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.     

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.     

Recovery of Burrowing Mayflies (Ephemeroptera: Ephemeridae: Hexagenia) in Western Lake Erie

Burrowing mayflies (Hexagenia spp.) are native to western Lake Erie and were abundant until the 1950s, when they disappeared due to degraded water and sediment quality. Nymphs were absent from the sediments of most of western Lake Erie after the 1950s, although small, widely disjunct populations apparently persisted near shore. Sediment samples collected in 1993 revealed several small populations near the western and southern shores and beyond the mouths of the Detroit and Maumee rivers. A larger population was found in the southern island area, but nymphs were absent in the middle of the basin. By 1995, nymphs had spread throughout the western half and eastern end of the basin but remained absent from the middle of the basin. These data indicate that Hexagenia began recolonizing nearshore areas before offshore areas. Increasingly large swarms of winged Hexagenia on shore and over the lake between 1992 and 1994 further indicate that mayflies are recolonizing the basin. Factors that have permitted Hexagenia recovery in western Lake Erie probably include improved sediment and water quality attributed to pollution abatement programs implemented after the early 1970s, and perhaps environmental changes in the early 1990s attributed to effects of the exotic zebra mussel (Dreissena polymorpha)     

A comparative examination of recent changes in nutrients and lower food web structure in Lake Michigan and Lake Huron

The lower food webs of Lake Huron and Lake Michigan have experienced similar reductions in the spring phytoplankton bloom and summer populations of Diporeia and cladocerans since the early 2000s. At the same time phosphorus concentrations have decreased and water clarity and silica concentrations have increased. Key periods of change, identified by using a method based on sequential t-tests, were 2003–2005 (Huron) and 2004–2006 (Michigan). Estimated filtration capacity suggests that dreissenid grazing would have been insufficient to directly impact phytoplankton in the deeper waters of either lake by this time (mid 2000s). Despite some evidence of decreased chlorophyll:TP ratios, consistent with grazing limitation of phytoplankton, the main impact of dreissenids on the offshore waters was probably remote, e.g., through interception of nutrients by nearshore populations. A mass balance model indicates that decreased phosphorus loading could not account for observed in-lake phosphorus declines. However, model-inferred internal phosphorus dynamics were strongly correlated between the lakes, with periods of increased internal loading in the 1990s, and increased phosphorus loss starting in 2000 in Lake Michigan and 2003 in Lake Huron, prior to dreissenid expansion into deep water of both lakes. This suggests a limited role for deep populations of dreissenids in the initial phosphorus declines in the lakes, and also suggests a role for meteorological influence on phosphorus dynamics. The high synchrony in lower trophic level changes between Lake Michigan and Lake Huron suggests that both lakes should be considered when investigating underlying causal factors of these changes.     

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.     

Distribution and Abundance of Burrowing Mayflies (Hexagenia spp.) in Lake Erie, 1997–2005

Burrowing mayflies (Hexagenia limbata and H. rigida) recolonized sediments of the western basin of Lake Erie in the 1990s following decades of pollution abatement. We predicted that Hexagenia would also disperse eastward or expand from existing localized populations and colonize large regions of the other basins. We sampled zoobenthos in parts of the western and central basins yearly from 1997–2005, along the north shore of the eastern basin in 2001–2002, and throughout the lake in 2004. In the island area of the western basin, Hexagenia was present at densities ≤ 1,278 nymphs/m² and exhibited higher densities in odd years than even years. By contrast, Hexagenia became more widespread in the central basin from 1997-2000 at densities ≤ 48 nymphs/m² but was mostly absent from 2001-2005. Nymphs were found along an eastern basin transect at densities≤ 382/m² in 2001 and 2002. During the 2004 lake-wide survey, Hexagenia was found at 63 of 89 stations situated throughout the western basin (≤ 1,636 nymphs/m², mean = 195 nymphs/m², SE = 32, N = 89) but at only 7 of 112 central basin stations, all near the western edge of the basin (≤ 708 nymphs/m²), and was not found in the eastern basin. Hexagenia was found at 2 of 62 stations (≤ 91 nymphs/m²) in harbors, marinas, and tributaries along the south shore of the central basin in 2005. Oxygen depletion at the sediment-water interface and cool temperatures in the hypolimnion are probably the primary factors preventing successful establishment throughout much of the central basin. Hexagenia can be a useful indicator of lake quality where its distribution and abundance are limited by anthropogenic causes.     

A hydrodynamic approach to modeling phosphorus distribution in Lake Erie

The purpose of this paper is to show how a high-resolution numerical circulation model of Lake Erie can be used to gain insight into the spatial and temporal variability of phosphorus (and by inference, other components of the lower food web) in the lake. The computer model simulates the detailed spatial and temporal distribution of total phosphorus in Lake Erie during 1994 based on tributary and atmospheric loading, hydrodynamic transport, and basin-dependent net apparent settling. Phosphorus loads to the lake in 1994 were relatively low, about 30% lower than the average loads for the past 30 years. Results of the model simulations are presented in terms of maps of 1) annually averaged phosphorus concentration, 2) temporal variability of phosphorus concentration, and 3) relative contribution of annual phosphorus load from specific tributaries. Model results illustrate that significant nearshore to offshore gradients occur in the vicinity of tributary mouths and their along-shore plumes. For instance, the annually averaged phosphorus concentration can vary by a factor of 10 from one end of the lake to the other. Phosphorus levels at some points in the lake can change by a factor of 10 in a matter of hours. Variance in phosphorus levels is up to 100 times higher near major tributary mouths than it is in offshore waters. The model is also used to estimate the spatial distribution of phosphorus variability and to produce maps of the relative contribution of individual tributaries to the annual average concentration at each point in the lake.     

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.     

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

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

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.     

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.     

Evidence for a Trophic Cascade Effect on North-shore Western Lake Erie Phytoplankton Prior to the Zebra Mussel Invasion

Increasing summer total phosphorus (TP) concentrations measured in samples from a municipal water intake off the north shore of western Lake Erie during 1976 to 1983 were inconsistent with TP loads to the western basin of Lake Erie and with phytoplankton densities in the intake samples, both of which declined over the same time interval. The long-term (1976 to 1988) summer TP data were inversely correlated (r = −0.858) with summer average maximum daily wind velocities, suggesting that low wind velocities contributed to anoxia at the sediment-water interface and high sediment TP release rates in summer. While TP loading reductions in the late 1960s and early 1970s likely contributed to phytoplankton declines, continued phytoplankton declines during the late 1970s to early 1980s could not have been caused by continued reductions in TP loadings while TP concentrations increased. The phytoplankton declines of the 1980s are more likely attributable to changes in the trophic cascade associated with dramatic declines in some species of zooplanktivorous fish during the 1970s and 1980s as a result of a restored walleye population. Long term phytoplankton densities were fit (R² = 0.902) to a multiple regression model with western Lake Erie TP loads and an index of zooplanktivore density as independent variables; the zooplanktivore component of this model was the most significant contributor to the prediction of phytoplankton density. The implications of these findings for maintenance of good lake water quality include the need to maintain strong piscivore populations as well as reduced phosphorus loads.     

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.     

Assessing and addressing the re-eutrophication of Lake Erie: Central basin hypoxia

Relieving phosphorus loading is a key management tool for controlling Lake Erie eutrophication. During the 1960s and 1970s, increased phosphorus inputs degraded water quality and reduced central basin hypolimnetic oxygen levels which, in turn, eliminated thermal habitat vital to cold-water organisms and contributed to the extirpation of important benthic macroinvertebrate prey species for fishes. In response to load reductions initiated in 1972, Lake Erie responded quickly with reduced water-column phosphorus concentrations, phytoplankton biomass, and bottom-water hypoxia (dissolved oxygen < 2 mg/l). Since the mid-1990s, cyanobacteria blooms increased and extensive hypoxia and benthic algae returned. We synthesize recent research leading to guidance for addressing this re-eutrophication, with particular emphasis on central basin hypoxia. We document recent trends in key eutrophication-related properties, assess their likely ecological impacts, and develop load response curves to guide revised hypoxia-based loading targets called for in the 2012 Great Lakes Water Quality Agreement. Reducing central basin hypoxic area to levels observed in the early 1990s (ca. 2000 km²) requires cutting total phosphorus loads by 46% from the 2003–2011 average or reducing dissolved reactive phosphorus loads by 78% from the 2005–2011 average. Reductions to these levels are also protective of fish habitat. We provide potential approaches for achieving those new loading targets, and suggest that recent load reduction recommendations focused on western basin cyanobacteria blooms may not be sufficient to reduce central basin hypoxia to 2000 km².     

The effects of dreissenid mussels on the survival and condition of burrowing mayflies (Hexagenia spp.) in western Lake Erie

Burrowing mayflies (Hexagenia limbata and H. rigida) are once again prominent members of the benthic community in western Lake Erie. However, this community is now dominated by dreissenid mussels. We conducted a laboratory experiment and field sampling to investigate whether survival and condition of Hexagenia were affected by the presence, density, and quality of dreissenid mussels. In a laboratory experiment, Hexagenia survival was higher in microcosms without dreissenid mussels. We also found Hexagenia density to be higher at field sites with low dreissenid density, suggesting that Hexagenia survival is higher in these areas as well. In microcosm treatments with low dreissenid density, Hexagenia survival was higher in treatments with live dreissenids than in treatments containing only dreissenid shells. These findings suggest that while dreissenid shells degrade the quality of soft sediments for Hexagenia, some of the negative effect is offset by the presence of live dreissenids. The positive effect of live dreissenids is likely due to additional food resources made available to Hexagenia by the filtering activity of dreissenids. Neither dreissenid density nor shell “type” (shells alone or live dreissenids in shells) had an effect on Hexagenia condition. Thus, the interactions between these dominant benthic invertebrates are complex. Recovery of Hexagenia populations in western Lake Erie is likely affected by both changing environmental conditions due to anthropogenic activities and the introduction of exotic species into the benthic community. The results are likely to be continued instability of the benthic food web and unpredictable consequences for human utilization of this ecosystem.     

Phosphorus retention and transformation in a dammed reservoir of the Thames River,Ontario: Impacts on phosphorus load and speciation

Extensive efforts are underway to reduce phosphorus (P) export from the Lake Erie watershed. On the Canadian side, the Thames River is the largest tributary source of P to Lake Erie’s western basin. However, the role of dams in retaining and modifying riverine P loading to the lake has not been comprehensively evaluated. We assessed whether Fanshawe Reservoir, the largest dam reservoir on the Thames River, acts as a source or sink of P, using year-round discharge and water chemistry data collected in 2018 and 2019. We also determined how in-reservoir processes alter P speciation by comparing the dissolved reactive P to total P ratio (DRP:TP) in upstream and downstream loads. Annually, Fanshawe Reservoir was a net sink for P, retaining 25% (36 tonnes) and 47% (91 tonnes) of TP in 2018 and 2019, respectively. Seasonally, the reservoir oscillated between a source and sink of P. Net P release occurred during the spring of 2018 and the summers of 2018 and 2019, driven by internal P loading and hypolimnetic discharge from the dam. The reservoir did not exert a strong influence on DRP:TP annually, but ratio increases occurred during both summers, concurrent with water column stratification. Our analysis demonstrates that Fanshawe Reservoir is not only an important P sink on the Thames River, but also modulates the timing and speciation of P loads. We therefore propose that the potential of using existing dam reservoirs to attenuate downstream P loads should be more thoroughly explored alongside source based P mitigation strategies.     

Quantification of internal phosphorus load in large,partially polymictic and mesotrophic Lake Simcoe,Ontario

Hypoxia and cyanobacteria still occur occasionally in large, mesotrophic Lake Simcoe, and total phosphorus (TP) concentration has remained relatively constant despite external nutrient load reduction. This may indicate a potential internal P source. Internal load as redox-dependent P release from bottom sediments is hard to determine in such a relatively shallow and mostly mixed lake. This study represents the first attempt to quantify internal P loading over many years for the three main sections of Lake Simcoe. Internal load was determined (a) as in situ estimate based on TP increases between July and October and (b) as gross estimate from the product of experimentally determined P release rates and hypoxic extent of sediment surfaces in space and time. Hypoxic extent was quantified (1) as the hypoxic factor determined from dissolved oxygen profiles below the level of 3.5 mg/L, and (2) as active sediment area release factor (AA) modeled from summer euphotic TP concentration, which is especially useful in the mixed sections. Annual internal load for the whole lake was determined as a near constant 62.2 metric tonnes/yr (86 mg/m²/yr) for 1980–2011 using the gross estimates of the AA approach and 88 t/yr before and 53 t/yr after external load abatement and zebra mussel invasion using in situ estimates. Means of in situ and AA-based estimates for 2000–2011 are in close agreement except for polymictic Cook's Bay. These estimates are 45 to 89% of external load, which suggests that internal loading is an important source of P in Lake Simcoe.     

DNA barcoding to confirm morphological traits and determine relative abundance of burrowing mayfly species in western Lake Erie

Burrowing mayfly species of the genus Hexagenia are well known indicators of environmental health in lakes and rivers. Two species, H. limbata and H. rigida, are indistinguishable as nymphs and as adult females. Our objectives were to develop a genetic technique to distinguish between the two species and identify morphological features that separate cryptic nymphs and adult females. Fifty nymphs were collected before emergence from 10 sites throughout the western basin of Lake Erie in 2004 and 2005. Using known specimens of adult aerial male H. limbata and H. rigida, we used the mitochondrial cytochrome c oxidase subunit 1 (COI) gene to identify a 16 base pair (bp) difference between species. DNA sequencing confirmed correct species identification based on differences in abdominal pigmentation patterns on adult female imagos in 19 of 20 cases; the lone exception was a female with very faint pigmentation. Pigmentation patterns between species were consistent on nymphs, subimagos and imagos of both sexes. Populations of both species are panmictic across the western basin of Lake Erie, but H. limbata is the numerically dominant species, representing 70 to 100% of nymphs at sites in both years. A separate lineage of H limbata was discovered in the samples. The ability to distinguish nymphs of the two species will aid in developing more sensitive ecosystem indicators.     

A 200 year chronology of burrowing mayflies (Hexagenia spp.) in Saginaw Bay

After an absence of 50 years, burrowing mayflies (Hexagenia spp.) colonized western Lake Erie which led to interest in whether this fauna can be used to measure recovery in nearshore waters throughout the Great Lakes. However, in many areas we do not know if mayflies were native/endemic and thus, whether recovery is a logical measure to assess progress of recovery. In the present study, we construct a chronologic record of relative abundance of burrowing mayflies in Saginaw Bay by the use of mayfly tusks and radionuclides in sediments (i.e., a paleoecologic record) and historic records of mayfly nymphs in the bay. These records reveal that mayflies: (1) were few before 1799, which indicates that nymphs were probably native/endemic in the bay, (2) increased between 1799 and 1807 and remained at relatively high levels between 1807 and 1965, probably in response to increased nutrient run-off from the watershed, (3) declined dramatically between 1965 and 1973, probably as a result of excessive eutrophication in the mid-1950s; and, (4) were few and highly variable between 1973 and 2001, probably as a result of low and unstable abundances of mayfly nymphs. Historic records verify that nymphs disappeared in the bay in the late-1950s to early-1960s which is in agreement with the paleoecologic record. Reoccurrence of low abundances of nymphs in the bay between 1991 and 2008 and comparison of chronologic records of nymphs in Saginaw Bay and western Lake Erie suggest that mayflies may return to Saginaw Bay in the early-21st century. Undoubtedly, watershed conservation and three decades of pollution abatement have set the stage for a recovery of burrowing mayflies in Saginaw Bay, and possibly in other areas of the Great Lakes.     

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.