A Comparative Examination of Recent Changes in Nutrients and Lower Food Web Structure in Lake Michigan and Lake Huron.

The lower food webs of Lake Huron and Lake Michigan have experienced similar reductions in the spring phytoplankton bloom and summer populations of Diporeia and cladocerans since the early 2000s. At the same time phosphorus concentrations have decreased and water clarity and silica concentrations have increased. Key periods of change, identified by using a method based on sequential t-tests, were 2003-2005 (Huron) and 2004-2006 (Michigan). Estimated filtration capacity suggests that dreissenid grazing would have been insufficient to directly impact phytoplankton in the deeper waters of either lake by this time (mid 2000s). Despite some evidence of decreased chlorophyll:TP ratios, consistent with grazing limitation of phytoplankton, the main impact of dreissenids on the offshore waters was probably remote, e.g., through interception of nutrients by nearshore populations. A mass balance model indicates that decreased phosphorus loading could not account for observed in-lake phosphorus declines. However, model-inferred internal phosphorus dynamics were strongly correlated between the lakes, with periods of increased internal loading in the 1990s, and increased phosphorus loss starting in 2000 in Lake Michigan and 2003 in Lake Huron, prior to dreissenid expansion into deep water of both lakes. This suggests a limited role for deep populations of dreissenids in the initial phosphorus declines in the lakes, and also suggests a role for meteorological influence on phosphorus dynamics. The high synchrony in lower trophic level changes between Lake Michigan and Lake Huron suggests that both lakes should be considered when investigating underlying causal factors of these changes.

Multidimensional approach to invasive species prevention.

Nonindigenous species (NIS) cause global biotic homogenization and extinctions, with commercial shipping being a leading vector for spread of aquatic NIS. To reduce transport of NIS by ships, regulations requiring ballast water exchange (BWE) have been implemented by numerous countries. BWE appears to effectively reduce risk for freshwater ports, but provides only moderate protection of marine ports. In the near future, ships may be required to undertake ballast water treatment (BWT) to meet numeric performance standards, and BWE may be phased out of use. However, there are concerns that BWT systems may not operate reliably in fresh or turbid water, or both. Consequently, it has been proposed that BWE could be used in combination with BWT to maximize the positive benefits of both management strategies for protection of freshwater ports. We compared the biological efficacy of "BWE plus BWT" against "BWT alone" at a ballast water treatment experimental test facility. Our comparative evaluation showed that even though BWT alone significantly reduced abundances of all tested organism groups except total heterotrophic bacteria, the BWE plus BWT strategy significantly reduced abundances for all groups and furthermore resulted in significantly lower abundances of most groups when compared to BWT alone. Our study clearly demonstrates potential benefits of combining BWE with BWT to reduce invasion risk of freshwater organisms transported in ships' ballast water, and it should be of interest to policy makers and environmental managers.

Asymmetric, biraphid diatoms from the Laurentian Great Lakes.

This taxonomic account of light micrographs from the coastal Laurentian Great lakes contains taxa from the diatom genera Amphora, Halamphora, Cymbella, Cymbopleura, Delicatophycus, Encyonema, Encyonopsis, Reimeria, Gomphonema, Gomphosphenia, Gomphonella, Gomphosinica, and Gomphoneis. A total of 207 samples of surface sediment and periphyton collected from 106 wetland, high-energy, embayment, and deeper nearshore locales are represented. Light micrographs of 154 taxa are presented. Of these, 76 could not be fully identified as known taxa from the existing literature and so are given tentative names, numbers or conferred assignments. Lake and habitat specificity, modeled autecological optima for phosphorus and chloride, and tolerance to anthropogenic stressors are described for 39 of the more common taxa.

Zooplankton-phytoplankton biomass and diversity relationships in the Great Lakes.

Quantifying the relationship between phytoplankton and zooplankton may offer insight into zooplankton sensitivity to shifting phytoplankton assemblages and the potential impacts of producer-consumer decoupling on the rest of the food web. We analyzed 18 years (2001-2018) of paired phytoplankton and zooplankton samples collected as part of the United States Environmental Protection Agency (U.S. EPA) Great Lakes Biology Monitoring Program to examine both the long-term and seasonal relationships between zooplankton and phytoplankton across all five Laurentian Great Lakes. We also analyzed effects of phytoplankton diversity on zooplankton biomass, diversity, and predator-prey (zooplanktivore/grazer) ratios. Across the Great Lakes, there was a weak positive correlation between total algal biovolume and zooplankton biomass in both spring and summer. The relationship was weaker and not consistently positive within individual lakes. These trends were consistent over time, providing no evidence of increasing decoupling over the study period. Zooplankton biomass was weakly negatively correlated with algal diversity across lakes, whereas zooplankton diversity was unaffected. These relationships did not change when we considered only the edible phytoplankton fraction, possibly due to the high correlation between total and edible phytoplankton biovolume in most of these lakes. Lack of strong coupling between these producer and consumer assemblages may be related to lagging responses by the consumers, top-down effects from higher-level consumers, or other confounding factors. These results underscore the difficulty in predicting higher trophic level responses, including zooplankton, from changes in phytoplankton assemblages.

Early presence of Bythotrephes cederströmii (Cladocera: Cercopagidae) in lake sediments in North America: evidence or artifact?

The spiny water flea (Bythotrephes cederströmii), a freshwater crustacean considered to be the world's best-studied invasive zooplankter, was first recorded in North America in the Laurentian Great Lakes during the 1980s. Its arrival is widely considered to be the result of ocean-going cargo ships that translocated contaminated ballast water from Eurasia to the Great Lakes during the 1970-1980s. The subsequent first discovery of the species in inland lakes is consistent with the hypothesis that propagules dispersed initially from established Great Lakes populations. Here we present evidence of exoskeletal remains, including mandibles, tail spines, and resting eggs, in 210Pb-dated lake sediment cores, which suggests that B. cederströmii was already resident in four inland North American lakes (two in Minnesota, USA; two in Ontario, Canada) by at least the early 1900s. Densities of exoskeletal remains were low and relatively steady from first appearance until about 1990, after which time they increased in all cores. The earliest evidence that we found was a mandible at 33-cm depth (pre-1650) in the sediments of Three Mile Lake, Ontario, Canada. These unexpected findings challenge the current paradigm of B. cederströmii invasion, renew uncertainty about the timing and sequence of its colonization of North American lakes, and potentially question our ability to detect invasive species with traditional sampling methods. We attempted to eliminate errors in the dated stratigraphies of the exoskeletal remains that might have been introduced either methodologically (e.g., core-wall smearing) or naturally (e.g., bioturbation). Nonetheless, given the very low numbers of subfossils encountered, questions remain about the possible artifactual nature of our observations and therefore we regard our results as 'preliminary findings' at this time.

Anthropocene geochemistry of metals in sediment cores from the Laurentian Great Lakes.

Geochemical analyses applied to lake sedimentary records can reveal the history of pollution by metals and the effects of remedial efforts. Lakes provide ideal environments for geochemical studies because they have steady deposition of fine grained material suitable for fixation of pollutants. The Laurentian Great Lakes are the most studied system in this field, and they have well-preserved chronological profiles. To date, this important system has been considered in parts for inorganic geochemistry, hampering basin-wide conclusions regarding metal contamination. We filled spatial and temporal gaps in a comprehensive geochemical analysis of 11 sediment cores collected from all five Great Lakes. Hierarchical cluster analysis of all Great Lakes samples divided the metal analytes into five functional groups: (1) carbonate elements; (2) metals and oxides with diverse natural sources, including a subgroup of analytes known to be anthropogenically enriched (Cd, Pb, Sn, Zn, and Sb); (3) common crustal elements; (4) metals related to coal and nuclear power generation; and (5) all of the co-occurring rare earth elements. Two contamination indices (Igeo and EF) applied to sedimentary metals indicated that Na, Co, Mn, Cd, Pb, Ta, and Cu were each, at some point during the Anthropocene, the most enriched metal pollutants in Great Lakes sediments. Land uses correlated with the metal analytes, such as increases in contaminant metals with the rise in catchment population and increases in carbonate elements (e.g. Ca) with agriculture. Certain contamination trends were observed basin-wide, such as for the atmospheric pollutant Pb, which followed a rise associated with fossil fuel combustion and a decline following the ban of leaded gasoline. Other trends were lake-specific, such as recent high concentrations of Na in Lake Superior, likely due to road salt applications, and a late-20th-century peak in Ca associated with algal whiting events in Lake Ontario. Some metals exceeded guidelines for sediment quality, in some cases prior to European settlement of the basin, indicating that a paleolimnological context is important for appropriate management of sediment contamination. The Great Lakes are sensitive to environmental changes such as pollution by metals, and it is clear that while there has been remedial success, results from the uppermost intervals of cores indicate ongoing problems.

Metallic elements and oxides and their relevance to Laurentian Great Lakes geochemistry.

The Laurentian Great Lakes are the most studied system in lake geochemistry and have well-preserved chronological profiles. Metals play numerous critical roles in natural and anthropogenic characteristics of lake ecosystems, so patterns in the historical records of metals from sedimentary cores provide important information about environmental baselines and human impacts. Relevant studies of Great Lakes geochemistry are listed, and we follow with encyclopedic descriptions of metals and their oxides in the lakes. These descriptions include likely natural and anthropogenic sources of elements, their known history from previous paleoecological studies, and their status as potential contaminants of concern. Despite the well-studied geology of the Great Lakes catchment, sourcing elements was sometimes difficult due to materials often being moved long distances by glaciation and the global prevalence of atmospheric pollutants. We summarized available information on metals and their roles as geochemical indicators in the Great Lakes.

Environmental control of diatom community size structure varies across aquatic ecosystems.

Changes in the size structure of photoautotrophs influence food web structure and the biogeochemical cycling of carbon. Decreases in the median size of diatoms within communities, in concert with climate warming and water column stratification, have been observed over the Cenozoic in the ocean and over the last 50 years in Lake Tahoe. Decreases in the proportion of larger plankton are frequently observed in response to reduced concentrations of limiting nutrients in marine systems and large stratified lakes. By contrast, we show a decrease in the median size of planktonic diatoms in response to higher nutrient concentrations in a set of intermediate-sized alkaline lakes. Climate-induced increases in the frequency, duration and strength of water column stratification may select smaller planktonic species in the ocean and larger lakes owing to a reduction in nutrient availability and sinking rates, while light limitation, stimulated by nutrient eutrophication and high chlorophyll concentrations, may select smaller species within a community owing to their high light absorption efficiencies and lower sinking rates. The relative importance of different physiological and ecological rates and processes on the size structure of communities varies in different aquatic systems owing to varying combinations of abiotic and biotic constraints.

Consideration of species-specific diatom indicators of anthropogenic stress in the Great Lakes.

Robust inferences of environmental condition come from bioindicators that have strong relationships with stressors and are minimally confounded by extraneous environmental variables. These indicator properties are generally assumed for assemblage-based indicators such as diatom transfer functions that use species abundance data to infer environmental variables. However, failure of assemblage approaches necessitates the interpretation of individual dominant taxa when making environmental inferences. To determine whether diatom species from Laurentian Great Lakes sediment cores have the potential to provide unambiguous inferences of anthropogenic stress, we evaluated fossil diatom abundance against a suite of historical environmental gradients: human population, agriculture, mining, atmospheric nutrient deposition, atmospheric temperature and ice cover. Several diatom species, such as Stephanodiscus parvus, had reliable relationships with anthropogenic stress such as human population. However, many species had little or no indicator value or had confusing relationships with multiple environmental variables, suggesting one should be careful when using those species to infer stress in the Great Lakes. Recommendations for future approaches to refining diatom indicators are discussed, including accounting for the effects of broad species geographic distributions to minimize region-specific responses that can weaken indicator power.

Historical watershed stressors for the Laurentian Great Lakes.

This report provides a detailed set of historical stressor data for 60 watersheds comprising the Laurentian Great Lakes basin. Archival records were transcribed from public records to create quantitative data on human activities: population, mining, deforestation, and agriculture. Yearly records of stressors are provided from 1780 through 2010. These data may be used to track historical impacts on Great Lakes coastal and open water conditions. They may further be used to examine corresponding effects on response variables such as biological communities quantified during monitoring and palaeoecological programmes. OPEN PRACTICES: This article has earned an Open Data badge for making publicly available the digitally-shareable data necessary to reproduce the reported results. The data is available at https://doi.org/10.1594/PANGAEA.885879. Learn more about the Open Practices badges from the Center for Open Science: https://osf.io/tvyxz/wiki.

Are harmful algal blooms becoming the greatest inland water quality threat to public health and aquatic ecosystems?

In this Focus article, the authors ask a seemingly simple question: Are harmful algal blooms (HABs) becoming the greatest inland water quality threat to public health and aquatic ecosystems? When HAB events require restrictions on fisheries, recreation, and drinking water uses of inland water bodies significant economic consequences result. Unfortunately, the magnitude, frequency, and duration of HABs in inland waters are poorly understood across spatiotemporal scales and differentially engaged among states, tribes, and territories. Harmful algal bloom impacts are not as predictable as those from conventional chemical contaminants, for which water quality assessment and management programs were primarily developed, because interactions among multiple natural and anthropogenic factors determine the likelihood and severity to which a HAB will occur in a specific water body. These forcing factors can also affect toxin production. Beyond site-specific water quality degradation caused directly by HABs, the presence of HAB toxins can negatively influence routine surface water quality monitoring, assessment, and management practices. Harmful algal blooms present significant challenges for achieving water quality protection and restoration goals when these toxins confound interpretation of monitoring results and environmental quality standards implementation efforts for other chemicals and stressors. Whether HABs presently represent the greatest threat to inland water quality is debatable, though in inland waters of developed countries they typically cause more severe acute impacts to environmental quality than conventional chemical contamination events. The authors identify several timely research needs. Environmental toxicology, environmental chemistry, and risk-assessment expertise must interface with ecologists, engineers, and public health practitioners to engage the complexities of HAB assessment and management, to address the forcing factors for HAB formation, and to reduce the threats posed to inland surface water quality.

Laurentian Great Lakes phytoplankton and their water quality characteristics, including a diatom-based model for paleoreconstruction of phosphorus.

Recent shifts in water quality and food web characteristics driven by anthropogenic impacts on the Laurentian Great Lakes warranted an examination of pelagic primary producers as tracers of environmental change. The distributions of the 263 common phytoplankton taxa were related to water quality variables to determine taxon-specific responses that may be useful in indicator models. A detailed checklist of taxa and their environmental optima are provided. Multivariate analyses indicated a strong relationship between total phosphorus (TP) and patterns in the diatom assemblages across the Great Lakes. Of the 118 common diatom taxa, 90 (76%) had a directional response along the TP gradient. We further evaluated a diatom-based transfer function for TP based on the weighted-average abundance of taxa, assuming unimodal distributions along the TP gradient. The r(2) between observed and inferred TP in the training dataset was 0.79. Substantial spatial and environmental autocorrelation within the training set of samples justified the need for further model validation. A randomization procedure indicated that the actual transfer function consistently performed better than functions based on reshuffled environmental data. Further, TP was minimally confounded by other environmental variables, as indicated by the relatively large amount of unique variance in the diatoms explained by TP. We demonstrated the effectiveness of the transfer function by hindcasting TP concentrations using fossil diatom assemblages in a Lake Superior sediment core. Passive, multivariate analysis of the fossil samples against the training set indicated that phosphorus was a strong determinant of historical diatom assemblages, verifying that the transfer function was suited to reconstruct past TP in Lake Superior. Collectively, these results showed that phytoplankton coefficients for water quality can be robust indicators of Great Lakes pelagic condition. The diatom-based transfer function can be used in lake management when retrospective data are needed for tracking long-term degradation, remediation and trajectories.

COMPARISON OF SIMPLE AND MULTIMETRIC DIATOM-BASED INDICES FOR GREAT LAKES COASTLINE DISTURBANCE(1).

Because diatom communities are subject to the prevailing water quality in the Great Lakes coastal environment, diatom-based indices can be used to support coastal-monitoring programs and paleoecological studies. Diatom samples were collected from Great Lakes coastal wetlands, embayments, and high-energy sites (155 sites), and assemblages were characterized to the species level. We defined 42 metrics on the basis of autecological and functional properties of species assemblages, including species diversity, motile species, planktonic species, proportion dominant taxon, taxonomic metrics (e.g., proportion Stephanodiscoid taxa), and diatom-inferred (DI) water quality (e.g., DI chloride [Cl]). Redundant metrics were eliminated, and a diatom-based multimetric index (MMDI) to infer coastline disturbance was developed. Anthropogenic stresses in adjacent coastal watersheds were characterized using geographic information system (GIS) data related to agricultural and urban land cover and atmospheric deposition. Fourteen independent diatom metrics had significant regressions with watershed stressor data; these metrics were selected for inclusion in the MMDI. The final MMDI was developed as the weighted sum of the selected metric scores with weights based on a metric's ability to reflect anthropogenic stressors in the adjacent watersheds. Despite careful development of the multimetric approach, verification using a test set of sites indicated that the MMDI was not able to predict watershed stressors better than some of the component metrics. From this investigation, it was determined that simpler, more traditional diatom-based metrics (e.g., DI Cl, proportion Cl-tolerant species, and DI total phosphorus [TP]) provide superior prediction of overall stressor influence at coastal locales.

Human influences on water quality in Great Lakes coastal wetlands.

A better understanding of relationships between human activities and water chemistry is needed to identify and manage sources of anthropogenic stress in Great Lakes coastal wetlands. The objective of the study described in this article was to characterize relationships between water chemistry and multiple classes of human activity (agriculture, population and development, point source pollution, and atmospheric deposition). We also evaluated the influence of geomorphology and biogeographic factors on stressor-water quality relationships. We collected water chemistry data from 98 coastal wetlands distributed along the United States shoreline of the Laurentian Great Lakes and GIS-based stressor data from the associated drainage basin to examine stressor-water quality relationships. The sampling captured broad ranges (1.5-2 orders of magnitude) in total phosphorus (TP), total nitrogen (TN), dissolved inorganic nitrogen (DIN), total suspended solids (TSS), chlorophyll a (Chl a), and chloride; concentrations were strongly correlated with stressor metrics. Hierarchical partitioning and all-subsets regression analyses were used to evaluate the independent influence of different stressor classes on water quality and to identify best predictive models. Results showed that all categories of stress influenced water quality and that the relative influence of different classes of disturbance varied among water quality parameters. Chloride exhibited the strongest relationships with stressors followed in order by TN, Chl a, TP, TSS, and DIN. In general, coarse scale classification of wetlands by morphology (three wetland classes: riverine, protected, open coastal) and biogeography (two ecoprovinces: Eastern Broadleaf Forest [EBF] and Laurentian Mixed Forest [LMF]) did not improve predictive models. This study provides strong evidence of the link between water chemistry and human stress in Great Lakes coastal wetlands and can be used to inform management efforts to improve water quality in Great Lakes coastal ecosystems.