Food of Deepwater Sculpin,Myoxocephalus Thompsoni,from Southeastern Lake Michigan

Pontoporeia hoyi was the major food consumed by most sizes of deepwater sculpins (Myoxocephalus thompsoni) during spring and summer, 1981–82, at a 100-m station in southeastern Lake Michigan. There was no apparent seasonality in choices of food or volumes consumed. Mysis relicta made up less than 9% of the total food volume and averaged second to Pontoporeia volumetrically—a result which differed from previous investigations for Lake Michigan. Reduced consumption of Mysis in our study compared to previous work may be a function of lower prey density or station depth; mysids may have been farther above the lake bottom than at previously studied, more shallow stations. Ostracods were important in 6- and 7-cm fish diets. Mysis, chironomids, and fish eggs were only found in sculpins larger than 8 cm. Chironomid larvae, which typically burrow into sediments, and copepod and cladoceran zooplankton constituted a small percentage of the diet of all sculpins. These results indicated that the deepwater sculpins examined were primarily epibenthic predators at the sediment-water interface and apparently did not burrow for food or feed higher up in the water column.     

Comparing seasonal dynamics of the Lake Huron zooplankton community between 1983–1984 and 2007 and revisiting the impact of Bythotrephes planktivory

Zooplankton community composition can be influenced by lake productivity as well as planktivory by fish or invertebrates. Previous analyses based on long-term Lake Huron zooplankton data from August reported a shift in community composition between the 1980s and 2000s: proportional biomass of calanoid copepods increased while that of cyclopoid copepods and herbivorous cladocerans decreased. Herein, we used seasonally collected data from Lake Huron in 1983–1984 and 2007 and reported similar shifts in proportional biomass. We also used a series of generalized additive models to explore differences in seasonal abundance by species and found that all three cyclopoid copepod species (Diacyclops thomasi, Mesocylops edax, Tropocyclops prasinus mexicanus) exhibited higher abundance in 1983–1984 than in 2007. Surprisingly, only one (Epischura lacustris) of seven calanoid species exhibited higher abundance in 2007. The results for cladocerans were also mixed with Bosmina spp. exhibiting higher abundance in 1983–1984, while Daphnia galeata mendotae reached a higher level of abundance in 2007. We used a subset of the 2007 data to estimate not only the vertical distribution of Bythotrephes longimanus and their prey, but also the consumption by Bythotrephes in the top 20 m of water. This epilimnetic layer was dominated by copepod copepodites and nauplii, and consumption either exceeded (Hammond Bay site) or equaled 65% (Detour site) of epilimnetic zooplankton production. The lack of spatial overlap between Bythotrephes and herbivorous cladocerans and cyclopoid copepod prey casts doubt on the hypothesis that Bythotrephes planktivory was the primary driver underlying the community composition changes in the 2000s.     

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.     

A Decade of Predatory Control of Zooplankton Species Composition of Lake Michigan

From 1983 to 1992, 71 species representing 38 genera from the Calanoida, Cladocera, Cyclopoida, Mysidacea, Rotifera, Mollusca and Harpacticoida comprised the offshore zooplankton community of Lake Michigan. Our data demonstrate that the composition and abundance of the calanoid community after 1983 is not unlike that of 1960s and that species diversity of the calanoid community is more diverse than the cladoceran community in the 1990s as compared to the early 1980s. Even though the relative biomass of the cladocerans has remained similar over the 1983–1993 period, the species diversity and evenness of the Cladocera community in the early 1990s is unlike anything that has been previously reported for Lake Michigan. Cladocera dominance is centered in one species, Daphnia galeata mendotae, and only three species of Cladocera were observed in the pelagic region of the lake in 1991 and 1992. Nutrient levels, phytoplankton biomass, and the abundance of planktivorous alewife and bloater chub and Bythotrephes are examined as possible causes of these changes in zooplankton species composition. The increase in Rotifera biomass, but not Crustacea, was correlated with an increase in relative biomass of unicelluar algae. Food web models suggest Bythotrephes will cause Lake Michigan's plankton to return to a community similar to that of the 1970s; that is Diaptomus dominated. Such a change has occurred. However, correlational analysis suggest that alewife and bloater chubs (especially juveniles) are affecting size and biomass of larger species of zooplankton as well as Bythotrephes.     

Historical trophic position of Limnocalanus macrurus in Lake Michigan

In Lake Michigan, oligotrophication and invasive species proliferation have led to dramatic changes in the planktonic food web. Limnocalanus macrurus is a native, glacial relict predatory copepod whose trophic role within the zooplankton community may help us better understand the dynamics behind those changes. We applied nitrogen isotope spectrometry to zooplankton from Lake Michigan to quantify the yearly and seasonal position of Limnocalanus and other taxa in the historical Lake Michigan planktonic food web. We found that Limnocalanus was positioned about one-half trophic level above Bythotrephes and about two levels above Daphnia during the summer in 1989, 1993, 1995 and 1997. It was unlikely that adult Limnocalanus encountered Bythotrephes during summer months because of vertical segregation during thermal stratification. Instead, Limnocalanus probably had greater access to copepod prey such as Diaptomus and Epischura. Limnocalanus became isotopically lighter seasonally in relation to Daphnia and either shifted its diet to one consisting of more phytoplankton, or its prey (e.g., Diaptomus) shifted to a more phytoplankton food base, thus indirectly resulting in decreased trophic status of Limnocalanus. This study serves as a historical, foundational basis for zooplankton food web relations in Lake Michigan that complements similar investigation in Lake Huron. Comparing the historical to the recent zooplankton food web may now elucidate how invasive species such as Bythotrephes and quagga mussels have altered zooplankton communities and bioenergetic relationships within the Great Lakes.     

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.     

Crustacean Zooplankton of Southwestern Lake Ontario in Spring 1973 and at the Niagara and Genesee River Mouth Areas in 1972 and Spring 1973

In southwestern Lake Ontario during three cruises in spring 1973, the common zooplankters were nauplii (78%), immature cyclopoid copepodids, Cyclops bicuspidatus thomasi CVI, and immature calanoid copepodids. Zooplankton was most abundant in late April-early May (second cruise) and least abundant at the end of May. There was more zooplankton than at the same stations a year earlier probably because of earlier copepod reproduction. In the river mouth areas during four cruises in 1972 and one cruise in May 1973, the common zooplaankters were nauplii, immature cyclopoid and calanoid copepodids, bosminids with mucro, Cyclops bicuspidatus thomasi CVI, Eubosmina coregoni, Ceriodaphnia lacustris, Daphnia retrocurva (Genesee River only), D. galeata mendotae (Niagara River only), Tropocyclops prasinus mexicanus CVI (Genesee River only), Diaptomus oregonensis CVI (Niagara River only). At the Genesee River the common zooplankters were most abundant in late August or November; at the Niagara River mouth copepods were most numerous in the spring and cladocerans in August and December. When compared to its nearest nonriver mouth IFYGL station in Lake Ontario, the Genesee River area supported less zooplankton and the Niagara River mouth area more. Several species of harpacticoid copepods were encountered which have not been reported previously from Lake Ontario. They were Bryocamptus zschokkei CVI, Epactophanes richardi CVI, Nitocra hibernica CVI, Nitocra spinepes CVI.     

A short-term look at potential changes in Lake Michigan slimy sculpin diets

Diporeia hoyi and Mysis relicta are the most important prey items of slimy sculpins (Cottus cognatus) in the Great Lakes. Slimy sculpins were collected from dreissenid-infested bottoms off seven Lake Michigan ports at depths of 27–73 m in fall 2003 to study their lake-wide diets. Relatively large dreissenid biomass occurred at depths of 37- and 46-m. Quagga mussels (Dreissena bugnesis) composed at least 50% of dreissenid biomass at Manistique, Saugatuck, and Sturgeon Bay. Mysis accounted for 82% of the sculpin diet by dry weight at eastern Lake Michigan while Diporeia composed 54–69% of the diet at western Lake Michigan and dominated the diets of slimy sculpins at all sites deeper than 46 m. In northern Lake Michigan, this diet study in new sites showed that slimy sculpin consumed more prey with low energy contents, especially chironomids, than Mysis and Diporeia in shallow sites (depth <55 m). We recommend diet studies on sedentary benthic fishes to be conducted along perimeters of the Great Lakes to observe changes in their diets that may be impacted by changing benthic macroinvertebrate communities.     

Prey selection by the Lake Superior fish community

Mysis diluviana is an important prey item to the Lake Superior fish community as found through a recent diet study. We further evaluated this by relating the quantity of prey found in fish diets to the quantity of prey available to fish, providing insight into feeding behavior and prey preferences. We describe the seasonal prey selection of major fish species collected across 18 stations in Lake Superior in spring, summer, and fall of 2005. Of the major nearshore fish species, bloater (Coregonus hoyi), rainbow smelt (Osmerus mordax), and lake whitefish (Coregonus clupeaformis) consumed Mysis, and strongly selected Mysis over other prey items each season. However, lake whitefish also selected Bythotrephes in the fall when Bythotrephes were numerous. Cisco (Coregonus artedi), a major nearshore and offshore species, fed largely on calanoid copepods, and selected calanoid copepods (spring) and Bythotrephes (summer and fall). Cisco also targeted prey similarly across bathymetric depths. Other major offshore fish species such as kiyi (Coregonus kiyi) and deepwater sculpin (Myoxocephalus thompsoni) fed largely on Mysis, with kiyi targeting Mysis exclusively while deepwater sculpin did not prefer any single prey organism. The major offshore predator siscowet lake trout (Salvelinus namaycush siscowet) consumed deepwater sculpin and coregonines, but selected deepwater sculpin and Mysis each season, with juveniles having a higher selection for Mysis than adults. Our results suggest that Mysis is not only a commonly consumed prey item, but a highly preferred prey item for pelagic, benthic, and piscivorous fishes in nearshore and offshore waters of Lake Superior.     

Seasonal Abundance of Planktonic Rotifers in a Nearshore Area of Central Lake Michigan

Thirty-four species of planktonic rotifers were recorded from east central Lake Michigan. Polyarthra vulgaris and Keratella cochlearis were the most abundant species recorded. Kellicottia longispina, Notholca squamula, Keratella earlinae, K. quadrata, and Synchaeta stylata were also important species. Maximum abundance occurred in June (360 Ind./L). July and August abundances exceeded 100 ind./L while spring and fall samples contained 10-20 ind./L. The species assemblage in east central Lake Michigan, western Lake Michigan, and Lake Ontario is similar. The eutrophic indicator Brachionus, abundant in western Lake Erie, was rare in this investigation.     

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

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

Dynamics of Lake Michigan Phytoplankton: Mechanisms Controlling Epilimnetic Communities

Lake Michigan epilimnetic algal and particulate organic carbon sedimentation rates, determined from moored and floating sediment traps, and algal loss rates due to crustacean zooplankton grazing, determined from manipulation experiments with natural assemblages, were estimated during 1983 and 1984 field seasons. Sedimentation was the most important algal loss in spring when colonial diatoms and calanoid copepods dominated the plankton. Grazing was the most important loss during summer when phytoflagellates and Daphnia spp. dominated. Comparisons between algal community production and loss due to sedimentation and grazing suggest that natural mortality may be significant, but it does not dominate control. An hypothesis is offered for overall control of phytoplankton dynamics in Lake Michigan.     

Total and methyl mercury accumulation in 1994–1995 Lake Michigan lake trout and forage fish

As part of the Lake Michigan Mass Balance Project, total and methyl mercury were determined for lake trout (Salvelinus namaycush) and five forage fish species collected from Lake Michigan near Saugatuck, Michigan, and Port Washington, Sheboygan Reef, and Sturgeon Bay, Wisconsin, between 1994 and 1995. With a mean concentration of 179 ng/g wet wt., whole lake trout total mercury (Hg_T) concentrations ranged between 27.6 and 348 ng/g wet wt. For combined sites, 1–4 yrs, 5–6 yrs, 7–11 yrs, and 12–15 yrs lake trout mean Hg_T concentrations were 73.7, 130, 212, and 280 ng/g, respectively. Forage fish species alewife (Alosa pseudoharengus), bloater (Coregonus hoyi), slimy sculpin (Cottus cognatus), deepwater sculpin (Myoxocephalus thompsoni), and rainbow smelt (Osmerus mordax) had mean Hg_T concentrations of 63.8, 55.3, 36.7, 51.4, and 35.2 ng/g wet wt., respectively. With the exception of alewife, bloater, and slimy sculpin, all fish species contained approximately 100% methyl mercury (MeHg). Field bioaccumulation factors (BAF) were consistent with a Lake Michigan food chain that is more efficient at transferring MeHg to higher trophic levels than some inland lakes. This and other studies of lake trout from Lake Michigan document decreasing Hg_T concentrations in lake trout from 1971 to 1985 and constant or increasing concentrations between 1985 and 2000. These observations were supported by a similar trend in Lake Michigan Hg sediment fluxes. To our knowledge, this is the most intense two year study of mercury in fish for any Great Lake or other large fresh water system and is one of the most complete studies of mercury cycling in the Lake Michigan food chain.     

Prey of Deep-water Hydra in Lake Superior

Specimens of Hydra attached to a soft-substrate bottom were collected from a manned submersible at a 275-m-deep station near the Keweenaw Peninsula in Lake Superior. Near-bottom zoo-plankton were collected to assess Hydra feeding preference. We identified and counted the zooplankton samples, and identified the zooplankton prey found in the Hydra. Of 86 Hydra, 21 contained remains of 23 prey of 4 zooplankton taxa. Compared with expected numbers based on the proportions of the net samples, these Hydra appeared to be feeding selectively, with a preference for larger zooplankton. The large calanoid copepod Limnocalanus macrurus was particularly selected. A very provisional estimate of the rate of Hydra predation on the bottom of Lake Superior is one prey per 4 days. The relatively constant environmental conditions, lack of commensals and predators, and the zooplankton food source suggest that the bottom of Lake Superior is a stable habitat for Hydra.     

The Status of Limnocalanus macrurus (Copepoda: Calanoida: Centropagidae) in Lake Erie

The calanoid copepod Limnocalanus macrurus showed large declines in abundance and a narrowing of spatial distribution with the onset of cultural eutrophication and increases in rainbow smelt (Osmerus mordax) abundances in Lake Erie in the mid 20th century. Since 1995, however, Limnocalanus macrurus appears to have repopulated in western Lake Erie to levels of abundance that have not been observed since the late 1930s. We hypothesize that phosphorus abatement and the subsequent decrease in low dissolved oxygen events have assisted this resurgence. However, Limnocalanus macrurus abundances have not increased in the central and eastern basins, even though water quality has improved there too. High densities of rainbow smelt and associated smelt predation pressure in the central and eastern basins may be responsible for the low numbers in these basins.     

Behavioral response of Lake Michigan Daphnia mendotae to Mysis relicta

We performed laboratory experiments to determine if Mysis relicta induce changes in the behavior of Daphnia mendotae collected from Lake Michigan. Laboratory results indicate that Daphnia perceived Mysis kairomones and responded by changing their vertical position in cylinders. Experiments using different resource levels, and two procedures to examine the potential effects of the chemical cues from Mysis or from particulate matter or bacteria associated with capture and defecation of prey, suggest that Daphnia detect Mysis via a chemical cue. This is the first laboratory study that we are aware of that indicates that a zooplankton species from the Great Lakes responds behaviorally to an invertebrate predator. Our findings support the hypothesis that changes in vertical distribution of zooplankton associated with changes in invertebrate predator density, observed in previous Great Lakes studies, is due to behavioral responses to reduce predation risk. It is important to understand and quantify such responses, because predator-induced changes in prey behavior represent trait-mediated interactions that can potentially strongly affect prey growth rates, and indirectly affect resources, competitors, and predators of the prey.     

Recent increases in the large glacial-relict calanoid Limnocalanus macrurus in Lake Michigan

Since 2004, population density of the large hypolimnetic calanoid Limnocalanus macrurus Sars. has increased dramatically in Lake Michigan. The average summer biomass of this species between 2004 and 2006 was roughly three times that of the period 1984–2003, and at levels unprecedented in our 22-year dataset, making L. macrurus the dominant zooplankter in the lake in terms of biomass. These increases have been accentuated by coincident population declines of the main daphnid, Daphnia mendotae, in the lake with the result that in 2006, L. macrurus accounted for 75% and 50% of the large (> 0.9 mm) crustacean biomass in the northern and southern basins of Lake Michigan, respectively. The increases in L. macrurus populations have closely coincided with equally dramatic increases in summer water clarity. Recent extinction coefficients are among the lowest recorded for the lake, and deepening light penetration has permitted increases in the size of the deep chlorophyll layer. In addition, planktivorous fish populations have declined coincidently with the increases in L. macrurus. It seems likely that an increase in sub-epilimnetic production has resulted in increased food resources for the deep-living L. macrurus, while low planktivore abundances have reduced predation loss, permitting L. macrurus to respond to these increases in sub-epilimnetic production.     

Some Fungi and Water Molds in Waters of Lake Michigan with Emphasis on Those Associated with the Benthic Amphipod Diporeia spp.

To determine types of fungi in the water and associated with the benthic amphipod Diporeia spp., samples were collected at various depths in Lake Michigan in an area where the Diporeia population was in a severe state of decline. No fungi were found associated with living, freshly-dead, or dried Diporeia cultured separately from Lake Michigan water. When dead Diporeia and other organic substrates (snake skin and hemp seeds) were used to grow fungi in Lake Michigan water, a rich and diverse fungal and water mold community was revealed. A total of 31 species were found, with the most common genera being Achlya, Aphanomyces, Myzocytium, and Pythium. In general, species were homogeneously distributed in the water; that is, few differences were found in species richness between nearshore (10–15 m) and offshore (60–80 m) waters, and between near-surface (1 m) and near-bottom waters (1 m off bottom). Sampling occurred during the unstratified period (April and October) to maximize the number of species collected, which may have contributed to the uniform spatial pattern observed. While conclusions must be placed in context with our methods of detection, we found no evidence that a fungal infestation was associated with Diporeia in this region of the lake.     

The Asian cyclopoid copepod Mesocyclops pehpeiensis Hu 1943 reported from the western basin of Lake Erie

The Asian cyclopoid copepod Mesocyclops pehpeiensis Hu, 1943 has been reported as an introduced species at several locations in the western hemisphere. In the United States, reports of this exotic species are restricted to localities in Louisiana, Mississippi, and Washington D.C. This report documents a new record of occurrence for M. pehpeiensis from the western basin of Lake Erie. The detection of M. pehpeiensis in Lake Erie constitutes the first record of this species from the Laurentian Great Lakes, and the northernmost record in the western hemisphere. The species was found in 2016, 2017 and 2018, including females with egg sacs, and can therefore be considered established in the area. The occurrence of M. pehpeiensis in Lake Erie suggests that this Asian copepod may be more widely distributed in North America than is currently understood.     

Lake-wide,annual status of the Mysis diluviana population in Lake Michigan in 2015

Mysis diluviana is one of the most abundant zooplankton by biomass in the Laurentian Great Lakes of North America, a predator of other zooplankton and an important prey for fishes. Studies of long-term trends in Lake Michigan have shown 2005–2016 densities to be 50–80% lower than 1990s densities, but these observations have been based on annual monitoring that is either spatially or seasonally limited. We combined Lake Michigan Mysis data from three annual programs and the 2015 Cooperative Science and Monitoring Initiative to achieve broad spatial coverage during spring, summer, and fall of 2015 and broad depth coverage during spring 2016. Lake-wide, annual density and biomass were 82 (SE: 10) Mysis/m² and 200 (SE: 36) mg dry mass/m². Density and biomass estimates were highest offshore, generally higher in the north basin, and seasonally highest in summer. Annual lake-wide averages for depths >30 m were better captured by seasonally-extensive annual programs than spatially-extensive annual programs, although spring sampling may bias annual values low. Mysis cohorts grew 0.026 mm/d (age-0) and 0.007 to 0.027 mm/d (age-1). Annual mortality was 81–98%. Reproduction was fall-spring and seasonal lake-wide estimates ranged from 0.6 to 19.1% females brooding, 13–20 embryos/brood, and 3–46 embryos/m². Annual production (423 mg dry mass/m²/yr, SE: 31) was lower than all but one previous estimate from lakes Michigan, Huron, and Ontario. While Mysis tend to persist, low Mysis production may be a concern for prey fishes that feed on Mysis.