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.     

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.     

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.     

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.     

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.     

Distribution of the Ponto-Caspian Amphipod Echinogammarus ischnus in the Great Lakes and Replacement of Native Gammarus fasciatus

The gammarid amphipod Echinogammarus ischnus was found to be widespread from the south end of Lake Huron, downstream in the St. Clair River and across Lake Erie to the Niagara River outlet into Lake Ontario. The presence of this exotic species was first reported in the Detroit River, where it now dominates; this species has been common in western Lake Erie since the summer of 1995. The species has replaced the native amphipod Gammarus fasciatus on rocky habitats in the St. Clair, Detroit, and Niagara rivers, and is the dominant amphipod on rocky shores in western Lake Erie. In one year, E. ischnus became the dominant amphipod at the Lake Ontario end of the Welland Canal, although the fecundity of E. ischnus is less than G. fasciatus. E. ischnus has not yet been reported from the north shore of Lake Ontario or the outlet into the St. Lawrence River but occurs 100 km further downstream at Prescott.     

Regional Differences in Size-at-age of the Recovering Burbot (Lota lota) Population in Lake Erie

The burbot Lota lota population in Lake Erie increased dramatically between 1995 and 2003, due mainly to control of the sea lamprey Petromyzon marinus, which began in the late 1980s. We estimated total length- and weight-at-age at capture for burbot caught in annual gillnet surveys of eastern Lake Erie during August 1994–2003. Mean total length was generally greater for burbot age 4–9 years that were caught in New York waters than in either Ontario or Pennsylvania waters of Lake Erie. Similarly, mean weight was greater for burbot at ages 4 through 6 years in New York waters than in either Ontario or Pennsylvania waters. Age-9 burbot caught in Ontario waters had greater mean weight and mean total length than did age-9 burbot caught in Pennsylvania waters. One possible explanation for greater length- and weight-at-age for New York burbot may be greater abundance of prey fishes, particularly rainbow smelt Osmerus mordax and round goby Neogobius melanostomus in New York waters. Total lengths at ages 4–10 years were generally greater for burbot caught in Lake Erie during 1994–2003 than those from published studies of other large lakes in North America that we considered, including for Lake Erie in 1946. The regional differences in size-at-age have important management ramifications, particularly because a commercial fishery targeting burbot has been considered for Ontario waters of Lake Erie.     

A novel method for tracking western Lake Erie Microcystis blooms, 2002–2011

After a period of improvement from the late 1970s through the mid 1990s, western Lake Erie has returned to eutrophic conditions and harmful algal blooms now dominated by the cyanobacterium Microcystis aeruginosa. The detection of long-term trends in Microcystis blooms would benefit from a convenient method for quantifying Microcystis using archived plankton tows. From 2002 to 2011, summer Microcystis blooms in western Lake Erie were quantified using plankton tows (N = 649). A flotation separation method was devised to quantify Microcystis biovolume in the tows, and the method was tested against whole water cell counts. Floating Microcystis biovolume (mL) in preserved tows was highly correlated with total Microcystis cells (R² = 0.84) and biomass (R² = 0.95) in whole water samples. We found that Microcystis annual biovolume was highly variable among years; the 2011 bloom was 2.4 times greater than the second largest bloom (2008) and 29.0 times greater than the smallest bloom (2002). Advantages of the method include use of archived samples, high sampling volume, and low effort and expense. Limitations include specificity for cyanobacterial blooms dominated by large Microcystis colonies and the need for site-specific validation. This study indicates that the flotation method can be used to rapidly assess past and present Microcystis in western Lake Erie and that there was high variability in the timing, duration, and intensity of the annual Microcystis blooms over a 10-year period. The data made possible by this method will aid further investigations into the underlying causal factors of blooms.     

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.     

Lipid Concentration and Size Variation of Bythotrephes (Cladocera: Cercopagidae) from Lakes Erie,Huron, and Michigan

Lipid concentrations of Bythotrephes cederstroemi were compared among three Great Lakes, Erie, Huron, and Michigan, in an effort to investigate the phenotypic plasticity in size displayed among the lakes. Four developmental stages were measured in Lakes Erie and Huron and two stages were studied in Lake Michigan. With a gravimetric extraction method, the total lipid concentration range (μg lipid μg dry weight⁻¹, expressed as percent) for Bythotrephes was estimated to be 10–19%. Statistically significant differences were found in lipid concentrations of Bythotrephes among lakes and developmental stages. Lake Erie had significantly higher lipid concentration values than Lake Huron for stages 2 through 4, and had similar values to Lake Michigan for the analyzed stages 1 and 4. The first instar had indistinguishable lipid concentrations among Lakes Erie, Huron,and Michigan. Even though animals from Lake Erie were significantly smaller, the data suggest that they were not less well nourished. We hypothesize that selective mortality imposed by visual predators on larger Bythotrephes and the lack of deep water refuges in Lake Erie has encouraged the smaller size of Bythotrephes found there in comparison to those found in Lakes Huron and Michigan.     

Detection and expression of genes for phosphorus metabolism in picocyanobacteria from the Laurentian Great Lakes

The pelagic regions of Lake Superior and eastern Lake Erie (Laurentian Great Lakes) are typically phosphorus (P)-limited environments, and picocyanobacteria of the genus Synechococcus spp. are prominent primary producers during the summer. As a proxy for their utilization of organic P, the expression of two genes, phnD and phoX, was monitored. The phnD gene encodes the phosphonate binding protein of the ABC-type phosphonate transporter, whereas the phoX gene encodes a calcium-dependent alkaline phosphatase. Furthermore, to assess the ability of freshwater Synechococcus spp. to substitute sulfolipids for phospholipids, sqdX gene (cyanobacterial sulfolipid synthase) expression was examined. We employed PCR primers to detect the presence of all three genes in the endemic Synechococcus spp., and RT-PCR assays of cultured freshwater strains and environmental samples to assess the degree of P-stress in the phytoplankton. We show that the phnD gene was constitutively expressed, suggesting that freshwater picocyanobacteria were metabolizing exogenous phosphonate compounds in Lakes Erie and Superior. By contrast, phoX was regulated by P bioavailability. We also provide evidence that sqdX is expressed during increased growth rates in phosphorus-replete conditions, suggesting that sulfolipid synthesis is not a P conservation mechanism for freshwater Synechococcus spp.     

Distribution of Freshwater Ostracodes in Lake Erie

Samples collected from the Lake Erie sediment-water interface in 1975 indicate the presence of few live shelled crustaceans (ostracodes). Of the 26 species identified only one, Candona caudata, can be considered as successful today in Lake Erie. Cytherissa lacustris and Candona subtriangulata, primarily recovered as empty shells in this study, indicate that these species have become extinct because of chemical and/or physical change sometime during the last 100 years in Lake Erie.     

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.     

Lakewide dominance does not predict the potential for spread of dreissenids

In recent years, quagga mussels (Dreissena rostriformis bugensis) have almost completely replaced zebra mussels (Dreissena polymorpha) in the Lower Great Lakes. As recreational boats are the main vector of spread for dreissenids in North America, this study examined whether lakes Erie and Ontario could still be sources for the spread of zebra mussels. In the summer–fall of 2010, the abundance of each species of Dreissena on 196 boats from 5 marinas in lakes Erie and Ontario was examined. Additional samples of Dreissena in 2010–2012 were collected in tributaries, bays, and in the upper littoral zones of these lakes. A total of 77 boats were fouled by Dreissena, and of those 61 were fouled by both species, 13 were fouled just by zebra mussels, and only 3 were fouled solely by quagga mussels. Although quagga mussels compose ~ 99% of dreissenids in eastern Lake Erie and in Lake Ontario, on boats at most marinas sampled, zebra mussels were usually more abundant and significantly larger than quagga mussels. Refugia for zebra mussels were found in bays, tributaries, and upper littoral zones with high wave activity. Thus, although quagga mussels are now more abundant than zebra mussels within the Lower Great Lakes, these waterbodies still have the potential to be a source for the spread of zebra mussels, and for some vectors, the propagule pressure from zebra mussels is likely greater than that from quagga mussels.     

Hydrology influences generalist–specialist bird-based indices of biotic integrity in Great Lakes coastal wetlands

Marsh bird habitats are influenced by water levels which may pose challenges for interpreting bird-based indices of wetland health. We determined how much fluctuating water levels and associated changes in emergent vegetation influence the Index of Marsh Bird Community Integrity (IMBCI) using data collected in Great Lakes coastal wetlands by participants in Bird Studies Canada's Great Lakes Marsh Monitoring Program. IMBCI scores for 90 wetlands in Lake Erie and 131 wetlands in Lake Ontario decreased with decreasing water levels due to decreasing number of marsh-dependent species in Lake Erie and perhaps also in Lake Ontario. The average magnitude of the decrease in scores between extremely high and low water periods for wetlands with sufficient data was 15% in Lake Erie where water dropped 0.9 m on average (n = 11 wetlands) and 18% in Lake Ontario where water dropped 0.5 m (n = 7). Scores in Lake Erie increased with increasing Typha due to increasing numbers of marsh-dependent species and decreased with increasing Phragmites due to increasing numbers of generalist species. The opposite was observed in Lake Ontario, perhaps due to denser Typha and sparser Phragmites. Scores were explained by the naturally fluctuating water levels of Lake Erie, which favored Phragmites expansion and the regulated water levels of Lake Ontario which promoted Typha expansion. Scores were influenced by fluctuating water levels and associated changes in emergent vegetation. Inter-annual water level fluctuations should be considered when interpreting any indicator of wetland health that is based on marsh-dependent bird species.     

Changes in the Crustacean Communities of the Central Basin of Lake Erie during the First Full Year of the Bythotrephes longimanus Invasion

The exotic predatory cladoceran Bythotrephes longimanus was first observed in Lake Erie on 19 September, 1985. During the early summer immediately prior to its appearance, the cladoceran community in the central basin of Lake Erie was characterized by large populations of Bosmina longirostris, Eubosmina coregoni, Daphnia mendotae, and Daphnia retrocurva, with the latter three species persisting throughout August and September, along with Diaphanosoma spp. Community composition during early summer 1986 was similar to that of the previous year, but densities of all cladocerans decreased dramatically coincident with the appearance of Bythotrephes in mid-July, and remained suppressed throughout August and September. Only D. mendotae was present in appreciable numbers during this period; from mid-July through the end of August, 86-98% of cladoceran biomass (exclusive of Bythotrephes and Leptodora) was contributed by D. mendotae. Densities of Leptodora and Bythotrephes showed a strikingly inverse relationship, both temporally and spatially, during 1986. Size frequency distributions of D. mendotae exhibited an immediate shift towards extremely large ( > 2.5 mm) individuals coincident with Bythotrephes’ appearance in 1986, suggesting an upper size limit to efficient prey utilization by Bythotrephes. These results suggest that Bythotrephes can have substantial impacts both on cladoceran community composition, and on the size distributions of individual species.     

Suitable habitat model for walleye (Sander vitreus) in Lake Erie: Implications for inter-jurisdictional harvest quota allocations

Models predicting habitat distributions can give insight into species–habitat requirements and anticipate how populations respond to environmental change. Despite the economic and ecological importance of walleye (Sander vitreus) in Lake Erie, no preferred-habitat model exists and the spatial extent of suitable habitat is poorly understood. Empirical species-habitat models for three groups of walleye (juveniles, adults, and all walleye) was developed using records from a long term gill net data base (21 years). We examined the degree to which habitat suitability varies with vertical stratum for each group and whether the new model yields different estimates of available walleye habitat when compared to the current depth-based approach. Walleye occurrence in gill nets was positively related to water temperature, negatively related to water depth and water clarity, and unrelated to dissolved oxygen concentration. A model that incorporated interaction terms among the independent variables performed better than the linear, quadratic, and cubic generalized linear models (GLMs) for all three groups. Our results indicate that the extent of suitable habitat varies spatially in Lake Erie and is greatest in the West basin. Weighted Habitat Suitability Areas (WHSA), a combination of habitat quality and quantity, differed significantly among basins and vertical strata in Lake Erie. The current quota allocation strategy for Lake Erie walleye is based on the proportional amount of preferred habitat by jurisdiction. However, the current depth-based definition of preferred habitat may not be an adequate representation of walleye suitable habitat shared by each jurisdiction.     

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.     

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

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

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.