Great Lakes Cladophora in the 21st century: same algae—different ecosystem

Nuisance growth of the attached, green alga Cladophora was considered to have been abated by phosphorus management programs mandated under the Great Lakes Water Quality Agreement. The apparent resurgence of nuisance growth in Lakes Erie, Michigan and Ontario has been linked conceptually to ecosystem alterations engineered by invasive dreissenid mussels (Dreissena polymorpha and Dreissenabugensis). Here, we apply contemporary modeling tools and historical water quality data sets in quantifying the impact of long-term changes in phosphorus loading and dreissenid-mediated changes in water clarity on the distribution and production of Cladophora. It is concluded that reductions in phosphorus loading in the pre-dreissenid period achieved the desired effect, as model simulations were consistent with the biomass declines reported from the early 1970s to the early 1980s. These declines were, however, largely offset by dreissenid-driven changes in water clarity that extended the depth of colonization by Cladophora, increasing total production. We were not able to isolate and quantify the significance of dreissenid mediation of phosphorus cycling using the historical database. Phosphorus management remains the appropriate mechanism for reducing nuisance levels of Cladophora growth. The development of action plans will require an improved understanding of nearshore phosphorus dynamics such as might be obtained through regular monitoring of soluble reactive phosphorus levels, internal phosphorus content and Cladophora biomass in impacted nearshore regions of the Great Lakes.     

Phytoplankton and benthic algal response to ecosystem engineers and multiple stressors in the nearshore of Lake Huron

Resurgence of nuisance benthic algae in the Great Lakes, despite substantial efforts to reduce phosphorus loading, has stimulated renewed interest in exploring the diverse drivers of near-shore water quality. Interestingly, broad similarity in the underlying causes of shore fouling by benthic algae in Lakes Ontario, Erie, and Michigan appear to contrast with Lake Huron where system productivity and dreissenid abundance are lower. While total phosphorus was the primary predictor of chlorophyll concentrations (70–90% of variation) in the water column, we identified a series of spatial patterns that underpin this relationship (up to 28% of variation) and which integrate catchment processes, tributary influences, shoreline complexity, and distance from shore. Dreissenid mussels were the most important predictors of benthic algae cover and biomass in our models, explaining between 20 and 52% of variation. Spatial patterns explained an additional 21–48% of the variation in benthic algae cover and biomass and highlight the importance of site-specific spatial heterogeneity in benthic algae growth. Our results are consistent with the nearshore shunt hypothesis, wherein higher algal cover and biomass coincided with higher mussel density and biomass, although correlative effects with lake depth and loss of algal and mussel biomass due to physical disturbance must also be considered. These results underscore the difficulty associated with identifying the potential drivers of nearshore water quality as the diverse processes of nutrient loading, changes in catchment land use, and ecosystem change associated with invasion by dreissenids all vary in relative influence over a range of spatial scales.     

The Great Lakes Cladophora Model: Development,testing, and application to Lake Michigan

A recent review of the Great Lakes Water Quality Agreement has concluded that while controls on phosphorus inputs to Lake Michigan achieved the desired effect in offshore waters, the nearshore region continues to suffer from elevated phosphorus levels. Failure to achieve trophic state goals in the nearshore is manifested in nuisance growth of Cladophora and attendant impacts on property owners, utilities, and the public health and welfare. This study focuses on a site in Lake Michigan near Milwaukee, Wisconsin, where nuisance growth of Cladophora and associated beach fouling occur regularly. A mechanistic model simulating Cladophora growth, suitable for guiding nutrient management in the Great Lakes nearshore, is presented. The model represents an update of the Canale and Auer framework, reflecting current understandings of Cladophora ecology and offering a user-friendly interface making the software more widely available to decision makers. This Great Lakes Cladophora Model (GLCM) is first validated for the Auer/Canale data set collected in 1979 at a site on Lake Huron and then for a data set developed in 2006 for a site on Lake Michigan. Model performance under the strikingly different forcing conditions (depth, light, phosphorus levels) characteristic of these two sites affirms the widespread applicability of the tool. The GLCM is then extended to examine the impacts of ecosystem perturbation (dreissenid colonization) on Cladophora growth and to future approaches to monitoring and management.     

Factors affecting Cladophora growth in the eastern basin of Lake Erie: Analysis of a monitoring dataset (2012–2019)

Cladophora is a naturally occurring benthic alga in the Great Lakes which can reach nuisance levels in the nearshore, leading to beach closures and other impacts. A monitoring program was initiated in 2012 in the eastern basin of Lake Erie to identify ecological factors driving its growth. Inflows from the Grand River, the largest river to the north shore, were generally positively associated with phosphorus concentrations in the nearshore and negatively associated with light reaching the lakebed. At the depths sampled (3 m–18 m), Cladophora was strongly influenced by light availability, and due to shading by the Grand River plume, an overall negative association was found between Cladophora biomass and phosphorus inputs. Phosphorus limitation was only observed at shallow sites farthest from the Grand River. Positive associations between dreissenid mussel coverage and both Cladophora biomass and tissue phosphorus suggest that nutrient cycling by dreissenids supports Cladophora growth. Our results indicate that i.) the Grand River has a strong influence on nearshore nutrient levels and water clarity; and ii.) Cladophora is limited by both phosphorus and light to varying degrees within the study area, although light appears to be the dominant factor, at least at these depths, years, and locations. The implication that phosphorus reductions could lead to increased Cladophora biomass by improving light conditions will need to be considered carefully against the known historical success of controlling nuisance algae through nutrient management.     

Changes in the Dreissenid Community in the Lower Great Lakes with Emphasis on Southern Lake Ontario

A field study was conducted in the lower Great Lakes to assess changes in spatial distribution and population structure of dreissenid mussel populations. More specifically, the westward range expansion of quagga mussel into western Lake Erie and toward Lake Huron was investigated and the shell size, density, and biomass of zebra and quagga mussel with depth in southern Lake Ontario in 1992 and 1995 were compared. In Lake Erie, quagga mussel dominated the dreissenid community in the eastern basin and zebra mussel dominated in the western basin. In southern Lake Ontario, an east to west gradient was observed with the quagga mussel dominant at western sites and zebra mussel dominant at eastern locations. Mean shell size of quagga mussel was generally larger than that of zebra mussel except in western Lake Erie and one site in eastern Lake Erie. Although mean shell size and our index of numbers and biomass of both dreissenid species increased sharply in southern Lake Ontario between 1992 and 1995, the increase in density and biomass was much greater for quagga mussels over the 3-year period. In 1995, zebra mussels were most abundant at 15 to 25 m whereas the highest numbers and biomass of quagga mussel were at 35 to 45 m. The quagga mussel is now the most abundant dreissenid in areas of southern Lake Ontario where the zebra mussel was once the most abundant dreisenid; this trend parallels that observed for dreissenid populations in the Dneiper River basin in the Ukraine.     

Dreissenid mussels are not a “dead end” in Great Lakes food webs

Dreissenid mussels have been regarded as a “dead end” in Great Lakes food webs because the degree of predation on dreissenid mussels, on a lakewide basis, is believed to be low. Waterfowl predation on dreissenid mussels in the Great Lakes has primarily been confined to bays, and therefore its effects on the dreissenid mussel population have been localized rather than operating on a lakewide level. Based on results from a previous study, annual consumption of dreissenid mussels by the round goby (Neogobius melanostomus) population in central Lake Erie averaged only 6 kilotonnes (kt; 1 kt = one thousand metric tons) during 1995–2002. In contrast, our coupling of lake whitefish (Coregonus clupeaformis) population models with a lake whitefish bioenergetics model revealed that lake whitefish populations in Lakes Michigan and Huron consumed 109 and 820 kt, respectively, of dreissenid mussels each year. Our results indicated that lake whitefish can be an important predator on dreissenid mussels in the Great Lakes, and that dreissenid mussels do not represent a “dead end” in Great Lakes food webs. The Lake Michigan dreissenid mussel population has been estimated to be growing more than three times faster than the Lake Huron dreissenid mussel population during the 2000s. One plausible explanation for the higher population growth rate in Lake Michigan would be the substantially higher predation rate by lake whitefish on dreissenid mussels in Lake Huron.     

Phosphorus Uptake Dynamics as Related to Mathematical Modeling of Cladophora at a Site on Lake Huron

Cladophora is a significant symptom of eutrophication in Lakes Erie and Ontario and is a local problem associated with nutrient perturbations in Lakes Huron, Michigan, and Superior. This paper presents results of measurements of phosphorus uptake rates as a function of internal phosphorus levels by Cladophora growing near Harbor Beach, Michigan. Cladophora collected near the Harbor Beach wastewater treatment plant had high levels of internal phosphorus and low (or even negative) phosphorus uptake rates. Cladophora distant from the wastewater treatment plant had low internal phosphorus levels and rapid phosphorus uptake rates. The experimental results are discussed in terms of quantitative kinetic formulations which may be incorporated into mathematical models useful for predicting the response of Cladophora to alternative management and control strategies. Preliminary model simulations of Cladophora biomass, internal phosphorus, and external phosphorus are qualitatively similar to observed field data.     

Abundance and distribution of benthic macroinvertebrates in offshore soft sediments in Western Lake Huron, 2001–2007

Invasive species have had major impacts on the Great Lakes. This is especially true of exotic dreissenid mussels which are associated with decreased abundance of native macroinvertebrates and changes in food availability for fish. Beginning in 2001, we added a benthic macroinvertebrate survey to the USGS-Great Lakes Science Center's annual fall prey fish assessment of Lake Huron to monitor abundance of macrobenthos. Mean abundance of Diporeia, the most abundant benthic taxon in Lake Huron reported by previous investigators, declined greatly between 2001 and 2007. Diporeia was virtually absent at 27-m sites by 2001, decreased and was lost completely from 46-m depths by 2006, but remained present at reduced densities at 73-m sites. Dreissenids in our samples were almost entirely quagga mussels Dreissena bugensis. Zebra mussels Dreissena polymorpha were virtually absent from our samples, suggesting that they were confined to nearshore areas shallower than we sampled. Loss of Diporeia at individual sites was associated with arrival of quagga mussels, even when mussel densities were low. Quagga mussel density peaked during 2002, then decreased thereafter. During the study quagga mussels became established at most 46-m sites, but remained rare at 73-m sites. Length frequency distributions suggest that initial widespread recruitment may have occurred during 2001–2002. Like other Great Lakes, Lake Huron quagga mussels were associated with decreased abundance of native taxa, but negative effects occurred even though dreissenid densities were much lower. Dreissenid effects may extend well into deep oligotrophic habitats of Lake Huron.     

Associations between Breeding Marsh Bird Abundances and Great Lakes Hydrology

We used Great Lakes hydrologic data and bird monitoring data from the Great Lakes Marsh Monitoring Program from 1995–2002 to: 1) evaluate trends and patterns of annual change in May-July water levels for Lakes Ontario, Erie, and Huron-Michigan, 2) report on trends of relative abundance for birds breeding in Great Lakes coastal marshes, and 3) correlate basin-wide and lake-specific annual indices of bird abundance with Great Lakes water levels. From 1995–2002, average May, June, and July water levels in all lake basins showed some annual variation, but Lakes Erie and Huron-Michigan had identical annual fluctuation patterns and general water level declines. No trend was observed in Lake Ontario water levels over this period. Abundance for five of seven marsh birds in Lake Ontario wetlands showed no temporal trends, whereas abundance of black tern (Chlidonias niger) declined and that of swamp sparrow (Melospiza georgiana) increased from 1995–2002. In contrast, abundances of American coot (Fulica americana), black tern, common moorhen (Gallinula chloropus), least bittern (Ixobrychus exilis), marsh wren (Cistorthorus palustris), pied-billed grebe (Podilymbus podiceps), sora (Porzana carolina), swamp sparrow, and Virginia rail (Rallus limicola) declined within marshes at Lakes Erie and Huron/Michigan from 1995–2002. Annual abundances of several birds we examined showed positive correlations with annual lake level changes in non-regulated Lakes Erie and Huron/Michigan, whereas most birds we examined in Lake Ontario coastal wetlands were not correlated with suppressed water level changes of this lake. Overall, our results suggest that long-term changes and annual water level fluctuations are important abiotic factors affecting abundance of some marsh-dependent birds in Great Lakes coastal marshes. For this reason, wetland bird population monitoring initiatives should consider using methods in sampling protocols, or during data analyses, to account for temporal and spatial components of hydrologic variability that affect wetlands and their avifauna.     

Monitoring Cladophora Growth Conditions and the Effect of Phosphorus Additions at a Shoreline Site in Northeastern Lake Erie

A study to measure environmental conditions, Cladophora standing crop, internal nutrient levels, and the effect of the addition of phosphorus to Cladophora growth at a single location on the north shore in the eastern basin of Lake Erie is described. In 1979, the mean standing crop for depths 0.5–3 m was 431 gDW/m² as measured at the time of maximum standing crop in early July. Thereafter, the alga was sloughed and carried ashore causing a rapid decline in standing crop. These events coincided with the attainment of lake temperatures exceeding 20° C. Total phosphorus concentrations averaged about 18 μg P/L while soluble reactive phosphorus levels remained near the limit of detection. Stoichiometric ratios of nitrate nitrogen to soluble reactive phosphorus approximated 150:1, suggestive of phosphorus limited conditions. Internal phosphorus and nitrogen levels averaged about 0.06% and 1.80%, respectively. In 1980, phosphorus (0.34 kg/day) was discharged at the 0.5 m depth commencing July 19. No response was noted until the water temperature dropped below 20° C in September when a rapid regrowth occurred, apparently in response to the nutrient addition. It is concluded that Cladophora grows in response to available phosphorus in the eastern basin of Lake Erie and that limitation of this nutrient may be expected to reduce Cladophora growth.     

Changes in mid-summer water temperature and clarity across the Great Lakes between 1968 and 2002

In recent decades, three important events have likely played a role in changing the water temperature and clarity of the Laurentian Great Lakes: 1) warmer climate, 2) reduced phosphorus loading, and 3) invasion by European Dreissenid mussels. This paper compiled environmental data from government agencies monitoring the middle and lower portions of the Great Lakes basin (lakes Huron, Erie and Ontario) to document changes in aquatic environments between 1968 and 2002. Over this 34-year period, mean annual air temperature increased at an average rate of 0.037 °C/y, resulting in a 1.3 °C increase. Surface water temperature during August has been rising at annual rates of 0.084 °C (Lake Huron) and 0.048 °C (Lake Ontario) resulting in increases of 2.9 °C and 1.6 °C, respectively. In Lake Erie, the trend was also positive, but it was smaller and not significant. Water clarity, measured here by August Secchi depth, increased in all lakes. Secchi depth increased 1.7 m in Lake Huron, 3.1 m in Lake Ontario and 2.4 m in Lake Erie. Prior to the invasion of Dreissenid mussels, increases in Secchi depth were significant (p < 0.05) in lakes Erie and Ontario, suggesting that phosphorus abatement aided water clarity. After Dreissenid mussel invasion, significant increases in Secchi depth were detected in lakes Ontario and Huron.     

Depth drives growth dynamics of dreissenid mussels in Lake Ontario

Understanding dreissenid mussel population dynamics and their impacts on lake ecosystems requires quantifying individual growth across a range of habitats. Most dreissenid mussel growth rates have been estimated in nutrient rich or nearshore environments, but mussels have continued to expand into deep, cold, low-nutrient habitats of the Great Lakes. We measured annual quagga mussel (Dreissena rostriformis bugensis) growth at 15 m, 45 m, and 90 m in Lake Ontario using caged mussels near Oswego, New York, USA from June 2018 to May 2019. Quagga mussel growth (starting size 12 mm) was greatest at 15 m (mean shell length increase = 10.2 mm), and was lower at 45 m (5.9 mm) and 90 m (0.7 mm). Caged mussels were obtained from near the 90-m site and those reared at 15 and 45 m developed thicker shells than those that were caged at 90 m. We observed relatively high colonization by quagga and, to a lesser degree, zebra mussels (Dreissena polymorpha) at 15 m, very few colonizers at 45 m, and none at 90 m. Higher growth potential, but low natural mussel densities observed at 15 m and 45 m suggest factors other than growth limit dreissenid abundance at these depths. The relatively slow dreissenid growth rates observed in offshore habitats are consistent with the gradual abundance increases documented in these zones across the Great Lakes and suggest new mussels that become established in these habitats may contribute to ecosystem effects for decades.     

Decadal changes in the benthic invertebrate community in western Lake Erie between 1981 and 2004

Faunal surveys of western Lake Erie showed a dramatic change in the species composition and abundance of the invertebrate mud-bottom community. Abundances of formerly dominant ostracods, tubificid and naidid oligochaetes, and chironomids declined 85%, 87%, 80%, and 72%, respectively, between 1982 and 2003. The majority of the declines occurred between 1982 and 1993, when dreissenid abundances were increasing rapidly. Hexagenia nymphs became important members of the infaunal community after 1993. Faunal declines were uncorrelated to weather-related variables. The frequency and magnitude of thermal stratification with accompanying hypoxic conditions during the study period remained within the normal range of variation observed in western Lake Erie. Significant wind events and storm driven seiches that might cause benthos mortality were not correlated with the decadal patterns of faunal decline. Nor were the faunal declines associated with the historical decrease in organic loading, because most of the decrease occurred prior to the study period. Nearly all abundant species declined significantly between 1982 and 1993, but deposit-feeding fauna, eutrophic indicator species, small organisms, and nearshore mud-bottom stations adjacent to dreissenid mussel habitat were most severely affected. These patterns suggest that a systemic change in the trophic structure of western Lake Erie occurred, due mostly to the invasion of dreissenid mussels nearshore and an accompanying diversion of organic matter away from deeper offshore muds. Trophic group amensalism may operate differently in marine and freshwater habitats, although dreissenids may exert ecosystem effects in the Great Lakes similar to oysters in Chesapeake Bay.     

The good,the bad,and the algae: Perceiving ecosystem services and disservices generated by zebra and quagga mussels

Dreissenid (zebra and quagga) mussels are widely recognized as having strong, adverse ecological and economic impacts, e.g., biofouling and loss of water column primary production. We assessed perceptions and values associated with two less often considered ecological outcomes of dreissenid mussel influences on coastal ecosystems along Lake Ontario and the western St. Lawrence River in New York State. One, the generation of water clarity through filtration, we define as an ecosystem service; the other, the production of large amounts of nuisance algae (e.g., Cladophora and Microcystis) is defined as an ecosystem disservice. Surveys of business owners and homeowners quantified their preferences and the formation of values regarding these products of zebra mussel influence. Water clarity increased greatly, particularly in the eastern portion of Lake Ontario, and algal problems increased throughout. Businesses attributed increases and decreases in revenues associated with water clarity and algae; homeowners reported analogous changes in property values. Water clarity was positively associated, and algae negatively associated, with changes in revenues and property values. Threshold responses of costs as functions of filamentous algae were evident. Given the likely continued influx of invasive species due to human activities, further development of the ecosystem service concept should consider potential “goods” and “bads” of invasives and their influence on ecosystem and social system resiliency.     

The Zebra Mussel Dreissena polymorpha Found in North America in 1986 and 1987

The zebra mussel Dreissena polymorpha was first detected in the western basin of Lake Erie, Ontario, Canada, on natural gas wellheads and well markers between April and November 1986. It was found again in 1987 on the north shore of Lake Erie in a water treatment plant, and in vessel fouling. The population increased in Lake Erie in 1988. Dreissena may have spread from Lake Erie to Lake St. Clair, where it was then discovered on 1 June 1988.     

Ecological Studies and Mathematical Modeling of Cladophora in Lake Huron: 5. Model Development and Calibration

The development of a mathematical model for calculating the spatial distribution and temporal variation in Cladophora biomass and selected forms of phosphorus at a site on Lake Huron is described. The model is intended for use in evaluating the utility of various phosphorus management strategies in reducing the nuisance growth of Cladophora in the Great Lakes. The model is composed of a transport component which accounts for bulk transfer and exchange of phosphorus within the system and a kinetic component which describes chemical and biological reactions. The kinetic component includes the effects of light, temperature, and internal phosphorus levels on Cladophora growth rate, aspects of phosphorus uptake, and the impact of sloughing and self-shading on standing crop. The numerical value of coefficients associated with kinetic equations have been independently determined through laboratory studies.The model has been calibrated to a data set collected in the vicinity of the Harbor Beach, Michigan, wastewater treatment plant. A satisfactory fit was achieved for model calculations and observed levels of Cladophora biomass, internal phosphorus, and soluble reactive phosphorus. The model has few degrees of freedom because the coefficient values were fixed from field and laboratory measurements. A standing crop of Cladophora biomass of 300 gDW/m² with an internal phosphorus level of 0.35%P was characteristic of stations in the vicinity of the nutrient source. Soluble reactive phosphorus levels fluctuated widely due to mixing of the effluent plume with offshore waters, but averaged approximately 30 μgP/L. A distinct spatial trend was observed with Cladophora biomass, internal phosphorus, and soluble reactive phosphorus decreasing with distance from the point source of nutrients. The opportunity for model verification by perturbation through phosphorus removal is discussed.     

Spatiotemporal patterns of water quality in Lake Ontario and their implications for nuisance growth of Cladophora

This study, motivated by a resurgence in Cladophora, investigates changes in the nutrient environment in the littoral zone of Lake Ontario. We measured nutrient concentrations from 2004 to 2008 at two littoral zone (2–12 m) sites on the north shore of Lake Ontario where Cladophora has experienced a resurgence and compared concentrations with data collected in the late 1970s. Spring total phosphorus (TP) and soluble reactive P (SRP) concentrations have significantly declined at these two sites. Furthermore, P loading from the major tributaries to our study sites declined between 1964 and 2008. Upwelling events were not detectably associated with increases in P concentrations at our sites. We conclude that a recent upsurge in nuisance Cladophora, at least at these sites, cannot be explained by deteriorating littoral zone water quality in terms of P concentrations or by changes in catchment loading. For additional context, we also examined trends in coastal (14–20 m) and offshore (> 50 m) nutrients using Environment Canada epilimnetic surveillance data, 1975–2008. Significant declines in TP and SRP concentrations have occurred in north coast waters, concurrent with declines in the offshore. However, nutrient concentrations, notably spring SRP, have not decreased among south coast stations, potentially reflecting greater coastal entrapment of catchment-derived waters. We infer that EC-monitored north coast stations reflect integrated interannual water quality, while south coast stations are more strongly influenced by catchment loading. The effects of higher nutrient concentrations along the south coast, which co-occur with lower water transparency, on benthic algal growth have yet to be determined.     

Post-dreissenid Increases in Transparency During Summer Stratification in the Offshore Waters of Lake Ontario: Is a Reduction in Whiting Events the Cause?

Since the dreissenid invasion of the lower Great Lakes, calcium concentrations in the offshore waters of Lake Ontario have decreased by approximately 4–5 mg/L. This decline has coincided with a three-fold reduction in August turbidity values and nearly a doubling of Secchi depths, presumably due to reduced summer calcite precipitation events in the lake. The reductions in calcium have followed a dramatic reduction in alkalinity in the central and eastern basins of Lake Erie, which provides most of the inflow to Lake Ontario. This reduction in alkalinity in Lake Erie corresponds to a period of rapid dreissenid growth in that lake, strongly suggesting calcium uptake by dreissenid mussels as a causative factor. The mass of calcium resident in the dreissenid population in Lake Erie, estimated from published lake-wide census data, is sufficient to account for the observed decreases in alkalinity. In addition, observed changes in alkalinity in Lake Ontario closely match those expected to result from inflows from Lake Erie, based on mass balance considerations. Considered in sum, our data strongly suggest that calcium uptake by dreissenid mussels in Lake Erie has resulted in decreases in the calcium concentration in Lake Ontario, reducing the frequency and/or intensity of whiting events in the latter lake. We believe this is the first report of an increase in transparency that can be reasonably attributed to a chemical change brought about by Dreissena. These increases in transparency may have very different consequences than those of dreissenid filtration activities. For example, rather than decreasing phytoplankton populations, the improved light climate might increase summer phytoplankton populations, particularly sub-epilimnetic ones.     

Ecosystem changes and nuisance benthic algae on the southeast shores of Lake Huron

We conducted studies of algal fouling along the southeastern shore of Lake Huron to determine the kinds and spatial distribution of benthic algae, the spatial extent and timing of beach fouling, and the possible influences of biological changes to Lake Huron. There was no change in the physical texture of the substratum, but coverage by algal turf increased from 11% of sites in 1977 to nearly 90% in 2007. Shoreline surveys showed that Chara was most common on flat rocky substrata at depths of 0.15 to 0.20 m. Algae stranded on beaches consisted of 62% periphyton turf, 30% Chara and 8% Cladophora and were not evenly distributed; the largest accumulations were found where shoreline irregularities interrupted longshore flow. Quantities of stranded algae partially reflected the height and duration of waves directed onshore. Macroinvertebrate densities were not correlated with benthic algal abundance in 2010 and were lower with fewer large grazers than in 1980. Densities of Dreissena spp. increased with depth, and small tufts of Cladophora were found on larger individuals. The proliferation of algae in the nearshore zone of central eastern Lake Huron appears to be the result of several recent changes. Phosphorus management and filtering by dreissenid mussels have reduced phytoplankton abundance, improving the light regime. Changes in land use may have increased loadings of phosphorus through shallow groundwater and tributary streams. Dreissenids have also redirected nutrients to the lakebed, further enhancing benthic primary production, and predation by round goby has reduced the numbers of grazing benthic invertebrates.     

Trends in Total Phosphorus in Canadian Near–Shore Waters of the Laurentian Great Lakes: 1976–1999

Beginning as early as 1976 at many locations, total phosphorus concentrations (TP) were measured weekly in samples collected year-round in the intake water of 18 municipal water treatment plants in Canadian (Ontario) waters of the Laurentian Great Lakes. No consistent long-term trends were evident at two north-shore Lake Superior sampling locations, but there were significant long-term declines in TP measured at all three Lake Huron locations; however, concentrations there have remained relatively constant during the past decade. Declines in TP averaging about 1 μg/L/yr during 1976 to 1990 were prevalent at lower Great Lakes sampling locations and by the early 1990s TP had declined to 15–25 μg/L in Lake Erie and 10–20 μg/L in Lake Ontario. Declines generally levelled out in Lake Ontario after 1990, but TP increased substantially at some Lake Erie locations in the late 1990s. Recent (1996 to 1999) total phosphorus concentrations in north-shore Lake Erie locations in the range of 20 to 30 μg/L were 2 to 3 times higher than at Lake Ontario near-shore locations in the 8 to 11 μg/L range. Rates of decline of TP were generally highest for the March–April period (−1.88, −1.61, and −1.34 μg/L/yr in Lakes Ontario, Erie, and Huron, respectively for 1976 to 1990). The March–April Lake Ontario near-shore rate of TP decline was nearly twice as high as that reported previously for off-shore Lake Ontario (attributed to proximity to P loading sources and to lower net sedimentation losses of P in the near-shore environment). There were substantial declines in chlorophyll-to-TP ratios and in the slopes and Y-intercepts of chlorophyll-TP regressions for both Lake Erie and Lake Ontario following the establishment of dreissenid mussels.