Status,causes and controls of cyanobacterial blooms in Lake Erie

The Laurentian Great Lakes are among the most prominent sources of fresh water in the world. Lake Erie's infamous cyanobacterial blooms have, however, threatened the health of this valuable freshwater resource for decades. Toxic blooms dominated by the cyanobacterium Microcystis aeruginosa have most recently been one of primary ecological concerns for the lake. These toxic blooms impact the availability of potable water, as well as public health and revenues from the tourism and fishery industries. The socioeconomic effects of these blooms have spurred research efforts to pinpoint factors that drive bloom events. Despite decades of research and mitigation efforts, these blooms have expanded both in size and duration in recent years. However, through continued joint efforts between the Canadian and United States governments, scientists, and environmental managers, identification of the factors that drive bloom events is within reach. This review provides a summary of historical and contemporary research efforts in the realm of Lake Erie's harmful cyanobacterial blooms, both in terms of experimental and management achievements and insufficiencies, as well as future directions on the horizon for the lake's research community.     

Fungal community dynamics associated with harmful cyanobacterial blooms in two Great Lakes

Harmful algal blooms (HABs) impose major costs on aquatic ecosystems worldwide, including the Laurentian Great Lakes. Microbial consumers, including fungi, can have important interactions with bloom-forming algae and cyanobacteria, although relatively few studies have investigated the relationship between fungi and HABs. We examined changes in the aquatic fungal community coincident with the occurrence of large cyanobacterial blooms in two areas of the Great Lakes (western Lake Erie and Saginaw Bay, Lake Huron). We collected water samples over the course of bloom development, peak, and decline from 3 sites in western Lake Erie on 11 dates and 2 sites in Saginaw Bay on 4 dates. Single molecule sequencing (PacBio RS II) with two molecular markers (the internal transcribed spacer (ITS) of the rRNA locus using fungal-specific primers and the 18S rRNA with primers targeting early-diverging lineages of fungi) was used to estimate fungal community composition. Results indicate a diverse fungal community within the lakes, including several major fungal phyla. The Chytridiomycota were particularly well-represented (54.8% and 45.4% of ITS and 18S sequences, respectively), and we also found representation from both Cryptomycota and Aphelidiomycota, which are putatively obligate intracellular parasites. Further, we found associations between the fungal community (alpha diversity; community composition) and measures of bloom magnitude (chlorophyll a, phycocyanin, and microcystin concentrations) in western Lake Erie. Our results suggest potentially important spatial and temporal heterogeneity in the fungal community that motivates further research on functional importance of fungi in the Great Lakes and consequences for HABs and freshwater ecosystems more broadly.     

Cyanobacterial blooms in the central basin of Lake Erie: Potentials for cyanotoxins and environmental drivers

Lake Erie western basin (WB) cyanobacterial blooms are a yearly summer occurrence; however, blooms have also been reported in the offshore waters of the central basin (CB), and very little is known about what drives these blooms or their potential for cyanobacterial toxins. Cyanobacteria Index was quantified using MODIS and MERIS data for the CB between 2003 and 2017, and water samples were collected between 2013 and 2017. The goals were to 1) quantify cyanobacteria, 2) determine environmental drivers of CB blooms, and 3) determine the potential for cyanobacterial toxins in the CB. Dolichospermum (Anabaena) occurred in the CB during July before the onset of the WB bloom, and then in August and September, the cyanobacteria community shifted towards Microcystis. The largest Dolichospermum blooms (2003, 2012, 2013, and 2015) were associated with reduced water clarity (Secchi disk depth?<?4?m), whereas large CB Microcystis blooms (2011 and 2015) were associated with large WB blooms. Dolichospermum blooms occurred in high nitrate concentrations (>20?μmol/L) and high nitrogen-to?phosphorus ratios (>100), which indicate nutrient concentrations or ratios did not select for Dolichospermum. Additionally, the sxtA gene, but not mcyE or microcystins, were detected in the CB during July 2016 and 2017. The mcyE gene and microcystins were detected in the CB during August 2016 and 2017. The results indicate the CB's potential for cyanotoxins shifts from saxitoxins to microcystins throughout the summer. Continued monitoring of cyanobacteria and multiple cyanobacterial toxins is recommended to ensure safe drinking water for CB coastal communities.     

Mapping cyanobacterial blooms in the Great Lakes using MODIS

Toxin-producing Cyanobacteria have been documented in Lake Erie and Ontario in the last several years. We developed algorithms to discriminate potentially toxic cyanobacterial blooms from other harmless phytoplankton blooms and to extract relative phycocyanin abundances from Moderate Resolution Imaging Spectrometer (MODIS) satellite data. Lee's quasi-analytical algorithm was used to calculate total absorption and backscatter from the 250 m, 500 m and 1 km bands of MODIS scenes. A non-negative least square algorithm was then utilized to discern relative concentrations of Chlorophyta (green algae), phycocyanin-rich Cyanobacteria (blue-green algae), and colored dissolved organic matter and suspended sediments combined in lake waters using published absorption spectra for these components. MODIS-derived cyanobacterial concentrations and/or bloom distributions from 10 scenes acquired in the summers of 2004 and 2005 were successfully verified against contemporaneous calibrated measurements of pigments that were acquired from measurements made using continuous fluorimetric measurements of surface water (1 m depth) from six cruises, and three additional cyanobacterial blooms reported in the scientific literature between 2002 and 2006. These results demonstrate that this methodology could be used to develop a cost-effective practical screening method for rapid detection and warning of potentially toxic cyanobacterial blooms in the lower Great Lakes.     

Status of the major aquaculture carps of China in the Laurentian Great Lakes Basin

There is concern of economic and environmental damage occuring if any of the four major aquacultured carp species of China, black carp Mylopharyngodon piceus, bighead carp Hypophthalmichthys nobilis, silver carp H. molitrix, or grass carp Ctenopharyngodon idella, were to establish in the Laurentian Great Lakes. All four are reproducing in the Mississippi River Basin. We review the status of these fishes in relation to the Great Lakes and their proximity to pathways into the Great Lakes, based on captures and collections of eggs and larvae. No black carp have been captured in the Great Lakes Basin. One silver carp and one bighead carp were captured within the Chicago Area Waterway System, on the Great Lakes side of electric barriers designed to keep carp from entering the Great Lakes from the greater Mississippi River Basin. Three bighead carp were captured in Lake Erie, none later than the year 2000. By December 2019, at least 650 grass carps had been captured in the Great Lakes Basin, most in western Lake Erie, but none in Lake Superior. Grass carp reproduction has been documented in the Sandusky and Maumee rivers in Ohio, tributaries of Lake Erie. We also discuss environmental DNA (eDNA) results as an early detection and monitoring tool for bighead and silver carps. Detection of eDNA does not necessarily indicate presence of live fish, but bigheaded carp eDNA has been detected on the Great Lakes side of the barriers and in a small proportion of samples from the western basin of Lake Erie.     

Assessment of nitrogen fixation rates in the Laurentian Great Lakes

Nitrogen fixation (N_Fix) is an important, yet understudied, microbial process in aquatic ecosystems, especially in the Laurentian Great Lakes (LGL). To date, a dearth of nitrogen fixation rate measurements exists in the LGL, are from temporally isolated studies, and were collected primarily from near-shore and surface water environments. Evidence of nitrogen accumulation across the Laurentian Great Lakes suggest that we do not have a firm grasp on nitrogen cycling in large lakes. Thus, we sought to quantify the spatial variability of N_Fix in the LGL. We found lakes are significantly different in N_Fix rates from one another and that rates are depth dependent. Overall mean surface N_Fix rates of Lakes Superior, Michigan, Huron, Erie and Ontario were 0.024, 0.020, 0.069, 0.145, and 0.078 (nmol N₂/L/hr), respectively. Likewise, we found the Western, Central and Eastern basins of Lake Erie are significantly different in N_Fix rates (0.1540, 0.1032, 0.0738 nmol N₂/L/hr). However, we found no significant difference in N_Fix rates between near and offshore sites in Lake Erie, which may have been biased due to a cyanobacterial bloom containing a nitrogen-fixing Dolichospermum sp. Linear regression models indicate N_Fix is generally positively correlated with chlorophyll-a concentration and negatively correlated with oxidized nitrogen species concentrations. However, Lakes Erie and Huron exhibited a positive linear relationship with oxidized nitrogen, suggesting that N_Fix may persist to meet cellular and community nitrogen demands. Together, our data highlight N_Fix is important despite the presence of abundant nitrogen in all LGL.     

Detection and expression of genes for phosphorus metabolism in picocyanobacteria from the Laurentian Great Lakes

The pelagic regions of Lake Superior and eastern Lake Erie (Laurentian Great Lakes) are typically phosphorus (P)-limited environments, and picocyanobacteria of the genus Synechococcus spp. are prominent primary producers during the summer. As a proxy for their utilization of organic P, the expression of two genes, phnD and phoX, was monitored. The phnD gene encodes the phosphonate binding protein of the ABC-type phosphonate transporter, whereas the phoX gene encodes a calcium-dependent alkaline phosphatase. Furthermore, to assess the ability of freshwater Synechococcus spp. to substitute sulfolipids for phospholipids, sqdX gene (cyanobacterial sulfolipid synthase) expression was examined. We employed PCR primers to detect the presence of all three genes in the endemic Synechococcus spp., and RT-PCR assays of cultured freshwater strains and environmental samples to assess the degree of P-stress in the phytoplankton. We show that the phnD gene was constitutively expressed, suggesting that freshwater picocyanobacteria were metabolizing exogenous phosphonate compounds in Lakes Erie and Superior. By contrast, phoX was regulated by P bioavailability. We also provide evidence that sqdX is expressed during increased growth rates in phosphorus-replete conditions, suggesting that sulfolipid synthesis is not a P conservation mechanism for freshwater Synechococcus spp.     

Determining remote sensing spatial resolution requirements for the monitoring of harmful algal blooms in the Great Lakes

Harmful algal blooms (HABs) have become a major health and environmental concern in the Great Lakes. In 2014, severe HABs prompted the State of Ohio to request NASA Glenn Research Center (GRC) to assist with monitoring algal blooms in Lake Erie. The most notable species of HAB is Microcystis aeruginosa, a hepatotoxin producing cyanobacteria that is responsible for liver complications for humans and other fauna that come in contact with these blooms. NASA GRC conducts semiweekly flights in order to gather up-to-date imagery regarding the blooms' spatial extents and concentrations. Airborne hyperspectral imagery is collected using two hyperspectral imagers, HSI-2 and HSI-3. Hyperspectral imagery is necessary in order to conduct experiments on differentiation of algal bloom types based on their spectral reflectance. In this analysis, imagery from September 19, 2016 was utilized to study the subpixel variability within the footprint of arbitrary sized pixels using several analysis techniques. This particular data set is utilized because it represents a worst case scenario where there is significant potential for public health concern due to high concentrations of microcystin toxin found in the water on this day and the concurrent observational challenges to accurately measure the algal bloom concentration variability with a remote sensing system due to the blooms high spatial variability. It has been determined that the optimal spatial resolution to monitor algal blooms in the Great Lakes is at most 50 m, and for much lower error 25 m, thus allowing for greater ease in identifying high concentration blooms near the surface. This resolution provides the best sensitivity to high concentration areas that are of significant importance in regard to human health and ecological damage.     

Investigation of viruses in Diporeia spp. from the Laurentian Great Lakes and Owasco Lake as potential stressors of declining populations

The benthic amphipod Diporeia is an ecologically and biogeochemically important constituent of deep freshwater lakes in North America. The proliferation of dreissenid mussels in the mid-1990s coincided with a sharp decrease in Diporeia populations in several Laurentian Great Lakes; however the ultimate cause and mechanisms of their decline are still unknown. Here we examined the composition of DNA viruses associated with Diporeia collected from populations of Lake Michigan that had declined and stable populations in Lake Superior and Owasco Lake (Finger Lake in central New York State). Viral metagenomic libraries from Owasco Lake and Lake Superior were comprised primarily of bacteriophages, which may infect bacteria within the amphipod microbiome. In contrast, the metagenomic library from Lake Michigan contained well-represented ssDNA circular viral genomes. The prevalence and viral load of one putative Type V ssDNA circular virus (LM29173) that recruited almost 30% of total viral sequence reads in the Lake Michigan library was analyzed by quantitative PCR. The prevalence of LM29173 was over two orders of magnitude greater in Lake Michigan compared to the other two lakes. Although further research is necessary to establish the pathology and epidemiological extent of viral-Diporeia interactions, our data suggest that viruses may be numerically significant constituents of the Diporeia microbiome, and if pathogenic some of these viruses may be a stressor of Great Lakes Diporeia populations. Our data further indicate that special attention should be given to the circovirus that was prevalent in the declining Michigan population but uncommon in the other two lakes.     

Brachionus leydigii (Monogononta: Ploima) reported from the western basin of Lake Erie

Several species of non-indigenous planktonic invertebrates have historically been introduced to the Laurentian Great Lakes. Previous introductions of non-indigenous planktonic invertebrates to the Great Lakes have been crustacean zooplankton, specifically Cladocera and Copepoda. This report documents the first known occurrence of Brachionus leydigii var. tridentatus (Zernov, 1901) in Lake Erie and possibly the first detection of a non-indigenous rotifer species in the Laurentian Great Lakes. The specimen was collected from a U.S. EPA monitoring station in the western basin of Lake Erie on April 4, 2016.     

Historical changes and current status of crayfish diversity and distribution in the Laurentian Great Lakes

Despite increasing recognition of the importance of invertebrates, and specifically crayfish, to nearshore food webs in the Laurentian Great Lakes, past and present ecological studies in the Great Lakes have predominantly focused on fishes. Using data from many sources, we provide a summary of crayfish diversity and distribution throughout the Great Lakes from 1882 to 2008 for 1456 locations where crayfish have been surveyed. Sampling effort was greatest in Lake Michigan, followed by lakes Huron, Erie, Superior, and Ontario. A total of 13 crayfish species occur in the lakes, with Lake Erie having the greatest diversity (n = 11) and Lake Superior having the least (n = 5). Five crayfish species are non-native to one or more lakes. Because Orconectes rusticus was the most widely distributed non-native species and is associated with known negative impacts, we assessed its spread throughout the Great Lakes. Although O. rusticus has been found for over 100 years in Lake Erie, its spread there has been relatively slow compared to that in lakes Michigan and Huron, where it has spread most rapidly since the 1990s and 2000, respectively. O. rusticus has been found in both lakes Superior and Ontario for 22 and 37 years, respectively, and has expanded little in either lake. Our broad spatial and temporal assessment of crayfish diversity and distribution provides a baseline for future nearshore ecological studies, and for future management efforts to restore native crayfish and limit non-native introductions and their impact on food web interactions.     

First report of tubenose goby (Proterorhinus semilunaris) in the eastern basin of Lake Erie

The tubenose goby (Proterorhinus semilunaris), native to the Ponto-Caspian region, was first discovered in the Laurentian Great Lakes in 1990 after it was introduced through ballast water discharge. Compared with Neogobius melanostomus, another exotic gobiid from the Ponto-Caspian, colonization of the Great Lakes by P. semilunaris has been slow, with reports of the species being largely confined to the Huron-Erie Corridor (HEC) and western portions of Lake Erie and Lake Superior. This is the first report of P. semilunaris in the Great Lakes east of the western basin of Lake Erie. Between 28 June and 27 July, 2012, 176 P. semilunaris were collected from shallow (< 1.2 m) water of Marina Lake, a 40 ha embayment in Presque Isle State Park (Erie, PA). The large number of P. semilunaris collected at the site and the presence of individuals as small as 17 mm total length suggest an established population. However, the mechanism by which P. semilunaris was introduced to Presque Isle Bay is not clear.     

European Valve Snail Valvata piscinalis (Müller) in the Laurentian Great Lakes Basin

Previously reported from the lower Great Lakes basin and St. Lawrence and Hudson rivers, the nonindigenous gastropod Valvata piscinalis was found for the first time in Superior Bay (Minnesota) of Lake Superior, Lake Michigan (Wisconsin), and Oneida Lake (New York) of the Lake Ontario basin. This snail was not abundant in Lakes Superior and Michigan, whereas in eutrophic Oneida Lake it reached a maximum density of 1,690 individuals/m² (mean density = 216 individuals/m²). Human-mediated disturbances could facilitate the range extension of this snail by providing dispersal opportunities (e.g., canals, shipping traffic) or increasing nutrients (e.g., eutrophication). A native of the Palaearctic region, V. piscinalis has colonized sites across the Great Lakes basin, suggesting that it will likely become common in disturbed Great Lakes areas.     

Toxaphene trends in the Great Lakes fish

As part of the U.S. Great Lakes Fish Monitoring and Surveillance Program (GLFMSP), more than 300 lake trout (Salvelinus namaycush) and walleye (Stizostedion vitreum vitreum) collected from the Laurentian Great Lakes each year from 2004 to 2009, have been analyzed for total toxaphene and eight selected congeners. The analytical results show fish toxaphene concentrations are quite different among lakes. Between 2004 and 2009, Lake Superior lake trout had the highest concentration (119 to 482 ng/g) and Lake Erie walleye had the lowest concentration (18 to 47 ng/g). Combining these results with the historical total toxaphene data (1977–2003), temporal changes were examined for each lake. Because of different analytical methods used in the previous studies, the historical data were adjusted using a factor of 0.56 based on a previous inter-method comparison in our laboratory. Trend analysis using an exponential decay regression showed that toxaphene in Great Lakes fish exhibited a significant decrease in all of the lakes with t_1/2 (confidence interval) of 0.9 (0.8–1.1) years for Lake Erie walleye, 3.8 (3.5–4.1) years for Lake Huron lake trout, 5.6 (5.1–6.1) years for Lake Michigan lake trout, 7.5 (6.7–8.4) years for Lake Ontario lake trout and 10.1 (8.2–13.2) years for Lake Superior lake trout. Parlars 26, 50 and 62 were the dominant toxaphene congeners accounting for 0.53% to 41.7% of the total toxaphene concentration. Concentrations of these congeners generally also decreased over time.     

Status of the amphipod Diporeia spp. in Lake Superior, 2006–2016

The amphipod Diporeia spp. has historically been an important component of the benthic food web of the Laurentian Great Lakes. The Great Lakes Water Quality Agreement included its population density as an indicator of ecological condition for Lake Superior, with target values of 220–320 m^?2 in nearshore areas (≤100 m depth) and 30–160 m^?2 in offshore areas (>100 m). To assess the status of Diporeia in Lake Superior, we used a probability-based lake-wide survey design to obtain estimates of Diporeia density and biomass in 2006, 2011 and 2016. A PONAR grab sampler was used to collect Diporeia at 50–53 sites each year, with approximately half in the nearshore (<100 m depth) region of the lake and half in the offshore. The mean area-weighted lake-wide density was 395 ± 56 (SE) m^?2 in 2006, 756 ± 129 m^?2 in 2011, and 502 ± 60 m^?2 in 2016. For all years, both density and biomass were greater in the nearshore than in the offshore stratum. The densities for 2006–2016 were 3–5 times higher than those reported from a lake-wide survey conducted in 1973 by the Canada Centre for Inland Waters. The severe declines in Diporeia populations observed in the other Great Lakes during recent decades have apparently not occurred in Lake Superior. Further research is needed to understand spatial and temporal variability of Diporeia populations in Lake Superior to enhance the utility of Diporeia density as an indicator of benthic condition.     

Algal bloom transport in Lake Erie using remote sensing and hydrodynamic modelling: Sensitivity to buoyancy velocity and initial vertical distribution

In this study, we simulate three-dimensional transport of algal blooms in Lake Erie using a combination of remote sensing and hydrodynamic modelling. The remote sensing algorithms use data from the Sentinel-3 OLCI satellite sensor to derive chlorophyll-a concentration from cyanobacteria blooms in Lake Erie. The derived chlorophyll-a concentration initializes an algal bloom transport model driven by the lake component of the Water Cycle Prediction System for the Great Lakes, a system of coupled atmosphere-lake-hydrological models operated out of Environment and Climate Change Canada. The bloom is modelled as Microcystis aeruginosa, a buoyant species that is often dominant in harmful algal blooms in western Lake Erie. Short-term (a few days) predictions of algal bloom transport from July 27 to October 8, 2017 are modelled in both Eulerian and Lagrangian frameworks. The Eulerian framework is used to evaluate the sensitivity of model results to the initial vertical distribution of the bloom. In this work, the Lagrangian framework is limited to two-dimensional surface confined particles. We use several error metrics to evaluate model predictions. We find that results are sensitive to the buoyancy velocity for cases where the bloom was initially distributed over a large portion of the water column. An initial vertical distribution selected from modelled chlorophyll-a half depth shows the highest accuracy for the entire range of buoyancy velocities tested. We also find that the Pierce skill score is difficult to interpret, particularly in cases where bloom intensity is greatly overpredicted by the model.     

Molecular characterization of bacterial communities associated with sediments in the Laurentian Great Lakes

The Laurentian Great Lakes freshwater ecosystem comprises a series of interconnected lakes exhibiting wide spatial differences in hydrogeochemistry, productivity, and allochthonous inputs of nutrients and xenobiotics. Bacterial communities associated with offshore sediments in this ecosystem have been rarely studied. We evaluated bacterial communities associated with sediments from 10 locations in four of the five Great Lakes (Lakes Superior, Michigan, Huron, and Ontario) by generating16S rRNA pyrosequence libraries. Pyrosequencing analysis revealed the presence of 26 bacterial phyla and proteobacterial classes among Great Lakes sediment samples. Actinobacteria, Acidobacteria, Betaproteobacteria, and Gammaproteobacteria were the most abundant groups of bacteria. Redundancy analysis was used to examine the role of sediment properties, including depth and chemical composition, in shaping bacterial community structure. One sample from Lake Huron was distinctly different from all other samples. Phylogenetic analysis revealed that the sample contained greater abundances of groups of bacteria associated with polluted environments. This study constitutes the most extensive examination of bacteria associated with Laurentian Great Lakes sediments and sheds useful insight into the microbial ecology of the Great Lakes.     

Changes in Fecundity and Egg Lipid Content of Lake Whitefish (Coregonus clupeaformis) in the Upper Laurentian Great Lakes between 1986–87 and 2003–05

Lake whitefish (Coregonus clupeaformis Mitchill), an important commercial species in the Laurentian Great Lakes, have experienced decreased growth and condition in regions of the upper Great Lakes over the past 20 years. Increases in lake whitefish density and decreases in the density of Diporeia spp., an energy rich and historically important part of the lake whitefish diet, have been implicated in the recent declines in lake whitefish growth and condition. The goal of this study was to describe lake whitefish fecundity, egg lipid content, and total ovary lipid content in selected regions of Lakes Huron, Michigan, and Superior in 1986–87 and 2003–05, two time periods with different lake whitefish and Diporeia densities. Under conditions of high lake whitefish density and low Diporeia density, female lake whitefish in the upper Laurentian Great Lakes generally produced fewer eggs. Egg lipid content was higher in 2003–05 than in 1986–87 at all sites, regardless of changes in lake whitefish or Diporeia densities. Total ovary lipid content and lake whitefish abundance were inversely related, while there was no significant relationship between total ovary lipid content and Diporeia density. The amount of energy that lake whitefish invested in egg production was more closely associated with lake whitefish abundance than with Diporeia density. This study provides evidence that recent changes in production dynamics of Great Lakes lake whitefish have not been driven solely by declines in Diporeia but have been significantly influenced by lake whitefish abundance.     

A cross-lake comparison of crustacean zooplankton communities in the Laurentian Great Lakes, 1997–2016

Spring and summer open-water crustacean zooplankton communities were examined across all five Laurentian Great Lakes from 1997 to 2016. Spring communities were dominated by calanoid (lakes Superior, Huron and Michigan) or cyclopoid (lakes Erie and Ontario) copepods. Volumetric biomass of summer communities increased along an assumed trophic gradient (Superior, Huron, Michigan, Ontario; eastern, central and western Erie), as did dominance by cyclopoids and cladocerans. Over the time series of the study, summer communities in lakes Michigan, Huron and Ontario shifted towards greater dominance by calanoids and greater similarity with Lake Superior. Trajectories of changes were different; however, reductions in cladocerans accounted for most of the change in lakes Michigan and Huron while reductions in cyclopoids and increases in Leptodiaptomus sicilis were behind the changes in Lake Ontario. Shifts in the predatory cladoceran community in Lake Ontario from Cercopagis pengoi to occasional dominance by Bythotrephes longimanus, a species much more vulnerable to planktivory, as well as the appearance of Daphnia mendotae in a daphnid community previously consisting almost exclusively of the smaller Daphnia retrocurva, suggest impacts of reduced vertebrate predation. In contrast, strong correlations between cladocerans and chlorophyll in lakes Michigan and Huron point to the possible importance of bottom-up forces in those lakes. Large interannual shifts in cladoceran community structure in the central and eastern basins of Lake Erie suggest intense but variable vertebrate predation pressure. The zooplankton communities of lakes Huron, Michigan and Ontario may be approaching a historic community structure represented by Lake Superior.