditions are rapidly changing in temperate ecosystems, particularly for those that experience periods of snow and ice cover. Relatively little is known of winter ecology in these systems, due to a historical research focus on summer 'growing seasons'. We executed the first global quantitative synthesis on under-ice lake ecology, including 36 abiotic and biotic variables from 42 research groups and 101 lakes, examining seasonal differences and connections as well as how seasonal differences vary with geophysical factors. Plankton were more abundant under ice than expected; mean winter values were 43.2% of summer values for chlorophyll a, 15.8% of summer phytoplankton biovolume and 25.3% of summer zooplankton density. Dissolved nitrogen concentrations were typically higher during winter, and these differences were exaggerated in smaller lakes. Lake size also influenced winter-summer patterns for dissolved organic carbon (DOC), with higher winter DOC in smaller lakes. At coarse levels of taxonomic aggregation, phytoplankton and zooplankton community composition showed few systematic differences between seasons, although literature suggests that seasonal differences are frequently lake-specific, species-specific, or occur at the level of functional group. Within the subset of lakes that had longer time series, winter influenced the subsequent summer for some nutrient variables and zooplankton biomass.

Foraging success under uncertainty: search tradeoffs and optimal space use.

Understanding the structural complexity and the main drivers of animal search behaviour is pivotal to foraging ecology. Yet, the role of uncertainty as a generative mechanism of movement patterns is poorly understood. Novel insights from search theory suggest that organisms should collect and assess new information from the environment by producing complex exploratory strategies. Based on an extension of the first passage time theory, and using simple equations and simulations, we unveil the elementary heuristics behind search behaviour. In particular, we show that normal diffusion is not enough for determining optimal exploratory behaviour but anomalous diffusion is required. Searching organisms go through two critical sequential phases (approach and detection) and experience fundamental search tradeoffs that may limit their encounter rates. Using experimental data, we show that biological search includes elements not fully considered in contemporary physical search theory. In particular, the need to consider search movement as a non-stationary process that brings the organism from one informational state to another. For example, the transition from remaining in an area to departing from it may occur through an exploratory state where cognitive search is challenged. Therefore, a more comprehensive view of foraging ecology requires including current perspectives about movement under uncertainty.