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.     

Sources and Sinks of Escherichia coli in Benthic and Pelagic Fish

Escherichia coli and fecal coliform bacteria were isolated from five benthic and four pelagic fish species to determine their role in the fecal contamination of recreational waters. All fish were collected during fall 2006 from Southworth Marsh in the Duluth-Superior Harbor, a public beach that is commonly posted to minimize water contact due to high E. coli levels. Although fecal coliform bacteria were isolated from each fish species, they were only isolated from 66% and 72% of the individual benthic and pelagic fish, respectively. While 42% of the fecal coliforms from benthic fish were E. coli, only 4% of these bacteria from pelagic fish were E. coli. Cluster analysis showed different fish species harbored identical strains of E. coli and some fish contained multiple E. coli strains. The potential source for 65% of the E. coli isolates obtained from fish were identified by using the HFERP DNA fingerprinting method and libraries of E. coli DNA fingerprints from warm-blooded animals and environmental isolates collected in the area. The E. coli strains whose source could be identified were most similar to strains isolated from sediments, Canada geese, mallard ducks, and wastewater. None of the fish E. coli had DNA fingerprints matching those from any water or beach sand isolates. Although our results demonstrate that benthic fish contain E. coli, it may be more appropriate to consider these fish as a vector of E. coli from other sources, rather than a new source of E. coli contamination in aquatic environments.     

Impact of the Alga Cladophora on the Survival of E. coli, Salmonella, and Shigella in Laboratory Microcosm

E. coli is an indicator of recent fecal contamination of freshwater beaches around the Great Lakes region. Elevated concentrations indicate that a fecal contamination has occurred, and that the risk for contact with fecal pathogenic organisms is heightened. The green algae, Cladophora, harbors populations of E. coli and potentially allows for prolonged survival and even replication of the bacterium in the lake environment. If presence of Cladophora mats on beaches is associated with persistence of E. coli in beach water, then E. coli would be a useful indicator organism only if pathogens also were able to survive and persist in the algae. This study utilized lab microcosms to study the persistence of E. coli, and of the fecal pathogens, Salmonella and Shigella, in lake water with and without the presence of Cladophora. E. coli was able to persist for extended periods in the presence of Cladophora (attached to algal mats for 45 days). Salmonella and Shigella, however, were unable to persist for this time period while in the presence of Cladophora (Salmonella attached to Cladophora was detectable for 10 days and Shigella was detectable for only 2 days). These data imply that E. coli is able to survive in the presence of Cladophora for greater times than are the fecal pathogens and that E. coli may not be an appropriate indicator organism for beaches with accumulations of algal material.     

Distribution and Fate of Escherichia coli in Lake Michigan Following Contamination with Urban Stormwater and Combined Sewer Overflows

Escherichia coli distribution and persistence in nearshore Lake Michigan were assessed following heavy rains and sanitary sewer overflow (SSO) and combined sewer overflow (CSO) events over a 5-year period, including an 18-day period following 25.4 cm of rainfall in which intensive studies were conducted following multiple CSO and SSO events. E. coli levels in the Milwaukee estuary and harbor following SSO and CSO events ranged from 10⁴ to nearly 10⁵ CFU/100 mL, which were significantly higher (p ≤ 0.05) than levels following rainfall alone. Sites outside of the breakwall but within the contamination plume (e.g., within 2 km of the harbor) were an order of magnitude lower. Locations 2–5 km from the harbor ranged from below detection limits, of < 1 to 5 CFU/100 mL. E. coli levels corrected for dilution based on specific conductivity measurements were lower than what would be expected for loss due to dilution alone, suggesting a combination of die-off and dilution, were responsible for the rapid disappearance of these organisms outside of the harbor. E. coli and fecal coliforms measured concurrently demonstrated that fecal coliforms could be recovered longer than E. coli in the open waters of the lake. E. coli isolated directly from sewage treatment plant influent were found to have a marked increase in antibiotic resistance traits for ten antibiotics commonly used in the human population compared with isolates from two animal sources of fecal pollution. However, E. coli obtained from sewage impacted water (n = 2,513) and from stormwater impacted water (n = 1,465) collected the previous year when there were no sewage overflows, were found to have no significant difference (p < 0.05) in the frequency of resistance when comparing the two conditions. E. coli survival characteristics and population dynamics are most likely influenced by multiple factors in complex systems such as the watershed/estuarine/lake environments of the Great Lakes.     

A model laboratory system to study the synergistic interaction and growth of environmental Escherichia coli with macrophytic green algae

Escherichia coli, an important fecal indicator bacterium, is known to persist and reproduce in association with Cladophora and other macrophytic algae and environmental substrates. Recent increases in the growth of Cladophora and other macrophytic algae in many of the Laurentian Great Lakes result in the accumulation of large amounts of algal biomass along the shoreline and on beaches. While the Cladophora–E. coli association may pose substantial public health risks, detailed laboratory-based studies have not been done to investigate the bases of the interaction. This is due, in large part, to past inabilities to culture many macrophytic algae under near axenic conditions. Here we describe the development and experimental use of laboratory microcosms to study the synergistic interaction and growth of the green alga Pithophora, a close relative of Cladophora, with environmental E. coli. In the absence of exogenous organic carbon supply, the E. coli population attached to algal filaments increased approximately three orders of magnitude within 72 h of inoculation. Growth of E. coli on Pithophora appeared to be limited by dissolved nitrogen with a concentration of ≥ 66 μg/mL N allowing maximal bacterial growth. In contrast, an environmental strain of Salmonella did not grow under identical conditions in the microcosms, suggesting that this bacterium requires additional growth factors not provided by Cladophora. Since the alga can be maintained in the laboratory for long periods of time, this system allows for further experimentation and understanding of macroalga–microbe interactions.     

Fecal indicator organism density in beach sands: Impact of sediment grain size,uniformity, and hydrologic factors on surface water loading

Fecal loading to beach sands and subsequent transport to surface water may result in the degradation of surface water quality. To better understand the relationship between Escherichia coli in sands, beach hydrology, and recreational water quality, we collected surface water, groundwater, and sand samples from three Lake Michigan beaches with varying substrates (fine sand to pebbles, July 2005–June 2006). Sediment cores collected within transects perpendicular to and at fixed distances from the shoreline (0 m, 10 m, 20 m) and surface water samples collected at a depth of 1 m were analyzed for E. coli. Grain size analysis was performed on duplicate core samples to assess the relationship between E. coli density and mean grain size and uniformity. Groundwater samples, collected from shallow drive-point piezometers within the test area, were also analyzed for E. coli. E. coli density in beach sands differed significantly with distance from shore with the highest density occurring at the berm crest (0 m). Mean grain size and uniformity accounted for variation in E. coli density with fine sand of uniform distribution having the highest content. E. coli density in surface water was correlated to E. coli density in beach sand samples at the berm crest. E. coli in groundwater was < 10 to 579 MPN/100 ml (2005); none was detected in 2006. Management interventions, including altered beach grooming practices and slope assessments, may be effective in reducing E. coli content at beaches comprised of fine sands of uniform grain size, hence reducing water quality advisories.     

The establishment of the nuisance cyanobacteria Lyngbya wollei in Lake St. Clair and its potential to harbor fecal indicator bacteria

Lyngbya wollei is a filamentous cyanobacterium which forms large nuisance mats and has infested eastern and southeastern U.S. Lakes and reservoirs for over 100 years. Lyngbya was recently identified in the Great Lakes system in the St. Lawrence River, and Western Lake Erie. Here we report on large deposits of L. wollei washing onshore at a popular recreational beach in Lake Saint Clair, part of the Great Lakes system. The amount of L. wollei deposited on shore was quantified and evaluated for the presence of fecal indicator bacteria (FIB). High concentrations of Escherichia coli, enterococci and Clostridium perfringens were found in the L. wollei in nearshore waters. The densities of E. coli (MPN), enterococci (MPN) and C. perfringens (CFU) attached to L. wollei averaged 3.5, 3.2 and 3.2 log/g, respectively. In contrast, nearshore waters contained nearly 10 times less FIB, averaging 2.6, 2.4 and 2.6 log/100 ml of E. coli (MPN), enterococci (MPN) and C. perfringens (CFU), respectively. DNA fingerprint analysis was used to examine the population structure of E. coli isolates obtained from L. wollei mats. The L. wollei-borne E. coli strains were genetically diverse, suggesting a causal relationship between E. coli and L. wollei. Results from this study indicate that in addition to the macroalga such as Cladophora, cyanobacteria like L. wollei also harbor FIB, potentially impacting water quality and human health in the Great Lakes.     

The Effects of Rainfall on Escherichia coli and Total Coliform Levels at 15 Lake Superior Recreational Beaches

Fifteen beaches along Lake Superior were monitored over the course of the 2003 and 2004 summer swimming seasons from mid-May through mid-September. Water samples were collected at these 15 beaches less than 24-h after a rainfall event of at least 6 mm. The effect of rainfall on bacterial concentrations along the Wisconsin shores of Lake Superior was investigated. Water samples were collected at a water depth of 24–30 inches as specified by the requirements of the WI BEACH Program. The Colisure® test was utilized for Escherichia coli (E. coli) and coliform determinations. No relationship between rainfall amount and bacterial concentrations at any of the 15 beaches tested was found. Although other researchers have observed a direct positive relationship between rainfall and E. coli levels in beach water, we found no significant relationship for Lake Superior beaches. This is an important finding given the fact that beach closures are often based upon rainfall alone, rather than on actual E. coli concentration measurements. This study reinforces the fact that the data obtained at one location should not necessarily be extrapolated to beach closure decisions at other locations.     

Evidence for Remote Effects of Dreissenid Mussels on the Amphipod Diporeia: Analysis of Lake Ontario Benthic Surveys, 1972–2003

The status of invasive dreissenid mussels (Dreissena polymorpha and D. bugensis) and native amphipods (Diporeia spp.) in Lake Ontario was assessed in 2003 and compared with historical data. D. polymorpha (zebra mussels) were rarely observed in 2003, having been displaced by D. bugensis (quagga mussels). D. bugensis expanded its depth range from 38 m depth in 1995 to 174 m in 2003 and this dreissenid reached densities averaging 8,000/m² at all sites < 90 m. During the same time period, Diporeia populations almost completely disappeared from 0–90 m depth, continuing a declining trend from 1994–1997 reported in previous studies. The average density of Diporeia in the 30–90 m depth interval decreased from 1,380/m² to 63/m² between 1997 and 2003. Prior to 2003, areas deeper than 90 m represented a refuge for Diporeia, but even these deep populations decreased, with densities declining from 2,181/m2 in 1999 to 545/m2 in 2003. Two common hypotheses for the decline of Diporeia in the Great Lakes are food limitation and a toxin/pathogen associated with dreissenid pseudofeces. The Diporeia decline in deep waters preceded the expansion of D. bugensis to these depths, and suggests that shallow dreissenid populations remotely influence profundal habitats. This pattern of decline is consistent with mechanisms that act from some distance including nearshore dreissenid grazing and downslope transport of pseudofeces.     

Is reduced benthic flux related to the Diporeia decline? Analysis of spring blooms and whiting events in Lake Ontario

Benthic monitoring by USGS off the southern shore of Lake Ontario from October 1993 to October 1995 provides a detailed view of the early stages of the decline of the native amphipod Diporeia. A loss of the 1994 and 1995 year classes of Diporeia preceded the disappearance of the native amphipod at sites near Oswego and Rochester at depths from 55 to 130 m. In succeeding years, Diporeia populations continued to decline in Lake Ontario and were nearly extirpated by 2008. Explanations for Diporeia's decline in the Great Lakes include several hypotheses often linked to the introduction and expansion of exotic zebra and quagga mussels (Dreissena sp.). We compare the timeline of the Diporeia decline in Lake Ontario with trends in two sources of organic matter to the sediments — spring diatom blooms and late summer whiting events. The 1994–95 decline of Diporeia coincided with localized dreissenid effects on phytoplankton in the nearshore and a year (April 1994 to May 1995) of decreased flux of organic carbon recorded by sediment traps moored offshore of Oswego. Later declines of profundal (> 90 m) Diporeia populations in 2003 were poorly associated with trends in spring algal blooms and late summer whiting events.     

Microbial Concentrations in Sand and their Effect on Beach Water in Door County,Wisconsin

The seasonal variations and patterns of Escherichia coli in Wisconsin's coastal waters have been closely studied in recent years due to increased beach monitoring activities. Patterns of distribution of the indicator organism, E. coli, in the sand at these beaches are now being investigated as a source of E. coli to adjacent beach water. This project investigates the concentrations of E. coli in beach sand, and the relationship between these sand-microbe concentrations and concentrations of microbes in the corresponding beach water. Weekly sampling of upshore, swash, and submerged sand at six beaches provided numbers of the indicator bacteria in each beach's sand substrate for two consecutive summers. Overall concentrations of E. coli were highest in the swash sand of the beach, with the highest numbers seen in the summer months and lowest numbers in the winter months. Each location had very different concentrations of E. coli in the beach sand from 1,800 CFU/100 g to 21,670 CFU/100 g sand. Each location had a very different relationship between the indicator organism found in the beach sand and that found in the beach water. These data suggest that sand may be a reservoir for E. coli at some locations, and another source of contamination that should be considered in beach monitoring programs. However, elevated levels of E. coli in beach sand were not universal and varied greatly from location to location.     

Lyngbya wollei in western Lake Erie

We report on the emergence of the potentially toxic filamentous cyanobacterium, Lyngbya wollei as a nuisance species in western Lake Erie. The first indication of heavy L. wollei growth along the lake bottom occurred in September 2006, when a storm deposited large mats of L. wollei in coves along the south shore of Maumee Bay. These mats remained intact over winter and new growth was observed along the margins in April 2007. Mats ranged in thickness from 0.2 to 1.2 m and we estimated that one 100-m stretch of shoreline along the southern shore of Maumee Bay was covered with approximately 200 metric tons of L. wollei. Nearshore surveys conducted in July 2008 revealed greatest benthic L. wollei biomass (591 g/m² ± 361 g/m² fresh weight) in Maumee Bay at depth contours between 1.5 and 3.5 m corresponding to benthic irradiance of approximately 4.0–0.05% of surface irradiance and sand/crushed dreissenid mussel shell-type substrate. A shoreline survey indicated a generally decreasing prevalence of shoreline L. wollei mats with distance from Maumee Bay. Surveys of nearshore benthic areas outside of Maumee Bay revealed substantial L. wollei beds north along the Michigan shoreline, but very little L wollei growth to the east along the Ohio shoreline.     

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.     

Stock assessment and fishery management of Henicorhynchus spp., Cyclocheilichthys enoplos and Channa micropeltes in Tonle Sap Great Lake,Cambodia

Tonle Sap Great Lake, in the lower Mekong River basin, contributes 60% of Cambodia's inland fisheries catch. There are four types of fisheries, including the middle-scale fishery. The major species comprising the middle-scale fishery are Henicorhynchus spp. (16%), a small-bodied fish caught mainly with small-mesh nets, and Cyclocheilichthys enoplos (13%), and Channa micropeltes (7%) which are caught with a variety of gear. Here we apply the Schaefer surplus production model to the middle-scale fisheries using monitoring data collected between 1995 and 1999. Model simulations determined that the maximum sustainable yield (F_MSY) for Henicorhynchus spp., was obtained with the use of 47,206 gillnets (mesh size < 50 mm) and 4269 and 1605 fishing boats for C. enoplos and C. microplestes, respectively. Over 1995–1999, there was a strong trend of increase in fishing effort and decrease of catch-per-unit-effort in the waters of Pursat, Siem Riep and Kampong Chnnang provinces. These provinces have large population centres located close to the fishing grounds; fishing in these areas should be more strictly regulated.     

Coexistence of the native benthic amphipod Diporeia spp. and exotic dreissenid mussels in the New York Finger Lakes

Populations of the benthic amphipod Diporeia spp. have sharply declined since the early 1990s in all North America's Great Lakes except Lake Superior. The onset and continued decline coincides with the invasion of these lakes by zebra (Dreissena polymorpha) and quagga (Dreissena rostriformis bugensis) mussels and the spread of quagga mussels to deep habitats. The six deepest Finger Lakes of central New York (Seneca, Cayuga, Skaneateles, Canandaigua, Keuka, and Owasco) have historically been Diporeia habitat and have had dreissenids for more than a decade. These lakes represent a wide range of trophic state, maximum depth, and dreissenid invasion history. We hypothesized that Diporeia abundance would be negatively impacted by dreissenid mussel expansion in the Finger Lakes. During 2006–2010, we sampled Diporeia and mussel populations in these six lakes. Diporeia was present in all six lakes, and was abundant (2000/m²) in Owasco Lake that has only zebra mussels and in Cayuga and Seneca Lakes that have had zebra and quagga mussels since 1994. Diporeia abundance was lowest (1000/m²) in Skaneateles, Canandaigua, and Keuka Lakes where quagga mussels have recently expanded. Productivity indicators explained much of the variability of Diporeia abundance. The persistence of Diporeia with quagga mussels in these lakes may be because of available alternative food resources. Fatty acid tracers indicate that Diporeia from Owasco Lake, the lake without quagga mussels, utilize diatoms, but Diporeia from Cayuga Lake that coexist with abundant quagga mussels also use food resources associated with terrestrial detritus that cannot be intercepted by dreissenids.     

Occurrence of the Great Lake's most recent invader, Hemimysis anomala, in the diet of fishes in southeastern Lake Ontario

The Ponto-Caspian mysid, Hemimysis anomala, was first observed in southeastern Lake Ontario in May 2006. During July and August 2007, gill nets were fished in 6 to 8 m of water at two locations of known Hemimysis colonization in southeastern Lake Ontario to determine if fish that consume macroinvertebrates were beginning to include this new invasive mysid in their diets. Of nine fish species captured in August, September, and October 2007, three species had consumed Hemimysis: alewife (Alosa pseudoharengus), rock bass (Ambloplites rupestris), and yellow perch (Perca flavescens); and six species had not: round goby Apollonia melanostoma, smallmouth bass Micropterus dolomieu, spottail shiner Notropis hudsonius, gizzard shad Dorosoma cepedianum, white perch Morone americana and log perch Percina caprodes. Diets of alewives from all samples were composed predominantly of Hemimysis (69.6% -100% frequency of occurrence, 46.0%–74.5% dry weight diet composition). Two of 6 rock bass stomachs sampled in August contained ≥ 98.9% Hemimysis (10 and 40 individuals each) and one of 61 yellow perch stomachs sampled in September contained 10.0% Hemimysis (6 individuals) and 90.0% fish. While Hemimysis were observed only sparsely in the diet of most nearshore fish, their predominance in alewife diets and their omnivorous feeding behavior indicated that they have the potential to alter energy flow in Great Lakes' foodwebs.     

Occurrence of a Cylindrospermopsis bloom in Louisiana

Cylindrospermopsis raciborskii occurs predominantly in tropical to subtropical freshwaters but appears to be spreading to temperate regions. After the hurricanes of 2005, a bloom of this organism took place in Lake Dauterive and Lake Fausse Pointe in Louisiana. The cyanobacterium dominated the phytoplankton community for three months. Of the three known morphotypes (straight, coiled and spiral) only coiled and spiral were found. In June 2006, 60% of the organisms were of the spiral morphotype but the relative abundance of this morphotype decreased dramatically within the month of June. However, the average density for both morphotypes remained the same until September. Densities (up to 160,000 cells/mL) significantly exceeded the threshold for toxicity. Because the sampled region offers optimal light and nutrient conditions, it is likely that C. raciborskii has been established in this region and future blooms can be expected. Therefore, a monitoring program should be implemented.     

Evaluating Sampling Strategies for Larval Cisco (Coregonus artedi)

To improve our ability to assess larval cisco (Coregonus artedi) populations in Lake Superior, we conducted a study to compare several sampling strategies. First, we compared density estimates of larval cisco concurrently captured in surface waters with a 2 × 1-m paired neuston net and a 0.5-m (diameter) conical net. Density estimates obtained from the two gear types were not significantly different, suggesting that the conical net is a reasonable alternative to the more cumbersome and costly neuston net. Next, we assessed the effect of tow pattern (sinusoidal versus straight tows) to examine if propeller wash affected larval density. We found no effect of propeller wash on the catchability of larval cisco. Given the availability of global positioning systems, we recommend sampling larval cisco using straight tows to simplify protocols and facilitate straightforward measurements of volume filtered. Finally, we investigated potential trends in larval cisco density estimates by sampling four time periods during the light period of a day at individual sites. Our results indicate no significant trends in larval density estimates during the day. We conclude estimates of larval cisco density across space are not confounded by time at a daily timescale. Well-designed, cost effective surveys of larval cisco abundance will help to further our understanding of this important Great Lakes forage species.     

Recent increases in the large glacial-relict calanoid Limnocalanus macrurus in Lake Michigan

Since 2004, population density of the large hypolimnetic calanoid Limnocalanus macrurus Sars. has increased dramatically in Lake Michigan. The average summer biomass of this species between 2004 and 2006 was roughly three times that of the period 1984–2003, and at levels unprecedented in our 22-year dataset, making L. macrurus the dominant zooplankter in the lake in terms of biomass. These increases have been accentuated by coincident population declines of the main daphnid, Daphnia mendotae, in the lake with the result that in 2006, L. macrurus accounted for 75% and 50% of the large (> 0.9 mm) crustacean biomass in the northern and southern basins of Lake Michigan, respectively. The increases in L. macrurus populations have closely coincided with equally dramatic increases in summer water clarity. Recent extinction coefficients are among the lowest recorded for the lake, and deepening light penetration has permitted increases in the size of the deep chlorophyll layer. In addition, planktivorous fish populations have declined coincidently with the increases in L. macrurus. It seems likely that an increase in sub-epilimnetic production has resulted in increased food resources for the deep-living L. macrurus, while low planktivore abundances have reduced predation loss, permitting L. macrurus to respond to these increases in sub-epilimnetic production.     

Distribution of nuisance Cladophora in the lower Great Lakes: Patterns with land use, near shore water quality and dreissenid abundance

Selected shorelines and offshore shoals in Lakes Erie, Huron and Ontario were surveyed with a high frequency hydroacoustic system to investigate current spatial patterns of nuisance benthic filamentous algal (e.g., Cladophora) cover and stand height. Cladophora reached nuisance levels at all sites in Lakes Erie and Ontario, but not in Lake Huron or Georgian Bay. Despite clear gradients in coastal land cover, near shore water quality gradients were generally weak, and for Lakes Erie and Ontario, measures of near shore water quality were similar to that at offshore shoals. Hierarchical partitioning analysis suggested that while dreissenid mussel abundance appeared to be important in determining the magnitude of Cladophora standing crop, the joint contribution of catchment land cover, near shore water quality (nutrient levels and suspended matter) and dreissenid mussel abundance explained nearly 95% of the total variance in nuisance Cladophora standing crop observed in this study. Although the results from this study are necessarily correlative in nature and definition of causal relationships is not possible, these results provide corroborating evidence from sites across a gradient within and across the lower Great Lakes that is consistent with the operation of the near shore shunt model.