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Freshwater ecosystems face ever increasing environmental and ecological challenges. Recognition of those challenges is not new—over 30 years ago, a series of workshops convened to develop a research agenda (The Freshwater Imperative) to address those challenges. It was recognized that a critical component of solving the problems that freshwater ecosystems face would require greater interdisciplinary collaboration. Here, we discuss the context for the “Revisiting the Freshwater Imperative” workshop and describe how a small‐workshop model can be used effectively to seed longer term collaborative partnerships in support of meeting the challenges freshwater ecosystems face. Some key recommendations include: (1) engage with participants prior to the workshop to maximize efficient use of time and prime divergence‐convergence thinking; (2) promote a bottom‐up participant driven structure; (2) develop an agenda with enough flexibility to accommodate participant driven changes (i.e., a “live” agenda); (3) utilize best practices for facilitating team science; and (4) provide coordinating structure for post‐workshop working group activities.

Lake surfaces are warming worldwide, raising concerns about lake organism responses to thermal habitat changes. Species may cope with temperature increases by shifting their seasonality or their depth to track suitable thermal habitats, but these responses may be constrained by ecological interactions, life histories or limiting resources. Here we use 32 million temperature measurements from 139 lakes to quantify thermal habitat change (percentage of non-overlap) and assess how this change is exacerbated by potential habitat constraints. Long-term temperature change resulted in an average 6.2% non-overlap between thermal habitats in baseline (1978–1995) and recent (1996–2013) time periods, with non-overlap increasing to 19.4% on average when habitats were restricted by season and depth. Tropical lakes exhibited substantially higher thermal non-overlap compared with lakes at other latitudes. Lakes with high thermal habitat change coincided with those having numerous endemic species, suggesting that conservation actions should consider thermal habitat change to preserve lake biodiversity.

Processes regulating the rate of oxygen depletion determine whether hypoxia occurs and the extent to which greenhouse gases accumulate in seasonally ice‐covered lakes. Here, we investigate the oxygen budget of four arctic lakes using high‐frequency data during two winters in three shallow lakes (9–13 m maximal depth) and four winters in 24 m deep main basin of Toolik Lake. Incubation experiments measured sediment metabolism. Volume‐averaged oxygen depletion measured in situ was independent of water temperature and duration of the ice‐covered period. Average rates were between 0.2 and 0.39 g O2m−2d−1in the shallow lakes and between 0.03 and 0.14 g O2m−2d−1in Toolik Lake, with higher rates in smaller lakes with their larger sediment area to volume ratio. Rates decreased to ~ 20%–50% of initial values in late winter in the shallow lakes but less or not at all in Toolik. The lack of a decline in Toolik Lake points to continued oxygen transport to the sediment–water interface where oxygen consumption occurs. In all lakes, lower in situ oxygen depletion than in incubation measurements points toward increasing anoxia in the lower water column depressing loss rates. In Toolik, oxygen loss during early winter was less in years with minimal snow cover. Penetrative convection occurred, which could mix downwards oxygen produced by photosynthesis or excluded during ice formation. Estimates of these terms exceeded photosynthesis measured in sediment incubations. Modeling under ice‐oxygen dynamics requires consideration of optical properties and biological and transport processes that modify oxygen concentrations and distributions.

The concentration of dissolved oxygen in aquatic systems helps to regulate biodiversity1,2, nutrient biogeochemistry3, greenhouse gas emissions4, and the quality of drinking water5. The long-term declines in dissolved oxygen concentrations in coastal and ocean waters have been linked to climate warming and human activity6,7, but little is known about the changes in dissolved oxygen concentrations in lakes. Although the solubility of dissolved oxygen decreases with increasing water temperatures, long-term lake trajectories are difficult to predict. Oxygen losses in warming lakes may be amplified by enhanced decomposition and stronger thermal stratification8,9or oxygen may increase as a result of enhanced primary production10. Here we analyse a combined total of 45,148 dissolved oxygen and temperature profiles and calculate trends for 393 temperate lakes that span 1941 to 2017. We find that a decline in dissolved oxygen is widespread in surface and deep-water habitats. The decline in surface waters is primarily associated with reduced solubility under warmer water temperatures, although dissolved oxygen in surface waters increased in a subset of highly productive warming lakes, probably owing to increasing production of phytoplankton. By contrast, the decline in deep waters is associated with stronger thermal stratification and loss of water clarity, but not with changes in gas solubility. Our results suggest that climate change and declining water clarity have altered the physical and chemical environment of lakes. Declines in dissolved oxygen in freshwater are 2.75 to 9.3 times greater than observed in the world’s oceans6,7and could threaten essential lake ecosystem services2,3,5,11.

The intensity and frequency of storms are projected to increase in many regions of the world because of climate change. Storms can alter environmental conditions in many ecosystems. In lakes and reservoirs, storms can reduce epilimnetic temperatures from wind‐induced mixing with colder hypolimnetic waters, direct precipitation to the lake's surface, and watershed runoff. We analyzed 18 long‐term and high‐frequency lake datasets from 11 countries to assess the magnitude of wind‐ vs. rainstorm‐induced changes in epilimnetic temperature. We found small day‐to‐day epilimnetic temperature decreases in response to strong wind and heavy rain during stratified conditions. Day‐to‐day epilimnetic temperature decreased, on average, by 0.28°C during the strongest windstorms (storm mean daily wind speed among lakes: 6.7 ± 2.7 m s−1, 1 SD) and by 0.15°C after the heaviest rainstorms (storm mean daily rainfall: 21.3 ± 9.0 mm). The largest decreases in epilimnetic temperature were observed ≥2 d after sustained strong wind or heavy rain (top 5thpercentile of wind and rain events for each lake) in shallow and medium‐depth lakes. The smallest decreases occurred in deep lakes. Epilimnetic temperature change from windstorms, but not rainstorms, was negatively correlated with maximum lake depth. However, even the largest storm‐induced mean epilimnetic temperature decreases were typically <2°C. Day‐to‐day temperature change, in the absence of storms, often exceeded storm‐induced temperature changes. Because storm‐induced temperature changes to lake surface waters were minimal, changes in other limnological variables (e.g., nutrient concentrations or light) from storms may have larger impacts on biological communities than temperature changes.

Few coupled lake‐watershed studies examine long‐term effects of climate on the ecosystem function of lakes in a hydrological context. We use 32 years of hydrological and biogeochemical data from a high‐elevation site in the Sierra Nevada of California to characterize variation in snowmelt in relation to climate variability and explore the impact on factors affecting phytoplankton biomass. The magnitude of accumulated winter snow, quantified through basinwide estimates of snow water equivalent (SWE), was the most important climate factor controlling variation in the timing and rate of spring snowmelt. Variations in SWE and snowmelt led to significant differences in lake flushing rate, water temperature, and nitrate concentrations across years. On average in dry years, snowmelt started 25 days earlier and proceeded 7 mm/day slower, and the lake began the ice‐free season with nitrate concentrations ~2 μM higher and water temperatures 9°C warmer than in wet years. Flushing rates in wet years were 2.5 times larger than those in dry years. Consequently, particulate organic matter concentrations, a proxy for phytoplankton biomass, were 5–6 μM higher in dry years. There was a temporal trend of increase in particulate organic matter across dry years that corresponded to lake warming independent of variation in SWE. These results suggest that phytoplankton biomass is increasing as a result of both interannual variability in precipitation and long‐term warming trends. Our study underscores the need to account for local‐scale catchment variability that may affect the accumulation of winter snowpack when predicting climate responses in lakes. The magnitude and water content of winter snowpack explain the majority of variation in timing and rate of snowmeltCatchment snowpack controls interannual variation in phytoplankton biomass by regulating nutrients, water temperature, and flushing ratesHigh interannual variation in mountain snowfall may mask weak long‐term climate‐related changes in phytoplankton biomass

The extent to which snowmelt flowing into ice‐covered lakes spreads horizontally and mixes vertically influences retention of solutes derived from the landscape. To quantify these transport processes and retention, we combine time series temperature and specific conductance measurements in Toolik Lake (Alaska) and its major inflow, with measurements of discharge and meteorology, and profiles of specific conductance, temperature, fluorescence, chlorophyll a, and dissolved organic carbon (DOC) in spring of 3 yr. During early snowmelt, the concentration of DOC in the stream was 750 μM, twice that in the lake. During slow melt (discharge (Q) < 4 m3s−1), the incoming solute‐rich intrusion spread lakewide below the ice. During melt with Q> 6 m3s−1, the incoming water partially flushed the inlet basin and the more dilute water flowed over the original intrusion with a preferential flowpath to the outlet. Penetrative convection was restricted by the increased density gradients from the incoming plume and initially constrained to shallow mixing zones associated with the step changes in density. As ice thickness decreased to less than 1 m, heating caused density instabilities at the base of the intrusions that mixed solutes ∼ 10 m vertically, contributing to retention. Near‐surface layers enriched with DOC persisted for ∼ 10 d during a rapid melt and for over 3 weeks when the melt was slow. Retention, of order 10–20%, also depended on the rapidity of melt and magnitude of discharge.

Our changing climate is having effects on freshwater ecosystems in all seasons, especially winter. High latitude lakes, wetlands, and rivers are experiencing shorter periods of ice cover, and lower latitudes systems that used to freeze are experiencing open water conditions throughout the winter. A 2019 AGU Chapman conference convened aquatic scientists to examine these changes and address the implications of changing winters to aquatic life, chemistry, and physics. Several studies demonstrate decreased ice cover duration than in the past. The removal of an ice “lid” from lakes and rivers impacts the exchange of gases with the atmosphere and the predominant types of metabolism occurring in the waters below, with the potential for more photosynthesis and an increase in oxic versus anoxic metabolism when the lid is removed. Multiple studies indicated an increase in the interannual variability of winters, especially in terms of ice‐cover duration and ice quality. Increased variability may simply be an outcome of a more variable winter climate or small differences in environmental conditions such as temperature that can have strong effects on gas exchange, light transmission, and turbulence when ice forms. A question that merits further consideration is whether and how winters of shorter duration and severity will change the dynamics of freshwater systems. Are there memory or legacy effects that carry over to the next season or year? There is much work to be done to understand how changing winters will impact the biogeochemical behavior of lakes and rivers in the coming decades. Winters at mid‐high latitudes are changing rapidly with important implications for freshwater ecosystems. An AGU Chapman conference, convened in 2019, brought together scientists to outline what we know about the changes that are occurring and what questions need further study. This special issue is a collection of the papers that developed from that workshop. These studies demonstrate that lakes and rivers at mid‐high latitudes are experiencing less ice cover duration and greater variability in year‐to‐year ice cover. Changing ice and snow cover impacts processes occurring under ice in winter by altering the light regime, mixing, biota present, and the availability of oxygen. There are many important research questions regarding winter dynamics that still need to be addressed but one of the most important ones has to do with how changing winters might influence the biota in other seasons. Several of the papers point to interannual variability in winter and seasonal ice cover along with decreased ice cover durationAvailable winter datasets have grown supporting conceptual developments, but increased efforts focused on ecological responses are neededFuture research should examine connections between seasons, deploy submersed sensors year‐round, and use remote sensing approaches Several of the papers point to interannual variability in winter and seasonal ice cover along with decreased ice cover duration Available winter datasets have grown supporting conceptual developments, but increased efforts focused on ecological responses are needed Future research should examine connections between seasons, deploy submersed sensors year‐round, and use remote sensing approaches

We characterized patterns of nighttime ecosystem respiration (ER) in 23 globally‐distributed lakes using free‐water changes in dissolved oxygen (DO). We considered three alternative models of ER to describe overnight changes in DO: constant ER, linearly declining ER, or logistically declining ER. Variation in ER was widespread. Each model of respiration was found to best fit observed overnight DO dynamics with some degree of frequency in every lake. Constant ER occurred with an average frequency of 62% across lakes and was most commonly the best description of DO dynamics. It never occurred with a frequency < 19% in any lake and ranged up to 90% in one dystrophic lake. The average frequency across lakes with which the linear and logistic models occurred was 21% and 17%, respectively, although they ranged as low as 3% and as high as 42% in some lakes. Although data limitations restricted the majority of our analysis to summer months, annual data records from five lakes suggest there is little seasonal variation in the frequency with which patterns occurred. As our conceptual model predicted, the frequency of constant ER among lakes increased and logistic ER decreased along a gradient of increasing terrestrial influence. However, despite significant correlations with total phosphorus and photosynthetically active radiation, the effect of lake productivity on the frequency of patterns of ER among lakes was less clear. This study suggests that diel variability in ER results from complex interactions between different components of respiration and the forcing factors that govern them.

The impacts of wildfire smoke on lake habitats remains unclear. We determined the metabolic response to smoke in the epi‐pelagic and two littoral habitats in Castle Lake, California. We compared light regime, gross primary production, ecosystem respiration, and net ecosystem production in years with and without smoke. During the smoke period incident ultraviolet‐B (UV‐B) radiation and photosynthetically active radiation (PAR) decreased by 53% and 28%, respectively, while the water column extinction coefficient of UV‐B and PAR increased by 20% and 18% respectively. Epi‐pelagic productivity increased during smoke cover because of decreased solar inputs. PAR values remained sufficient to saturate productivity, suggesting observed differences were primarily the result of changes in UV‐B. Littoral‐benthic productivity did not change, possibly reflecting adaptation to high‐intensity UV‐B light in these habitats. Our results highlight the importance of understanding how prolonged wildfire smoke alters the amount of energy produced from specific habitats in lakes. Large wildfires are increasing worldwide, but the impacts of wildfire smoke on lakes are unknown. We determined how photosynthetic activity and organism's respiration respond to smoke in the offshore and nearshore habitats in a mountain lake. Smoke limited the damage to algae caused by ultraviolet‐B radiation, resulting in increased offshore photosynthesis. Photosynthesis in the nearshore habitat did not change during smoke cover, possibly because organisms in shallow habitats have adapted to the higher intensity of damaging ultraviolet‐B radiation. Our results highlight the importance of understanding how prolonged smoke conditions can alter lakes' energy flows and food webs. Wildfire smoke reduced ultraviolet‐B and photosynthetically active radiationLake habitats respond differently to changes in light from smoke, pelagic primary production increased with no change to littoral metabolismSmoke alters energy flows with implications for food web structure in lakes Wildfire smoke reduced ultraviolet‐B and photosynthetically active radiation Lake habitats respond differently to changes in light from smoke, pelagic primary production increased with no change to littoral metabolism Smoke alters energy flows with implications for food web structure in lakes

AbstractWe characterized the effect of an extreme rain event on the biogeochemistry and ecosystem metabolism of an oligotrophic Sierra Nevada (California) lake. During a 10 hour period of an autumn season rainfall event, lake discharge increased from <1.0 L s−1to over 3000 L s−1, reaching a rate one order of magnitude higher than found during peak snow-tmelt. Large quantities of terrestrial particulate and dissolved organic matter were washed into the lake. An entire season of autochthonous dissolved organic carbon (DOC) was flushed and replaced by allochthonous DOC as light attenuation increased by >300%. The resulting truncation of the photic zone, reduction of water column chlorophyll-a, and increase in particulate and dissolved organic matter available to microbes resulted in a 47% reduction in whole lake gross primary production and 30% increase in respiration relative to average autumn values. As a consequence, the lake went from being slightly autotrophic to strongly heterotrophic. If rain events increase in frequency, as many climate change models predict, increased terrestrial inputs to Sierran lakes may result in more frequent periods of reduced primary production, increased periods of hypoxia and anoxia, and an ecosystem shift toward net heterotrophy during the ice-free season.

Warming winters will reduce ice cover and change under‐ice conditions in temperate mountain lakes, where snow contributes most of winter cover on lakes. Snow‐dominated mountain lakes are abundant and highly susceptible to climate warming, yet we lack an understanding of how climate variation and local attributes influence winter processes. We investigated climatic and intrinsic controls on ice phenology, water temperature, and bottom‐water dissolved oxygen (DO) in 15 morphologically diverse lakes in the Sierra Nevada and Klamath Mountains of California, USA, using high‐frequency measurements from multiple (2–5) winters. We found that ice phenology was determined by winter climate variables (snowfall and air temperature) that influence ice‐off timing, whereas ice‐on timing was relatively invariant among years. Lake size and morphology mediated the effect of climate on lake temperature and DO dynamics in early and late winter. Rates of hypolimnetic DO decline were highest in small, shallow lakes, and were unrelated to water temperature. Temperature and oxygen dynamics were more variable in small lakes because heavy snowfall caused ice submergence, mixing, and DO replenishment that affected the entire water column. As the persistence of snow declines in temperate mountain regions, autumn, and spring climatic conditions are expected to gain importance in regulating lake ice phenology. Water temperature and DO will likely increase in most lakes during winter as snowpack declines, but morphological attributes such as lake size will determine the sensitivity of ice phenology and under‐ice processes to climate change. Mountain lakes in temperate regions are at risk of losing ice cover as winters become warmer and snowfall decreases. Small mountain lakes are numerous and provide important habitats for aquatic species, yet we currently lack an understanding of what controls ice duration and under‐ice conditions in these snow‐dominated systems because most are remote and inaccessible. We used hourly measurements within a diverse set of California mountain lakes across multiple winters to explore how climate variables and lake or watershed characteristics influence the duration and timing of ice cover, as well as under‐ice temperature and oxygen concentrations. We found that winter snowfall and air temperature controlled the timing of ice breakup and the length of ice duration. Lake size and shape controlled oxygen and temperature immediately after ice formation, but late winter conditions depended on winter climate variables that dictated ice cover duration. Oxygen depletion rates and the prevalence of low oxygen conditions were greater in small, shallow lakes than in large, deep lakes. As winters warm, most mountain lakes will have shorter ice cover, warmer water temperatures, and increased oxygen concentrations, but small and large lakes will respond differently to the loss of snow and ice cover. Winter air temperature and snowfall drive ice‐off timing and ice phenology of temperate mountain lakesLake size and morphology mediate how climate affects water temperature and dissolved oxygen dynamics during early and late winterRates of hypolimnetic oxygen decline are highest in shallow lakes and do not vary with water temperature Winter air temperature and snowfall drive ice‐off timing and ice phenology of temperate mountain lakes Lake size and morphology mediate how climate affects water temperature and dissolved oxygen dynamics during early and late winter Rates of hypolimnetic oxygen decline are highest in shallow lakes and do not vary with water temperature

The extent to which terrestrial organic matter supports aquatic consumers remains uncertain because factors regulating resource flows are poorly understood. We sampled 12 lakes throughout the Sierra Nevada (California, USA) spanning large gradients in elevation and size to evaluate how watershed attributes and lake morphometry influence resource flows to lake carbon pools and zooplankton. We found that the size and composition of carbon pools in lakes were often more strongly determined by watershed or lake features rather than by elevational position. Using three different tracers of resource origin (δ13C, Δ14C, C:N ratio), we found terrestrial contributions to most lake resource pools (dissolved organic carbon, particulate organic matter (POM), sediments) and pelagic consumers (zooplankton) were more strongly related to local‐scale watershed features such as vegetation cover or watershed area: lake area rather than to elevation. Landscape patterns in multiple tracers indicated consistent contribution of within‐lake C sources to bulk resource pools across elevations (POM, sediments, zooplankton). δ13C‐enrichment of lake C pools and overlap with δ13C of terrestrial resources can arise due to reduced fractionation of 13C by phytoplankton under CO2limitation, therefore we recommend careful consideration of potential environmental drivers when interpreting among‐lake patterns in δ13C. Our findings emphasize the importance of local‐scale variation in mediating terrestrial contributions to lake food webs. Lake and landscape features are better predictors of the composition of lake resource pools than elevationAllochthony gradients in mountain systems can vary substantially among different lake resource poolsAquatic producers are the dominant contribution to lake carbon pools across a large gradient in landscape vegetation cover Lake and landscape features are better predictors of the composition of lake resource pools than elevation Allochthony gradients in mountain systems can vary substantially among different lake resource pools Aquatic producers are the dominant contribution to lake carbon pools across a large gradient in landscape vegetation cover