First Occurrence of Cercopagis pengoi in Lake Michigan

Adults and juveniles of the predaceous cladoceran Cercopagis pengoi, native to the Ponto- Caspian region, were found in nearshore waters of southern Lake Michigan during August and September 1999. This invasive cladoceran was previously reported from Lake Ontario, but this is the first report of C. pengoi from Lake Michigan. In Lake Michigan, densities of C. pengoi peaked at about 75/m3 on 22 September when water temperature was 16°C, and first and second instars comprised up to 90% of the samples. Of the third and fourth instar reproductive forms, parthenogenetic females were most abundant (up to 33% of all individuals sampled) and were present on all sampling dates. Males and females with resting eggs appeared in low abundance on the last two sampling dates perhaps in response to decreasing temperatures. The impact of this invader on the Lake Michigan food web is yet undetermined, but C. pengoi individuals were found in the stomachs of alewife on 22 September indicating that fishes will feed on C. pengoi.     

A cross-lake comparison of crustacean zooplankton communities in the Laurentian Great Lakes, 1997–2016

Spring and summer open-water crustacean zooplankton communities were examined across all five Laurentian Great Lakes from 1997 to 2016. Spring communities were dominated by calanoid (lakes Superior, Huron and Michigan) or cyclopoid (lakes Erie and Ontario) copepods. Volumetric biomass of summer communities increased along an assumed trophic gradient (Superior, Huron, Michigan, Ontario; eastern, central and western Erie), as did dominance by cyclopoids and cladocerans. Over the time series of the study, summer communities in lakes Michigan, Huron and Ontario shifted towards greater dominance by calanoids and greater similarity with Lake Superior. Trajectories of changes were different; however, reductions in cladocerans accounted for most of the change in lakes Michigan and Huron while reductions in cyclopoids and increases in Leptodiaptomus sicilis were behind the changes in Lake Ontario. Shifts in the predatory cladoceran community in Lake Ontario from Cercopagis pengoi to occasional dominance by Bythotrephes longimanus, a species much more vulnerable to planktivory, as well as the appearance of Daphnia mendotae in a daphnid community previously consisting almost exclusively of the smaller Daphnia retrocurva, suggest impacts of reduced vertebrate predation. In contrast, strong correlations between cladocerans and chlorophyll in lakes Michigan and Huron point to the possible importance of bottom-up forces in those lakes. Large interannual shifts in cladoceran community structure in the central and eastern basins of Lake Erie suggest intense but variable vertebrate predation pressure. The zooplankton communities of lakes Huron, Michigan and Ontario may be approaching a historic community structure represented by Lake Superior.     

Distribution,Fecundity, and Genetics of Cercopagis pengoi (Ostroumov) (Crustacea,Cladocera) in Lake Ontario

Two distinctive forms of cercopagids, first detected in 1998 and identified as Cercopagis pengoi and C. ossiani using taxonomic keys, were observed to co-occur in Lake Ontario. C. ossiani was the predominant form in western Lake Ontario in mid-June 1999 but was then replaced by C. pengoi-like animals over the rest of the season. Mitochondrial DNA analyses revealed that these forms were genetically identical at the ND5 gene and that they are morphologically distinctive forms of C. pengoi. In 1999, Cercopagis reached a maximum abundance of 1,759 individuals/m³ (average abundance = 281 individuals/ m³, average biomass = 5.2 mg/m³). In August, Cercopagis biomass was lowest at nearshore and embayment sites and highest at offshore sites. Body length of parthenogenetic females was lower at nearshore (1.16 mm) and embayment (1.19 mm) sites relative to offshore (1.32 mm) ones. Maximal clutch size of parthenogenetic females was 24 embryos per individual. Cercopagis has already spread to Lake Michigan and five Finger Lakes. Although waterfowl may disperse Cercopagis, these invasions likely resulted from human activities.     

Recent changes in the offshore crustacean zooplankton community of Lake Ontario

Crustacean zooplankton communities in the offshore of Lake Ontario have undergone substantial changes between 1997 and 2011. A shift was apparent in 2004 from an initial assemblage dominated by cyclopoid copepods (mostly Diacyclops thomasi), Daphnia retrocurva and bosminids, and with Cercopagis pengoi (which invaded the lake in 1998) the dominant predatory cladoceran, to one characterized by reduced numbers of cyclopoids, a more varied predatory cladoceran community, and increased biomass of calanoid copepods. These changes represented a shift from a community which has been relatively stable in the offshore of the lake for at least 40 years. A further change was seen in 2008, marked by increased biomass of Leptodiaptomus sicilis, Bythotrephes longimanus and Daphnia mendotae. Unlike the somewhat similar changes seen recently in the zooplankton communities of Lakes Huron and Michigan, the shifts in Lake Ontario have not been accompanied by a trend towards increased oligotrophy. A more likely explanation for the observed shifts in Lake Ontario is decreased vertebrate predation, likely due to alewife declines, and changes in the predatory invertebrate community.     

A New Biological Marker Layer in the Sediments of the Great Lakes: Bythotrephes cederstroemi (Schödler) Spines

The European cladoceran, Bythotrephes cederstroemi (Schödler), recently invaded the Laurentian Great Lakes. Based on recent zooplankton records, it most likely appeared first in 1984 in Lakes Ontario, Erie, and Huron, and in 1985 in Lake Michigan. It has yet to be reported from Lake Superior. This species is a relatively large-bodied predatory form that possesses a long, caudal, latterally barbed spine. B. cederstroemi spines and spine fragments were found in the upper fractions (predominantly 0–4 cm) of 35 sediment cores collected from seven areas of deposition in the eastern basin of Lake Erie. All remains were well preserved and easy to identify. Very few to 0 spines were found in core depths greater than 4 cm suggesting that the invasion of this species has resulted in a new, readily distinguishable time horizon marker.     

Origins of Rainbow Smelt in Lake Ontario

The first rainbow smelt (Osmerus mordax) to enter Lake Ontario were probably migrants from an anadromous strain introduced into New York's Finger Lakes. Since the upper Great Lakes were originally stocked with a landlocked strain from Green Lake, Maine, subsequent migration to Lake Ontario from Lake Erie makes Lake Ontario unique among the Great Lakes in probably having received introductions from two distinct populations.     

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.     

Monitoring Round Goby (Neogobius melanostomus) Population Expansion in Eastern and Northern Lake Michigan

In the Great Lakes basin, round gobies (Neogobius melanostomus) were first reported from the St. Clair River in 1990. Reported here are the details of range extension and establishment of round gobies in eastern Lake Michigan at Grand Haven, Michigan and in northern Green Bay at Escanaba, Michigan. Round gobies were first collected in survey samples at Grand Haven in 1997 (0.0 to 0.5 fish per trawl hour). Catch rates increased in 1998 (1.5 to 3.0 fish/h) and 1999 (38.0 to 69.0 fish/h). Individuals collected by trawling (N = 207) ranged in total length from 18 to 94 mm. Round gobies were first collected in survey (trawl) samples at Escanaba in 1998 (7.5 fish/h; total length range 43 to 58 mm). Round gobies were also collected during 1999 in trawls at Muskegon and in gill nets at Grand Haven and Saugatuck. No round gobies were collected at five other eastern Lake Michigan ports sampled during this same time period (1995 to 1999). Numerous adult round gobies were captured by anglers at Grand Haven and Escanaba, indicating that these populations had been established for some time prior to being detected in survey sampling. Additional populations were verified from angler reports at Charlevoix (1998) and Kipling (1999), Michigan. Round gobies collected by anglers were generally larger, and taken in areas that are difficult to sample using conventional inshore sampling gear. These results indicate that survey and angler samples are complementary, but that up-to-date angler reports of exotic species that bite readily (round gobies) or are otherwise collected by anglers (Bythotrephes cederstroemi, Cercopagis pengoi) are critical to tracking the spread of these organisms.     

Delivery of nutrients and seston from the Muskegon River Watershed to near shore Lake Michigan

Drowned river mouth lakes are major features of coastal Great Lakes habitats and may influence nutrient and organic matter contributions from watersheds to near shore coastal zones. In May through October 2003, we measured loads of nutrients, surficial sediment, and seston to track the delivery of riverine-derived materials from the lower Muskegon River Watershed (MRW) into the near shore area of southeast Lake Michigan. Nutrient flux data indicated that seasonal loads of 1800 metric tons (MT) of particulate organic carbon, 3400 MT of dissolved organic carbon, and 24 MT of total phosphorus were discharged from the lower Muskegon River, with approximately 33% of TP load and 53% of the POC load intercepted within the drowned river mouth terminus, Muskegon Lake. Carbon: phosphorus molar ratios of seston in Muskegon River (C:P = 187) and Muskegon Lake (C:P = 176) were lower than in Lake Michigan (C:P = 334), indicating phosphorus limitation of phytoplankton in near shore Lake Michigan. Isotopic signatures of seston collected in Muskegon Lake were depleted in δ¹³C (− 30.8 ± 1.6‰) relative to the isotope signatures of seston from Lake Michigan (− 26.2 ± 1.3‰) or the mouth of the Muskegon River (− 28.1 ± 0.5‰), likely due to the presence of biogenic methane in Muskegon Lake. Seston δ¹⁵N increased on a strong east-to-west gradient within Muskegon Lake, indicating significant microbial processing of nutrients. The extent of nutrient uptake in Muskegon Lake altered the chemical and isotopic characterization of seston flowing into Lake Michigan from Muskegon River.     

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.     

Associations between Breeding Marsh Bird Abundances and Great Lakes Hydrology

We used Great Lakes hydrologic data and bird monitoring data from the Great Lakes Marsh Monitoring Program from 1995–2002 to: 1) evaluate trends and patterns of annual change in May-July water levels for Lakes Ontario, Erie, and Huron-Michigan, 2) report on trends of relative abundance for birds breeding in Great Lakes coastal marshes, and 3) correlate basin-wide and lake-specific annual indices of bird abundance with Great Lakes water levels. From 1995–2002, average May, June, and July water levels in all lake basins showed some annual variation, but Lakes Erie and Huron-Michigan had identical annual fluctuation patterns and general water level declines. No trend was observed in Lake Ontario water levels over this period. Abundance for five of seven marsh birds in Lake Ontario wetlands showed no temporal trends, whereas abundance of black tern (Chlidonias niger) declined and that of swamp sparrow (Melospiza georgiana) increased from 1995–2002. In contrast, abundances of American coot (Fulica americana), black tern, common moorhen (Gallinula chloropus), least bittern (Ixobrychus exilis), marsh wren (Cistorthorus palustris), pied-billed grebe (Podilymbus podiceps), sora (Porzana carolina), swamp sparrow, and Virginia rail (Rallus limicola) declined within marshes at Lakes Erie and Huron/Michigan from 1995–2002. Annual abundances of several birds we examined showed positive correlations with annual lake level changes in non-regulated Lakes Erie and Huron/Michigan, whereas most birds we examined in Lake Ontario coastal wetlands were not correlated with suppressed water level changes of this lake. Overall, our results suggest that long-term changes and annual water level fluctuations are important abiotic factors affecting abundance of some marsh-dependent birds in Great Lakes coastal marshes. For this reason, wetland bird population monitoring initiatives should consider using methods in sampling protocols, or during data analyses, to account for temporal and spatial components of hydrologic variability that affect wetlands and their avifauna.     

Contaminant Residues in Tissues of Adult and Prefledged Herring Gulls from the Great Lakes in Relation to Diet in the Early 1990s

In the early 1990s, herring gulls (Larus argentatus) were collected in 15 breeding colonies throughout the Great Lakes basin and in two reference colonies on Lake Winnipeg and the Bay of Fundy. Organochlorine and metal concentrations, and stable isotope ratios (¹⁵N/¹⁴N and ¹³C/¹²C) were measured in their tissues, and we qualitatively assessed their diet. Breast muscle δ¹⁵N suggested that adults fed on planctivorous or insectivorous fish at six colonies, on piscivorous fish at four, and at a lower trophic level at the remaining three. The concentrations of Co, Ni, Al, Cr, Sn, Fe, and Pb in kidneys of adults suggested anthropogenic enrichment in the Great Lakes basin. Concentrations of contaminants were highest most often in tissues of gulls from Lake Ontario and northern Lake Michigan colonies. Concentrations of Pb in adults from Hamilton Harbour and the Detroit River, and of Se in adults from the southern Lake Huron colony, were similar to published toxicity thresholds. Tissue levels of Cd have increased, while those of Pb have decreased markedly since 1983. DDE, dieldrin, mirex, and ΣPCB concentrations in livers collected from nine of these colonies revealed declines of 16 to 87% at most locations since the early 1980s. 2,3,7,8-tetrachlorodibenzo-p-dioxin equivalents were highest in adults from the offshore colonies in western Lake Erie and northern Lake Michigan, where gulls feed on piscivorous fish, and were driven by non-ortho PCBs. There was evidence of an unusually high bioavailablity of organochlorines, especially dieldrin, near the northern Lake Michigan colony during the period of chick growth, and of an ongoing loading of mercury to eastern Lake Ontario. Tissues of adult gulls from colonies on Lakes Ontario, Erie, and Michigan best reflect local conditions whereas those from Lake Superior and northern Lake Huron reflect contaminants accumulated from time spent on the lower lakes.     

Diel Vertical Migration of Round Goby Larvae in the Great Lakes

One hypothesis for the transcontinental and intra-Great Lakes basin transfer of round gobies (Neogobius melanostomus) has been that round gobies were pumped into the ballast water of ships. During June 2005 in Lake Erie, we obtained evidence of a vertical migration of round goby larvae, when we collected 167 round goby larvae in surface ichthyoplankton net tows at night and zero during day. These results complemented similar findings from the Muskegon River estuary of Lake Michigan during 2003 and 2004, documenting diel vertical migration for the first time in larval round gobies. We suggest vertical migration behavior may have allowed larval round gobies to be transported to and within the Great Lakes via ballast water and dispersed in the Great Lakes via advection of 6.5–8.5-mm long larvae at the surface. Based on our results, if ballast water was only taken on near the surface during daylight hours from May through September when larval round gobies were present, it would have mitigated the spread of round gobies throughout the Great Lakes.     

Blueback Herring (Alosa aestivalis) in Lake Ontario: First Record,Entry Route,and Colonization Potential

Two juvenile blueback herring (Alosa aestivalis) were caught in Lake Ontario in October 1995, the first record of this anadromous marine clupeid in the Great Lakes. Blueback herring most likely gained entry to Lake Ontario via the Erie Barge Canal, a navigation canal that links the Mohawk-Hudson rivers, which drain to the Atlantic Ocean, to Oneida Lake, which drains to Lake Ontario through the Oneida-Oswego rivers. Blueback herring ascend the Hudson River to spawn and were first reported from the upper Mohawk River in 1978. They currently spawn in several of the upper Mohawk's tributaries, including one about 430 km from the ocean but only 25 km from Oneida Lake. They were first found in Oneida Lake in 1982 and, in fall 1994, large numbers of juvenile blueback herring were found moving down the Oswego River. In the southern United States, blueback herring established self-reproducing populations in several reservoirs, and thus they have the potential to colonize Lake Ontario. If blueback herring become established in Lake Ontario, they could spread to other Great Lakes and impede recovery of depressed populations of indigenous fishes, like lake herring (Coregonus artedi) and lake trout (Salvelinus namaycush), through competition with, or predation on, their larvae.     

Quantification of the Lake Trophic Typologies of Naumann (Surface Quality) and Thienemann (Oxygen) with Special Reference to the Great Lakes

Separate trophic scales and indices are developed for two of the most significant symptoms of eutrophication: surface water quality and hypolimnetic dissolved oxygen depletion. The scales are made comparable by expressing them in dimensionless form with a lower bound of zero and a mesotrophic range from 5 to 10. In this way, the two symptoms can be compared and their relative importance judged. This is done for the Great Lakes with the result that for both scales Lakes Superior, Huron, and Michigan are classified as oligotrophic. However, while central and eastern Lake Erie and Lake Ontario are classified as mesotrophic in terms of surface water quality, they range from eutrophic (central Lake Erie) to oligotrophic (Lake Ontario) on the oxygen scale. This is because, although these lakes are similar in surface water quality, their hypolimnion thicknesses range from approximately 4 m for central Erie to 70 m for Lake Ontario. Because of its shallowness, western Lake Erie does not have a persistent oxygen problem. In terms of surface quality it is classified as eutrophic.We have attempted to relate the two scales by correlating surface primary production and areal depletion rate. The results indicate that for lakes of similar primary production, areal oxygen depletion is directly proportional to hypolimnion thickness.     

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.     

Trends in Mysis diluviana abundance in the Great Lakes, 2006–2016

With the large Diporeia declines in lakes Michigan, Huron, and Ontario, there is concern that a similar decline of Mysis diluviana related to oligotrophication and increased fish predation may occur. Mysis density and biomass were assessed from 2006 to 2016 using samples collected by the Great Lakes National Program Office's biomonitoring program in April and August in all five Great Lakes. Summer densities and biomasses were generally greater than spring values and both increased with bottom depth. There were no significant time trends during these 10–11 years in lakes Ontario, Michigan, or Huron, but there was a significant increase in Lake Superior. Density and biomass were highest in lakes Ontario and Superior, somewhat lower in Lake Michigan, and substantially lower in Lake Huron. A few Mysis were collected in eastern Lake Erie, indicating a small population in the deep basin of that lake. On average, mysids contributed 12–18% (spring-summer, Michigan), 18–14% (spring-summer, Superior), 30–13% (spring-summer, Ontario), and 3% (Huron) of the total open-water crustacean biomass. Size distributions consisted of two peaks, indicating a 2-year life cycle in all four of the deep lakes. Mysis were larger in Lake Ontario than in lakes Michigan, Superior, and Huron. Comparisons with available historic data indicated that mysid densities were higher in the 1960s–1990s (5 times higher in Huron, 2 times higher in Ontario, and around 40% higher in Michigan and Superior) than in 2006–2016.     

European Valve Snail Valvata piscinalis (Müller) in the Laurentian Great Lakes Basin

Previously reported from the lower Great Lakes basin and St. Lawrence and Hudson rivers, the nonindigenous gastropod Valvata piscinalis was found for the first time in Superior Bay (Minnesota) of Lake Superior, Lake Michigan (Wisconsin), and Oneida Lake (New York) of the Lake Ontario basin. This snail was not abundant in Lakes Superior and Michigan, whereas in eutrophic Oneida Lake it reached a maximum density of 1,690 individuals/m² (mean density = 216 individuals/m²). Human-mediated disturbances could facilitate the range extension of this snail by providing dispersal opportunities (e.g., canals, shipping traffic) or increasing nutrients (e.g., eutrophication). A native of the Palaearctic region, V. piscinalis has colonized sites across the Great Lakes basin, suggesting that it will likely become common in disturbed Great Lakes areas.     

Mean Circulation in the Great Lakes

In this paper new maps are presented of mean circulation in the Great Lakes, employing long-term current observations from about 100 Great Lakes moorings during the 1960s to 1980s. Knowledge of the mean circulation in the Great Lakes is important for ecological and management issues because it provides an indication of transport pathways of nutrients and contaminants on longer time scales. Based on the availability of data, summer circulation patterns in all of the Great Lakes, winter circulation patterns in all of the Great Lakes except Lake Superior, and annual circulation patterns in Lakes Erie, Michigan, and Ontario were derived. Winter currents are generally stronger than summer currents, and, therefore, annual circulation closely resembles winter circulation. Circulation patterns tend to be cyclonic (counterclockwise) in the larger lakes (Lake Huron, Lake Michigan, and Lake Superior) with increased cyclonic circulation in winter. In the smaller lakes (Lake Erie and Lake Ontario), winter circulation is characterized by a two-gyre circulation pattern. Summer circulation in the smaller lakes is different; predominantly cyclonic in Lake Ontario and anticyclonic in Lake Erie.     

Predicting the potential distribution of the non-native Red Swamp Crayfish Procambarus clarkii in the Laurentian Great Lakes

The ongoing threat of introduction of invasive species, including crayfish, to the Laurentian Great Lakes has motivated the development of predictive models to inform where these invaders are likely to establish. Our study is among the first to apply regional freshwater-specific GIS layers to species occurrence data to predict ecosystem suitability to invasions, specifically for the red swamp crayfish, Procambarus clarkii, in the Great Lakes. We combined a database of crayfish species occurrences with the Great Lakes Aquatic Habitat Framework (GLAHF) GIS layers to model habitats suitable to invasion by P. clarkii using boosted regression trees and physiological information for this species. We developed a model of all suitable crayfish habitat across the Great Lakes, then constrained this habitat to areas anticipated to be suitable for P. clarkii based on known physiological limitations of this species. Specifically, P. clarkii requires a minimum temperature of 15?°C for copulation and oviposition, with peak reproduction occurring at temperatures of 20–23?°C. We identified 2% of the Great Lakes as suitable for P. clarkii establishment and 0.88% as optimal for this crayfish, primarily located on the southern coastlines of lakes Michigan and Erie and shallow bays including Saginaw Bay (Lake Huron), Green Bay (Lake Michigan), and Henderson Bay (Lake Ontario). These predictions of where P. clarkii is likely to establish populations can be used to identify areas where education, outreach, compliance, and law enforcement efforts should seek to prevent new introductions of this crayfish and help prioritize locations for surveillance to detect newly established populations.