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.     

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.     

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.     

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.     

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.     

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.     

A modeling study to determine the contribution of interbasin versus intrabasin phosphorus loads on the southwestern nearshore of Lake Ontario

Elevated phosphorus and nuisance algae such as Cladophora have been persistent environmental concerns in the coastal areas of Lake Ontario. Phosphorus is regarded as one of the drivers of nearshore Cladophora and the most likely mitigation that can be used to control levels of this nuisance algae in the lakes. The Niagara River, carrying the Lake Erie interbasin load, is the major contributor of the overall phosphorus load to Lake Ontario. Due to circulation patterns in the lake, this contribution is especially significant in the southwestern nearshore areas. Here we apply a mathematical model to provide insight into the relative contribution of the Niagara River versus loadings from local rivers (intrabasin loads) on the nearshore phosphorus concentrations in this region. We performed numerical experiments to determine to what extent the Niagara, Genesee and smaller local rivers impact the nearshore (<20 m depth) phosphorus concentrations. Our model results show that the Niagara River dominates the nearshore region between its discharge location and the Genesee River’s mouth, but the Genesee River strongly impacts the nearby Ontario Beach region in the very nearshore (<5 m depth). Smaller rivers have some impact close to their discharge locations. However, uncertainty with the Niagara River phosphorus load is the limiting factor in making any credible nearshore phosphorus predictions. Model accuracy is also impacted by insufficient short time scale phosphorus loads for all of the rivers, the dynamic nature of the lake circulation in shallow nearshore areas, and the simplified assumptions of the model.     

Historical changes and current status of crayfish diversity and distribution in the Laurentian Great Lakes

Despite increasing recognition of the importance of invertebrates, and specifically crayfish, to nearshore food webs in the Laurentian Great Lakes, past and present ecological studies in the Great Lakes have predominantly focused on fishes. Using data from many sources, we provide a summary of crayfish diversity and distribution throughout the Great Lakes from 1882 to 2008 for 1456 locations where crayfish have been surveyed. Sampling effort was greatest in Lake Michigan, followed by lakes Huron, Erie, Superior, and Ontario. A total of 13 crayfish species occur in the lakes, with Lake Erie having the greatest diversity (n = 11) and Lake Superior having the least (n = 5). Five crayfish species are non-native to one or more lakes. Because Orconectes rusticus was the most widely distributed non-native species and is associated with known negative impacts, we assessed its spread throughout the Great Lakes. Although O. rusticus has been found for over 100 years in Lake Erie, its spread there has been relatively slow compared to that in lakes Michigan and Huron, where it has spread most rapidly since the 1990s and 2000, respectively. O. rusticus has been found in both lakes Superior and Ontario for 22 and 37 years, respectively, and has expanded little in either lake. Our broad spatial and temporal assessment of crayfish diversity and distribution provides a baseline for future nearshore ecological studies, and for future management efforts to restore native crayfish and limit non-native introductions and their impact on food web interactions.     

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.     

Mapping and monitoring the extent of submerged aquatic vegetation in the Laurentian Great Lakes with multi-scale satellite remote sensing

A satellite-based algorithm intended to map submerged aquatic vegetation (SAV), which was mostly the nuisance algae Cladophora, for the Laurentian Great Lakes has been developed and successfully demonstrated in test areas in Lakes Michigan and Ontario. The new Submerged Aquatic Vegetation Mapping Algorithm (SAVMA) first utilizes deep water (opaque) radiance values to correct shallow water values (transparent) so that depth invariant reflectance values for all three visible Landsat bands of the lake bottom can be classified. Combinations of two bands are then used to generate a depth invariant bottom type index. The algorithm then maps the lake bottom into three types: sand, dense SAV, less dense SAV by thresh-holding the depth invariant reflectance values. The SAVMA also generates a biomass estimate by assigning an average dry weight obtained by field sampling to both the dense and less dense SAV areas identified by the algorithm.The algorithm performance was successfully evaluated on Lake Michigan at the Sleeping Bear Dunes National Lakeshore (SBDNL) using Cladophora locations provided by diver survey as well as from an independent National Park Service monitoring. The SAVMA correctly mapped Cladophora to an approximate accuracy of 85%, where the misclassification was a result of mixed pixels due to the resolution of the Landsat data and imprecise atmospheric corrections. The algorithm was also successfully evaluated in Lake Ontario near Pickering, ON. The SAVMA was then used to generate both short and long term time-series analyses of Cladophora extent at SBDNL.     

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.     

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.     

Phosphorus Nutrition of Two Potentially Competing Filamentous Algae, Cladophora Glomerata (L.) Kütz. and Stigeoclonium Tenue (Agardh) Kütz. from Lake Ontario

Cladophora glomerata and Stigeoclonium tenue were isolated from Lake Ontario. Batch culture growth experiments at 20° C and 175 μE/m²?sec continuous light show the maximum growth rate of Stigeoclonium (2.0 day^?1) to be about 2.5 times that of Cladophora (0.8 day^?1). The maximum phosphorus uptake rate for Stigeoclonium is 20 times greater than that for Cladophora and the half-saturation coefficient is larger by a factor of three. From growth experiments, the level of minimum cellular phosphorus of declining-phase cells is similar for both species but Stigeoclonium exhibits a greater luxury maximum level (in lag-phase cells) by a factor of over two. This, plus the fact that the growth rate of Stigeoclonium exceeds that of Cladophora at all levels of external and cellular phosphorus, allows Stigeoclonium to be more successful at phosphate-based competition. This is further confirmed by the facts that (compared to Stigeoclonium) the half-saturation growth constant is three times greater in Cladophora and the ratio of this constant to maximum growth rate is also eight times greater in Cladophora. It is hypothesized that the present-day dominance by Cladophora in the littoral region of the lower Great Lakes may not be determined by phosphate limitation but by certain competitive aspects of the life cycle, growth form, and physiology.     

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

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

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.     

Discovery of the Asiatic Clam,Corbicula Fluminea,in Lake Michigan

Ten live specimens of the Asiatic clam, Corbicula fluminea (Müller) were collected 22 November 1983 in southeastern Lake Michigan offshore from the J. H. Campbell Power Plant. The site is 7.6 m deep with a substrate of coarse sand and gravel. This represents the second record of Corbicula from the Laurentian Great Lakes and the first from Lake Michigan.     

Dynamics of fecal indicator bacteria,bacterial pathogen genes,and organic wastewater contaminants in the Little Calumet River–Portage Burns Waterway,Indiana

Little information exists on the co-occurrence of fecal indicator bacteria (FIB), bacterial pathogens, and organic wastewater-associated chemicals (OWCs) within Great Lakes tributaries. Fifteen watershed sites and one beach site adjacent to the Little Calumet River–Portage Burns Waterway (LCRPBW) on Lake Michigan were tested on four dates for pH, dissolved oxygen, specific conductance, chloride, color, ammonia- and nitrate-nitrogen, soluble phosphorus, sulfate, turbidity, and atrazine; for concentrations of FIB; and for genes indicating the presence of human-pathogenic enterococci (ENT) and of Shiga-toxin producing Escherichia coli (EC) from various animal sources. Nineteen samples were also tested for 60 OWCs. Half of the watershed samples met EC recreational water quality standards; none met ENT standards. Human-wastewater-associated OWC detections were correlated with human-influence indicators such as population/km², chloride concentrations, and the presence of WWTP effluents, but EC and ENT concentrations were not. Bacterial pathogen genes indicated rural human and several potential animal sources. OWCs of human or ecosystem health concern (musk fragrances AHTN and HHCB, alkylphenols, carbamazepine) and 3 bacterial pathogen genes were detected at the mouth of the LCRPBW, but no such OWCs and only 1 pathogen gene were detected at the beach. The LCRPBW has significant potential to deliver FIB, potential bacterial pathogens, and OWCs of human or ecosystem health concern to the nearshore of Lake Michigan, under conditions enhancing nearshore transport of the river plume. Nearshore mixing of lake and river water, and the lack of relationship between OWCs and FIB or pathogen genes, pose numerous challenges for watershed and nearshore assessment and remediation.