It Takes Two to Tango: When and Where Dual Nutrient (N & P) Reductions Are Needed to Protect Lakes and Downstream Ecosystems.

Preventing harmful algal blooms (HABs) is needed to protect lakes and downstream ecosystems. Traditionally, reducing phosphorus (P) inputs was the prescribed solution for lakes, based on the assumption that P universally limits HAB formation. Reduction of P inputs has decreased HABs in many lakes, but was not successful in others. Thus, the "P-only" paradigm is overgeneralized. Whole-lake experiments indicate that HABs are often stimulated more by combined P and nitrogen (N) enrichment rather than N or P alone, indicating that the dynamics of both nutrients are important for HAB control. The changing paradigm from P-only to consideration of dual nutrient control is supported by studies indicating that (1) biological N fixation cannot always meet lake ecosystem N needs, and (2) that anthropogenic N and P loading has increased dramatically in recent decades. Sediment P accumulation supports long-term internal loading, while N may escape via denitrification, leading to perpetual N deficits. Hence, controlling both N and P inputs will help control HABs in some lakes and also reduce N export to downstream N-sensitive ecosystems. Managers should consider whether balanced control of N and P will most effectively reduce HABs along the freshwater-marine continuum.

Ecological responses of a large shallow lake (Okeechobee, Florida) to climate change and potential future hydrologic regimes.

We considered how Lake Okeechobee, a large shallow lake in Florida, USA, might respond to altered hydrology associated with climate change scenarios in 2060. Water budgets and stage hydrographs were provided from the South Florida Water Management Model, a regional hydrologic model used to develop plans for Everglades restoration. Future scenarios include a 10% increase or decrease in rainfall (RF) and a calculated increase in evapotranspiration (ET), which is based on a 1.5 °C rise in temperature. Increasing RF and ET had counter-balancing effects on the water budget and when changing concurrently did not affect hydrology. In contrast, when RF decreased while ET increased, this resulted in a large change in hydrology. The surface elevation of the lake dropped by more than 2 m under this scenario compared to a future base condition, and extreme low elevation persisted for multiple years. In this declining RF/increasing ET scenario, the littoral and near-shore zones, areas that support emergent and submerged plants, were dry 55% of the time compared to less than 4% of the time in the future base run. There also were times when elevation increased as much as 3 m after intense RF events. Overall, these changes in hydrologic conditions would dramatically alter ecosystem services. Uncertainty about responses is highest at the pelagic-littoral interface, in regard to whether an extremely shallow lake could support submerged vascular plants, which are critical to the recreational fishery and for migratory birds. Along with improved regional climate modeling, research in that interface zone is needed to guide the adaptive process of Everglades restoration.

Predicting ecological responses of the Florida Everglades to possible future climate scenarios: introduction.

Florida's Everglades stretch from the headwaters of the Kissimmee River near Orlando to Florida Bay. Under natural conditions in this flat landscape, water flowed slowly downstream as broad, shallow sheet flow. The ecosystem is markedly different now, altered by nutrient pollution and construction of canals, levees, and water control structures designed for flood control and water supply. These alterations have resulted in a 50% reduction of the ecosystem's spatial extent and significant changes in ecological function in the remaining portion. One of the world's largest restoration programs is underway to restore some of the historic hydrologic and ecological functions of the Everglades, via a multi-billion dollar Comprehensive Everglades Restoration Plan. This plan, finalized in 2000, did not explicitly consider climate change effects, yet today we realize that sea level rise and future changes in rainfall (RF), temperature, and evapotranspiration (ET) may have system-wide impacts. This series of papers describes results of a workshop where a regional hydrologic model was used to simulate the hydrology expected in 2060 with climate changes including increased temperature, ET, and sea level, and either an increase or decrease in RF. Ecologists with expertise in various areas of the ecosystem evaluated the hydrologic outputs, drew conclusions about potential ecosystem responses, and identified research needs where projections of response had high uncertainty. Resource managers participated in the workshop, and they present lessons learned regarding how the new information might be used to guide Everglades restoration in the context of climate change.

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.

Hurricanes, El Niño and harmful algal blooms in two sub-tropical Florida estuaries: Direct and indirect impacts.

Future increases in the intensity of hurricanes and El Niño periods predicted by climate change models have focused attention on their role in stimulating harmful algal blooms (HABs). A series of hurricanes that recently impacted Florida (USA) provided a unique opportunity to explore the relationships between hurricanes, El Niño and HABs in two Florida estuaries subject to repeated intense ecosystem disruptive HABs, the Indian River Lagoon and the St. Lucie Estuary. The roles that hurricanes and El Niño play in contributing to HAB events are examined in the context of key structural and functional features of each estuary and their watersheds, including morphology, water residence time and hydrology, such as the influence of Lake Okeechobee discharges into the St. Lucie Estuary. The most direct impact was the increase in rainfall associated with hurricanes and El Niño, resulting in enhanced nutrient loads which drive HABs in the Indian River Lagoon and Lake Okeechobee. Major HABs in Lake Okeechobee also present an indirect threat of freshwater HAB blooms in the St. Lucie Estuary via mandated discharges from the lake into the estuary during high rainfall periods. Conversely, during the absence of HABs in Lake Okeechobee, short water residence times produced by discharges into the St. Lucie Estuary can result in lower bloom intensities.

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.

Cyanobacteria blooms: effects on aquatic ecosystems.

Cyanobacteria become increasingly dominant as concentrations of TP and TN increase during eutrophication of lakes, rivers and estuaries. Temporal dynamics of cyanobacteria blooms are variable--in some systems persistent blooms occur in summer to fall, whereas in other systems blooms are more sporadic. Cyanobacteria blooms have a wide range of possible biological impacts including potential toxic effects on other algae, invertebrates and fish, impacts to plants and benthic algae due to shading, and impacts to food web function as large inedible algae produce a bottleneck to C and energy flow in the plankton food web. In lakes with dense blooms of cyanobacteria, accumulation of organic material in lake sediments and increased bacterial activity also may lead to anoxic conditions that alter the structure of benthic macro-invertebrates. Diffusive internal P loading may increase, and hypolimnetic anoxia may lead to a loss of piscivorous fish that require a summer cold water refuge in temperate lakes. Ecosystem changes associated with frequent blooms may result in delayed response of lakes, rivers and estuaries to external nutrient load reduction. Despite numerous case studies and a vast literature on species-specific responses, community level effects of cyanobacterial blooms are not well understood--in particular the realized impacts of toxins and changes in food web structure/function. These areas require additional research given the prevalence of toxic blooms in the nation's lakes, rivers and coastal waters--systems that provide a wide range of valued ecosystem services.

Cyanobacterial toxins: a qualitative meta-analysis of concentrations, dosage and effects in freshwater, estuarine and marine biota.

This paper reviews the rapidly expanding literature on the ecological effects of cyanobacterial toxins. The study employs a qualitative meta-analysis from the literature examining results from a large number of independent studies and extracts general patterns from the literature or signals contradictions. The meta-analysis is set up by putting together two large tables--embodying a large and representative part of the literature (see Appendix A). The first table (Table A.1) reviews the presence (concentrations) of different cyanobacterial toxins in the tissues of various groups of aquatic biota after exposure via different routes, experimentally in the lab or via natural routes in the environment. The second table (Table A.2) reviews the dose dependent effect of toxins on biota. The great majority of studies deal with the presence and effects of microcystin, especially of the MC-LR congener. Although this may partly be justified--MC-LR is an abundant and highly toxic protein--our review also emphasizes what is known about (i) other MC congeners (a number of studies showed a preferred accumulation of the less toxic variant MC-RR in animal tissues), (ii) nodularin (data on a range of biota from studies on the Baltic Sea), (iii) neurotoxins like anatoxin-a(s), which are conspicuously often present at times when mass mortalities of birds occur, (iv) a few studies on the presence and effects of cylindrospermposin, as well as (v) the first examples of ecological effects of newly identified bioactive compounds, like microviridin-J. Data were reorganized to assess to what extent bioconcentration (uptake and concentration of toxins from the water) or biomagnification (uptake and concentration via the food) of cyanobacterial toxins occurs in ecosystems. There is little support for the occurrence of biomagnification, and this reduces the risk for biota at higher trophic levels. Rather than biomagnification biodilution seems to occur in the foodweb with toxins being subject to degradation and excretion at every level. Nevertheless toxins were present at all tropic levels, indicating that some vectorial transport must take place, and in sufficient quantities for effects to possibly occur. Feeding seemed to be the most important route for exposure of aquatic biota to cyanobacterial toxins. A fair number of studies focus on dissolved toxins, but in those studies purified toxin typically is used, and biota do not appear very sensitive to this form of exposure. More effects are found when crude cyanobacterial cell lysates are used, indicating that there may be synergistic effects between different bioactive compounds. Aquatic biota are by no means defenseless against toxic cyanobacteria. Several studies indicate that those species that are most frequently exposed to toxins in their natural environment are also the most tolerant. Protection includes behavioral mechanisms, detoxication of MC and NODLN by conjugation with glutathione, and fairly rapid depuration and excretion. A common theme in much of the ecological studies is that of modulating factors. Effects are seldom straightforward, but are dependent on factors like the (feeding) condition of the animals, environmental conditions and the history of exposure (acclimation and adaptation to toxic cyanobacteria). This makes it harder to generalize on what is known about ecological effects of cyanobacterial toxins. The paper concludes by summarizing the risks for birds, fish, macroinvertebrates and zooplankton. Although acute (lethal) effects are mentioned in the literature, mass mortalities of--especially--fish are more likely to be the result of multiple stress factors that co-occur during cyanobacterial blooms. Bivalves appear remarkably resistant, whilst the harmful effects of cyanobacteria on zooplankton vary widely and the specific contribution of toxins is hard to evaluate.

A reevaluation of the cultural eutrophication of Lake Okeechobee using multiproxy sediment records.

Lake Okeechobee, the hydrological lynchpin of the Everglades ecosystem, is the subject of an ambitious, multiagency restoration effort aimed at reducing phosphorus inputs and resulting algal blooms and impaired water clarity. This restoration is predicated on returning the lake to something closer to its predisturbance condition, but that goal has been challenged on the premise that the lake has always been eutrophic. The resolution of this debate and the appropriateness of the nutrient reduction goals thus depend on obtaining a reliable sediment record of past limnological conditions--the aim of this study. Because of the potential for severe sediment mixing from tropical storms, this investigation used multiple dating tools to examine the integrity of the sediment record and then analyzed proxies for nutrient enrichment, phytoplankton composition, and paleoproductivity. Sediment profiles for atmospheric pollutants, fertilizer contaminants, and radiocesium from three widely spaced cores showed good preservation of stratigraphic detail and coherence with the 210Pb chronologies. These results demonstrated that sediment stratigraphy is largely intact and retains a reliable record of limnological change. Geochemical proxies provide strong evidence of increased nutrient loading beginning ca. 1950. Concentrations of sediment P double, and N:P and C:N ratios drop, while those for N isotopes (delta15N) increase. At the same time, tracers of phosphate fertilizers (uranium, vanadium, and arsenic) rise. These changes are synchronous among cores and constitute a robust, internally consistent record of increasing water-column P. Biotic responses are manifested in higher concentrations and in changing composition of fossil algal pigments, including (1) large increases in the concentrations of chemically robust carotenoids, (2) corresponding decreases in the ratios of pigments from diatoms to chlorophyte and cyanobacterial algae, and (3) increases in UVR-photo-protective compounds indicating greater prevalence of surface algal blooms. This study provides strong evidence that Lake Okeechobee has experienced accelerated eutrophication linked with post-1950s land use changes in its watershed, a conclusion consistent with the nutrient reduction goals of the Lake Okeechobee Protection Program. The results contradict recent claims that the lake's trophic state has not changed over time, as well as the assertion that sediments of large shallow lakes cannot support a reliable chronology of past events.

Plankton food web responses to experimental nutrient additions in a subtropical lake.

During two controlled enclosure experiments using water from a subtropical lake, the plankton food web displayed a highly variable response to combined addition of nitrogen and phosphorus. In July, the nutrients stimulated growth of Cylindrospermopsis raciborskii, and the biomass of macrozooplankton and microbial food web components did not increase. In October, the same addition of nutrients stimulated growth of small edible Lyngbya spp., and there were coincident increases in biomass of macrozooplankton and components of the microbial web. Past generalizations that cyanobacteria blooms inhibit growth of other food web components may not always hold true.

Mitigating cyanobacterial harmful algal blooms in aquatic ecosystems impacted by climate change and anthropogenic nutrients.

Mitigating the global expansion of cyanobacterial harmful blooms (CyanoHABs) is a major challenge facing researchers and resource managers. A variety of traditional (e.g., nutrient load reduction) and experimental (e.g., artificial mixing and flushing, omnivorous fish removal) approaches have been used to reduce bloom occurrences. Managers now face the additional effects of climate change on watershed hydrologic and nutrient loading dynamics, lake and estuary temperature, mixing regime, internal nutrient dynamics, and other factors. Those changes favor CyanoHABs over other phytoplankton and could influence the efficacy of control measures. Virtually all mitigation strategies are influenced by climate changes, which may require setting new nutrient input reduction targets and establishing nutrient-bloom thresholds for impacted waters. Physical-forcing mitigation techniques, such as flushing and artificial mixing, will need adjustments to deal with the ramifications of climate change. Here, we examine the suite of current mitigation strategies and the potential options for adapting and optimizing them in a world facing increasing human population pressure and climate change.

A global database of lake surface temperatures collected by in situ and satellite methods from 1985-2009.

Global environmental change has influenced lake surface temperatures, a key driver of ecosystem structure and function. Recent studies have suggested significant warming of water temperatures in individual lakes across many different regions around the world. However, the spatial and temporal coherence associated with the magnitude of these trends remains unclear. Thus, a global data set of water temperature is required to understand and synthesize global, long-term trends in surface water temperatures of inland bodies of water. We assembled a database of summer lake surface temperatures for 291 lakes collected in situ and/or by satellites for the period 1985-2009. In addition, corresponding climatic drivers (air temperatures, solar radiation, and cloud cover) and geomorphometric characteristics (latitude, longitude, elevation, lake surface area, maximum depth, mean depth, and volume) that influence lake surface temperatures were compiled for each lake. This unique dataset offers an invaluable baseline perspective on global-scale lake thermal conditions as environmental change continues.

N:P ratios, light limitation, and cyanobacterial dominance in a subtropical lake impacted by non-point source nutrient pollution.

A long-term (28-year) data set was used to investigate historical changes in concentrations of phosphorus (P), nitrogen (N), N:P ratios, and Secchi disk transparency in a shallow subtropical lake (Lake Okeechobee, Florida, USA). The aim was to evaluate changes in the risk of N2-fixing cyanobacterial blooms, which have infrequently occurred in the lake's pelagic zone. Predictions regarding bloom risk were based on previously published N:P ratio models. Temporal trends in the biomass of cyanobacteria were evaluated using phytoplankton data collected in 1974, 1989-1992, and 1997-2000. Concentrations of pelagic total P increased from near 50 microg l-1 in the mid-1970s to over 100 microg l-1 in the late 1990s. Coincidentally, the total N:P (mass) ratio decreased from 30:1 to below 15:1, and soluble N:P ratio decreased from 15:1 to near 6:1, in the lake water. Published empirical models predict that current conditions favor cyanobacteria. The observations confirm this prediction: cyanobacteria presently account for 50-80% of total phytoplankton biovolume. The historical decrease in TN:TP ratio in the lake can be attributed to a decreased TN:TP ratio in the inflow water and to a decline in the lake's assimilation of P, relative to N. Coincident with these declines in total and soluble N:P ratios, Secchi disk transparency declined from 0.6 m to near 0.3 m, possibly due to increased mineral turbidity in the lake water. Empirical models predict that under the turbid, low irradiance conditions that prevail in this lake, non-heterocystous cyanobacteria should dominate the phytoplankton. Our observations confirmed this prediction: non-N2-fixing taxa (primarily Oscillatoria and Lyngbya spp.) typically dominated the cyanobacteria community during the last decade. The only exception was a year with very low water levels, when heterocystous N2-fixing Anabaena became dominant. In the near-shore regions of this shallow lake, low N:P ratios potentially favor blooms of N2-fixing cyanobacteria, but their occurrence in the pelagic zone is restricted by low irradiance and lack of stable stratification.

Zooplankton structure and potential food web interactions in the plankton of a subtropical chain-of-lakes.

This study evaluates the taxonomic and size structure of macro-zooplankton and its potential role in controlling phytoplankton in the Kissimmee Chain-of-Lakes, six shallow interconnected lakes in Florida, U.S. Macro-zooplankton species biomass and standard limnological attributes (temperature, pH, total phosphorus [TP], chlorophyll a [Chl a], and Secchi transparency) were quantified on a bimonthly basis from April 1997 to February 1999. Concentrations of TP ranged from below 50 to over 150 microg l(-1). Peak concentrations of particulate P coincided with maximal Chl a, and in one instance a high concentration of soluble reactive P followed. The cladoceran zooplankton was dominated by small species, including Eubosmina tubicen, Ceriodaphnia rigaudi, and Daphnia ambigua. The exotic daphnid, D. lumholtzii, periodically was abundant. The copepods were strongly dominated by Diaptomus dorsalis, a species previously shown to be highly resistant to fish predation. These results, and findings of controlled experiments on a nearby lake with a nearly identical zooplankton species complement, suggest that fish predation may be a major factor structuring the macro-zooplankton assemblage. Zooplankton biomass, on the other hand, may be affected by resource availability. There was a significant positive relationship between average biomass of macro-zooplankton and the average concentration of TP among the six lakes. No such relationship existed between zooplankton biomass and Chl a, suggesting that the predominant food web in these systems may be based on bacteria-plankton, as has been documented in nearby Lake Okeechobee. All of the zooplankton taxa encountered in the Kissimmee Chain-of-Lakes (except Mesocyclops edax) are known bacteria grazers in Florida lakes. Phytoplankton biomass, measured as Chl a, was strongly associated with TP, both within and across lakes. Phytoplankton biomass was not associated with the biomass of zooplankton. These results, when considered in the context of nutrient-addition, zooplankton-exclosure studies on Lake Okeechobee, support the hypothesis that phytoplankton biomass in subtropical lakes is regulated by "bottom-up," rather than "top-down" forces.

The International Editorship of Freshwater Systems.

It is my pleasure to announce that two distinguished internationalscientists have joined the editorship of the FreshwaterSystems domain of TheScientificWorldJOURNAL - Professor BrijGopal of Jawaharlal Nehru University (India) and Dr. Manual Graça of the Universityof Coimbra (Portugal). Professor Gopal is the Secretary General of the NationalInstitute of Ecology, Editor of the InternationalJournal of Ecology & Environmental Science,and Chairman of the SIL (International Association of Theoretical and AppliedLimnology) Committee on Limnology in Developing Countries. His research interestsinclude the ecology, biogeochemistry and biodiversity of wetland ecosystems,the management of wetlands as an integral part of the watershed, and wetlandwater policy-related issues. Dr. Graça is a stream ecologist whose researchinterests include the two general areas of organic matter decomposition andbiological monitoring. His specific areas of research focus include quantificationof organic matter and other chemical changes in decomposing leaves, the ecologyof aquatic hyphomycetes, and the ecology of animals feeding on detritus. Hisresearch dealing with biological monitoring is carried out in close cooperationwith the paper and mining industries, facilitating the practical applicationof his work.

Stable isotope food web analysis of a large subtropical lake: alternative explanations for 15N enrichment of pelagic vs. littoral fisheries.

The food webs of littoral, pelagic, and littoral-pelagic ecotone (interface) regions of a large subtropical lake were investigated using stable isotope ratio methods, expanding the focus of a previous fish-only study to include other food web components such as primary producers and invertebrates. In these food webs, delta13C increased approximately 4 per thousand and delta15N increased approximately 10 per thousand from primary producers to fish. The delta15N of fish was approximately 9 per thousand in the littoral zone, approximately 10 per thousand in the ecotone, and approximately 12 per thousand in the pelagic zone. The cross-habitat enrichment in fish 15N corresponded with both an increase in the size of fish and an increase in the d15N of primary consumers (mollusks). Despite larger body size in the pelagic zone, fish in all three habitats appear to occur at the same average trophic level (TL = 4), assuming an enrichment factor of 3.4 per thousand per trophic level, and normalizing to the delta15N of primary consumers.

Welcome to Freshwater Systems, A Domain of TheScientificWorld.

With an ever-increasing human population size, increasing human impacts on the natural environment, and a growing scarcity of freshwater resources, the study of freshwater systems has become one of the most critical areas of focus for environmental scientists. Never before has there been such a need for an integrated understanding of lakes, wetlands, and flowing waters. This understanding must cross the boundaries of traditional disciplines, reach a wide international audience, and strongly facilitate the linkage between scientific discovery and its real-world application.

Large-scale mapping and predictive modeling of submerged aquatic vegetation in a shallow eutrophic lake.

A spatially intensive sampling program was developed for mapping the submerged aquatic vegetation (SAV) over an area of approximately 20,000 ha in a large, shallow lake in Florida, U.S. The sampling program integrates Geographic Information System (GIS) technology with traditional field sampling of SAV and has the capability of producing robust vegetation maps under a wide range of conditions, including high turbidity, variable depth (0 to 2 m), and variable sediment types. Based on sampling carried out in August-September 2000, we measured 1,050 to 4,300 ha of vascular SAV species and approximately 14,000 ha of the macroalga Chara spp. The results were similar to those reported in the early 1990s, when the last large-scale SAV sampling occurred. Occurrence of Chara was strongly associated with peat sediments, and maximal depths of occurrence varied between sediment types (mud, sand, rock, and peat). A simple model of Chara occurrence, based only on water depth, had an accuracy of 55%. It predicted occurrence of Chara over large areas where the plant actually was not found. A model based on sediment type and depth had an accuracy of 75% and produced a spatial map very similar to that based on observations. While this approach needs to be validated with independent data in order to test its general utility, we believe it may have application elsewhere. The simple modeling approach could serve as a coarse-scale tool for evaluating effects of water level management on Chara populations.