Experimental Ecosystem Eutrophication Causes Offsetting Effects on Emissions of CO2, CH4, and N2O from Agricultural Reservoirs.

Despite decades of research and management efforts, eutrophication remains a persistent threat to inland waters. As nutrient pollution intensifies in the coming decades, the implications for aquatic greenhouse gas (GHG) emissions are poorly defined, particularly the responses of individual GHGs: carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O). The biogeochemical controls of each gas can differ, making it difficult to predict the overall effect of nutrient pollution on the net radiative forcing of aquatic ecosystems. Here, we induced eutrophication of small nitrogen (N)-limited agricultural reservoirs and measured changes in diffusive GHG emissions within a before-after-control-impact (BACI) study design during June to September 2021. Each gas exhibited a unique response to 300% increases in primary production, with a shift from an overall CO2 source to a sink, a modest increase in N2O flux, and, unexpectedly, no significant change in CH4 emissions. The lack of net directional change in CO2-equivalent GHG emissions in fertilized reservoirs during the summer contrasts findings from empirical studies of eutrophic lakes. Our findings illustrate the difficulty in extrapolating among different sized ecosystems and suggest that forecast 2-fold increases in agricultural N fertilization by 2050 may not result in consistently elevated GHG emissions during summer, at least from small reservoirs in continental grassland regions.

Salinity causes widespread restriction of methane emissions from small inland waters.

Inland waters are one of the largest natural sources of methane (CH4), a potent greenhouse gas, but emissions models and estimates were developed for solute-poor ecosystems and may not apply to salt-rich inland waters. Here we combine field surveys and eddy covariance measurements to show that salinity constrains microbial CH4 cycling through complex mechanisms, restricting aquatic emissions from one of the largest global hardwater regions (the Canadian Prairies). Existing models overestimated CH4 emissions from ponds and wetlands by up to several orders of magnitude, with discrepancies linked to salinity. While not significant for rivers and larger lakes, salinity interacted with organic matter availability to shape CH4 patterns in small lentic habitats. We estimate that excluding salinity leads to overestimation of emissions from small Canadian Prairie waterbodies by at least 81% ( ~ 1 Tg yr-1 CO2 equivalent), a quantity comparable to other major national emissions sources. Our findings are consistent with patterns in other hardwater landscapes, likely leading to an overestimation of global lentic CH4 emissions. Widespread salinization of inland waters may impact CH4 cycling and should be considered in future projections of aquatic emissions.

Impact of wastewater treatment upgrade and nitrogen removal on bacterial communities and their interactions in eutrophic prairie streams.

Eutrophication can impact bacteria by altering fluxes and processing of nutrients and organic matter. However, relatively little is known of how bacterial communities, diversity, and interactions with phytoplankton might respond to nutrient management. We used 16S rRNA amplicon sequencing to compare bacterial assemblages in the water column upstream (control) and downstream (impact) of a wastewater treatment plant (WWTP) located on a eutrophic prairie stream. Sampling occurred before (2012) and after (2018) the 2016 biological nutrient removal (BNR) upgrade that removed >90% of nitrogen (N, mainly NH4+). Multivariate ordination suggested that effluent-impacted bacterial communities were associated mainly with elevated NH4+ concentrations before the upgrade, whereas those after BNR were characteristic of reference systems (low NO3-, diverse regulation). Genera such as Betaproteobacteria and Rhodocyclacea were abundant at impacted sites in 2012, whereas Flavobacterium and a potential pathogen (Legionella) were common at impacted sites in 2018. Nitrifier bacteria (Nitrospira and Nitrosomonas) were present but rare at all sites in 2012, but recorded only downstream of the WWTP in 2018. Generalized additive models showed that BNR reduced bacterial diversity, with ∼70% of the deviance in diversity explained by hydrology, pH, nutrients, and phytoplankton abundance. Overall, NH4+ removal reduced symptoms of cultural eutrophication in microbe assemblages.

Controls of thermal response of temperate lakes to atmospheric warming.

Atmospheric warming heats lakes, but the causes of variation among basins are poorly understood. Here, multi-decadal profiles of water temperatures, trophic state, and local climate from 345 temperate lakes are combined with data on lake geomorphology and watershed characteristics to identify controls of the relative rates of temperature change in water (WT) and air (AT) during summer. We show that differences in local climate (AT, wind speed, humidity, irradiance), land cover (forest, urban, agriculture), geomorphology (elevation, area/depth ratio), and water transparency explain >30% of the difference in rate of lake heating compared to that of the atmosphere. Importantly, the rate of lake heating slows as air warms (P < 0.001). Clear, cold, and deep lakes, especially at high elevation and in undisturbed catchments, are particularly responsive to changes in atmospheric temperature. We suggest that rates of surface water warming may decline relative to the atmosphere in a warmer future, particularly in sites already experiencing terrestrial development or eutrophication.

The varved succession of Crawford Lake, Milton, Ontario, Canada as a candidate Global boundary Stratotype Section and Point for the Anthropocene series.

An annually laminated succession in Crawford Lake, Ontario, Canada is proposed for the Global boundary Stratotype Section and Point (GSSP) to define the Anthropocene as a series/epoch with a base dated at 1950 CE. Varve couplets of organic matter capped by calcite precipitated each summer in alkaline surface waters reflect environmental change at global to local scales. Spheroidal carbonaceous particles and nitrogen isotopes record an increase in fossil fuel combustion in the early 1950s, coinciding with early fallout from nuclear and thermonuclear testing - 239+240Pu and 14C:12C, the latter more than compensating for the effects of old carbon in this dolomitic basin. Rapid industrial expansion in the North American Great Lakes region led to enhanced leaching of terrigenous elements by acid precipitation during the Great Acceleration, and calcite precipitation was reduced, producing thin calcite laminae around the GSSP that is marked by a sharp decline in elm pollen (Dutch Elm disease). The lack of bioturbation in well-oxygenated bottom waters, supported by the absence of fossil pigments from obligately anaerobic purple sulfur bacteria, is attributed to elevated salinities and high alkalinity below the chemocline. This aerobic depositional environment, highly unusual in a meromictic lake, inhibits the mobilization of Pu, the proposed primary stratigraphic guide for the Anthropocene.

Anthropogenic eutrophication of shallow lakes: Is it occasional?

Understanding and managing the susceptibility of lakes to anthropogenic eutrophication has been a primary goal of limnological research for decades. To achieve United Nations' Sustainable Development Goals, scientists have attempted to understand why shallow lakes appear to be prone to eutrophication and resistant to restoration. A rich data base of 1151 lakes (each ≥ 0.5 km2) located within the Europe and the United States of America offers a rare opportunity to explore potential answers. Analysis of sites showed that lake depth integrated socio-ecological systems and reflected potential susceptibility to anthropogenic stressors, as well as lake productivity. In this study, lakes distributed in agricultural plain and densely populated lowland areas were generally shallow and subjected to intense human activities with high external nutrient inputs. In contrast, deep lakes frequently occurred in upland regions, dominated by natural landscapes with little anthropogenic nutrient input. Lake depth appeared to not only reflect external nutrient load to the lake, but also acted as an amplifier that increased shallow lake susceptibility to anthropogenic disturbance. Our findings suggest that shallow lakes are more susceptible to human forcing and their eutrophication may be not an occasional occurrence, and that societal expectations, policy goals, and management plans should reflect this observation.

Breathing space: deoxygenation of aquatic environments can drive differential ecological impacts across biological invasion stages.

The influence of climate change on the ecological impacts of invasive alien species (IAS) remains understudied, with deoxygenation of aquatic environments often-overlooked as a consequence of climate change. Here, we therefore assessed how oxygen saturation affects the ecological impact of a predatory invasive fish, the Ponto-Caspian round goby (Neogobius melanostomus), relative to a co-occurring endangered European native analogue, the bullhead (Cottus gobio) experiencing decline in the presence of the IAS. In individual trials and mesocosms, we assessed the effect of high, medium and low (90%, 60% and 30%) oxygen saturation on: (1) functional responses (FRs) of the IAS and native, i.e. per capita feeding rates; (2) the impact on prey populations exerted; and (3) how combined impacts of both fishes change over invasion stages (Pre-invasion, Arrival, Replacement, Proliferation). Both species showed Type II potentially destabilising FRs, but at low oxygen saturation, the invader had a significantly higher feeding rate than the native. Relative Impact Potential, combining fish per capita effects and population abundances, revealed that low oxygen saturation exacerbates the high relative impact of the invader. The Relative Total Impact Potential (RTIP), modelling both consumer species' impacts on prey populations in a system, was consistently higher at low oxygen saturation and especially high during invader Proliferation. In the mesocosm experiment, low oxygen lowered RTIP where both species were present, but again the IAS retained high relative impact during Replacement and Proliferation stages at low oxygen. We also found evidence of multiple predator effects, principally antagonism. We highlight the threat posed to native communities by IAS alongside climate-related stressors, but note that solutions may be available to remedy hypoxia and potentially mitigate impacts across invasion stages. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s10530-021-02542-3.

Global data set of long-term summertime vertical temperature profiles in 153 lakes.

Climate change and other anthropogenic stressors have led to long-term changes in the thermal structure, including surface temperatures, deepwater temperatures, and vertical thermal gradients, in many lakes around the world. Though many studies highlight warming of surface water temperatures in lakes worldwide, less is known about long-term trends in full vertical thermal structure and deepwater temperatures, which have been changing less consistently in both direction and magnitude. Here, we present a globally-expansive data set of summertime in-situ vertical temperature profiles from 153 lakes, with one time series beginning as early as 1894. We also compiled lake geographic, morphometric, and water quality variables that can influence vertical thermal structure through a variety of potential mechanisms in these lakes. These long-term time series of vertical temperature profiles and corresponding lake characteristics serve as valuable data to help understand changes and drivers of lake thermal structure in a time of rapid global and ecological change.

Widespread deoxygenation of temperate lakes.

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 stratification8,9 or 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 oceans6,7 and could threaten essential lake ecosystem services2,3,5,11.

Deeper waters are changing less consistently than surface waters in a global analysis of 102 lakes.

Globally, lake surface water temperatures have warmed rapidly relative to air temperatures, but changes in deepwater temperatures and vertical thermal structure are still largely unknown. We have compiled the most comprehensive data set to date of long-term (1970-2009) summertime vertical temperature profiles in lakes across the world to examine trends and drivers of whole-lake vertical thermal structure. We found significant increases in surface water temperatures across lakes at an average rate of + 0.37 °C decade-1, comparable to changes reported previously for other lakes, and similarly consistent trends of increasing water column stability (+ 0.08 kg m-3 decade-1). In contrast, however, deepwater temperature trends showed little change on average (+ 0.06 °C decade-1), but had high variability across lakes, with trends in individual lakes ranging from - 0.68 °C decade-1 to + 0.65 °C decade-1. The variability in deepwater temperature trends was not explained by trends in either surface water temperatures or thermal stability within lakes, and only 8.4% was explained by lake thermal region or local lake characteristics in a random forest analysis. These findings suggest that external drivers beyond our tested lake characteristics are important in explaining long-term trends in thermal structure, such as local to regional climate patterns or additional external anthropogenic influences.

Effects of a century of mining and industrial production on metal contamination of a model saline ecosystem, Great Salt Lake, Utah.

Effects of mining and metals production have been reported in freshwater lake sediments from around the world but are rarely quantified in saline lake sediments, despite the importance of these lake ecosystems. Here we used dated sediment cores from Great Salt Lake, Utah, USA, a large saline lake adjacent to one of the world's largest copper mines, to measure historical changes in the deposition of 22 metals. Metal concentrations were low prior to the onset of mining in the catchment in 1860 CE. Concentrations of copper, lead, zinc, cadmium, mercury, and other metals began increasing in the late 1800s, with peaks in the 1950s, concomitant with enhanced mining and smelting activities. Sedimentary metal concentrations in the 1950s were 20-40-fold above background levels for copper, lead, silver, and molybdenum. Concentrations of most metals in surficial sediments have decreased 2-5-fold, reflecting: 1) storage and mineralization of sedimenting materials in a deep brine layer, thereby reducing metal transport to the sediments; 2) improved pollution control technologies, and; 3) reduction in mining activity beginning in the 1970s and 1980s. Despite reductions, concentrations of many metals in surficial sediments remain above acceptable contamination thresholds for aquatic ecosystems with migratory birds, and consumption advisories for mercury have been placed on three waterfowl species. The research also highlights that metal deposition in saline lakes is complicated by effects of hypersaline brines and deep-water anoxia in regulating sediment redox and release of metals to surface waters. Given the importance of saline lakes to migratory birds, metals contamination from mining and metals production should be a focus of saline lake remediation.

Nutrients and warming interact to force mountain lakes into unprecedented ecological states.

While deposition of reactive nitrogen (N) in the twentieth century has been strongly linked to changes in diatom assemblages in high-elevation lakes, pronounced and contemporaneous changes in other algal groups suggest additional drivers. We explored the origin and magnitude of changes in two mountain lakes from the end of the Little Ice Age at ca 1850, to ca 2010, using lake sediments. We found dramatic changes in algal community abundance and composition. While diatoms remain the most abundant photosynthetic organisms, concentrations of diatom pigments decreased while pigments representing chlorophytes increased 200-300% since ca 1950 and total algal biomass more than doubled. Some algal changes began ca 1900 but shifts in most sedimentary proxies accelerated ca 1950 commensurate with many human-caused changes to the Earth System. In addition to N deposition, aeolian dust deposition may have contributed phosphorus. Strong increases in summer air and surface water temperatures since 1983 have direct and indirect consequences for high-elevation ecosystems. Such warming could have directly enhanced nutrient use and primary production. Indirect consequences of warming include enhanced leaching of nutrients from geologic and cryosphere sources, particularly as glaciers ablate. While we infer causal mechanisms, changes in primary producer communities appear to be without historical precedent and are commensurate with the post-1950 acceleration of global change.

Factors regulating primary producers' assemblages in Posidonia oceanica (L.) Delile ecosystems over the past 1800 years.

Posidonia oceanica (L.) Delile meadows are highly productive coastal marine ecosystems that provide multiple ecosystem services. The seagrass is not always the major contributor to total primary production, however, little is known about long-term changes in the composition of primary producers within seagrass meadows. Understanding compositional shifts within the community of primary producers is crucial to evaluate how climate and anthropogenic change affect the functioning of seagrass ecosystems. Here we analysed marker pigment composition in seagrass cores from two bays of the Cabrera Island (Balearic Islands, Spain) to asses long-term changes in phototrophic community composition and production in seagrass meadows, and identify the environmental factors triggering those changes. The proxy dataset was explored using principal component analyses (PCA): one including the pigment dataset to look for associations between producers' groups, and another one combining the pigment dataset with plausible local and global regulatory factors to assess the environmental drivers of change. Analyses of characteristic pigments and morphological fossils (cysts) showed that the abundance of dinoflagellates increased over the last 150-300 years, coeval with a rise in solar irradiance and air temperature. When compared among embayments, pigments from cyanobacteria predominated in seagrass meadows located at Es Port, a sheltered bay receiving higher terrestrial runoff; whereas pigments from diatoms, seagrasses and rodophytes were more common at Santa Maria, an exposed bay with clearer waters. Water depth also played a role in controlling the phototrophic community composition, with greater abundance of diatoms in the shallowest waters (<5 m). Overall, our results suggested that historical and spatial variation in seagrass meadows' phototrophic community composition was influenced by the interaction between local factors (catchment-bay characteristics) and global climate processes (energy influx). Together these patterns forecast how marine primary producers and seagrass ecosystem structure may respond to future global warming.

Unabated Nitrogen Pollution Favors Growth of Toxic Cyanobacteria over Chlorophytes in Most Hypereutrophic Lakes.

Human release of reactive nitrogen (N) to the environment has increased 10-fold since 1860 and is expected to increase by a further ∼75% by 2050. Much of this N enters phosphorus (P)-rich, eutrophic lakes in agricultural and urbanized watersheds. While N pollution of eutrophic lakes can promote toxic cyanobacterial growth, some cases of extreme N loading have led to the dominance of chlorophytes (green algae). As N loads required to shift communities from cyanobacterial to chlorophyte dominance are unclear, we experimentally tested phytoplankton responses to a gradient of N loading in a P-rich lake. Low-to-moderate doses (1-3 mg N L-1 week-1) promoted toxic cyanobacterial dominance and elevated concentrations of the hepatotoxin microcystin. Conversely, loads characteristic of pure urban or agricultural effluents (up to 18 mg N L-1 week-1) led to the dominance of chlorophytes over cyanobacteria and lower microcystin content. This indicates that N loads needed to sustain chlorophyte dominance are uncommon, likely restricted to select shallow lakes directly exposed to urban or agricultural effluents. As most N pollution regimes in P-rich lakes will favor toxic cyanobacterial dominance, restricting future N pollution will help curb further cyanobacterial dominance in lakes both directly and by constraining the capacity for future P loading and climate warming to drive cyanobacterial growth.

Widespread nitrous oxide undersaturation in farm waterbodies creates an unexpected greenhouse gas sink.

Nitrogen pollution and global eutrophication are predicted to increase nitrous oxide (N2O) emissions from freshwater ecosystems. Surface waters within agricultural landscapes experience the full impact of these pressures and can contribute substantially to total landscape N2O emissions. However, N2O measurements to date have focused on flowing waters. Small artificial waterbodies remain greatly understudied in the context of agricultural N2O emissions. This study provides a regional analysis of N2O measurements in small (<0.01 km2) artificial reservoirs, of which an estimated 16 million exist globally. We show that 67% of reservoirs were N2O sinks (-12 to -2 µmol N2O⋅m-2⋅d-1) in Canada's largest agricultural area, despite their highly eutrophic status [99 ± 289 µg⋅L-1 chlorophyll-a (Chl-a)]. Generalized additive models indicated that in situ N2O concentrations were strongly and nonlinearly related to stratification strength and dissolved inorganic nitrogen content, with the lowest N2O levels under conditions of strong water column stability and high algal biomass. Predicted fluxes from previously published models based on lakes, reservoirs, and agricultural waters overestimated measured fluxes on average by 7- to 33-fold, challenging the widely held view that eutrophic N-enriched waters are sources of N2O.

Regional versus local drivers of water quality in the Windermere catchment, Lake District, United Kingdom: The dominant influence of wastewater pollution over the past 200 years.

Freshwater ecosystems are threatened by multiple anthropogenic stressors acting over different spatial and temporal scales, resulting in toxic algal blooms, reduced water quality and hypoxia. However, while catchment characteristics act as a 'filter' modifying lake response to disturbance, little is known of the relative importance of different drivers and possible differentiation in the response of upland remote lakes in comparison to lowland, impacted lakes. Moreover, many studies have focussed on single lakes rather than looking at responses across a set of individual, yet connected lake basins. Here we used sedimentary algal pigments as an index of changes in primary producer assemblages over the last ~200 years in a northern temperate watershed consisting of 11 upland and lowland lakes within the Lake District, United Kingdom, to test our hypotheses about landscape drivers. Specifically, we expected that the magnitude of change in phototrophic assemblages would be greatest in lowland rather than upland lakes due to more intensive human activities in the watersheds of the former (agriculture, urbanization). Regional parameters, such as climate dynamics, would be the predominant factors regulating lake primary producers in remote upland lakes and thus, synchronize the dynamic of primary producer assemblages in these basins. We found broad support for the hypotheses pertaining to lowland sites as wastewater treatment was the main predictor of changes to primary producer assemblages in lowland lakes. In contrast, upland headwaters responded weakly to variation in atmospheric temperature, and dynamics in primary producers across upland lakes were asynchronous. Collectively, these findings show that nutrient inputs from point sources overwhelm climatic controls of algae and nuisance cyanobacteria, but highlights that large-scale stressors do not always initiate coherent regional lake response. Furthermore, a lake's position in its landscape, its connectivity and proximity to point nutrients are important determinants of changes in production and composition of phototrophic assemblages.

Effects of experimental nitrogen fertilization on planktonic metabolism and CO2 flux in a hypereutrophic hardwater lake.

Hardwater lakes are common in human-dominated regions of the world and often experience pollution due to agricultural and urban effluent inputs of inorganic and organic nitrogen (N). Although these lakes are landscape hotspots for CO2 exchange and food web carbon (C) cycling, the effect of N enrichment on hardwater lake food web functioning and C cycling patterns remains unclear. Specifically, it is unknown if different eutrophication scenarios (e.g., modest non point vs. extreme point sources) yield consistent effects on auto- and heterotrophic C cycling, or how biotic responses interact with the inorganic C system to shape responses of air-water CO2 exchange. To address this uncertainty, we induced large metabolic gradients in the plankton community of a hypereutrophic hardwater Canadian prairie lake by adding N as urea (the most widely applied agricultural fertilizer) at loading rates of 0, 1, 3, 8 or 18 mg N L-1 week-1 to 3240-L, in-situ mesocosms. Over three separate 21-day experiments, all treatments of N dramatically increased phytoplankton biomass and gross primary production (GPP) two- to six-fold, but the effects of N on autotrophs plateaued at ~3 mg N L-1. Conversely, heterotrophic metabolism increased linearly with N fertilization over the full treatment range. In nearly all cases, N enhanced net planktonic uptake of dissolved inorganic carbon (DIC), and increased the rate of CO2 influx, while planktonic heterotrophy and CO2 production only occurred in the highest N treatments late in each experiment, and even in these cases, enclosures continued to in-gas CO2. Chemical effects on CO2 through calcite precipitation were also observed, but similarly did not change the direction of net CO2 flux. Taken together, these results demonstrate that atmospheric exchange of CO2 in eutrophic hardwater lakes remains sensitive to increasing N loading and eutrophication, and that even modest levels of N pollution are capable of enhancing autotrophy and CO2 in-gassing in P-rich lake ecosystems.

Daphniid zooplankton assemblage shifts in response to eutrophication and metal contamination during the Anthropocene.

Human activities during the Anthropocene result in habitat degradation that has been associated with biodiversity loss and taxonomic homogenization of ecological communities. Here we estimated effects of eutrophication and heavy metal contamination, separately and in combination, in explaining zooplankton species composition during the past 125-145 years using analysis of daphniid diapausing egg banks from four lakes in the northeastern USA. We then examined how these community shifts influenced patterns of diversity and homogenization. Analysis of past lake production (via subfossil pigments) and metal contamination (via sedimentary metals) demonstrated that eutrophication alone (19-39%) and in combination with metal pollution (17-54%) explained 36-79% of historical variation in daphniid species relative abundances in heavily fertilized lakes. In contrast, metal pollution alone explained the majority (72%) of historical variation in daphniid assemblages at the oligotrophic site. Several species colonization events in eutrophying lakes resulted in increased species richness and gamma diversity through time. At the same time, daphniid assemblages in three eutrophied lakes became more similar to each other (homogenized), but this pattern was only seen when accounting for species presence/absence. We did not observe consistent patterns of divergence between the assemblages in the eutrophying lakes and the low-nutrient reference site. Given the pervasive nature of fertilization and metal pollution, and the sensitivity of cladocerans to these factors, we suggest that many inhabited lake districts may already exhibit similar patterns of daphniid assemblage shifts.