Distribution and abundance of freshwater polychaetes, Manayunkia speciosa (Polychaeta), in the Great Lakes with a 70-year case history for western Lake Erie

Manayunkia speciosa has been a taxonomic curiosity for 150 years with little interest until 1977 when it was identified as an intermediate host of a fish parasite (Ceratomyxa shasta) responsible for fish mortalities (e.g., chinook salmon). Manayunkia was first reported in the Great Lakes in 1929. Since its discovery, the taxon has been reported in 50% (20 of 40 studies) of benthos studies published between 1960 and 2007. When found, Manayunkia comprised < 1% of benthos in 70% of examined studies. In one extensive study, Manayunkia occurred in only 26% of 378 sampled events (1991–2009). The taxon was found at higher densities in one area of Lake Erie (mean = 3658/m²) and Georgian Bay (1790/m²) than in five other areas (mean = 60 to 553/m²) of the lakes. A 70-year history of Manayunkia in western Lake Erie indicates it was not found in 1930, was most abundant in 1961 (mean = 8039, maximum = 67,748/m²), and decreased in successive periods of 1982 (3529, 49,639/m²), 1993 (1876, 25,332/m²), and 2003 (79, 2583/m²). It occurred at 48% of stations in 1961, 58% in 1982, 52% in 1993, and 6% of stations in 2003. In all years, Manayunkia was distributed primarily near the mouth of the Detroit River. Causes for declines in distribution and abundance are unknown, but may be related to pollution-abatement programs that began in the 1970s, and invasion of dreissenid mussels in the late-1980s which contributed to de-eutrophication of western Lake Erie. At present, importance of the long-term decline of Manayunkia in Lake Erie is unknown.     

First evidence of grass carp recruitment in the Great Lakes Basin

We use aging techniques, ploidy analysis, and otolith microchemistry to assess whether four grass carp Ctenopharyngodon idella captured from the Sandusky River, Ohio were the result of natural reproduction within the Lake Erie Basin. All four fish were of age 1 +. Multiple lines of evidence indicate that these fish were not aquaculture-reared and that they were most likely the result of successful reproduction in the Sandusky River. First, at least two of the fish were diploid; diploid grass carp cannot legally be released in the Great Lakes Basin. Second, strontium:calcium (Sr:Ca) ratios were elevated in all four grass carp from the Sandusky River, with elevated Sr:Ca ratios throughout the otolith transect, compared to grass carp from Missouri and Arkansas ponds. This reflects the high Sr:Ca ratio of the Sandusky River, and indicates that these fish lived in a high-strontium environment throughout their entire lives. Third, Sandusky River fish were higher in Sr:Ca ratio variability than fish from ponds, reflecting the high but spatially and temporally variable strontium concentrations of southwestern Lake Erie tributaries, and not the stable environment of pond aquaculture. Fourth, Sr:Ca ratios in the grass carp from the Sandusky River were lower in their 2011 growth increment (a high water year) than the 2012 growth increment (a low water year), reflecting the observed inverse relationship between discharge and strontium concentration in these rivers. We conclude that these four grass carp captured from the Sandusky River are most likely the result of natural reproduction within the Lake Erie Basin.     

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.     

Trends in Diporeia populations across the Laurentian Great Lakes, 1997-2009

Benthic communities in the Laurentian Great Lakes have been in a state of flux since the arrival of dreissenid mussels, with the most dramatic changes occurring in population densities of the amphipod Diporeia. In response, the US EPA initiated an annual benthic macroinvertebrate monitoring program on all five Great Lakes in 1997. Although historically the dominant benthic invertebrate in all the lakes, no Diporeia have been found in Lake Erie during the first 13 years of our study, confirming that Diporeia is now effectively absent from that lake. Populations have almost entirely disappeared from our shallow (< 90 m) sites in lakes Ontario, Huron, and Michigan. In Lake Ontario, three of our four deep (> 90 m) sites still supported Diporeia populations in 2009, with densities at those sites ranging between 96 and 198/m². In Lake Michigan, populations were still found at six of our seven deep sites in 2009, with densities ranging from 57 to 1409/m². Densities of Diporeia in 2009 at the four deep sites in Lake Huron were somewhat lower than those in Lake Michigan, ranging from 191 to 720/m². Interannual changes in population size in Lake Huron and Lake Michigan have shown a degree of synchrony across most sites, with periods of rapid decline (1997-2000, 2003-2004) alternating with periods of little change or even increase (2001-2002, 2005-2009). There has been no evidence of directional trends at any sites in Lake Superior, although substantial interannual variability was seen.     

Influences on Bythotrephes longimanus life-history characteristics in the Great Lakes

We compared Bythotrephes population demographics and dynamics to predator (planktivorous fish) and prey (small-bodied crustacean zooplankton) densities at a site sampled through the growing season in Lakes Michigan, Huron, and Erie. Although seasonal average densities of Bythotrephes were similar across lakes (222/m² Erie, 247/m² Huron, 162/m² Michigan), temporal trends in abundance differed among lakes. In central Lake Erie where Bythotrephes' prey assemblage was dominated by small individuals (60%), where planktivorous fish densities were high (14,317/ha), and where a shallow water column limited availability of a deepwater refuge, the Bythotrephes population was characterized by a small mean body size, large broods with small neonates, allocation of length increases mainly to the spine rather than to the body, and a late summer population decline. By contrast, in Lake Michigan where Bythotrephes' prey assemblage was dominated by large individuals (72%) and planktivorous fish densities were lower (5052/ha), the Bythotrephes population was characterized by a large mean body size (i.e., 37–55% higher than in Erie), small broods with large neonates, nearly all growth in body length occurring between instars 1 and 2, and population persistence into fall. Life-history characteristics in Lake Huron tended to be intermediate to those found in Lakes Michigan and Erie, reflecting lower overall prey and predator densities (1224/ha) relative to the other lakes. Because plasticity in life history can affect interactions with other species, our findings point to the need to understand life-history variation among Great Lakes populations to improve our ability to model the dynamics of these ecosystems.     

Escherichia coli toxin and attachment genes in sand at Great Lakes recreational beaches

The United States Environmental Protection Agency recommends density thresholds for the fecal indicator organism Escherichia coli in order to ensure the safety of recreational waters. A number of studies published over the past ten years indicate that E. coli is encountered frequently in sand at recreational beaches. While a majority of the sand-associated E. coli may be commensal or environmental strains, the potential for pathogenic strains of E. coli to be present exists. Therefore, the aim of this study was to assess the presence of attachment and virulence genes associated with enteropathogenic and enterohemorrhagic strains of E. coli (EPEC and EHEC) in populations of E. coli recovered from swash zone sand from seven recreational beaches along Lake Huron and Lake St. Clair in eastern Michigan, USA. Genes coding for attachment proteins in EPEC and EHEC were very prevalent in sand E. coli, but genes coding for toxin genes were uncommon. The paucity of genes associated with E. coli toxins suggests that the EPEC and EHEC pathotypes are not common in sand; however, the high prevalence of genes associated with attachment in E. coli pathotypes suggests that these genes are being retained within the beach sand E. coli population.     

Early observations on an emerging Great Lakes invader Hemimysis anomala in Lake Ontario

Hemimysis anomala, a Ponto-Caspian littoral mysid, is an emerging Great Lakes invader that was discovered in Lakes Michigan and Ontario in 2006. Similar to the native mysid Mysis diluviana, Hemimysis exhibits a diel vertical migration pattern but generally inhabits shallower and warmer waters than M. diluviana. Because basic information on the distribution, habitat use, and biology of Hemimysis in the Great Lakes is scarce, the potential for food web disruption by Hemimysis cannot easily be predicted. Preliminary observations indicate widespread invasion of Hemimysis in Lake Ontario. In this study, we confirm the presence of Hemimysis at sites spanning the northern and southern shores of Lake Ontario and the presence of the individuals during winter months. In one horizontal tow in November 2007, over 26,000 individuals were collected with a length range of 4.4 to 9.0 mm and an average caloric density of 611 cal/g wet weight. The most effective methods for sampling Hemimysis were horizontal tows with either a zooplankton net in the water column or a benthic sled near the lake bottom. Although more quantitative data on the life history and distribution of this species is necessary, our preliminary observations support the prediction that the potential for Hemimysis to impact the nearshore food web in Lake Ontario appears high.     

Shifting status and distribution of range margin chorus frog (Pseudacris) populations in eastern Great Lakes watersheds

Contiguous with their range across major rivers into Canada, two different species of chorus frogs are now thought to inhabit the Great Lakes watersheds of New York. Pseudacris triseriata is found along the western Lake Ontario and Lake Erie plains while P. maculata (tentatively a new frog species in NY) inhabits the lowlands of eastern Lake Ontario and the St. Lawrence River. Both species are on their extreme range margins in NY. In 2010 we detected distributional declines of both putative chorus frog species based on a broad survey following standardized occupancy detection protocols. Causes are unclear but could relate to reforestation and urbanization of formerly more extensive agricultural lands, climate change, pathogenic fungal outbreaks and/or the contaminant effects of intensive agriculture. On the other hand, the prior overestimation of ranges because of misidentification may have inflated earlier perceived distributions (positive survey bias) because false positives are problematical for this cryptic frog. At broad geographical scales, chorus frog (meta)populations are highly dynamic and are likely shifting their ranges in response to rapidly changing overall environmental conditions in the northeastern U.S. and Canada.     

Relationship of road density and marsh condition to turtle assemblage characteristics in the Laurentian Great Lakes

Human-induced degradation of coastal wetlands often leads to altered trophic dynamics and species assemblages. Here we use data from 77 coastal marshes in three Laurentian Great Lakes collected between 2001 and 2007 to examine the relationship between human disturbance (road density and wetland quality) and characteristics of aquatic turtle assemblages, including species richness and abundances. Painted turtles (Chrysemys picta) were encountered disproportionately in degraded wetlands and the probability of occurrence decreased with improved site quality. Abundance of painted turtles peaked, however, at intermediate road density in surrounding 1- and 2-km buffers. Across all sites, species richness was highest and common snapping turtles (Chelydra serpentina) were most abundant in wetlands with intermediate water quality. The common musk turtle (Sternotherus odoratus) was absent from degraded wetlands in the lower lakes (Erie and Ontario) that fell within their historical range, but reached high abundances in marshes of Georgian Bay and the North Channel, a region with relatively low human disturbance. Analysis of sex ratios in painted turtles revealed a significant male bias in an area with high road density, while the sex ratio did not differ significantly from 1:1 in a less developed region, consistent with reports of high female mortality in urban areas.     

An analysis of long-term phytoplankton dynamics in Muskegon Lake, a Great Lakes Area of Concern

Long-term monitoring of aquatic ecosystems is essential to distinguish the effects of human-induced stressors from natural patterns of ecologic variation, especially in Great Lakes Area of Concern such as Muskegon Lake. Samples collected between 2003-2009, as part of a continuing long-term monitoring study of the lake, were analyzed to: 1) detect spatial and temporal patterns in the phytoplankton biovolume and species composition; 2) evaluate the environmental variables that regulate phytoplankton community composition changes; and 3) determine the ecological conditions under which toxin-producing Microcystis species occur. Distinct patterns in Muskegon Lake phytoplankton were not evident among sites, which can be explained, at least in part, by the well-mixed waters in this drowned river mouth system. However, surface and bottom samples within sites had significantly different biovolume and species composition, suggesting that horizontal mixing did not extend throughout the water column. Surface samples had greater phytoplankton biovolume than bottom samples, possibly because of greater irradiance. Seasonally, the least biovolume was recorded in spring samples, which were dominated by diatoms. Phytoplankton biovolume was greatest in the summer when cyanobacteria were abundant. Environmental variables that correlated highly with the ordination space defined by species composition included sulfate, specific conductance, total dissolved solids, and chloride. Results from regression tree analysis predicted increasing biovolume of Microcystis aeruginosa with increasing concentrations of the total Kjeldahl nitrogen. The seven-year period of this study did not reveal major changes in the lake's environment and phytoplankton communities, but the presence of invasive and toxin-producing species warrants their continued monitoring.     

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.     

A New Record of Pinaciophora Fluviatilis Greeff Sensu Penard from the Laurentian Great Lakes

This report clarifies the presence of Pinaciophora fluviatilis Greeff sensu Penard in the Laurentian Great Lakes. Transmission electron microscopic observations were made of a shadowcast carbon replica of a single scale. Taxonomic problems of this organism are discussed.     

Spatial and temporal variation in the distribution of burrowing mayfly nymphs (Ephemeroptera: Hexagenia limbata and H. rigida) in western Lake Erie

In the early 1990s, burrowing mayfly species reappeared in sediments of the western basin of Lake Erie after an absence of over 30 years due to episodic hypoxia at the sediment–water interface. Long-term monitoring of adult mayflies at shoreline areas had revealed that Hexagenia rigida was more abundant than Hexagenia limbata during the initial recolonization period, but was gradually replaced by H. limbata. We hypothesized that this shift in dominance would be confirmed by the distribution and abundance of nymphs. We identified nymphs collected each spring throughout western Lake Erie from 1997 to 2004. The relative abundances of H. rigida and H. limbata nymphs exhibited the same temporal sequence as adults. Furthermore, the number of sites in the western basin in which H. rigida occurred decreased as the occurrence frequency of H. limbata increased. H. limbata were dominant in the basin by 2004. Hexagenia limbata nymphs persisted in the center-most part of the basin, whereas H. rigida did not, possibly due to differences in tolerance to hypoxia. There were no significant differences in body size between the two populations. Differences in dispersal distance from source populations and the timing and success of egg hatching likely accounted for the initial colonizing success of H. rigida, but the differential ability of H. limbata eggs to overwinter in sediments and possible tolerance of nymphs to hypoxia has possibly led to its current dominance in the western basin.     

Burrowing mayfly (Ephemeroptera: Ephemeridae: Hexagenia spp.) bioturbation and bioirrigation: A source of internal phosphorus loading in Lake Erie

Traditional lake eutrophication models predict lower phosphorus concentrations with decreased external loads. However, in lakes where decreased external phosphorus loads are accompanied by increasing phosphorus concentrations, a seeming “trophic paradox” exists. Western Lake Erie is an example of such a paradox. Internal phosphorus loads may help explain this paradox. We examined bioturbation and bioirrigation created from burrowing mayfly, Hexagenia spp., as a possible source of internal phosphorus loading. Phosphorus concentrations of experimental microcosms containing lake sediments, filtered lake water, and nymphs (417/m²) collected from western Lake Erie were compared to control microcosms containing sediments and lake water over a 7-day period. Phosphorus concentrations in microcosms containing Hexagenia were significantly greater than microcosms without nymphs. Further, we estimate the soluble reactive phosphorus flux from the sediments due to Hexagenia is 1.03 mg/m²/day. Thus, Hexagenia are a source of internal phosphorus loading. High densities of Hexagenia nymphs in western Lake Erie may help explain the “trophic paradox.” Furthermore, Hexagenia may be a neglected source of internal phosphorus loading in any lake in which they are abundant. Future studies of phosphorus dynamics in lakes with Hexagenia must account for the ability of these organisms to increase lake internal phosphorus loading.     

Nutritive Substitution of Zinc by Cadmium and Cobalt in Phytoplankton Isolated from the Lower Great Lakes

Low ambient concentrations of trace metals including Zn may limit phytoplankton productivity in the North American Great Lakes. The microalgae Chlorella sp. UTCC 522 (Chlorophyta) and Cyclotella sp. UTCC 520 (Heterokontophyta), indigenous to the Great Lakes, were assayed for their Zn requirement and the ability for Co and Cd to metabolically substitute for Zn under conditions in which Zn bioavailability was limiting cell growth. Bioassays were conducted in the laboratory using chemically defined media and the metal buffer EDTA to control the free-ion concentrations of Zn²⁺ (10^−15.3−10⁻¹⁰ mol/L), Cd²⁺ (10^−15.1−10⁻¹⁰ mol/L), and Co²⁺ (10⁻¹⁴−10⁻¹⁰ mol/L). Influence of the metals on the phytoplankton was measured by the effect on specific growth rate. Both microalgae required Zn and could use Co and Cd as Zn substitutes; Cd was used less-effectively, although to a greater extent by the diatom Cyclotella than by the chlorophyte, Chlorella. The observed ability of the phytoplankton to use Cd and Co as Zn substitutes suggests that microalgae play an important role in the geochemical cycling of Zn, Cd and Co in large lakes, and the mobilization of Cd in lake ecosystems impacted by pollution.     

Satellite ocean color algorithms: A review of applications to the Great Lakes

We review the literature dealing with retrievals of chlorophyll concentrations in the Great Lakes from satellite observations. Most studies show that the satellite estimates of chlorophyll concentrations are linearly related to the observed concentrations, though they tend to overestimate concentrations at lower values and underestimate them at higher values. Deviations from a consistent, accurate, linear relationship can be attributed to temporal and spatial variations in the inherent optical properties of the color producing agents in the water as well as to varying concentrations of non-algal substances that interfere with the retrievals. We confirmed these results by using a simple optical model to examine the sensitivity of the retrieved chlorophyll values to the concentrations of interfering substances and to differences in model parameters. Because the spatial and temporal optical properties of the Great Lakes are unpredictable, no retrieval method is likely to produce accurate results all the time. The papers we reviewed show that simple band ratio algorithms can provide chlorophyll estimates that are proportional to in situ concentrations. The published literature suggests that the band ratio methods will be of most value in regions where the concentrations of non-algal interfering substances are minimal. Because of these limitations we recommend that future papers presenting chlorophyll analysis based on satellite data provide confirming field observations that include measurements of chlorophyll, suspended particles and dissolved organic carbon. We also recommend that Great Lakes scientists explore novel methods for retrieving chlorophyll concentrations from satellite observations that have proven useful in other optically complex waters.     

Heavy metals in sediments and uptake by burrowing mayflies in western Lake Erie basin

During the past two decades, burrowing Hexagenia mayflies have returned to the western basin of Lake Erie. Because of their importance as a prey resource for higher trophic levels and their extensive residence time in potentially contaminated sediment, Hexagenia may be a source of heavy metal transfer. To better understand the distribution and transfer of heavy metals in sediment and mayflies, sediment and mayfly nymphs were collected from 24 locations across the western basin of Lake Erie in May 2007. Following USEPA protocols, samples were analyzed for 16 elements using ICP-OES or ICP-MS. Metal concentrations in the sediments exceeded the Threshold Effect Level for at least one metal at all sample sites. Sediment heavy metal distribution profiles indicate metal concentrations are correlated with organic matter content, and the highest heavy metal concentrations were found in the central deeper region of the western basin where organic content in the sediments was greatest. Hexagenia were distributed throughout the western basin, with greatest density (1350/m²) within the Detroit River plume. The Cd and Zn levels in mayflies were on average approximately 4 and 2 times greater, respectively, than sediment levels, and the Cd concentrations in the sediments exceeded the Threshold Effect Level at 27 of 28 sites and exceeded the Probable Effect Level at 9 of 28 sites. Spatial representation of heavy metal concentrations in mayflies exhibited a similar pattern to the spatial distribution of heavy metals and organic matter in the sediments with higher concentrations of metals found in mayflies residing in the central deeper region of the western basin.     

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.     

Metazoan Parasites of Introduced Round and Tubenose Gobies in the Great Lakes: Support for the “Enemy Release Hypothesis”

Recent invasion theory has hypothesized that newly established exotic species may initially be free of their native parasites, augmenting their population success. Others have hypothesized that invaders may introduce exotic parasites to native species and/or may become hosts to native parasites in their new habitats. Our study analyzed the parasites of two exotic Eurasian gobies that were detected in the Great Lakes in 1990: the round goby Apollonia melanostoma and the tubenose goby Proterorhinus semilunaris. We compared our results from the central region of their introduced ranges in Lakes Huron, St. Clair, and Erie with other studies in the Great Lakes over the past decade, as well as Eurasian native and nonindigenous habitats. Results showed that goby-specific metazoan parasites were absent in the Great Lakes, and all but one species were represented only as larvae, suggesting that adult parasites presently are poorly-adapted to the new gobies as hosts. Seven parasitic species are known to infest the tubenose goby in the Great Lakes, including our new finding of the acanthocephalan Southwellina hispida, and all are rare. We provide the first findings of four parasite species in the round goby and clarified two others, totaling 22 in the Great Lakes—with most being rare. In contrast, 72 round goby parasites occur in the Black Sea region. Trematodes are the most common parasitic group of the round goby in the Great Lakes, as in their native Black Sea range and Baltic Sea introduction. Holarctic trematode Diplostomum spathaceum larvae, which are one of two widely distributed species shared with Eurasia, were found in round goby eyes from all Great Lakes localities except Lake Huron proper. Our study and others reveal no overall increases in parasitism of the invasive gobies over the past decade after their establishment in the Great Lakes. In conclusion, the parasite “load” on the invasive gobies appears relatively low in comparison with their native habitats, lending support to the “enemy release hypothesis.”     

Diatoms abound in ice-covered Lake Erie: An investigation of offshore winter limnology in Lake Erie over the period 2007 to 2010

The limnology of offshore Lake Erie during periods of extensive (> 70%) ice cover was examined from ship borne sampling efforts in 2007 to 2010, inclusive. Dense and discrete accumulations of the centric filamentous diatom Aulacoseria islandica (> 10 μg Chl-a/L) were located in the isothermal (< 1 °C) water column directly below the ice and only detectable in the ship wake; viable phytoplankton were also observed within ice. Evidence from these surveys supports the notions that winter blooms of diatoms occur annually prior to the onset of ice cover and that the phytoplankton from these blooms are maintained in the surface waters of Lake Erie and reduce silicate concentrations in the lake prior to spring. The mechanisms by which high phytoplankton biomass rise at this time of year requires further investigation, but these winter blooms probably have consequences for summer hypoxia and how the lake responds to climate change.