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International audience; Zooplankton can spend winter actively under the ice cover of lakes. However, dietary resources under lake ice are both quantitatively and qualitatively limited, and feeding might not be energetically rewarding for most zooplankton species. Many zooplankters are expected to fast throughout the winter, exhausting their previously accumulated fat storage. We hypothesised that only a fraction of the actively overwintering zooplankton contributes to an active food web under lake ice, leading to few trophic linkages within the planktonic community.2. Zooplankton habitats and feeding were investigated under the ice of Lake Onego. Zooplankton habitats and migrations were studied by coupling zooplankton sam-pling around the clock to measurements of particle movement using an acoustic Doppler current profiler. Secondly, fatty acid-specific stable isotope compositions were used to determine whether and which zooplankton fatty acids ultimately came from the assimilation of under-ice seston.3. The algal biomass was low under ice and mostly dominated by large diatoms. Copepods dominated the zooplankton community. Species present as late cope-podite and adult instars were confined to the deeper layers, while nauplii occupied the surface layer. Diel vertical migration by Cyclops was the most tangible ob-servation of persistent feeding under the ice. Previously accumulated fat storage represented most of zooplankton fatty acids, with few, yet detectable, fatty acids recently acquired by feeding under the ice.4. Although some zooplankton taxa maintained feeding activity under the ice of Lake Onego, the food source available beneath the ice was not sufficiently rewarding to leave an isotopic imprint upon the dominant fatty acids of bulk zooplankton. The seston fatty acids that were passed on to zooplankton from feeding under ice were not provided by diatoms, although they made up most of the phytoplank-ton biovolume. Instead, the zooplankton food web was supported by mixotrophic phytoplankton (i.e. cryptophytes and chrysophytes) that represented

Winter is a long period of the annual cycle of many lakes in the northern hemisphere. Low irradiance, ice, and snow cover cause poor light penetration into the water column of these lakes. Therefore, in northern lakes, respiration often exceeds primary production leading to low dissolved oxygen concentrations. This study aimed to quantify under-ice metabolic processes during winter in an arid zone lake with little snow cover. This study was carried out in a mid-latitude lake in Inner Mongolia, northern China. The study lake receives relatively high incoming solar radiation on the ice in mid-winter, and radiation can penetrate down to the bottom sediment as the lake is shallow and the ice lacks snow cover. Primary production and respiration were estimated during two winters using high-frequency sensor measurements of dissolved oxygen. To quantify under-ice metabolic processes, sensors were deployed to different depths. During both winters, sensors collected data every 10 min over several weeks. The amount of solar radiation controlled photosynthesis under ice; temperature and photosynthesis together appeared to control respiration. The balance between gross primary production and ecosystem respiration was especially sensitive to changes in snow cover, and the balance between P and R decreased. Our data suggest that photosynthesis by plankton, submerged plants, and epiphytic algae may continue over winter in shallow lakes in mid-latitudes when there is no snow cover on the ice, as may occur in arid climates. The continuation of photosynthesis under ice buffers against dissolved oxygen depletion and prevents consequent harmful ecosystem effects.

We investigated radiatively driven under-ice convection in Lake Onego (Russia) during 3 consecutive late winters. In ice-covered lakes, where the temperature of water is below the temperature of maximum density, radiatively driven heating in the upper water column induces unstable density distributions leading to gravitational convection. In this work, we quantified the key parameters to characterise the radiatively driven under-ice convection: (1) the effective buoyancy flux, B∗ (driver), and its vertical distribution; (2) the convective mixed-layer thickness, hCML (depth scale); and (3) the convective velocity,w∗(kinematic scale). We compared analytical w∗ scaling estimates to in situ observations from high-resolution acoustic Doppler current profilers. The results show a robust correlation between w∗and the direct observations, except during the onset and decay of the solar radiation. Our results highlight the importance of accurately defining the upper limit of hCML in highly turbid water and the need for spectrally resolving solar radiation measurements and their attenuation for accurate B∗ estimates. Uncertainties in the different parameters were also investigated. We finally examined the implications of under-ice convection for the growth rate of nonmotile phytoplankton and provide a simple heuristic model as a function of easily measurable parameters.

Among its many impacts, climate warming is leading to increasing winter air temperatures, decreasing ice cover extent, and changing winter precipitation patterns over the Laurentian Great Lakes and their watershed. Understanding and predicting the consequences of these changes is impeded by a shortage of winter-period studies on most aspects of Great Lake limnology. In this review, we summarize what is known about the Great Lakes during their 3–6 months of winter and identify key open questions about the physics, chemistry, and biology of the Laurentian Great Lakes and other large, seasonally frozen lakes. Existing studies show that winter conditions have important effects on physical, biogeochemical, and biological processes, not only during winter but in subsequent seasons as well. Ice cover, the extent of which fluctuates dramatically among years and the five lakes, emerges as a key variable that controls many aspects of the functioning of the Great Lakes ecosystem. Studies on the properties and formation of Great Lakes ice, its effect on vertical and horizontal mixing, light conditions, and biota, along with winter measurements of fundamental state and rate parameters in the lakes and their watersheds are needed to close the winter knowledge gap. Overcoming the formidable logistical challenges of winter research on these large and dynamic ecosystems may require investment in new, specialized research infrastructure. Perhaps more importantly, it will demand broader recognition of the value of such work and collaboration between physicists, geochemists, and biologists working on the world's seasonally freezing lakes and seas.

Long-term lake ice phenological records from around the Northern Hemisphere provide unique sensitive indicators of climatic variations, even prior to the existence of physical meteorological measurement stations. Here, we updated ice phenology records for 60 lakes with time-series ranging from 107–204 years to provide the first re-assessment of Northern Hemispheric ice trends since 2004 by adding 15 additional years of ice phenology records and 40 lakes to our study. We found that, on average, ice-on was 11.0 days later, ice-off was 6.8 days earlier, and ice duration was 17.0 days shorter per century over the entire record for each lake. Trends in ice-on and ice duration were six times faster in the last 25-year period (1992–2016) than previous quarter centuries. More extreme events in recent decades, including late ice-on, early ice-off, shorter periods of ice cover, or no ice cover at all, contribute to the increasing rate of lake ice loss. Reductions in greenhouse gas emissions could limit increases in air temperature and abate losses in lake ice cover that would subsequently limit ecological, cultural, and socioeconomic consequences, such as increased evaporation rates, warmer water temperatures, degraded water quality, and the formation of toxic algal blooms.

Lakes are traditionally classified based on their thermal regime and trophic status. While this classification adequately captures many lakes, it is not sufficient to understand seasonally ice-covered lakes, the most common lake type on Earth. We describe the inverse thermal stratification in 19 highly varying lakes and derive a model that predicts the temperature profile as a function of wind stress, area, and depth. The results suggest an additional subdivision of seasonally ice-covered lakes to differentiate underice stratification. When ice forms in smaller and deeper lakes, inverse stratification will form with a thin buoyant layer of cold water (near 0°C) below the ice, which remains above a deeper 4°C layer. In contrast, the entire water column can cool to ∼0°C in larger and shallower lakes. We suggest these alternative conditions for dimictic lakes be termed “cryostratified” and “cryomictic.” The authors thank the Global Lake Ecological Observatory Network for organizing the GLEON 21 meeting, where this work was initially conceived. B. Yang and M. G. Wells acknowledges support from the Ontario Ministry of Environment, Conservation, and Parks (Grant LS-14-15-011) and the NSERC Discovery program (Grant RGPIN-2016-06542). The Lake Superior observations were supported by National Science Foundation Division of Ocean Sciences grant OCE-0825633 and NSF RAPID (Grant OCE-1445567). A. Laas acknowledges support from the Estonian Research Council Grant PSG32. G. A. Weyhenmeyer acknowledges the Swedish Infrastructure for Ecosystem Science (SITES) for provisioning of data from Lake Erken. SITES receives funding through the Swedish Research Council under the grant 2017-00635. Lake Mendota and Crystal Bog observations were supported by the US National Science Foundation grant DEB-1440297 and grant DEB-1856224. RMP acknowledges support from US National Science Foundation grants DEB 17276 and DEB 1950170. C. C. Carey acknowledges support from US National Science Foundation grant DEB 1753639 and 1933016. RMP acknowledges support from US National Science Foundation grants DEB 1754265, DEB 1754276, DEB 1950170. The data for the Pocono lakes were collected with support from Craig Williamson's Global Change Limnology Laboratory and Miami University, and from Kevin Rose's Global Water Laboratory at Rensselaer Polytechnic Institute. Alexie Lake data collections were supported by DeBeers Canada and Fisheries & Oceans Canada. The authors thank the Global Lake Ecological Observatory Network for organizing the GLEON 21 meeting, where this work was initially conceived. B. Yang and M. G. Wells acknowledges support from the Ontario Ministry of Environment, Conservation, and Parks (Grant LS-14-15-011) and the NSERC Discovery program (Grant RGPIN-2016-06542). The Lake Superior observations were supported by National Science Foundation Division of Ocean Sciences grant OCE-0825633 and NSF RAPID (Grant OCE-1445567). A. Laas acknowledges support from the Estonian Research Council Grant PSG32. G. A. Weyhenmeyer acknowledges the Swedish Infrastructure for Ecosystem Science (SITES) for provisioning of data from Lake Erken. SITES receives funding through the Swedish Research Council under the grant 2017-00635. Lake Mendota and Crystal Bog observations were supported by the US National Science Foundation grant DEB-1440297 and grant DEB-1856224. RMP acknowledges support from US National Science Foundation grants DEB 17276 and DEB 1950170. C. C. Carey acknowledges support from US National Science Foundation grant DEB 1753639 and 1933016. RMP acknowledges support from US National Science Foundation grants DEB 1754265, DEB 1754276, DEB 1950170. The data for the Pocono lakes were collected with support from Craig Williamson's Global Change Limnology Laboratory and Miami University, and from Kevin Rose's Global Water Laboratory at Rensselaer Polytechnic Institute. Alexie Lake data collections were supported by DeBeers Canada and Fisheries & Oceans Canada.

As the global climate warms, the fate of lacustrine fish is of huge concern, especially given their sensitivity as ectotherms to changes in water temperature. The Arctic charr (Salvelinus alpinusL.) is a salmonid with a Holarctic distribution, with peripheral populations persisting at temperate latitudes, where it is found only in sufficiently cold, deep lakes. Thus, warmer temperatures in these habitats particularly during early life stages could have catastrophic consequences on population dynamics. Here, we combined lake temperature observations, a 1-D hydrodynamic model, and a multi-decadal climate reanalysis to show coherence in warming winter water temperatures in four European charr lakes near the southernmost limit of the species’ distribution. Current maximum and mean winter temperatures are on average ~ 1 °C warmer compared to early the 1980s, and temperatures of 8.5 °C, adverse for high charr egg survival, have frequently been exceeded in recent winters. Simulations of winter lake temperatures toward century-end showed that these warming trends will continue, with further increases of 3–4 °C projected. An additional 324 total accumulated degree-days during winter is projected on average across lakes, which could impair egg quality and viability. We suggest that the perpetuating winter warming trends shown here will imperil the future status of these lakes as charr refugia and generally do not augur well for the fate of coldwater-adapted lake fish in a warming climate.

Nutrient transport in seasonally ice-covered lakes is an important factor affecting spring algal blooms in eutrophic waters; because phase changes during the ice growth process redistribute the nutrients. In this study, nutrient transport under static conditions was simulated by using two ice thickness models in combination with an indoor freezing experiment under different segregation coefficient conditions for nutrients. A real-time prediction model for nutrient and pollutant concentrations in ice-covered lakes was established to explore the impact of the ice-on period in eutrophic shallow lakes. The results demonstrated that the empirical degree-day model and the high-resolution thermodynamic snow and sea-ice model (HIGHTSI) could both be used to simulate lake ice thickness. The empirical degree-day model performed better at predicting the maximum ice thickness (measured thickness 0.22-0.55 m; simulated thickness 0.48 m), whereas the HIGHTSI model was more accurate when estimating the mean thickness (5-6% error). When simulating ice growth, the HIGHTSI model considered more meteorological factors impacting ice cover ablation; hence, it performed better during the ablation stage relative to the empirical degree-day model. Two non-dynamic nutrient transport models were developed by combining the segregation coefficient model and the ice thickness prediction model. The HIGHTSI nutrient transport model can be used to predict real-time changes in nutrient concentrations under ice cover, and the degree-day model can be used to predict changes in the lake water ecosystem.

Permafrost thawing results in the formation of thermokarst lakes, which are biogeochemical hotspots in northern landscapes and strong emitters of greenhouse gasses to the atmosphere. Most studies of thermokarst lakes have been in summer, despite the predominance of winter and ice-cover over much of the year, and the microbial ecology of these waters under ice remains poorly understood. Here we first compared the summer versus winter microbiomes of a subarctic thermokarst lake using DNA- and RNA-based 16S rRNA amplicon sequencing and qPCR. We then applied comparative metagenomics and used genomic bin reconstruction to compare the two seasons for changes in potential metabolic functions in the thermokarst lake microbiome. In summer, the microbial community was dominated by Actinobacteria and Betaproteobacteria, with phototrophic and aerobic pathways consistent with the utilization of labile and photodegraded substrates. The microbial community was strikingly different in winter, with dominance of methanogens, Planctomycetes, Chloroflexi and Deltaproteobacteria, along with various taxa of the Patescibacteria/Candidate Phyla Radiation (Parcubacteria, Microgenomates, Omnitrophica, Aminicenantes). The latter group was underestimated or absent in the amplicon survey, but accounted for about a third of the metagenomic reads. The winter lineages were associated with multiple reductive metabolic processes, fermentations and pathways for the mobilization and degradation of complex organic matter, along with a strong potential for syntrophy or cross-feeding. The results imply that the summer community represents a transient stage of the annual cycle, and that carbon dioxide and methane production continue through the prolonged season of ice cover via a taxonomically distinct winter community and diverse mechanisms of permafrost carbon transformation.

The annual minimum of lake surface water temperature influences ecological and biogeochemical processes, but variability and change in this extreme have not been investigated. Here, we analysed observational data from eight European lakes and investigated the changes in annual minimum surface water temperature. We found that between 1973 and 2014, the annual minimum lake surface temperature has increased at an average rate of + 0.35 degrees Cdecade(-1), comparable to the rate of summer average lake surface temperature change during the same period (+ 0.32 degrees C decade(-1)). Coherent responses to climatic warming are observed between the increase in annual minimum lake surface temperature and the increase in winter air temperature variations. As a result of the rapid warming of annual minimum lake surface temperatures, some of the studied lakes no longer reach important minimum surface temperature thresholds that occur in winter, with complex and significant potential implications for lakes and the ecosystem services that they provide.

Despite the rapidly changing winter conditions in temperate ecosystems, little attention has been devoted to the effects of these changes on lake ecology. Few studies on the seasonal changes in abundance and biomass of the major groups of the metazooplankton community (i.e,. rotifers, cladocerans and copepods) in northern oligotrophic lakes include data from the ice-covered winter months. This study reports monthly variation in zooplankton abundance and biomass from June 2010 to October 2011, including winter, in an oligotrophic, subalpine lake in southeastern Norway (Lake Atnsjøen). Changes in rotifer, cladoceran, copepod, and total zooplankton abundances and biomass were related to seasonal variation in water temperature and phytoplankton biomass by means of ordination analysis. The zooplankton abundance and biomass were much lower in winter than during the open water season. However, an underice phytoplankton bloom occurred during the final winter months, when snow cover and ice thickness were reduced and (presumably) light penetration increased, leading to an increase in abundance of copepod nauplii. Winter zooplankton abundance was dominated by copepods and rotifers, while winter zooplankton biomass was dominated by copepods and cladocerans. Both phytoplankton and zooplankton had two biomass peaks in 2010 and one peak in 2011. Rotifers dominated zooplankton abundance with a peak in August and total zooplankton abundance followed a similar pattern. In contrast, cladocerans dominated zooplankton biomass with a peak in July and total zooplankton biomass also peaked at this time. Rotifer and total zooplankton abundance and rotifer biomass were most closely correlated to water temperature. However, cladoceran biomass and total biomass were most closely correlated with phytoplankton biomass, but also appeared to be dependent on other carbon sources. Estimates of non-phytoplankton particulate organic carbon indicated that this part of the carbon pool could be an additional food source for zooplankton particularly in early and mid-winter. The longer growing season in 2011 than in 2010, owing to earlier ice-off in 2011, may have contributed to higher phytoplankton and zooplankton biomass in 2011. With climate warming, this is an expected change in temperate lake ecosystems.

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. National Science Foundation (NSF DEB) [1431428, 1136637]; Washington State University; Russian Science Foundation [14-14-00400]; Ministry of education and science of Russia Gos-Zasanie project [1354-2014/51]; Natural Environment Research Council [NE/J00829X/1, 1230750, NE/G019622/1, NE/J010227/1] Funding was provided by the National Science Foundation (NSF DEB #1431428; NSF DEB #1136637) and Washington State University. M. Timofeyev and E. Silow were partially supported by Russian Science Foundation project No 14-14-00400 and Ministry of education and science of Russia Gos-Zasanie project No 1354-2014/51. We are grateful to Marianne Moore, Deniz Ozkundakci, Chris Polashenski and Paula Kankaala for discussions that greatly improved this work. We also gratefully acknowledge the following individuals for contributing to this project: John Anderson, Jill Baron, Rick Bourbonniere, Sandra Brovold, Lluis Camarero, Sudeep Chandra, Jim Cotner, Laura Forsstom, Guillaume Grosbois, Chris Harrod, Klaus D. Joehnk, T.Y. Kim, Daniel Langenhaun, Reet Laugaste, Suzanne McGowan, Virginia Panizzo, Giampaolo Rossetti, R.E.H. Smith, Sarah Spaulding, Helen Tammert, Steve Thackeray, Kyle Zimmer, Priit Zingel and two anonymous reviewers. Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the US Government.

Northern lakes are ice-covered for considerable portions of the year, where carbon dioxide (CO2) can accumulate below ice, subsequently leading to high CO2 emissions at ice-melt. Current knowledge on the regional control and variability of below ice partial pressure of carbon dioxide (pCO(2)) is lacking, creating a gap in our understanding of how ice cover dynamics affect the CO2 accumulation below ice and therefore CO2 emissions from inland waters during the ice-melt period. To narrow this gap, we identified the drivers of below ice pCO(2) variation across 506 Swedish and Finnish lakes using water chemistry, lake morphometry, catchment characteristics, lake position, and climate variables. We found that lake depth and trophic status were the most important variables explaining variations in below ice pCO(2) across the 506 lakes(.) Together, lake morphometry and water chemistry explained 53% of the site-to-site variation in below ice pCO(2). Regional climate (including ice cover duration) and latitude only explained 7% of the variation in below ice pCO(2). Thus, our results suggest that on a regional scale a shortening of the ice cover period on lakes may not directly affect the accumulation of CO2 below ice but rather indirectly through increased mobility of nutrients and carbon loading to lakes. Thus, given that climate-induced changes are most evident in northern ecosystems, adequately predicting the consequences of a changing climate on future CO2 emission estimates from northern lakes involves monitoring changes not only to ice cover but also to changes in the trophic status of lakes.

Lakes play a major role in the global carbon (C) cycle, despite making up a small area of earth’s surface. Lakes receive, transport and process sizable amounts of C, emitting a substantial amount of the greenhouse gases, carbon dioxide (CO2) and methane (CH4), into the atmosphere. Ice-covered lakes are particularly sensitive to climate change, as future reductions to the duration of lake ice cover will have profound effects on the biogeochemical cycling of C in lakes. It is still largely unknown how reduced ice cover duration will affect CO2 and CH4 emissions from ice-covered lakes. Thus, the primary aim of this thesis was to fill this knowledge gap by monitoring the spatial and temporal dynamics of CO2 and CH4 in ice-covered lakes. The results of this thesis demonstrate that below ice CO2 and CH4 were spatially and temporally variable. Nutrients were strongly linked to below ice CO2 and CH4 oxidation variations across lakes. In addition, below ice CO2 was generally highest in small shallow lakes, and in bottom waters. Whilst below ice CH4 was elevated in surface waters near where bubbles from anoxic lake sediment were trapped. During the ice-cover period, CO2 accumulation below ice was not linear, and at ice-melt incomplete mixing of lake waters resulted in a continued CO2 storage in bottom waters. Further, CO2 transported from the catchment and bottom waters contributed to high CO2 emissions. The collective findings of this thesis indicate that CO2 and CH4 emissions from ice-covered lakes will likely increase in the future. The strong relationship between nutrients and C processes below ice, imply that future changes to nutrient fluxes within lakes will influence the biogeochemical cycling of C in lakes. Since catchment and lake sediment C fluxes play a considerable role in below ice CO2 and CH4 dynamics, changes to hydrology and thermal stability of lakes will undoubtedly alter CO2 and CH4 emissions. Nevertheless, ice-covered lakes constitute a significant component of the global C cycle, and as such, should be carefully monitored and accounted for when addressing the impacts of global climate change.