Freshwater mussel community response to warm water discharge in western Lake Erie

Native unionid mussels are endangered in the Laurentian Great Lakes due to habitat degradation and biofouling by invasive dreissenids. However, a robust community was discovered living within the thermal discharge of a power plant at Oregon, Ohio, on the south shore of Lake Erie. Our study compared this community to nearby communities outside the thermal plume, and examined habitat characteristics that may affect unionids. Unionids were sampled from the exposed lake bed at three sites during a seiche in 2011: (1) within the thermal plume, (2) at Bayshore Park (2.0 km east of the plant), and (3) at the University of Toledo's Lake Erie Center (4.0 km east). In 2010, sediment samples were collected along a 2 km transect extending east from the plant discharge roughly parallel to the south shore of Lake Erie. Results indicated that the community within the thermal plume had higher densities, higher diversity (H′), more small individuals but overall larger sizes than communities outside the plume. Both the rate and intensity of fouling by dreissenids were lower within the plume. Dry mass of coarse surface sediment and sediment organic matter content were negatively related to distance from the plant (R² = 0.497, and 0.479, respectively). An unexpected discovery was that the bulk of the coarse sediment was comprised of shell material from Asian clams and dreissenid mussels, suggesting a contribution of these exotic species to sediment accumulation. In total, our results suggest that several habitat characteristics close to the power plant are favorable to unionids.     

Lyngbya wollei in western Lake Erie

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

Glyphosate influence on phytoplankton community structure in Lake Erie

In this study we investigated the effect of the phosphonate herbicide glyphosate (N-(phosphonomethyl)glycine) on the phytoplankton community structure in Lake Erie using lake water incubations, laboratory growth experiments and phylogenetic analysis of phosphonate metabolism genes. In microcosms, addition of glyphosate to Sandusky Bay water resulted in a significant increase in phytoplankton abundance, specifically causing an increase in the abundance of Planktothrix spp. In microcosms using Maumee Bay water, glyphosate did not stimulate phytoplankton growth but caused a decrease in Microcystis spp. abundance. The difference in the ability of Planktothrix spp. and Microcystis spp. to grow in the presence of glyphosate was confirmed in laboratory growth experiments. Further, an examination of the molecular pathways involved in phosphonate metabolism demonstrated that heterotrophic bacteria may be critical in allowing this proliferation. The results indicate that glyphosate has both positive and negative influences on phytoplankton community structure, serving as a nutrient source to microbes able to tolerate the herbicidal effects of the compound while killing those less tolerant. Moreover, this work highlights that in natural environments microorganisms function as communities, and the metabolic abilities of individual species are often less important than the collective ability of the community.     

From River to Lake: Phosphorus partitioning and algal community compositional changes in Western Lake Erie

The Maumee River is an important source of phosphorus (P) loading to western Lake Erie and potentially a source of Microcystis seed colonies contributing to the development of harmful algal blooms in the lake. Herein, we quantified P forms and size fractions, and phytoplankton community composition in the river–lake coupled ecosystem before (June), during (August), and after (September) a large Microcystis bloom in 2009. Additionally, we determined the distribution and density of a newly emergent cyanobacterium, Lyngbya wollei, near Maumee Bay to estimate potential P sequestration. In June, dissolved organic phosphorus (DOP) was the most abundant P form whereas particulate P (partP) was most abundant in August and September. Green algae dominated in June (44% and 60% of total chlorophyll in river and lake, respectively) with substantial Microcystis (17%) present only in the river. Conversely, in August, Microcystis declined in the river (3%) but dominated (32%) the lake. Lake phytoplankton sequestered < 6% of water column P even during peak Microcystis blooms; in all lake samples < 112 μm non-algal particles dominated partP. Lyngbya density averaged 19.4 g dry wt/m², with average Lyngbya P content of 15% (to 75% maximum) of water column P. The presence of Microcystis in the river before appearing in the lake indicates that the river is a potential source of Microcystis seed colonies for later lake blooms, that DOP is an important component of early summer total P, and that L. wollei blooms have the potential to increase P retention in nearshore areas.     

Heavy metals in sediments and uptake by burrowing mayflies in western Lake Erie basin

During the past two decades, burrowing Hexagenia mayflies have returned to the western basin of Lake Erie. Because of their importance as a prey resource for higher trophic levels and their extensive residence time in potentially contaminated sediment, Hexagenia may be a source of heavy metal transfer. To better understand the distribution and transfer of heavy metals in sediment and mayflies, sediment and mayfly nymphs were collected from 24 locations across the western basin of Lake Erie in May 2007. Following USEPA protocols, samples were analyzed for 16 elements using ICP-OES or ICP-MS. Metal concentrations in the sediments exceeded the Threshold Effect Level for at least one metal at all sample sites. Sediment heavy metal distribution profiles indicate metal concentrations are correlated with organic matter content, and the highest heavy metal concentrations were found in the central deeper region of the western basin where organic content in the sediments was greatest. Hexagenia were distributed throughout the western basin, with greatest density (1350/m²) within the Detroit River plume. The Cd and Zn levels in mayflies were on average approximately 4 and 2 times greater, respectively, than sediment levels, and the Cd concentrations in the sediments exceeded the Threshold Effect Level at 27 of 28 sites and exceeded the Probable Effect Level at 9 of 28 sites. Spatial representation of heavy metal concentrations in mayflies exhibited a similar pattern to the spatial distribution of heavy metals and organic matter in the sediments with higher concentrations of metals found in mayflies residing in the central deeper region of the western basin.     

Assessment of Microcystis growth rate potential and nutrient status across a trophic gradient in western Lake Erie

Plankton tow samples collected from 2002 through 2009 indicate that Microcystis biovolume in western Lake Erie is often most dense in transition zone (TZ) waters between Maumee Bay and the center of the western basin. TZ waters are generally high in nutrients and turbidity, and concentrations of each decrease with distance from Maumee Bay. High Microcystis biovolume in the TZ suggests the possibility that the conditions in these waters support a greater Microcystis growth rate relative to the open lake. To test this hypothesis, during the 2008 bloom, Microcystis was collected from western Lake Erie for measurements of total protein content (TPC) as an indicator of growth rate potential and cellular nutrient content to indicate nutrient deficiencies. TPC results indicate that Microcystis in the TZ had a higher potential growth rate compared to offshore waters. TPC values in Maumee Bay were intermediate but not significantly different from the TZ and offshore. Nitrogen content of Microcystis remained high over the summer at all sites, despite very low dissolved nitrate concentrations and low total nitrogen-to-total phosphorus ratio in late summer in the lake. Ammonium level in the lake was constant during the summer, and likely provided the nitrogen source for Microcystis. Cellular phosphorus content varied between site and sample date suggesting that Microcystis was moderately phosphorus deficient. Quotas of micronutrient indicated that Microcystis was not deficient of micronutrients. Results of this study suggest the waters in and adjacent to Maumee Bay provide more favorable growth conditions for Microcystis than offshore waters.     

Variability of sedimentary phosphorus fractions in the western and Sandusky basins of Lake Erie

Surface sediments and three sediment cores from the western basin and one sediment core from the Sandusky basin were analyzed to document spatial and temporal changes in five phosphorus fractions and total phosphorus (TP). The areal distributions of the bioavailable fractions NaCl-Pi, NaBD-Pi, and NaOH-Pi and the refractory organic fraction Res-P were broadly consistent and contrasted with those of the detrital fraction HCl-Pi which showed that high concentrations occurred mostly in high-energy littoral zones and low concentrations largely in profundal depositional areas. The contrasting distributions were induced by interactions among tributary inputs, wave action, circulation, and biogeochemical cycling and transfer in the basin. As revealed by the Sandusky basin sediment record, the detrital fraction HCl-Pi was dominant (70% of TP) during European settlement and decreased rapidly by 28.0% in the early 1910s due largely to impoundments of the Maumee and Sandusky Rivers. While HCl-Pi has ever since remained relatively constant, NaCl-Pi, NaBD-Pi, and NaOH-Pi increased significantly between 1950 and 1970 in the two basins. However, the post-regulation sediment records differed considerably among these coring sites. There was a marked increase of TP in two cores, corresponding to recent return of eutrophication and massive harmful algal blooms but contrasting with a relatively constant, low loading into the lake. This signified the role of internal loading as derived partialy from legacy pollution. Furthermore, NaCl-Pi has increased progressively throughout all the records. We conclude that the increased levels of NaCl-Pi in surface sediments may have altered the internal loading and contributed to the resurgences of harmful algal blooms in Lake Erie.     

Status,causes and controls of cyanobacterial blooms in Lake Erie

The Laurentian Great Lakes are among the most prominent sources of fresh water in the world. Lake Erie's infamous cyanobacterial blooms have, however, threatened the health of this valuable freshwater resource for decades. Toxic blooms dominated by the cyanobacterium Microcystis aeruginosa have most recently been one of primary ecological concerns for the lake. These toxic blooms impact the availability of potable water, as well as public health and revenues from the tourism and fishery industries. The socioeconomic effects of these blooms have spurred research efforts to pinpoint factors that drive bloom events. Despite decades of research and mitigation efforts, these blooms have expanded both in size and duration in recent years. However, through continued joint efforts between the Canadian and United States governments, scientists, and environmental managers, identification of the factors that drive bloom events is within reach. This review provides a summary of historical and contemporary research efforts in the realm of Lake Erie's harmful cyanobacterial blooms, both in terms of experimental and management achievements and insufficiencies, as well as future directions on the horizon for the lake's research community.     

Diatoms abound in ice-covered Lake Erie: An investigation of offshore winter limnology in Lake Erie over the period 2007 to 2010

The limnology of offshore Lake Erie during periods of extensive (> 70%) ice cover was examined from ship borne sampling efforts in 2007 to 2010, inclusive. Dense and discrete accumulations of the centric filamentous diatom Aulacoseria islandica (> 10 μg Chl-a/L) were located in the isothermal (< 1 °C) water column directly below the ice and only detectable in the ship wake; viable phytoplankton were also observed within ice. Evidence from these surveys supports the notions that winter blooms of diatoms occur annually prior to the onset of ice cover and that the phytoplankton from these blooms are maintained in the surface waters of Lake Erie and reduce silicate concentrations in the lake prior to spring. The mechanisms by which high phytoplankton biomass rise at this time of year requires further investigation, but these winter blooms probably have consequences for summer hypoxia and how the lake responds to climate change.     

Spatial and temporal variation in the distribution of burrowing mayfly nymphs (Ephemeroptera: Hexagenia limbata and H. rigida) in western Lake Erie

In the early 1990s, burrowing mayfly species reappeared in sediments of the western basin of Lake Erie after an absence of over 30 years due to episodic hypoxia at the sediment–water interface. Long-term monitoring of adult mayflies at shoreline areas had revealed that Hexagenia rigida was more abundant than Hexagenia limbata during the initial recolonization period, but was gradually replaced by H. limbata. We hypothesized that this shift in dominance would be confirmed by the distribution and abundance of nymphs. We identified nymphs collected each spring throughout western Lake Erie from 1997 to 2004. The relative abundances of H. rigida and H. limbata nymphs exhibited the same temporal sequence as adults. Furthermore, the number of sites in the western basin in which H. rigida occurred decreased as the occurrence frequency of H. limbata increased. H. limbata were dominant in the basin by 2004. Hexagenia limbata nymphs persisted in the center-most part of the basin, whereas H. rigida did not, possibly due to differences in tolerance to hypoxia. There were no significant differences in body size between the two populations. Differences in dispersal distance from source populations and the timing and success of egg hatching likely accounted for the initial colonizing success of H. rigida, but the differential ability of H. limbata eggs to overwinter in sediments and possible tolerance of nymphs to hypoxia has possibly led to its current dominance in the western basin.     

The effect of mayfly (Hexagenia spp.) burrowing activity on sediment oxygen demand in western Lake Erie

Previous studies support the hypothesis that large numbers of infaunal burrow-irrigating organisms in the western basin of Lake Erie may increase significantly the sediment oxygen demand, thus enhancing the rate of hypolimnetic oxygen depletion. We conducted laboratory experiments to quantify burrow oxygen dynamics and increased oxygen demand resulting from burrow irrigation using two different year classes of Hexagenia spp. nymphs from western Lake Erie during summer, 2006. Using oxygen microelectrodes and hot film anemometry, we simultaneously determined oxygen concentrations and burrow water flow velocities. Burrow oxygen depletion rates ranged from 21.7 mg/nymph/mo for 15 mm nymphs at 23 °C to 240.7 mg/nymph/mo for 23 mm nymphs at 13 °C. Sealed microcosm experiments demonstrated that mayflies increase the rate of oxygen depletion by 2–5 times that of controls, depending on size of nymph and water temperature, with colder waters having greater impact. At natural population densities, nymph pumping activity increased total sediment oxygen demand 0.3–2.5 times compared to sediments with no mayflies and accounted for 22–71% of the total sediment oxygen demand. Extrapolating laboratory results to the natural system suggest that Hexagenia spp. populations may exert a significant control on oxygen depletion during intermittent stratification. This finding may help explain some of the fluctuations in Hexagenia spp. population densities in western Lake Erie and suggests that mayflies, by causing their own population collapse irrespective of other environmental conditions, may need longer term averages when used as a bio-indicator of the success of pollution-abatement programs in western Lake Erie and possibly throughout the Great Lakes.     

Distribution and abundance of freshwater polychaetes, Manayunkia speciosa (Polychaeta), in the Great Lakes with a 70-year case history for western Lake Erie

Manayunkia speciosa has been a taxonomic curiosity for 150 years with little interest until 1977 when it was identified as an intermediate host of a fish parasite (Ceratomyxa shasta) responsible for fish mortalities (e.g., chinook salmon). Manayunkia was first reported in the Great Lakes in 1929. Since its discovery, the taxon has been reported in 50% (20 of 40 studies) of benthos studies published between 1960 and 2007. When found, Manayunkia comprised < 1% of benthos in 70% of examined studies. In one extensive study, Manayunkia occurred in only 26% of 378 sampled events (1991–2009). The taxon was found at higher densities in one area of Lake Erie (mean = 3658/m²) and Georgian Bay (1790/m²) than in five other areas (mean = 60 to 553/m²) of the lakes. A 70-year history of Manayunkia in western Lake Erie indicates it was not found in 1930, was most abundant in 1961 (mean = 8039, maximum = 67,748/m²), and decreased in successive periods of 1982 (3529, 49,639/m²), 1993 (1876, 25,332/m²), and 2003 (79, 2583/m²). It occurred at 48% of stations in 1961, 58% in 1982, 52% in 1993, and 6% of stations in 2003. In all years, Manayunkia was distributed primarily near the mouth of the Detroit River. Causes for declines in distribution and abundance are unknown, but may be related to pollution-abatement programs that began in the 1970s, and invasion of dreissenid mussels in the late-1980s which contributed to de-eutrophication of western Lake Erie. At present, importance of the long-term decline of Manayunkia in Lake Erie is unknown.     

Assessing the application of SeaWiFS ocean color algorithms to Lake Erie

The feasibility of satellite-based monitoring of phytoplankton chlorophyll a concentrations in Lake Erie is assessed by applying globally calibrated, ocean-derived color algorithms to spatially and temporally collocated measurements of SeaWiFS remote sensing reflectance. Satellite-based chlorophyll a retrievals were compared with fluorescence-based measurements of chlorophyll a from 68 field samples collected across the lake between 1998 and 2002. Twelve ocean-derived color algorithms, one regional algorithm derived for the Baltic Sea's Case 2 waters, and a set of regional algorithms developed for the western, central and eastern basins of Lake Erie were considered. While none of the ocean-derived algorithms performed adequately, the outlook for the success of regionally calibrated and validated algorithms, with forms similar to the ocean-derived algorithms, is promising over the eastern basin and possibly the central basin of the lake. In the western basin, each of the regional algorithms considered performed poorly, indicating that alternative approaches to algorithm development, or to satellite data screening and analysis procedures will be needed.     

Cylindrospermopsis in Lake Erie: Testing its Association with Other Cyanobacterial Genera and Major Limnological Parameters

We report the first documented observation of the potentially toxic cyanobacterium Cylindrospermopsis in Lake Erie and Sandusky Bay in 2005 (0.043–1.326 mg L^-1 wet weight, 16–1,942 trichomes mL^-1) and quantify the physical and chemical parameters and the cyanobacterial community composition contemporaneous to its occurrence. We hypothesize that the high temperature, low light intensity, and high nutrient content of Sandusky Bay, a shallow, drowned river mouth along the southwestern shore of Lake Erie, provides an ideal habitat for Cylindrospermopsis. This is consistent with published laboratory and field studies that show these physical and chemical conditions facilitate Cylindrospermopsis growth. Using multivariate statistics, we found that Cylindrospermopsis biomass correlated with high temperatures and shallow depths, conditions often found in Sandusky Bay. Light climate and nutrient concentrations were not associated with Cylindrospermopsis biomass, most likely because the light climate did not systematically change during the season and because nutrients exceeded demand. We propose that Cylindrospermopsis will increase in importance in Lake Erie, as previous research on climate change in the Great Lakes region predicts future higher water temperatures and lower water levels.     

Observations of Sediment Transport in Lake Erie during the Winter of 2004–2005

Time series measurements of current velocity, wave action, and water transparency were made at two sites—one in 24 m of water and the other in 53 m—in Lake Erie during the fall and winter of 2004–2005. The observations at the shallow site show that bottom resuspension occurred several times during the deployment. Although local resuspension did not occur at the deeper station, several advection episodes were observed. The storms during the observation period were not unusually large, so the processes observed are probably typical of those that occur on a yearly basis. The observations agree reasonably well with previous estimates for both the bottom shear stress during storms, and for the critical shear stress needed to resuspend bottom sediment, but previous estimates of the particle settling velocity are probably too low, while previous estimates of the sediment entrainment rate are too high. The results show that bottom material in the central basin is reworked numerous times before it is finally buried. Deposition in the eastern basin is a more continuous process, but the events observed were not sufficient to match the long-term accumulation rate, so deposition at this site is probably also due in part to larger, more infrequent storms.     

Spatial-temporal patterns of recolonizing adult mayflies in Lake Erie after a major disturbance

This 12-year study of Hexagenia male imagos documents the recovery of two species of burrowing mayflies, Hexagenia limbata and Hexagenia rigida in western Lake Erie after a 30-year absence due to hypoxia, resulting from cultural eutrophication. Annual adult mayfly collections were made at night during the peak emergence period at four sites along the north shore of the western basin of Lake Erie, 1997 to 2008. H. rigida, the dominant species in upstream riverine waterbodies, was the early colonizer, representing about 90% of all male imagos sampled in 1997. In 2000, when the two species were co-dominant, both inland aerial dispersal (5.5 km) and lakeward (0.25 to 4 km) oviposition patterns confirmed species co-existence. Twice weekly collections throughout the extended emergence period at one site confirmed that H. rigida was the dominant species in 1997, H. limbata and H. rigida were co-dominant in 2000, and H. limbata was dominant in 2002. Once H. limbata became the dominant species (> 90%) in 2000 to 2002 (depending on the site), it remained so. Both species followed a similar inland dispersal pattern, decreasing in density with increasing distance from shore; most mayflies were present within 1 km from shore. There was no significant difference in mean egg density of the two species among the sites extending lakeward in 2000 when the two species were equally abundant. The transition from the dominance of H. rigida to H. limbata may have resulted from several factors, including differential competition and growth between species or predation effects.     

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.     

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.     

The Invasive New Zealand Mud Snail (Potamopyrgus antipodarum) in Lake Erie

The New Zealand mud snail (Potamopyrgus antipodarum) is an invasive species in Europe, Japan, Australia, and North America. In the western United States it is a species of special concern where population densities in some rivers and streams are very large (∼300,000 per m²) and considerable ecological effects of its presence have been reported. Much less about the effects of this species is known in the Great Lakes, where the snail was found in Lake Ontario and the St. Lawrence River in 1991. Here we report the occurrence of the snail in Lake Erie. Two P. antipodarum were collected in 18 m deep water (sampling range 5–18 m) in Lake Erie off shore of Presque Isle State Park near Erie, Pennsylvania in the summer of 2005 and others were collected off of Sturgeon Point in Lake Erie (sampling range 5–20 m) south of Buffalo, NY and in the central basin of Lake Erie (18 m) in 2006. This finding demonstrates that this species continues to expand its range in the Great Lakes. The range expansion increases the likelihood that it may become established in rivers and streams emptying into the Great Lakes where higher densities and greater ecological damage may result.