Freshwater salinization syndrome on a continental scale.

Salt pollution and human-accelerated weathering are shifting the chemical composition of major ions in fresh water and increasing salinization and alkalinization across North America. We propose a concept, the freshwater salinization syndrome, which links salinization and alkalinization processes. This syndrome manifests as concurrent trends in specific conductance, pH, alkalinity, and base cations. Although individual trends can vary in strength, changes in salinization and alkalinization have affected 37% and 90%, respectively, of the drainage area of the contiguous United States over the past century. Across 232 United States Geological Survey (USGS) monitoring sites, 66% of stream and river sites showed a statistical increase in pH, which often began decades before acid rain regulations. The syndrome is most prominent in the densely populated eastern and midwestern United States, where salinity and alkalinity have increased most rapidly. The syndrome is caused by salt pollution (e.g., road deicers, irrigation runoff, sewage, potash), accelerated weathering and soil cation exchange, mining and resource extraction, and the presence of easily weathered minerals used in agriculture (lime) and urbanization (concrete). Increasing salts with strong bases and carbonates elevate acid neutralizing capacity and pH, and increasing sodium from salt pollution eventually displaces base cations on soil exchange sites, which further increases pH and alkalinization. Symptoms of the syndrome can include: infrastructure corrosion, contaminant mobilization, and variations in coastal ocean acidification caused by increasingly alkaline river inputs. Unless regulated and managed, the freshwater salinization syndrome can have significant impacts on ecosystem services such as safe drinking water, contaminant retention, and biodiversity.

Reversal of a cyanobacterial bloom in response to early warnings.

Directional change in environmental drivers sometimes triggers regime shifts in ecosystems. Theory and experiments suggest that regime shifts can be detected in advance, and perhaps averted, by monitoring resilience indicators such as variance and autocorrelation of key ecosystem variables. However, it is uncertain whether management action prompted by a change in resilience indicators can prevent an impending regime shift. We caused a cyanobacterial bloom by gradually enriching an experimental lake while monitoring an unenriched reference lake and a continuously enriched reference lake. When resilience indicators exceeded preset boundaries, nutrient enrichment was stopped in the experimental lake. Concentrations of algal pigments, dissolved oxygen saturation, and pH rapidly declined following cessation of nutrient enrichment and became similar to the unenriched lake, whereas a large bloom occurred in the continuously enriched lake. This outcome suggests that resilience indicators may be useful in management to prevent unwanted regime shifts, at least in some situations. Nonetheless, a safer approach to ecosystem management would build and maintain the resilience of desirable ecosystem conditions, for example, by preventing excessive nutrient input to lakes and reservoirs.

Response of plankton to nutrients, planktivory and terrestrial organic matter: a model analysis of whole-lake experiments.

Terrestrial organic matter can be assimilated by aquatic consumers but implications for biomass and production are unresolved. An ecosystem model was fit to estimate effects of phosphorus (P) load, planktivory, and supply rate of terrestrial particulate organic carbon (TPOC) on phytoplankton and zooplankton in five whole-lake experiments. Phytoplankton biomass increased with P load and planktivory and decreased with TPOC supply rate. Zooplankton biomass increased with P load and responded weakly to planktivory and TPOC supply rate. Zooplankton allochthony (proportion of carbon from terrestrial sources) decreased with P load and planktivory and increased with TPOC supply rate. Lakes with low allochthony (< 0.3) had wide ranges of phytoplankton and zooplankton biomass and production, depending on P load and planktivory. Lakes with high allochthony (> 0.3) had low biomass and production of both phytoplankton and zooplankton. In summary, terrestrial OC inhibits primary production and is a relatively low-quality food source for zooplankton.

Changes in ecosystem resilience detected in automated measures of ecosystem metabolism during a whole-lake manipulation.

Environmental sensor networks are developing rapidly to assess changes in ecosystems and their services. Some ecosystem changes involve thresholds, and theory suggests that statistical indicators of changing resilience can be detected near thresholds. We examined the capacity of environmental sensors to assess resilience during an experimentally induced transition in a whole-lake manipulation. A trophic cascade was induced in a planktivore-dominated lake by slowly adding piscivorous bass, whereas a nearby bass-dominated lake remained unmanipulated and served as a reference ecosystem during the 4-y experiment. In both the manipulated and reference lakes, automated sensors were used to measure variables related to ecosystem metabolism (dissolved oxygen, pH, and chlorophyll-a concentration) and to estimate gross primary production, respiration, and net ecosystem production. Thresholds were detected in some automated measurements more than a year before the completion of the transition to piscivore dominance. Directly measured variables (dissolved oxygen, pH, and chlorophyll-a concentration) related to ecosystem metabolism were better indicators of the approaching threshold than were the estimates of rates (gross primary production, respiration, and net ecosystem production); this difference was likely a result of the larger uncertainties in the derived rate estimates. Thus, relatively simple characteristics of ecosystems that were observed directly by the sensors were superior indicators of changing resilience. Models linked to thresholds in variables that are directly observed by sensor networks may provide unique opportunities for evaluating resilience in complex ecosystems.

Strong evidence for terrestrial support of zooplankton in small lakes based on stable isotopes of carbon, nitrogen, and hydrogen.

Cross-ecosystem subsidies to food webs can alter metabolic balances in the receiving (subsidized) system and free the food web, or particular consumers, from the energetic constraints of local primary production. Although cross-ecosystem subsidies between terrestrial and aquatic systems have been well recognized for benthic organisms in streams, rivers, and the littoral zones of lakes, terrestrial subsidies to pelagic consumers are more difficult to demonstrate and remain controversial. Here, we adopt a unique approach by using stable isotopes of H, C, and N to estimate terrestrial support to zooplankton in two contrasting lakes. Zooplankton (Holopedium, Daphnia, and Leptodiaptomus) are comprised of ≈ 20-40% of organic material of terrestrial origin. These estimates are as high as, or higher than, prior measures obtained by experimentally manipulating the inorganic (13)C content of these lakes to augment the small, natural contrast in (13)C between terrestrial and algal photosynthesis. Our study gives credence to a growing literature, which we review here, suggesting that significant terrestrial support of pelagic crustaceans (zooplankton) is widespread.

Increased river alkalinization in the Eastern U.S.

The interaction between human activities and watershed geology is accelerating long-term changes in the carbon cycle of rivers. We evaluated changes in bicarbonate alkalinity, a product of chemical weathering, and tested for long-term trends at 97 sites in the eastern United States draining over 260,000 km(2). We observed statistically significant increasing trends in alkalinity at 62 of the 97 sites, while remaining sites exhibited no significant decreasing trends. Over 50% of study sites also had statistically significant increasing trends in concentrations of calcium (another product of chemical weathering) where data were available. River alkalinization rates were significantly related to watershed carbonate lithology, acid deposition, and topography. These three variables explained ~40% of variation in river alkalinization rates. The strongest predictor of river alkalinization rates was carbonate lithology. The most rapid rates of river alkalinization occurred at sites with highest inputs of acid deposition and highest elevation. The rise of alkalinity in many rivers throughout the Eastern U.S. suggests human-accelerated chemical weathering, in addition to previously documented impacts of mining and land use. Increased river alkalinization has major environmental implications including impacts on water hardness and salinization of drinking water, alterations of air-water exchange of CO2, coastal ocean acidification, and the influence of bicarbonate availability on primary production.

Does the 11-year solar cycle affect lake and river ice phenology?

Records of ice-on and ice-off dates are available for lakes and rivers across the Northern Hemisphere spanning decades and in some cases centuries. This data provides an opportunity to investigate the climatic processes that may control ice phenology. Previous studies have reported a trend toward shorter ice-covered seasons with global warming, as well as links between ice phenology and several modes of natural climate variability such as the North Atlantic Oscillation, the Pacific-North American Pattern, the El Niño-Southern Oscillation, the Pacific Decadal Oscillation, and the Atlantic Multidecadal Oscillation. The 11-year sunspot cycle has also been proposed as a driver of ice phenology, which is somewhat surprising given that this cycle's strongest impacts are in the stratosphere. In this study, we use a large data set of lakes and rivers across the Northern Hemisphere to test this potential link. We find little or no connection between the sunspot cycle and either ice-on or ice-off dates. We conclude that while many well-known climate cycles do impact ice phenology, we are able to rule out any strong impact of the solar cycle.

Five state factors control progressive stages of freshwater salinization syndrome.

Factors driving freshwater salinization syndrome (FSS) influence the severity of impacts and chances for recovery. We hypothesize that spread of FSS across ecosystems is a function of interactions among five state factors: human activities, geology, flowpaths, climate, and time. (1) Human activities drive pulsed or chronic inputs of salt ions and mobilization of chemical contaminants. (2) Geology drives rates of erosion, weathering, ion exchange, and acidification-alkalinization. (3) Flowpaths drive salinization and contaminant mobilization along hydrologic cycles. (4) Climate drives rising water temperatures, salt stress, and evaporative concentration of ions and saltwater intrusion. (5) Time influences consequences, thresholds, and potentials for ecosystem recovery. We hypothesize that state factors advance FSS in distinct stages, which eventually contribute to failures in systems-level functions (supporting drinking water, crops, biodiversity, infrastructure, etc.). We present future research directions for protecting freshwaters at risk based on five state factors and stages from diagnosis to prognosis to cure.

Conditional heteroscedasticity as a leading indicator of ecological regime shifts.

Regime shifts are massive, often irreversible, rearrangements of nonlinear ecological processes that occur when systems pass critical transition points. Ecological regime shifts sometimes have severe consequences for human well-being, including eutrophication in lakes, desertification, and species extinctions. Theoretical and laboratory evidence suggests that statistical anomalies may be detectable leading indicators of regime shifts in ecological time series, making it possible to foresee and potentially avert incipient regime shifts. Conditional heteroscedasticity is persistent variance characteristic of time series with clustered volatility. Here, we analyze conditional heteroscedasticity as a potential leading indicator of regime shifts in ecological time series. We evaluate conditional heteroscedasticity by using ecological models with and without four types of critical transition. On approaching transition points, all time series contain significant conditional heteroscedasticity. This signal is detected hundreds of time steps in advance of the regime shift. Time series without regime shifts do not have significant conditional heteroscedasticity. Because probability values are easily associated with tests for conditional heteroscedasticity, detection of false positives in time series without regime shifts is minimized. This property reduces the need for a reference system to compare with the perturbed system.

Terrestrial, benthic, and pelagic resource use in lakes: results from a three-isotope Bayesian mixing model.

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections.

Climate change drives warming in the Hudson River Estuary, New York (USA).

Estuaries may be subject to warming due to global climate change but few studies have considered the drivers or seasonality of warming empirically. We analyzed temperature trends and rates of temperature change over time for the Hudson River estuary using long-term data, mainly from daily measures taken at the Poughkeepsie Water Treatment Facility. This temperature record is among the longest in the world for a river or estuary. The Hudson River has warmed 0.945 °C since 1946. Many of the warmest years in the record occurred in the last 16 years. A seasonal analysis of trends indicated significant warming for the months of April through August. The warming of the Hudson is primarily related to increasing air temperature. Increasing freshwater discharge into the estuary has not mitigated the warming trend.

Mudflat geomorphology determines invasive macroalgal effect on invertebrate prey and shorebird predators.

Impacts of invasive species are often context specific due to varying ecological interactions. Physical structure of environments hosting invaders is also potentially important but has received limited attention. An invasive macroalga, Agarophyton vermiculophyllum, has spread across the northern hemisphere with mixed positive, neutral and negative effects on resident species. Agarophyton colonizes mudflats that vary in topography due to interactions of sediments with hydrodynamic forces. We tested the hypothesis that mudflat geomorphology moderates the effect of Agarophyton on shorebirds and invertebrates. We surveyed 30 mudflats in the Virginia Coast Reserve quantifying elevation and topography. Invertebrate and bird abundances were also quantified. Mudflat geomorphology ranged from smooth to hummocky and was correlated with invertebrate and shorebird abundance and interactions based on piecewise structural equation models. After accounting for geomorphology, Agarophyton had little effect on invertebrate abundance. Shorebird numbers were differentially influenced by mudflat topography, with positive correlations to invertebrates (worms) on smooth mudflats, and to macroalgae on hummocky mudflats. These differences are likely to be due to sediment properties in interaction with structural changes induced by Agarophyton mats that affect prey accessibility for birds. Even on apparently simple mudflats, geomorphic structure emerged as important, modifying invasive species impacts and differentially influencing consumers.

Whole-lake carbon-13 additions reveal terrestrial support of aquatic food webs.

Ecosystems are supported by organic carbon from two distinct sources. Endogenous carbon is produced by photosynthesis within an ecosystem by autotrophic organisms. Exogenous carbon is produced elsewhere and transported into ecosystems. Consumers may use exogenous carbon with consequent influences on population dynamics, predator-prey relationships and ecosystem processes. For example, exogenous inputs provide resources that may enhance consumer abundance beyond levels supported by within-system primary production. Exogenous fluxes of organic carbon to ecosystems are often large, but this material is recalcitrant and difficult to assimilate, in contrast to endogenously produced organic matter, which is used more easily. Here we show, by the experimental manipulation of dissolved inorganic (13)C in two lakes, that internal primary production is insufficient to support the food webs of these ecosystems. Additions of NaH(13)CO(3) enriched the (13)C content of dissolved inorganic carbon, particulate organic carbon, zooplankton and fish. Dynamics of (13)C indicate that 40-55% of particulate organic carbon and 22-50% of zooplankton carbon are derived from terrestrial sources, showing that there is significant subsidy of these ecosystems by organic carbon produced outside their boundaries.

Impact of exotic macroalga on shorebirds varies with foraging specialization and spatial scale.

Exotic species may increase or decrease native biodiversity. However, effects of exotic species are often mixed; and indirect pathways and compensatory changes can mask effects. Context-specific assessments of the indirect impacts of exotic species are also needed across multiple spatial scales. Agarophyton vermiculophyllum (previously Gracilaria vermiculophylla), an exotic, invasive macroalga, has established throughout the western hemisphere with reported positive or neutral impacts on biodiversity. Shorebirds are an important group for conservation in areas invaded by A. vermiculophyllum. We assess the impacts of this invader on shorebirds by measuring behavior and habitat selection at spatial scales ranging from algal patches to the entire study region. Birds were considered either flexible-foragers that used diverse foraging techniques, or specialized-foragers that employed fewer, more specialized foraging techniques. Responses were scale dependent, with patterns varying between spatial scales, and between behavior and habitat selection. However, a general pattern of habitat selection emerged wherein flexible-foraging shorebirds preferred A. vermiculophyllum habitat, and for specialized-foragers, habitat selection of A. vermiculophyllum was mixed. Meanwhile, flexible-foraging birds tended to neutrally use or avoid uninvaded habitat, and specialized-foraging birds mostly preferred uninvaded habitat. Shorebird behavioral response was less clear; with flexible-foragers spending less time on bare sediment than expected, the only significant response. Shorebird response to A. vermiculophyllum differed by foraging mode; likely because flexible, opportunistic species more readily use invaded habitat. Increases in A. vermiculophyllum could result in functional homogenization if the bare habitat preferred by specialized-foragers is reduced too greatly. We hypothesize the effect of scale is driven by differences among tidal flats. Thus, tidal flat properties such as sediment grain size and microtopography would determine whether foraging from A. vermiculophyllum was optimal for a shorebird. Specialization and spatial scale are important when assessing the biodiversity conservation impacts of invasive A. vermiculophyllum.

The influence of environmental water on the hydrogen stable isotope ratio in aquatic consumers.

Aquatic food webs are subsidized by allochthonous resources but the utilization of these resources by consumers can be difficult to quantify. Stable isotope ratios of hydrogen (deuterium:hydrogen; deltaD) potentially distinguish allochthonous inputs because deltaD differs between terrestrial and aquatic primary producers. However, application of this tracer is limited by uncertainties regarding the trophic fractionation of deltaD and the contributions of H from environmental water (often called "dietary water") to consumer tissue H. We addressed these uncertainties using laboratory experiments, field observations, modeling, and a literature synthesis. Laboratory experiments that manipulated the deltaD of water and food for insects, cladoceran zooplankton, and fishes provided strong evidence that trophic fractionation of deltaD was negligible. The proportion of tissue H derived from environmental water was substantial yet variable among studies; estimates of this proportion, inclusive of lab, field, and literature data, ranged from 0 to 0.39 (mean 0.17 +/- 0.12 SD). There is a clear need for additional studies of environmental water. Accounting for environmental water in mixing models changes estimates of resource use, although simulations suggest that uncertainty about the environmental water contribution does not substantially increase the uncertainty in estimates of resource use. As long as this uncertainty is accounted for, deltaD may be a powerful tool for estimating resource use in food webs.

Hydrology and grazing jointly control a large-river food web.

Inputs of fresh water and grazing both can control aquatic food webs, but little is known about the relative strengths of and interactions between these controls. We use long-term data on the food web of the freshwater Hudson River estuary to investigate the importance of, and interactions between, inputs of fresh water and grazing by the invasive zebra mussel (Dreissena polymorpha). Both freshwater inputs and zebra mussel grazing have strong, pervasive effects on the Hudson River food web. High flow tended to reduce population size in most parts of the food web. High grazing also reduced populations in the planktonic food web, but increased populations in the littoral food web, probably as a result of increases in water clarity. The influences of flow and zebra mussel grazing were roughly equal (i.e., within a factor of 2) for many variables over the period of our study. Zebra mussel grazing made phytoplankton less sensitive to freshwater inputs, but water clarity and the littoral food web more sensitive to freshwater inputs, showing that interactions between these two controlling factors can be strong and varied.

A synthesis of modern organic carbon accumulation rates in coastal and aquatic inland ecosystems.

Organic carbon accumulation in the sediments of inland aquatic and coastal ecosystems is an important process in the global carbon budget that is subject to intense human modification. To date, research has focused on quantifying accumulation rates in individual or groups of aquatic ecosystems to quantify the aquatic carbon sinks. However, there hasn't been a synthesis of rates across aquatic ecosystem to address the variability in rates within and among ecosystems types. Doing so would identify gaps in our understanding of accumulation rates and potentially reveal carbon sinks vulnerable to change. We synthesized accumulation rates from the literature, compiling 464 rate measurements from 103 studies of carbon accumulated in the modern period (ca. 200 years). Accumulation rates from the literature spanned four orders of magnitude varying substantially within and among ecosystem categories, with mean estimates for ecosystem categories ranging from 15.6 to 73.2 g C m-2 y-1 within ecosystem categories. With the exception of lakes, mean accumulation rates were poorly constrained due to high variability and paucity of data. Despite the high uncertainty, the estimates of modern accumulation rate compiled here are an important step for constructing carbon budgets and predicting future change.

Novel 'chemical cocktails' in inland waters are a consequence of the freshwater salinization syndrome.

Widespread changes in water temperatures, salinity, alkalinity and pH have been documented in inland waters in North America, which influence ion exchange, weathering rates, chemical solubility and contaminant toxicity. Increasing major ion concentrations from pollution, human-accelerated weathering and saltwater intrusion contribute to multiple ecological stressors such as changing ionic strength and pH and mobilization of chemical mixtures resulting in the freshwater salinization syndrome (FSS). Here, we explore novel combinations of elements, which are transported together as chemical mixtures containing salts, nutrients and metals as a consequence of FSS. First, we show that base cation concentrations have increased in regions primarily in North America and Europe over 100 years. Second, we show interactions between specific conductance, pH, nitrate and metals using data from greater than 20 streams located in different regions of the USA. Finally, salinization experiments and routine monitoring demonstrate mobilization of chemical mixtures of cations, metals and nutrients in 10 streams draining the Washington, DC-Baltimore, MD metropolitan regions. Freshwater salinization mobilizes diverse chemical mixtures influencing drinking water quality, infrastructure corrosion, freshwater CO2 concentrations and biodiversity. Most regulations currently target individual contaminants, but FSS requires managing mobilization of multiple chemical mixtures and interacting ecological stressors as consequences of freshwater salinization.This article is part of the theme issue 'Salt in freshwaters: causes, ecological consequences and future prospects'.

Terrestrial support of lake food webs: Synthesis reveals controls over cross-ecosystem resource use.

Widespread evidence that organic matter exported from terrestrial into aquatic ecosystems supports recipient food webs remains controversial. A pressing question is not only whether high terrestrial support is possible but also what the general conditions are under which it arises. We assemble the largest data set, to date, of the isotopic composition (δ2H, δ13C, and δ15N) of lake zooplankton and the resources at the base of their associated food webs. In total, our data set spans 559 observations across 147 lakes from the boreal to subtropics. By predicting terrestrial resource support from within-lake and catchment-level characteristics, we found that half of all consumer observations that is, the median were composed of at least 42% terrestrially derived material. In general, terrestrial support of zooplankton was greatest in lakes with large physical and hydrological connections to catchments that were rich in aboveground and belowground organic matter. However, some consumers responded less strongly to terrestrial resources where within-lake production was elevated. Our study shows that multiple mechanisms drive widespread cross-ecosystem support of aquatic consumers across Northern Hemisphere lakes and suggests that changes in terrestrial landscapes will influence ecosystem processes well beyond their boundaries.

LAGOS-NE: a multi-scaled geospatial and temporal database of lake ecological context and water quality for thousands of US lakes.

Understanding the factors that affect water quality and the ecological services provided by freshwater ecosystems is an urgent global environmental issue. Predicting how water quality will respond to global changes not only requires water quality data, but also information about the ecological context of individual water bodies across broad spatial extents. Because lake water quality is usually sampled in limited geographic regions, often for limited time periods, assessing the environmental controls of water quality requires compilation of many data sets across broad regions and across time into an integrated database. LAGOS-NE accomplishes this goal for lakes in the northeastern-most 17 US states.LAGOS-NE contains data for 51 101 lakes and reservoirs larger than 4 ha in 17 lake-rich US states. The database includes 3 data modules for: lake location and physical characteristics for all lakes; ecological context (i.e., the land use, geologic, climatic, and hydrologic setting of lakes) for all lakes; and in situ measurements of lake water quality for a subset of the lakes from the past 3 decades for approximately 2600-12 000 lakes depending on the variable. The database contains approximately 150 000 measures of total phosphorus, 200 000 measures of chlorophyll, and 900 000 measures of Secchi depth. The water quality data were compiled from 87 lake water quality data sets from federal, state, tribal, and non-profit agencies, university researchers, and citizen scientists. This database is one of the largest and most comprehensive databases of its type because it includes both in situ measurements and ecological context data. Because ecological context can be used to study a variety of other questions about lakes, streams, and wetlands, this database can also be used as the foundation for other studies of freshwaters at broad spatial and ecological scales.