Biomass and Distribution of Cladophora Glomerata in Relation to Some Physical-Chemical Variables at Two Sites in Lake Erie

Cladophora was collected from 0.25-m² quadrats on alternate weeks at two sites located in Lake Erie near Hamburg, New York, and Walnut Creek near Erie, Pennsylvania, between June and November 1979. Cladophora biomass and internal nutrients (total Kjeldahl nitrogen and total phosphorus) were determined. Water samples were analyzed for total phosphorus, soluble reactive phosphorus, nitrate + nitrite nitrogen, total Kjeldahl nitrogen, and ammonia nitrogen. Temperature and Secchi disk transparency were also recorded.The peak standing crop for Cladophora was 211 gDW/m² (July) at the Hamburg site and 57 gDW/m² (July) at Walnut Creek. The depth distribution of Cladophora at the sample sites was influenced by wave activity and drifting sand. As a result of sand scouring at the shallower depths, Cladophora biomass generally was greatest at the 3-m depth.Reductions in Cladophora internal nutrients corresponded to decreased nutrients in the water. Cladophora detached from its substrate during periods when low internal nutrient levels were observed. Internal total phosphorus concentrations were low and approached the minimum cell quota (0.05–0.06% –P) most of the time. Soluble reactive phosphorus values were also low (1–4 μgP/L). Levels of soluble reactive phosphorus and internal total phosphorus were low and almost always limiting, and may be responsible for the low biomass values observed. In addition, internal total phosphorus levels showed a significant positive correlation with Cladophora biomass. Internal total nitrogen values were generally less than 1%-N at Walnut Creek and ammonia nitrogen measurements were near detection limits. Cladophora was nitrogen-limited at the Walnut Creek site during part of the study period. Nitrate + nitrite nitrogen showed a significant positive correlation with Cladophora biomass at that site.     

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.     

Environmental Factors Controlling Physiological Changes in Cladophora in Lake Erie

Several physiological characteristics of Cladophora glomerata from eastern Lake Erie were monitored frequently during the summer of 1977. Soluble reactive phosphorus and nitrate-nitrogen in the lake water were also measured.Cladophora biomass generally increased throughout June, although there were periodic declines. In mid-July, biomass sharply decreased to a low level. Chlorophyll content tended to be high during periods of increasing biomass but was lower after each biomass decline. Changes in biomass did not correlate with changes in either phosphate-phosphorus or nitrate-nitrogen in the lake water. Cellular nitrogen levels fluctuated independent of lake water nitrate-nitrogen concentrations. Cellular nitrogen levels remained above the critical concentration, 1.1%-N. Luxury phosphorus levels were observed to increase following pulses of soluble orthophosphate in the lake water. Measurements of cellular total phosphorus indicate that levels remain well above the critical concentration of 0.06%-P. It is concluded that neither nitrogen nor phosphorus is limiting the growth of Cladophora at the sites sampled. The mid-summer die-off does not appear to be due to a simple nutrient deficiency.     

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.     

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.     

Growth Dynamics of Cladophora Glomerata in Western Lake Erie in Relation to Some Environmental Factors

Cladophora was monitored at two sites in western Lake Erie during 1979 and 1980 as part of a lake-wide Lake Erie Cladophora Surveillance Program. Two distinctive zones within the littoral region were colonized by the alga, the eulittoral (splash zone) and infralittoral (defined in the present study as the 0.5–4 m depth zone). Cladophora of the eulittoral zone became established in May and remained present until late fall. The infralittoral zone Cladophora exhibited a bimodal growth pattern related to the seasonal temperature regime, with growth occurring from April to July and again from September to November. The infralittoral zone supported the largest share of biomass, which resulted in nuisance accumulations upon the beaches in the island region. Peak biomass was observed from mid-June to early July, obtaining maximum values of 102 gDW/m² and 214 gDW/m² for the 1979 and 1980 seasons, respectively. The depth to which Cladophora colonized was limited by light availability; maximum depth of growth occurred between 2 and 4 m in western Lake Erie due to the turbid nature of the basin. Phosphorus and nitrogen were not limiting to Cladophora growth in western Lake Erie; tissue nutrients remained above the critical levels defined by Gerloffand Fitzgerald (1976) throughout the season.     

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.     

Energy Relations in Cladophora Glomerata from Lake Erie

Photosynthesis and respiration measurements were made at lake temperatures with Cladophora glomerata collected from the eastern basin of Lake Erie. Although water temperatures varied from 10°C in May to almost 25°C in July, neither photosynthesis nor respiration showed any general increase in rates, even though there were wide daily fluctuations. Starch levels were low during the period of rapid growth but then increased until the time of sloughing. These data suggest that photosynthesis and respiration in Cladophora temperature-adapt and that a negative energy balance is not the cause of the mid-summer die-off.     

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.     

Re-eutrophication of Lake Erie: Correlations between tributary nutrient loads and phytoplankton biomass

Both abiotic and biotic explanations have been proposed to explain recent recurrent nuisance/harmful algal blooms in the western basin and central basin of Lake Erie. We used two long-term (> 10 years) datasets to test (1) whether Lake Erie total phytoplankton biomass and cyanobacterial biomass changed over time and (2) whether phytoplankton abundance was influenced by soluble reactive phosphorus or nitrate loading from agriculturally-dominated tributaries (Maumee and Sandusky rivers). We found that whereas total phytoplankton biomass decreased in Lake Erie's western basin from 1970 to 1987, it increased starting in the mid-1990s. Total phytoplankton and cyanobacterial seasonal (May–October) arithmetic mean wet-weight biomasses each significantly increased with increased water-year total soluble reactive phosphorus load from the Maumee River and the sum of soluble reactive phosphorus load from the Maumee and Sandusky rivers, but not for the Sandusky River alone during 1996–2006. During this same time period, neither total phytoplankton nor cyanobacterial biomass was correlated with nitrate load. Consequently, recently increased tributary soluble reactive phosphorus loads from the Maumee River likely contributed greatly to increased western basin and (central basin) cyanobacterial biomass and more frequent occurrence of harmful algal blooms. Managers thus must incorporate the form of and source location from which nutrients are delivered to lakes into their management plans, rather than solely considering total (both in terms of form and amount) nutrient load to the whole lake. Further, future studies need to address the relative contributions of not only external loads, but also sources of internal loading.     

Environmental Controls of Cladophora Growth Dynamics in Eastern Lake Erie: Application of the Cladophora Growth Model (CGM)

The Cladophora growth model (CGM) was used to estimate the importance of light, temperature, phosphorus, and self-shading on the spatial and temporal variability of Cladophora growth rates and biomass accrual in eastern Lake Erie during 2002. The CGM predicted that Cladophora growth was highly sensitive to spatial and temporal variations in soluble phosphorous concentration (SRP). Specifically the CGM predicted that: 1) Site-to-site differences in SRP concentration resulted in a 2× difference in depth-integrated biomass; 2) maximum growth rates were strongly influenced by SRP concentrations during periods of rapid biomass accrual (mid-June to mid-July); 3) inter-annual differences in SRP concentration during the spring period (∼ 1 μg/L) could result in up to a 3.5× difference in depth integrated biomass; 4) Spatial variations in water clarity could result in a 2× difference in depth-integrated biomass between sites, with variations betweens sites occurring primarily between 2–6 m depth; 5) the mid-summer sloughing phenomenon likely resulted from self-shading by the algal canopy; and 6) the seasonal growth pattern of Cladophora was strongly regulated by temperature.     

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.     

Cladophora (green algae) and dreissenid mussels over a nutrient loading gradient on the north shore of Lake Ontario

Benthic algae, much of it the green alga Cladophora, blanket the nearshore lakebed of the north shore of Lake Ontario. Nearshore field studies in Toronto, Ajax, Oshawa and Cobourg in 2012 and 2013, and Toronto in 2015 examined the distribution of Cladophora over a nutrient gradient on the north shore of the lake. Concentrations of total phosphorus, dissolved phosphorus and total inorganic nitrogen in the water column decreased from west to east over the gradient corresponding with decreasing watershed development and size. However, high surface cover and nuisance levels of Cladophora biomass were found across the gradient, including the least developed study area where total phosphorus concentrations were similar to those in the open lake. The abundance of Dreissena, high in all areas, was measured concurrently with Cladophora biomass and correlated positively at depths of 6 to 10 m. External loading of phosphorus provides a basis for abatement of Cladophora; however, the influences of enrichment along the shoreline, internal loading at the lakebed and lake trophic status in sustaining growth remain obscure. Nuisance levels of Cladophora are not isolated to locations experiencing elevated external loading and should be viewed in the context of interacting area-specific and lake-wide nutrient supplies.     

Potential Strategies for Recovery of Lake Whitefish and Lake Herring Stocks in Eastern Lake Erie

Lake Erie sustained large populations of ciscoes (Salmonidae: Coregoninae) 120 years ago. By the end of the 19^th century, abundance of lake whitefish (Coregonus clupeaformis) had declined drastically. By 1925, the lake herring (a cisco) population (Coregonus artedii) had collapsed, although a limited lake herring fishery persisted in the eastern basin until the 1950s. In the latter part of the 20^th century, the composition of the fish community changed as oligotrophication proceeded. Since 1984, a limited recovery of lake whitefish has occurred, however no recovery was evident for lake herring. Current ecological conditions in Lake Erie probably will not inhibit recovery of the coregonine species. Recovery of walleye (Sander vitreus) and efforts to rehabilitate the native lake trout (Salvelinus namaycush) in Lake Erie will probably assist recovery because these piscivores reduce populations of alewife (Alosa psuedoharengus) and rainbow smelt (Osmerus mordax), which inhibit reproductive success of coregonines. Although there are considerable spawning substrates available to coregonine species in eastern Lake Erie, eggs and fry would probably be displaced by storm surge from most shoals. Site selection for stocking or seeding of eggs should consider the reproductive life cycle of the stocked fish and suitable protection from storm events. Two potential sites in the eastern basin have been identified. Recommended management rocedures, including commercial fisheries, are suggested to assist in recovery. Stocking in the eastern basin of Lake Erie is recommended for both species, as conditions are adequate and the native spawning population in the eastern basin is low. For lake herring, consideration should be given to match ecophenotypes as much as possible. Egg seeding is recommended. Egg seeding of lake whitefish should be considered initially, with fingerling or yearling stocking suggested if unsuccessful. Spawning stocks of whitefish in the western basin of Lake Erie could be utilized.     

Phytoplankton Composition and Biomass in the Offshore Waters of Lake Erie: Pre- and Post-Dreissena Introduction (1983–1993)

Phytoplankton was collected in all basins of Lake Erie during 42 cruises during the spring and summer from 1983 to 1993—a period that spans the Dreissena mussel invasion. Two potential impacts of Dreissena on the phytoplankton community of the western, central, and eastern basins of Lake Erie were evaluated: Was selective feeding occurring as observed in Saginaw Bay and were reductions in biomass evident in the offshore regions of the three basins of Lake Erie? In the western basin, significant summer decreases in Chlorophyta, Bacillariophyta, Cyanobacteria, and total phytoplankton biomass were observed after Dreissena introduction. Similarly in the spring, Bacillariophyta and total phytoplankton biomass and chlorophyll a concentrations decreased significantly. Since several divisions of phytoplankton did not decrease in phytoplankton biomass in the western basin, and spring Cyanobacteria biomass increased significantly while other divisions decreased in biomass, selective feeding on the phytoplankton community was suggested. Where significant reductions in biomass were observed in the offshore waters of the western basin, they were approximately 50% of the reduction observed at the nearshore sites in Lake Erie by other workers.Dreissena impact on the phytoplankton community of the pelagic waters of the central and eastern basin appeared to be minimal. Pre- and post-Dreissena total phytoplankton biomass and chlorophyll a concentrations were not significantly different or increased significantly after the Dreissena invasion. Biomass of several divisions of phytoplankton significantly increased after Dreissena introduction in the central and eastern basins. These included Bacillariophyta (central basin), Cyanobacteria (central and eastern basin), Chrysophyta (eastern basin), Chlorophyta biomass (eastern basin) and phytoplankton biomass (central basin) and chlorophyll a (central basin) in the spring, and Chrysophyta (eastern basin) and Cryptophyta biomass (central basin) in the summer. Generally, a reduction in phytoplankton biomass would be expected as a result of Dreissena grazing, not an increase in biomass. Dreissena-mediated changes in phytoplankton have generally occurred in shallow, well-mixed lakes, ponds, and embayments, not in deeper waters such as the central and eastern basins of Lake Erie.     

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.     

Cladophora,mass transport,and the nearshore phosphorus shunt

The nearshore phosphorus shunt hypothesis and the potential for mussels to excrete phosphorus sufficient to meet the growth requirements of Cladophora are now well accepted by scientists studying Great Lakes biogeochemistry. The response of algal growth to near bottom water column phosphorus concentrations and the interplay between excretion and mass transport in yielding those concentrations have, however, not been elucidated. Here we present soluble reactive phosphorus profiles from the near bottom environment of Lake Michigan at a site near Good Harbor Bay, Michigan, where both mussels and Cladophora were present. Soluble reactive phosphorus was observed to accumulate under quiescent conditions, establishing a concentration boundary layer (CBL), 5–15 cm thick, with near bottom concentrations on the order of 2–8 μg P/L. A one-dimensional model was applied to determine mass transport conditions mediating the transition from CBL formation to CBL destruction. Significant wave height (SWH) was used as an indicator of mass transport intensity, and it was determined that the formation/destruction transition occurred at a SWH of 0.2 m at the 8-m study site depth. The Great Lakes Cladophora Model was applied to determine the time intervals required to saturate (1 day with the CBL present) and deplete (14 days with the CBL absent) algal internal P stores. A review of SWH conditions at the study site indicated that a CBL would be expected to form at a frequency sufficient to support the phosphorus nutrition of Cladophora over the entire May to August interval.