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The metabolic regimes of flowing waters

The processes and biomass that characterize any ecosystem are fundamentally constrained by the total amount of energy that is either fixed within or delivered across its boundaries. Ultimately, ecosystems may be understood and classified by their rates of total and net productivity and by the seasonal patterns of photosynthesis and respiration. Such understanding is well developed for terrestrial and lentic ecosystems but our understanding of ecosystem phenology has lagged well behind for rivers. The proliferation of reliable and inexpensive sensors for monitoring dissolved oxygen and carbon dioxide is underpinning a revolution in our understanding of the ecosystem energetics of rivers. Here, we synthesize our current understanding of the drivers and constraints on river metabolism, and set out a research agenda aimed at characterizing, classifying and modeling the current and future metabolic regimes of flowing waters.

Limnology and Oceanography↗

Associations between cyanobacteria and indices of secondary production in the western basin of Lake Erie

Large lakes provide a variety of ecological services to surrounding cities and communities. Many of these services are supported by ecological processes that are threatened by the increasing prevalence of cyanobacterial blooms which occur as aquatic ecosystems experience cultural eutrophication. Over the past 10 yr, Lake Erie experienced cyanobacterial blooms of increasing severity and frequency, which have resulted in impaired drinking water for the surrounding communities. Cyanobacterial blooms may impact ecological processes that support other services, but many of these impacts have not been documented. Secondary production (production of primary consumers) is an important process that supports economically important higher trophic levels. Cyanobacterial blooms may influence secondary production because cyanobacteria are a poor‐quality food resource and cyanotoxins may be harmful to consumers. Over 3 yr at 34 sites across the western basin of Lake Erie, we measured three indices of secondary production that focus on the dominant bivalve taxa: (1) growth of a native unionid mussel, (2) the size of young‐of‐year dreissenid mussels, and (3) the mass of colonizing animals on a Hester‐Dendy sampler. Associations between these indices and cyanobacterial data were estimated to assess whether cyanobacteria are associated with variation in secondary production in the western basin of Lake Erie. The results suggest cyanobacterial abundance alone is only weakly associated with secondary production, but that cyanotoxins have a larger effect on secondary production. Given recurring late‐summer cyanobacterial blooms, this impact on secondary production has the potential to undermine Lake Erie's ability to sustain important ecosystem services.

Lake Erie↗

Evidence for regional nitrogen stress on chlorophyll a in lakes across large landscape and climate gradients

Nitrogen (N) and phosphorus (P) commonly stimulate phytoplankton production in lakes, but recent observations from lakes from an agricultural region suggest that nitrate may have a subsidy‐stress effect on chlorophyll a (Chl a ). It is unclear, however, how generalizable this effect might be. Here, we analyzed a large water quality dataset of 2385 lakes spanning 60 regions across 17 states in the Northeastern and Midwestern U.S. to determine if N subsidy‐stress effects on phytoplankton are common and to identify regional landscape characteristics promoting N stress effects in lakes. We used a Bayesian hierarchical modeling framework to test our hypothesis that Chl a –total N (TN) threshold relationships would be common across the central agricultural region of the U.S. (“the Corn Belt”), where lake N and P concentrations are high. Data aggregated across all regions indicated that high TN concentrations had a negative effect on Chl a in lakes with concurrent high total P. This large‐scale pattern was driven by relationships within only a subset of regions, however. Eight regions were identified as having Chl a –TN threshold relationships, but only two of these regions located within the Corn Belt clearly demonstrated this subsidy‐stress relationship. N stress effects were not consistent across other intense agricultural regions, as we hypothesized. These findings suggest that interactions among regional land use and land cover, climate, and hydrogeology may be important in determining the synergistic conditions leading to N subsidy‐stress effects on lake phytoplankton.

Limnology and Oceanography↗

Why large cells dominate estuarine phytoplankton

Surveys across the world oceans have shown that phytoplankton biomass and production are dominated by small cells (picoplankton) where nutrient concentrations are low, but large cells (microplankton) dominate when nutrient-rich deep water is mixed to the surface. I analyzed phytoplankton size structure in samples collected over 25 yr in San Francisco Bay, a nutrient-rich estuary. Biomass was dominated by large cells because their biomass selectively grew during blooms. Large-cell dominance appears to be a characteristic of ecosystems at the land–sea interface, and these places may therefore function as analogs to oceanic upwelling systems. Simulations with a size-structured NPZ model showed that runs of positive net growth rate persisted long enough for biomass of large, but not small, cells to accumulate. Model experiments showed that small cells would dominate in the absence of grazing, at lower nutrient concentrations, and at elevated (+5°C) temperatures. Underlying these results are two fundamental scaling laws: (1) large cells are grazed more slowly than small cells, and (2) grazing rate increases with temperature faster than growth rate. The model experiments suggest testable hypotheses about phytoplankton size structure at the land–sea interface: (1) anthropogenic nutrient enrichment increases cell size; (2) this response varies with temperature and only occurs at mid-high latitudes; (3) large-cell blooms can only develop when temperature is below a critical value, around 15°C; (4) cell size diminishes along temperature gradients from high to low latitudes; and (5) large-cell blooms will diminish or disappear where planetary warming increases temperature beyond their critical threshold.

Limnology and Oceanography↗

Combining nutrient, productivity, and landscape-based regressions improves predictions of lake nutrients and provides insight into nutrient coupling at macroscales

Empirical nutrient models that describe lake nutrient, productivity, and water clarity relationships among lakes play a prominent role in limnology. Landscape-based regressions are also used to understand macroscale variability of lake nutrients, clarity, and productivity (hereafter referred to as nutrient-productivity). Predictions from both models are used to inform eutrophication management globally. To date, these two classes of models are generally conducted separately, which ignores the known dependencies among nutrient-productivity variables. We present a statistical model that integrates nutrient-productivity and landscape-based regressions—where lake nutrients, productivity, and clarity variables are modeled jointly. We fitted a joint nutrient-productivity model to over 7000 lakes with three nutrients (total phosphorus, total nitrogen, nitrate concentrations), chlorophyll a concentrations, and Secchi disk depth as response variables and landscape features as predictor variables. Because lakes in different regions respond to landscape features differently, we focused our analysis on two subregions with different dominant land uses, the agricultural Midwest and the forested Northeast U.S. Predictive performance was enhanced by modeling nutrient-productivity variables jointly. We also found strong evidence that nutrient-productivity variables were coupled, and that only nitrate may be decoupled from other nutrient-productivity variables in the forested region. We speculate that these regional differences may be related to differences in the strength of biogeochemical cycles and stoichiometric controls between these regions. Jointly modeling nutrient-productivity variables in lakes effectively integrates the two dominant approaches for studying lakes nutrient-productivity relationships and provides novel insight into macroscale patterns of the coupling of nutrients, chlorophyll, and water clarity in lakes.

Limnology and Oceanography↗

Changes in long-term water quality of Baltimore streams are associated with both gray and green infrastructure

The steadily rising global urban population has placed substantial strain on urban water quality, and this strain is projected to increase for the foreseeable future. Considerable attention has been given to the hydrological and physico-chemical effects of urbanization on stream ecosystems. However, due to the relative infancy of the field of urban ecology, long-term water quality analyses in urban streams are sparse. Using a 15-year stream chemistry monitoring record from Baltimore, MD, USA, we quantified long-term trends in nitrate, phosphate, total nitrogen, total phosphorus, chloride, and sulfate export at several sites along a rural-urban gradient. We found no significant change in solute export at most sites, although we did find specific patterns of interest for certain solutes. For example, nitrogen export declined at the most headwater urban site, while phosphorus export declined at the most downstream urban site. Coupling long-term monitoring with data on gray and green infrastructure management throughout the landscape, we established relationships between solute export at the most downstream urban monitoring site and sanitary sewer overflows (SSOs), best management practice (BMP) implementation, and road salt application rates. Phosphorus export was correlated with BMP implementation in the watershed, whereas nitrogen export was related to SSOs. Despite highly urbanized watersheds, water quality does not appear to be declining at most of these sites, suggesting that current management may have limited further impairment. Results of our study suggest that both gray and green infrastructure are key for maintaining and improving water quality in this highly urbanized watershed.

Maryland↗

Patterns, pace and processes of water-quality variability: Examples from a long-studied estuary

Environmental time series have rich information content that is invaluable for measuring and understanding changes over time and guiding policies to manage change. I extracted information from measurements of 10 water‐quality constituents in upper San Francisco Bay from 1975 to 2016, one of the longest observational records in a U.S. estuary. Changes were detected at every time scale captured by monthly sampling. Long‐term trends included increased ammonium (+53%), nitrate + nitrate (+50%), silicate (+14%), Secchi depth (+42%), and decreased chlorophyll a (Chl a ) (−74%) and suspended particulate matter (−45%). Changes at the decadal scale included abrupt shifts (Chl a , nitrate + nitrite) and oscillations between shorter trends of increase and decrease (Secchi depth, phosphate). Long‐term trends were not expressed equally across all seasons, and seasonal patterns of change varied across constituents. These examples illustrate key features of environmental variability at the land–sea interface: (1) water‐quality components change continually at time scales from months to decades; (2) patterns of seasonal, multiyear, and multidecadal change are complex and vary across constituents; (3) primary drivers of change are freshwater inflow, the master regulator of estuarine dynamics, and human activities such as river damming, water diversions, wastewater discharge, environmental policies, and species introductions; (4) extracting the full information content of time series requires multiple analyses, each revealing a different layer of insight into how changes develop over time; (5) water‐quality variability is nonstationary, so future changes cannot be forecast reliably; (6) repeated observation is an essential method of Earth system science with applications in the design and performance measures of environmental policies.

California↗

Estimating lake–climate responses from sparse data: An application to high elevation lakes

Although many studies demonstrate lake warming, few document trends from lakes with sparse data. Diel and seasonal variability of surface temperatures limit conventional trend analyses to datasets with frequent repeated observations. Thus, remote lakes, including many high elevation lakes, are underrepresented in trend analyses. We used a Bayesian technique to analyze sparse data that explicitly incorporated diel and seasonal variability. This approach allowed us to estimate lake warming in a region of limited knowledge: high elevation lakes (> 2100 m ASL) of the Southern Rocky Mountains, U.S.A. The analysis allowed for inclusion of lakes with few repeated measurements, and observations made before 1980 when more intensive lake monitoring began. We accumulated the largest dataset of high elevation lake temperatures analyzed to date. Data from 590 high elevation lakes in the Southern Rocky Mountains showed a 0.13°C decade −1 increase in surface temperatures and a 14% increase in seasonal degree days since 1955. This result is lower than other regional and global estimates of lake warming; however, it is similar to other high elevation lake studies. Our approach can be applied to other understudied regions, increasing our overall understanding of the effects of climate change on lakes and their temporal dynamics.

Colorado↗

Metabolic rhythms in flowing waters: An approach for classifying river productivity regimes

Although seasonal patterns of ecosystem productivity have been extensively described and analyzed with respect to their primary forcings in terrestrial and marine systems, comparatively little is known about these same processes in rivers. However, it is now possible to perform a large‐scale synthesis on the patterns and drivers of river productivity regimes because of the recent sensor advances allowing for near‐continuous estimates of river productivity. Here, we explore a dataset of 47 U.S. rivers to examine whether there are characteristic river productivity regimes. We use classification approaches to develop a typology of productivity regimes and then use these regimes to examine differences with respect to potential controls of productivity. We identified two distinct metabolic regimes, which we named Summer Peak and Spring Peak Rivers, within our dataset. These regimes meaningfully differed in both the timing and magnitude of productivity and were robust to different approaches to classification. We also found that several variables, including watershed area and characteristics of water temperature or discharge, were able to predict the class membership of these regimes with modest accuracy. Our results support the presence of characteristic metabolic regimes and suggests that these regimes may have common sets of environmental controls. We present classification as one approach to begin exploring the productivity regimes of rivers. The strength of our approach is that it fully leverages these newly available high‐frequency productivity estimates to create classes that can be used to draw inferences about how the controls of river productivity differ between or within systems.

Limnology and Oceanography↗

Improving estimates of coral reef construction and erosion with in-situ measurements

The decline in living coral since the 1970s has conspicuously slowed reef construction on a global scale, but the related process of reef erosion is less visible and not often quantified. Here we present new data on the constructional and deconstructional side of the carbonate-budget equation in the Florida Keys, U.S.A. We documented Orbicella spp. calcification rates at four offshore reefs and quantified decadal-scale rates of Orbicella-reef erosion at a mid-shore patch reef. Using Orbicella coral heads fitted with permanent markers in 1998, we measured reef-elevation loss at 28 stations over 17.3 years to estimate a mean erosion rate of -5.5 (± 3.2, SD) mm yr-1. This loss equates to an erosion rate of -8.2 (± 4.8, SD) kg m-2 yr-1 on dead Orbicella colonies, or -6.6 kg m-2 yr-1 when adjusted reef-wide. Calculating net carbonate production using a census-based approach on the same patch reef in 2017, we estimated a reef-wide bioerosion rate of -1.9 (± 2.0, SD) kg m-2 yr-1, and a net carbonate production rate of 0.5 (± 0.3, SD) kg m-2 yr-1. Substituting the erosion rate we estimated with the markers would suggest that net carbonate production at this patch reef was lower and negative, -4.2 kg m-2 yr-1. This divergence could be a function of high erosion rates measured on the tops of Orbicella colonies, which may be preferentially targeted by parrotfish. Nonetheless, our study suggests the need for new field data to improve estimates of reef-structure persistence as coral reefs continue to degrade.

Florida↗

Light exposure along particle flowpaths in large rivers

Sunlight is a critical resource in aquatic systems driving photosynthesis, photodegradation of organic matter and contaminants, animal behavior, and the activity of human pathogens. In rivers, solutes, materials, and organisms are turbulently mixed across the water column during downstream transport and exposed to highly variable sunlight. However, there are no measurements of suspended particles' sunlight exposure during downstream transport to characterize this variability, and it is unclear if current measurement approaches and optical theory capture the light exposure of suspended particles. We deployed neutrally buoyant drifters and stationary buoys in the Upper Mississippi (WI, U.S.A.) and Neuse Rivers (NC, U.S.A.) to measure underwater sunlight from the perspective of suspended particles. In our study sites, underwater sunlight varied more along flowpaths measured by drifters than over time measured by fixed‐site buoys; sunlight exposure along flowpaths was dominated by bursts of light (sunflecks) that accounted for 62–99% of the cumulative sunlight exposure; and modeled sunlight exposure using optical theory was consistently 56–1700% higher than measured sunlight exposure along flowpaths. Our results suggested that suspended particles in the study reaches experienced darker conditions than predicted and have important implications for how to quantify underwater sunlight in rivers.

North Carolina, Wisconsin↗

Calcite precipitation in Lake Powell reduces alkalinity and total salt loading to the Lower Colorado River Basin

Reservoirs can retain and transform carbon, nitrogen, phosphorus, and silica, but less is known about their effects on other biogeochemically relevant solutes. The salinization of freshwater ecosystems is a growing concern in many regions, and the role of reservoirs in salinity transport is an important research frontier. Here, we examine how a large desert southwest reservoir, Lake Powell, has altered the downstream transport of total dissolved solids (TDSs) as well as the dominant cations and anions comprising the TDS pool ( , , and Ca 2+ ). Average downstream TDS concentrations have declined significantly since river impoundment and seasonal fluctuations in TDS concentrations have become more modulated, but year to year variation in TDS concentrations has remained similar. While some of the reductions in TDS concentration can be attributed to watershed management, we find that Lake Powell retains about 10% of the TDS loaded to the system (1991 Mg TDS d −1 ). Much of this retention is occurring in the forms of calcium and bicarbonate, likely via calcite precipitation, and is equivalent to an average burial of 522 mg C m −2 d −1 , thus reducing the alkalinity of downstream water. Flow‐weighted modeling suggests that, in the absence of Lake Powell, downstream salinity limits would be surpassed at the outflow to Lake Powell 41% of the time (vs. 0% of the time currently). Understanding the dominant mechanisms regulating solute transport through the reservoir is important given the relevance for downstream drinking water and irrigation concerns, biogeochemical cycling, and the high potential for reduced flows in the future.

Arizona, Colorado, New Mexico, Utah, Wyoming↗

Hydrologic connectivity determines dissolved organic matter biogeochemistry in northern high-latitude lakes

Northern high‐latitude lakes are undergoing climate‐induced changes including shifts in their hydrologic connectivity with terrestrial ecosystems. How this will impact dissolved organic matter (DOM) biogeochemistry remains uncertain. We examined the drivers of DOM composition for lakes in the Yukon Flats Basin in Alaska, an arid region of low relief that is characteristic of over one‐quarter of circumpolar lake area. Utilizing the vascular plant biomarker lignin, chromophoric dissolved organic matter (CDOM), and ultrahigh‐resolution mass spectrometry, we interpreted DOM compositional changes using lake‐water stable isotope (δ 18 O‐H 2 O) composition as a proxy for lake hydrologic connectivity with the landscape. We observed a relative decrease in CDOM in more hydrologically isolated lakes (enriched δ 18 O‐H 2 O) without a corresponding decrease in dissolved organic carbon (DOC) concentration. Although DOC and CDOM were weakly correlated, a significant positive relationship between lignin and CDOM ( r 2 = 0.67) demonstrates that optical parameters are useful for estimating lignin concentration and thus vascular plant contribution to lake DOM. Indicators of allochthonous DOM, including lignin carbon normalized yields, CDOM aromaticity proxies, and relative abundances of polyphenolic and condensed aromatic compound classes, were negatively correlated with δ 18 O‐H 2 O ( r 2 > 0.45), suggesting there is little allochthonous DOM supplied to many of these hydrologically isolated lakes. We conclude that decreased lake hydrologic connectivity, driven by ongoing climate change (i.e., decreased precipitation, warming temperatures), will reduce allochthonous DOM contributions and shift lakes toward lower CDOM systems with ecosystem‐scale ramifications for heat transfer, photochemical reactions, productivity, and ultimately their biogeochemical function.

Alaska↗

Aufeis fields as novel groundwater-dependent ecosystems in the arctic cryosphere

River aufeis (ow′ fīse) are widespread features of the arctic cryosphere. They form when river channels become locally restricted by ice, resulting in cycles of water overflow and freezing and the accumulation of ice, with some aufeis attaining areas of ~ 25 + km 2 and thicknesses of 6+ m. During winter, unfrozen sediments beneath the insulating ice layer provide perennial groundwater‐habitat that is otherwise restricted in regions of continuous permafrost. Our goal was to assess whether aufeis facilitate the occurrence of groundwater invertebrate communities in the Arctic. We focused on a single aufeis ecosystem (~ 5 km 2 by late winter) along the Kuparuk River in arctic Alaska. Subsurface invertebrates were sampled during June and August 2017 from 50 3.5‐cm diameter PVC wells arranged in a 5 × 10 array covering ~ 40 ha. Surface invertebrates were sampled using a quadrat approach. We documented a rich assemblage of groundwater invertebrates (49 [43–54] taxa, [95% confidence limits]) that was distributed below the sediment surface to a mean depth of ~ 69 ± 2 cm ( ± 1 SE) throughout the entire well array. Although community structure differed significantly between groundwater and surface habitats, the taxa richness from wells and surface sediments (43 [35–48] taxa) did not differ significantly, which was surprising given lower richness in subsurface habitats of large, riverine gravel‐aquifer systems shown elsewhere. This is the first demonstration of a rich and spatially extensive groundwater fauna in a region of continuous permafrost. Given the geographic extent of aufeis fields, localized groundwater‐dependent ecosystems may be widespread in the Arctic.

Alaska↗

Drivers and projections of ice phenology in mountain lakes in the western United States

Climate change is causing rapid warming and altered precipitation patterns in mountain watersheds, both of which influence the timing of ice breakup in mountain lakes. To enable predictions of ice breakup in the future, we analyzed a dataset of mountain lake ice breakup dates derived from remote sensing and historical downscaled climate data. We evaluated drivers of ice breakup, constructed a predictive statistical model, and developed projections of mountain lake ice breakup date with global climate models. Using Random Forest analysis, we determined that winter and spring cumulative snow fraction (portion of precipitation falling as snow) and air temperature are the strongest predictors of ice breakup on mountain lakes. Interactions between precipitation, cumulative winter air temperature and lake surface area indicate that shifts in air temperature and precipitation affect smaller lakes (< 2 km 2 ) more than larger lakes (> 2–10 km 2 ). A linear mixed effects model (RMSE of 18 d), applied with an ensemble of 15 global climate models, projected that end-of-century ice breakup in mountain lakes will be earlier by 25 ± 4 and 61 ± 5 (mean ± SE) days for representative concentration pathways 4.5 and 8.5, respectively.

California, Idaho, Oregon, Washington↗

From forests to fish: Mercury in mountain lake food webs influenced by factors at multiple scales

Mountain lakes, while seemingly pristine, have been subjected to historical fish stocking practices and exposure to atmospherically deposited contaminants like mercury. Mercury bioaccumulation in these ecosystems varies widely due to strong environmental gradients, and there are complex, hierarchical factors that affect mercury transport and loading, methylmercury production, and food web biomagnification. We sought to assess how representative variables associated with watershed, lake, and food web‐scale processes—specifically, catchment tree cover, lake benthic primary production, and fish diet, respectively—are associated with mercury concentrations in mountain lake fish. Mean fish mercury concentrations varied threefold between lakes, with nearshore tree cover and fish diet accounting for the most variance in fish mercury. Tree cover was likely positively correlated to fish Hg due to its contributions to local deposition and its effect on lake biogeochemistry. Fish with benthic diets tended to have higher mercury concentrations, illustrating that food web processes are an important consideration when investigating drivers of contaminant bioaccumulation. Our results suggest that both landscape and ecological factors are determinants of fish mercury bioaccumulation, and thus variables at multiple scales should be considered when managing mountain lake food webs for mercury exposure risk.

Washington↗

Pore water exchange-driven inorganic carbon export from intertidal salt marshes

Respiration in intertidal salt marshes generates dissolved inorganic carbon (DIC) that is exported to the coastal ocean by tidal exchange with the marsh platform. Understanding the link between physical drivers of water exchange and chemical flux is a key to constraining coastal wetland contributions to regional carbon budgets. The spatial and temporal (seasonal, annual) variability of marsh pore water exchange and DIC export was assessed from a microtidal salt marsh (Sage Lot Pond, Massachusetts). Spatial variability was constrained from 224 Ra : 228 Th disequilibria across two hydrologic units within the marsh sediments. Disequilibrium between the more soluble 224 Ra and its sediment-bound parent 228 Th reveals significant pore water exchange in the upper 5 cm of the marsh surface (0–36 L m −2 d −1 ) that is most intense in low marsh elevation zones, driven by tidal overtopping. Surficial sediment DIC transport ranges from 0.0 to 0.7 g C m −2 d −1 . The sub-surface sediment horizon intersected by mean low tide was disproportionately impacted by tidal pumping (20–80 L m −2 d −1 ) and supplied a seasonal DIC flux of 1.7–5.4 g C m −2 d −1 . Export exceeded 10 g C m −2 d −1 for another marsh unit, demonstrating that fluxes can vary substantially across salt marshes under similar conditions within the same estuary. Seasonal and annual variability in marsh pore water exchange, constrained from tidal time-series of radium isotopes, was driven in part by variability in mean sea level. Rising sea levels will further inundate high marsh elevation zones, which may lead to greater DIC export.

Massachusetts↗

The extent and variability of storm‐induced temperature changes in lakes measured with long‐term and high‐frequency data

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 5 th percentile 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.

Limnology and Oceanography↗