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.     

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.     

Seasonal Abundance Patterns of Diatoms on Cladophora in Lake Huron

Rocks bearing Cladophora were collected from May to November 1979 at two locations near Harbor Beach, Michigan, in Lake Huron to document seasonal patterns of epiphytic diatom abundance and diatom proportion of the Cladophora-epiphyte assemblage biomass in an area receiving effluent from a municipal wastewater treatment plant. Data were examined for evidence of interactions between epiphytic diatoms and Cladophora.Cladophora first appeared in May at which time epiphytic diatoms comprised about 30% of the Cladophora-epiphytic assemblage biomass. Cladophora growth was greatest in June and July, accumulating much faster than diatoms. Peak Cladophora-epiphyte assemblage biomass was maintained from July to September. Cladophora biomass apparently decreased after August while diatom abundance increased to a September maximum. Diatoms were responsible for the sustained peak of Cladophora-epiphyte assemblage biomass as diatoms comprised over 60% of the assemblage biomass in September. The Cladophora-epiphyte assemblage collapsed by October.Low diversity of the epiphytic diatom taxocene and low diatom proportion of the Cladophora-epiphyte assemblage biomass indicated Cladophora may have outcompeted diatom epiphytes during the June-July Cladophora growth pulse. Subsequently epiphytic diatoms may have enhanced Cladophora sloughing by shading and successful nutrient competition.     

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.     

Abundance Patterns of Diatoms on Cladophora in Lake Huron with Respect to a Point Source of Wastewater Treatment Plant Effluent

Epiphytic diatoms on Cladophora were collected from July to September, 1979, at various distances from a point source of municipal wastewater treatment plant effluent to Lake Huron. Diatom accumulation rates were positively related to distance from the point source. Despite higher nutrient concentrations nearest the point source, low accumulation rates of Cladophora-epiphyte assemblage biomass and epiphytic diatoms indicated moderate growth inhibition by an undetermined factor. Evaluation of epiphytic diatom population abundance and ecology plus diatom taxocene species diversity supported our conclusions. A curvilinear relationship between diatom species diversity and impacts of growth inhibition is proposed and discussed. The diatom proportion of Cladophora-epiphyte assemblage biomass was positively related to distance from the point source of sewage effluent.     

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.     

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.     

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.     

Daphnia lumholtzi Sars a new alien species in Lake St. Clair

Daphnia lumholtzi Sars, an exotic tropical/subtropical cladoceran from Australia, southeast Asia, and Africa, was newly found in Lake St. Clair in a vertical tow sample taken at 3 m depth on 25 July 2007. The species was previously found in 1990/1991 in some reservoirs in the southern United States from where it colonized many waters north to the Great Lakes. In 1999, it was found in Lake Erie. This cladoceran had a density of 117 individuals/m³ when we collected it from Lake St. Clair and it was represented by both females (95.12 %) and males (4.88 %). It seems that D. lumholtzi will continue to expand its distribution area in the Great Lakes.     

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.     

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.     

Fate of phosphorus from a point source in the Lake Michigan nearshore zone

Nutrient loading into Lake Michigan can produce algal blooms which in turn can lead to hypoxia, beach closures, clogging of water intakes, and reduced water quality. The Great Lakes Water Quality Agreement targets for Lake Michigan are 5600 MT annually for total phosphorus (TP) loading, 7 μg L^?1 lake-wide mean TP concentration, and a chlorophyll-a concentration of 1.8 μg L^?1. However, in light of the recent resurgence of nuisance algal (Cladophora sp.) growth in the nearshore zone, the validity of these targets is now uncertain. The occurrence and abundance of Cladophora in the nearshore area depends primarily on the availability of dissolved phosphorus, light, and temperature. The availability of dissolved phosphorus is a potentially useful indicator of nearshore areas susceptible to excessive Cladophora growth and impaired water quality. Regulating agencies are looking for guidance in determining phosphorus loading rates that minimize local exceedance of the lake target concentration. In this study, the lake assimilative capacity was quantified by applying a biophysical model to estimate the area required for mixing and diluting wastewater treatment plant outfall TP loadings to the level of the lake target concentration during the Cladophora growing season. Model results compared well with empirical measurements of particulate and dissolved phosphorus as well as Cladophora biomass and phosphorus content. The model was applied to test scenarios of wastewater treatment plant phosphorus loading in two different years, in order to help establish phosphorus discharge limits for the plant.     

Contaminants Associated with Suspended Sediments in Lakes Erie and Ontario, 1997–2000

Sediment traps were installed at individual index stations in the western basin of Lake Erie and the Mississauga (central) basin of Lake Ontario, and refurbished seasonally during the period 1997–2000. In Lake Ontario, sediment down flux rates and corresponding contaminant down flux rates were highest in winter and increased with depth due to the influence of resuspended bottom sediments. Sediment down flux rates in western Lake Erie (22 to 160 g m⁻² d⁻¹) were far greater than in Lake Ontario (0.19–3.0 g m⁻² d⁻¹). Suspended material in western Lake Erie was characterized as predominately resuspended bottom sediments; down flux rates were roughly 5- to 10-fold higher in spring and fall, compared to summer. Suspended sediment concentrations of PCBs and other organochlorine contaminants, represented by both annual means and individual seasonal values, were higher in Lake Ontario throughout the duration of the study, compared to Lake Erie. The mean annual concentration of PCBs in suspended sediments over the period 1997–2000 was 330 ng/g in western Lake Erie and 530 ng/g in Lake Ontario. Based on a comparison with historical data from Lake Ontario, mean contaminant concentrations over the period 1997–2000 for PCBs, hexachlorobenzene, and mirex corresponded to decreases of 38%, 74%, and 40%, respectively, since the mid-1980s. Corresponding down flux rates for PCBs, hexachlorobenzene, and mirex decreased by approximately 70%, 90%, and 80%, respectively, since the 1980s.     

Twenty five years of changes in Dreissena spp. populations in Lake Erie

Lake Erie has the longest history of colonization by both Dreissena polymorpha and Dreissena rostriformis bugensis in North America and is therefore optimal for the study of long-term dynamics of dreissenid species. In addition, the morphometry of Lake Erie basins varies dramatically from the shallow western to the deep eastern basin, making this waterbody a convenient model to investigate patterns of Dreissena distribution, as well as interspecies interactions among dreissenids. We compare our data on the distribution, density and wet biomass of both dreissenid species in Lake Erie collected in 2009 and 2011–2012 with previous data. We found that Dreissena spp. distribution in Lake Erie varied depending on the time since the initial invasion, collection depth, and lake basin. In 2009–2012, zebra mussels were smaller than in 1992 and were consistently smaller than quagga mussels. During 2009–2012, quagga mussels were found at all depths and in all basins, while zebra mussels were common in the western basin only, and in the central and eastern basins were limited to shallow depths, resulting in an almost complete replacement of D. polymorpha with D. rostriformis bugensis. In the shallowest western basin of Lake Erie, zebra mussels represented > 30% of the combined dreissenid density even after more than 20 years of coexistence, providing strong evidence that, even in lakes as large as Lake Erie (or at least its western basin), D. polymorpha may sustain a significant presence for decades without being displaced by quagga mussels.     

Ecological Interactions Among Bangia, Cladophora, and Ulothrix Along the Lake Erie Shoreline

Bangia atropurpurea, a recent invader of Lake Erie, has altered the community structure of the littoral zone, which previously had been dominated by Cladophora or Ulothrix. Vertical and horizontal zonation patterns of these algae suggest that competition and differing physical tolerances may be involved. To determine the relative importance of these factors, seasonal changes in littoral algal distribution patterns and environmental conditions were monitored. Additionally, Bangia was examined in culture to relate growth and reproductive mechanisms to distribution patterns and competitive potential.Ulothrix established extensive cover immediately below mean water level (MWL) after thaw in mid-April along moderately exposed shores. By early May, Bangia colonized from +0.5–1.0 m to just below MWL coexisting with Ulothrix near MWL. As water temperature neared l5° C in late May, the distribution of Cladophora expanded from patches at ?0.5–1.0 m upward into the Ulothrix band. In early June, Ulothrix sporulated, the remaining empty filaments disintegrated, and the alga was not observed for the remainder of the summer. Cladophora continued upward colonization, eventually dominating the substrate to the +0.5–1.0 m level. Bangia was dominant above the Cladophora zone, up to +2.0 m. During periods of moderate wave wash, Cladophora dominated up to the +1.0 m level. Heavy wave wash resulted in the detachment of Cladophora, allowing domination by Bangia from +0.5 m to 2.0 m.Bangia shares substrate with Ulothrix in the limited areas where their distributions overlap, and perhaps through greater desiccation tolerance can persist in areas where Cladophora does not grow. This partial niche segregation provides Bangia with refugia in time as well as space. Coexistence between Bangia and the competitively dominant Cladophora occurs as a result of partial niche segregation as well as the impact of wave action in initiating the detachment of Cladophora.     

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.     

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.     

Colonization and Growth of Attached Algae at the Lake Michigan Water Line

Changes in populations of attached algae growing near the water line along the west shore of Lake Michigan were observed. Emphasis was placed on harbor populations with particular focus on the algae of Milwaukee Harbor during 1979 and 1980. Vertical zonation on rocks and harbor structures is documented and seasonal growth patterns and selected interrelationships are described. Of the three major filamentous algal forms present in this habitat, Bangia appears as a permanent resident, while Ulothrix may be considered a seasonal and opportunistic plant, responding rapdily to changes in water level. Cladophora is a perennial, but plants above a depth of approximately 30 cm tend to grow as annuals due to conditions of ice scour. It is suggested that seasonal and spatial variation in the abundance of particular species is related to the physical growth requirements of the species.