Occurrence of Zebra Mussels in Near-shore Areas of Western Lake Erie

We measured biomass, percent coverage, and length-frequency of zebra mussels in near-shore areas of western Lake Erie between 16 September and 10 November 1993 as part of a larger study on the ecological relationship between diving ducks and zebra mussels. Wet weight biomass of zebra mussels, determined by SCUBA diving, ranged from 0 to 3,611 g/m² and averaged ( ± 1 SE) 1,270 ± 380 g/m² (n = 11). Percent coverage of lake bottom by zebra mussels ranged from 0 to 70% and averaged 17 ± 4.0% (n = 27). Percent coverage of zebra mussels was relatively high in the northern portion (28–70% coverage) and in the southwestern portion (18–40%), but relatively low ( < 5%) in the southeastern portion of the study area. Percent coverage by zebra mussels, determined from underwater videography, was highly correlated (r² = 0.96) with zebra mussel biomass. Analysis of length-frequency data indicated that there was prominent recruitment of juvenile zebra mussels at only three of eight sites. Average shell length ranged from 11 mm to 15 mm at the other five sites. The non-uniform distribution of zebra mussels, as determined from biomass and videography, may have important ramifications when assessing zebra mussel impacts on waterfowl. These data may also be used when assessing impact of zebra mussels on other aquatic organisms in the near-shore areas of western Lake Erie.     

Quantifying a shift in benthic dominance from zebra (Dreissena polymorpha) to quagga (Dreissena rostriformis bugensis) mussels in a large,inland lake

Dreissenid mussels are aggressive invasive species that are continuing to spread across North America and co-occur in the same waterbodies with increasing frequency, yet the outcome and implications of this competition are poorly resolved. In 2009 and 2015, detailed (700 + sample sites) surveys were undertaken to assess the impacts of invasive dreissenid mussels in Lake Simcoe (Ontario, Canada). In 2009, zebra mussels were dominant, accounting for 84.3% of invasive mussel biomass recorded. In 2015, quagga mussels dominated (88.5% of invasive mussel biomass) and had expanded into profundal (> 20 m water depth) sites and onto soft (mud/silt) substrates with a mean profundal density of 887 mussels/m² (2015) compared to ~ 39 mussels/m² in 2009. Based on our annual benthos monitoring, at a subset of ~ 30 sites, this shift from zebra to quagga mussels occurred ~ 2010 and is likely related to a population decline of zebra mussels in waterbodies where both species are present, as recorded elsewhere in the Great Lakes Region. As the initial invasion of dreissenid mussels caused widespread ecological changes in Lake Simcoe, we are currently investigating the effects this change in species dominance, and their expansion into the profundal zone, will have on the lake; and our environmental management strategies. Areas of future study will include: changes in the composition of benthos, fish, or phytoplankton communities; increased water clarity and reduction of the spring phytoplankton bloom; energy/nutrient cycling; and fouling of anthropogenic in-lake infrastructures (e.g. water treatment intakes) built at depths > 25 m to avoid previous zebra mussel colonization.     

Microzooplankton distribution,dynamics, and trophic interactions relative to phytoplankton and quagga mussels in Saginaw Bay,Lake Huron

Invasive quagga mussels have recently replaced zebra mussels as the dominant filter-feeding bivalves in the Great Lakes. This study examined microzooplankton (i.e., grazers < 200 μm) and their trophic interactions with phytoplankton, bacteria, and bivalve mussels in Saginaw Bay, Lake Huron, following the zebra to quagga mussel shift. Microzooplankton distribution displayed strong spatial and temporal variability (1.73–28.5 μg C/L) relative to phytoplankton distribution. Ciliates were the dominant component, especially in the spring and early summer. Rotifers and dinoflagellates increased toward late summer/fall in the inner and outer parts of the bay, respectively. Microzooplankton grazing matched bacterial growth rates and removed ca. 30% of the phytoplankton standing stock in the < 100 μm size fraction per day. The greatest herbivory occurred at the site dominated by colonial cyanobacteria. Microzooplankton, which comprised < 4% of the quagga mussels prey field (i.e. available prey), contributed 77% and 34% to the quagga carbon-based diet during Microcystis and diatom blooms, respectively. Feeding on microzooplankton could buffer mussels during lean periods, or supplement other consumed resources, particularly during noxious cyanobacterial blooms. The results of this study demonstrate that microzooplankton are a resilient and critical component of the Saginaw Bay ecosystem.     

Effects of Zebra Mussels (Dreissena polymorpha) on Bacterioplankton: Evidence for Both Size-Selective Consumption and Growth Stimulation

Zebra mussels had significant direct and possible indirect effects on heterotrophic bacteria in two contrasting sites in Saginaw Bay. At a eutrophic site in the inner portion of Saginaw Bay, mussels fed directly on bacterial-sized particles and had a negative impact on bacterial abundances. Mussels removed large bacteria (> 0.9 μm) more effectively than small bacteria at this site. Individual mussels cleared from 37–89 ml per day. Results using different sizes of fluorescent microspheres suggest that zebra mussels have a lower limit for particle size removal that is less than 0.4 μm. Contradictory to inner bay results, mussels at an outer bay oligotrophic site had a positive impact on heterotrophic bacterial abundance, perhaps as a result of indirect effects, such as nutrient or organic carbon excretion by the mussels. Differences in the impact of mussels on the bacterial communities of the inner bay and outer bay probably result from differences in trophic state and bacterial community structure. A hypothesized smaller size of bacteria at outer bay sites may enable them to escape heavy predation pressure from mussels and the high rates of mussel nutrient excretion may facilitate their growth in these nutrient depleted conditions.     

Changes in Distribution and Abundance of Submerged Macrophytes in the Inner Bay at Long Point,Lake Erie: Implications for Foraging Waterfowl

Submerged aquatic macrophytes in Long Point Bay provide food for several species of migrating waterfowl, particularly diving ducks. The percent occurrence of several plant species in the diets of six species (207 birds) of diving ducks were studied between 1992 and 1994. Of 29 plant species consumed, Vallisneria americana was the most commonly occurring in the diet. Chara vulgaris, Potamogeton spp., Polygonum spp., and Najas flexilis/quadalupensis were also important dietary items. The seeds of a number of emergent macrophyte species were also consumed. Submerged macrophytes were sampled at 312 locations in the Inner Bay of Long Point, Lake Erie, in 1991, 1992, and 1995. Distribution and percent abundance were compared with a similar study in 1976 and with observations of macrophytes in 1962. In comparison with the 1976 study, Chara vulgaris is still dominant over much of the Bay. V. americana and N. flexilis/N. guadalupensis have expanded their ranges, probably a result of increasing light penetration associated with the filtering of suspended material by introduced zebra mussels Dreissena polymorpha. The continued widespread distribution of C. vulgaris, and the expansion of V. americana and N. flexilis/N. guadalupensis, has probably benefitted waterfowl, as these species have a high nutritive value and are readily consumed by diving ducks at Long Point. Myriophyllum spicatum, an exotic species introduced in the early 1970s, increased in distribution between 1976 and 1991 but decreased in distribution and percent abundance by 1995. Increased light transparency may have contributed to this decline, as M. spicatum typically grows best in turbid, eutrophic environments. The decline of this species has probably been beneficial to waterfowl because it commonly out-competes native plants and it is underutilized by ducks at Long Point. Ceratophyllum demersum appears to be a more prominent member of the submerged macrophyte communities in the 1990s, as it was not reported previously. In comparison with 1962, major changes have occurred in species composition and relative abundance: M. spicatum, Najas spp., and Elodea canadensis have increased, Potamogeton spp. and Nitella spp. have declined. Macrophyte distribution and abundance on the Inner Bay is influenced by environmental, biological, geomorphological and geological variables. However, the anthropogenic introduction of two exotic species, M. Spicatum and the zebra mussel, appears to have had a pronounced influence on the community structure of submerged macrophytes in Long Point Bay.     

Dreissenid mussels are not a “dead end” in Great Lakes food webs

Dreissenid mussels have been regarded as a “dead end” in Great Lakes food webs because the degree of predation on dreissenid mussels, on a lakewide basis, is believed to be low. Waterfowl predation on dreissenid mussels in the Great Lakes has primarily been confined to bays, and therefore its effects on the dreissenid mussel population have been localized rather than operating on a lakewide level. Based on results from a previous study, annual consumption of dreissenid mussels by the round goby (Neogobius melanostomus) population in central Lake Erie averaged only 6 kilotonnes (kt; 1 kt = one thousand metric tons) during 1995–2002. In contrast, our coupling of lake whitefish (Coregonus clupeaformis) population models with a lake whitefish bioenergetics model revealed that lake whitefish populations in Lakes Michigan and Huron consumed 109 and 820 kt, respectively, of dreissenid mussels each year. Our results indicated that lake whitefish can be an important predator on dreissenid mussels in the Great Lakes, and that dreissenid mussels do not represent a “dead end” in Great Lakes food webs. The Lake Michigan dreissenid mussel population has been estimated to be growing more than three times faster than the Lake Huron dreissenid mussel population during the 2000s. One plausible explanation for the higher population growth rate in Lake Michigan would be the substantially higher predation rate by lake whitefish on dreissenid mussels in Lake Huron.     

Replacement of Zebra Mussels by Quagga Mussels in the Canadian Nearshore of Lake Ontario: the Importance of Substrate,Round Goby Abundance,and Upwelling Frequency

The invasion of the Great Lakes by zebra mussels (Dreissena polymorpha) and quagga mussels (Dreissena bugensis) has been accompanied by tremendous ecological change. In this paper we characterize the extent to which dreissenids dominate the nearshore of the Canadian shoreline of Lake Ontario and examine mussel distribution in relation to environmental factors. We surveyed 27 5-m sites and 25 20-m sites in late August 2003. Quagga mussels dominated all sites (mean: 9,404/m²; range 31–24,270), having almost completely replaced zebra mussels. Round gobies (Neogobius melanostomus) were associated with quagga populations dominated by large mussels. Quagga mussel total mass was low at 5-m sites with high upwelling frequency; we believe this is the first documentation of reduced benthic biomass in areas of upwelling in Lake Ontario. Overall, we estimated 6.32×10¹² quagga mussels weighing 8.13×10¹¹ g dry weight and carpeting ∼66% of the nearshore benthic habitat. Quagga mussels are a dominant and defining feature of the Lake Ontario nearshore, and must be accounted for in management planning.     

Increased Abundance and Depth of Submersed Macrophytes in Response to Decreased Turbidity in Saginaw Bay,Lake Huron

Submersed macrophyte communities and turbidity near shore were measured from 1991 to 1993 to determine if more light resulting from colonization of zebra mussels (Dreissena polymorpha Pallas) into Saginaw Bay in 1990 corresponded with changes in macrophyte distribution. Turbidity was sampled along five transects distributed at intervals perpendicular to the perimeter of inner Saginaw Bay to monitor changes in light available to plants in Saginaw Bay. Vegetation was sampled in July along these transects to determine the distribution and composition of the macrophyte communities each summer. We also measured the maximum depth of colonization and the area of plant coverage by use of 31 transects evenly distributed around Saginaw Bay in August. Turbidity decreased (P ≤ 0.097) at transects in northern littoral regions from 1991 to 1993 over submersed plant communities and uncolonized sediments, but not in southern littoral regions. The relative abundance of submersed macrophytes increased (P ≤ 0.0001) at all transects from 1991 to 1993, especially at transects where turbidity decreased significantly. Maximum depth of colonization (2.0 m) and the area of macrophyte coverage (101.3 km²) increased in Saginaw Bay, especially in the northwestern littoral region of the bay. Macrophytic chlorophytes, charophytes, and Vallisneria americana increased (P ≤ 0.003) in relative abundance most at transects where turbidity decreased significantly. These results demonstrate that even in a large well-mixed lacustrine environment, zebra mussels have the capacity to reduce turbidity sufficiently to allow submersed macrophytes to expand their distribution and abundance.     

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.     

Depth drives growth dynamics of dreissenid mussels in Lake Ontario

Understanding dreissenid mussel population dynamics and their impacts on lake ecosystems requires quantifying individual growth across a range of habitats. Most dreissenid mussel growth rates have been estimated in nutrient rich or nearshore environments, but mussels have continued to expand into deep, cold, low-nutrient habitats of the Great Lakes. We measured annual quagga mussel (Dreissena rostriformis bugensis) growth at 15 m, 45 m, and 90 m in Lake Ontario using caged mussels near Oswego, New York, USA from June 2018 to May 2019. Quagga mussel growth (starting size 12 mm) was greatest at 15 m (mean shell length increase = 10.2 mm), and was lower at 45 m (5.9 mm) and 90 m (0.7 mm). Caged mussels were obtained from near the 90-m site and those reared at 15 and 45 m developed thicker shells than those that were caged at 90 m. We observed relatively high colonization by quagga and, to a lesser degree, zebra mussels (Dreissena polymorpha) at 15 m, very few colonizers at 45 m, and none at 90 m. Higher growth potential, but low natural mussel densities observed at 15 m and 45 m suggest factors other than growth limit dreissenid abundance at these depths. The relatively slow dreissenid growth rates observed in offshore habitats are consistent with the gradual abundance increases documented in these zones across the Great Lakes and suggest new mussels that become established in these habitats may contribute to ecosystem effects for decades.     

A mesocosm investigation of the effects of quagga mussels (Dreissena rostriformis bugensis) on Lake Michigan zooplankton assemblages

Dreissenid mussels are known to disrupt the base of the food web by filter feeding on phytoplankton; however, they may also directly ingest zooplankton thereby complicating their effects on plankton communities. The objective of this study was to quantify the effects of quagga mussel feeding on the composition and size structure of Lake Michigan zooplankton assemblages. Two mesocosm (six 946 L tanks) experiments were conducted in summer 2013, using quagga mussels and zooplankton collected near Beaver Island, MI, to examine the response of zooplankton communities to the presence and absence of mussels (experiment 1) and varying mussel density (experiment 2). Mesocosms were sampled daily and zooplankton taxa were enumerated and sized using microscopy and FlowCAM® imaging. In experiment 1, the presence of quagga mussels had a rapid negative effect on veliger and copepod nauplii abundance, and a delayed negative effect on rotifer abundance. In experiment 2, mussel density had a negative effect on veliger, nauplii, and copepodite abundance within 24 h. Multivariate analyses revealed a change in zooplankton community composition with increasing mussel density. Ten zooplankton taxa decreased in abundance and frequency as quagga mussel density increased: except for the rotifer Trichocerca sp., treatments with higher mussel densities (i.e., 1327, 3585, and 5389 mussels/m²) had the greatest negative effect on small-bodied zooplankton (≤ 128 μm). This study confirms results from small-scale (≤ 1 L) experiments and demonstrates that quagga mussels can alter zooplankton communities at mesoscales (~ 1000 L), possibly through a combination of direct consumption and resource depletion.     

Veliger density and environmental conditions control quagga mussel colonization rates in two perialpine lakes

Quagga and zebra mussels (Dreissena bugensis and D. polymorpha) are spreading across lakes in Europe and North America. In particular, quagga mussels colonize lakes to great depths (>200 m). To better understand the colonization pattern of quagga mussels in deep lakes, we studied the settlement of quagga mussels along a depth gradient on colonization plates at multiple depths (1–140 m) in the pelagic zone of two recently invaded perialpine lakes, Lake Constance and Lake Geneva. We measured colonization rates every three months over one year on colonization plates deployed in both lakes at defined depths. We also assessed long-term population dynamics from abundance and size distribution using repeated photogrammetry of colonization plates. Highest colonization rates and largest mussel sizes occurred above 8 m depth, and almost no zebra mussels were found. Colonization rates decreased to almost zero below 30 m. Colonization rates on plates were associated with variation in environmental conditions as well as veliger densities in the plankton across season and depth. Temperature was the most important environmental parameter that influenced colonization. Our results will help to better understand the seasonal colonization patterns of invasive quagga mussels in deep lakes.     

A Seasonal Comparison of Suspended Sediment Filtration by Quagga (Dreissena bugensis) and Zebra (D. polymorpha) Mussels

Field evidence suggests a shift in the dreissenid population from zebra (Dreissena polymorpha) to quagga (D. bugensis) mussels is occurring within the lower Great Lakes. This laboratory study directly compared per-mussel and per-dry-weight filtration rates (volume per time) of both species, gauged by the clearance of resuspended natural sediments (1 to 12 mg/L) from gently mixed, 1-L static vessels. Mussels of 15- and 20-mm lengths were collected together from the Lake Ontario drainage basin at Oak Orchard Creek, Medina, NY, and maintained and tested in ambient Niagara River water. A 2 × 4 factorial design was employed, with species and season as independent factors. Season significantly influenced filtration rate of both size classes, and winter rates were about half those measured during the rest of the year. Species significantly influenced filtration of 20-mm mussels. Quagga mussels of this size filtered up to 37% faster than zebra mussels (data for spring: 309 vs. 226 mL/h/mussel, n = 18 and 20 individuals, respectively). Species was not a significant factor alone for 15-mm mussels, but a species x season interaction was significant. The zebra mussels employed here had 16 to 22% more ash-free dry weight (AFDW) than the quagga mussels, accentuating filtration differences when expressed per-mg-AFDW.     

Cladophora,mass transport,and the nearshore phosphorus shunt

The nearshore phosphorus shunt hypothesis and the potential for mussels to excrete phosphorus sufficient to meet the growth requirements of Cladophora are now well accepted by scientists studying Great Lakes biogeochemistry. The response of algal growth to near bottom water column phosphorus concentrations and the interplay between excretion and mass transport in yielding those concentrations have, however, not been elucidated. Here we present soluble reactive phosphorus profiles from the near bottom environment of Lake Michigan at a site near Good Harbor Bay, Michigan, where both mussels and Cladophora were present. Soluble reactive phosphorus was observed to accumulate under quiescent conditions, establishing a concentration boundary layer (CBL), 5–15 cm thick, with near bottom concentrations on the order of 2–8 μg P/L. A one-dimensional model was applied to determine mass transport conditions mediating the transition from CBL formation to CBL destruction. Significant wave height (SWH) was used as an indicator of mass transport intensity, and it was determined that the formation/destruction transition occurred at a SWH of 0.2 m at the 8-m study site depth. The Great Lakes Cladophora Model was applied to determine the time intervals required to saturate (1 day with the CBL present) and deplete (14 days with the CBL absent) algal internal P stores. A review of SWH conditions at the study site indicated that a CBL would be expected to form at a frequency sufficient to support the phosphorus nutrition of Cladophora over the entire May to August interval.     

Nitrogen Dynamics in Sandy Freshwater Sediments (Saginaw Bay,Lake Huron)

Sediment-water nitrogen fluxes and transformations were examined at two sites in Saginaw Bay, Lake Huron, as a model for sandy freshwater sediments. Substantial ammonium release rates (74 to 350 μmole NH₄⁺/m²/h¹) were observed in flow-through cores and in situ benthic chamber experiments. Sediment-water ammonium fluxes were similar at the inner and outer bay stations even though inner bay waters are enriched with nutrients from the Saginaw River. The high net flux of remineralized ammonium into the overlying water from these sandy sediments resembles typical data for marine systems (11 to 470 μmole NH₄⁺/m²/h¹) but were higher than those reported for depositional freshwater sediments (0 to 15 μmole NH₄⁺/m²/h¹; Seitzinger 1988). Addition of montmorillonite clay (ca. 1 kg dry weight/m²) to the top of the sandy cores reduced ammonium flux. Mean “steady-state” ammonium flux following clay addition was 46 ± 2 (SE) % of the initial rates as compared to 81 ± 8% of the initial rates without clay addition. Zebra mussel excretion dominanted ammonium regeneration in the inner bay where the bivalve was abundant, but addition of zebra mussel feces/psuedofeces (3.0 g dw/m²) to sediments did not increase ammonium or nitrate flux. Partial nitrification of ammonium at the sediment-water interface was suggested by removal of added ¹⁵NH₄⁺ from lake water passing over dark sediment cores. Sediment-water fluxes of nitrogen obtained from flow-through sediment cores resembled those from in situ benthic chambers. However, extended static incubations in gas-tight denitrification chambers caused more of the regenerated nitrogen to be nitrified and denitrified than occurred with the other two measurement systems.     

Modeling the Role of Zebra Mussels in the Proliferation of Blue-green Algae in Saginaw Bay,Lake Huron

Between 1991 and 1993, Saginaw Bay experienced an invasion by zebra mussels, Dreissena polymorpha, which caused a significant perturbation to the ecosystem. Blooms of Microcystis, a toxin-producing blue-green alga, became re-established in the bay after the zebra mussel invasion. Microcystis blooms had all but been eliminated in the early 1980s with controls on external phosphorus loadings, but have re-occurred in the bay most summers since 1992. An apparent paradox is that these recent Microcystis blooms have not been accompanied by increases in external phosphorus loadings. An ecosystem model was used to investigate whether the re-occurrence of Microcystis could be due to changes caused by zebra mussels that impacted phytoplankton community structure and/or internal phosphorus dynamics. The model was first used to establish baseline conditions in Saginaw Bay for 1991, before zebra mussels significantly impacted the system. The baseline model was then used to investigate: (1) the composite impacts of zebra mussels with average 1991–1995 densities; (2) sensitivity to changes in zebra mussel densities and external phosphorus loadings; and (3) three hypotheses on potential causative factors for proliferation of blue-green algae. Under the model assumptions, selective rejection of blue-green algae by zebra mussels appears to be a necessary factor in the enhancement of blue-green production in the presence of zebra mussels. Enhancement also appears to depend on the increased sediment-water phosphorus flux associated with the presence of zebra mussels, the magnitude of zebra mussel densities, and the distribution of zebra mussel densities among different age groups.     

Trends of PCB concentrations in Lake Michigan coho and chinook salmon, 1975–2010

The manufacture and use of polychlorinated biphenyls (PCBs) was banned in the United States in 1977 after it was determined that these compounds adversely affect animals and humans. The Wisconsin Department of Natural Resources has quantified total PCB concentrations in Lake Michigan chinook (n = 765) and coho (n = 393) salmon (Oncorhynchus tshawytscha and Oncorhynchus kisutch, respectively) filets since 1975. We analyzed these data to estimate trends in PCB concentrations in these fish (1975–2010). We used generalized linear models with a gamma error distribution and log link fit to the untransformed concentrations. Trend patterns were examined using graphical smoothing and generalized additive models. We identified a candidate set of models that included time trend and other predictor variables. Using the Akaike Information Criterion to select among models we found the best models for both species included piecewise linear time trends, total body length, % lipid, and collection season as predictor variables. The intersection of the two trends was 1985 for chinook salmon and 1984 for coho salmon. PCB concentrations in both species increased with body length and % lipid, and were higher for individuals caught in the fall. Our data reveals a dramatic decline in PCB concentrations of − 16.7% and − 23.9% per year for chinook and coho, respectively, up until the intersection year likely reflecting implementation of restrictions on Aroclor-based PCBs. After the intersection year to 2010, PCB concentrations declined at an annual rate of − 4.0% (95% CI: − 4.4% to − 3.6%) and − 2.6% (95% CI: − 3.3% to − 1.9%) for chinook and coho, respectively.     

Effect of the urbanized embayment Toronto Harbour on the composition and production of zooplankton

To better understand zooplankton dynamics in Lake Ontario’s Toronto Harbour and adjacent coastal area (CA), we sampled zooplankton, phytoplankton, nutrients and physical parameters on six dates in 2016. Despite higher levels of nutrients, chlorophyll and primary production in the inner harbor (IH), the areas supported similar May to November zooplankton biomass (IH = 32 ± 7 and CA = 42 ± 10 mg/m³). IH values were much lower than other nutrient-enriched embayments in Lake Ontario, yet CA biomass was twice that of nearshore sites away from Toronto. Small zooplankton such as rotifers and Bosmina dominated IH; and large taxa (Daphnia, calanoids and predatory cladocerans) were more important in the CA. Daphnia, Bosmina, cyclopoids and calanoids were larger in the CA, and adult cyclopoids had higher egg ratios. This led to low annual IH production estimates for both cyclopoid and calanoid copepods. Total phosphorus and chlorophyll did not appear to regulate zooplankton biomass, but positive relationships were found with bacterial biomass in the IH and with temperature in the cool season. Atypically high fish planktivory rates likely suppressed larger IH zooplankton in 2016, allowing small, resilient Bosmina to flourish and contribute 84% of total production in the IH. Comparing 2016 data to previous zooplankton surveys revealed considerable inter-annual variation in proportions of Daphnia, Bosmina and predatory cladocerans over the 1994 to 2016 period, and the strong top-down controls observed in 2016 were not typical. Elevated microbial production may serve as an important alternate trophic pathway supporting cladoceran populations in Toronto Harbour.     

New Records of Freshwater Sponges (Porifera) for Southern Lake Michigan

Four species of freshwater sponge were identified in southern Lake Michigan: Spongilla lacustris (Linneaus), Eunapius fragilis (Leidy), Ephydatia fluviatilis (Linneaus), and Ephydatia muelleri (Lieberkuhn). Samples were collected from artificial substrates in Calumet (IL), Hammond (IN), and Michigan City (IN) harbors and represent the first reports of sponges in these waters. Numerical densities for all species combined were as high as 698 sponge colonies/m² on the iron hull of the permanently moored 110-m-long ship Milwaukee Clipper in Hammond Harbor. These densities were lowest at 1-m depth and greater at 2- and 3-m depths. A positive correlation (r² = 0.74) was found between the density of sponge colonies and the angle of the hull as it changed from near vertical at the water line to overhanging at greater depth (toward the keel). Ephydatia muelleri was the most common species based on frequency of occurrence and percent surface area covered. Observations of epizoic growth of sponges on live zebra mussels (Dreissena polymorpha Pallas) suggest that an ecological interaction exists between these two groups of organisms on these artificial substrates.     

In situ reciprocal transplants reveal species-specific growth pattern and geographic population differentiation among zebra and quagga mussels

For bivalves, somatic growth is often inferred from shell measurements alone. However, shell growth may not always reflect changes in soft tissue due to confounding factors such as seasonal or ontogenetic asynchrony between shell and tissue, flexible energy allocation, or population differences. This study compares the relationship between shell growth, changes in soft tissue mass, and RNA/DNA ratio in the zebra mussel (Dreissena polymorpha) and quagga mussel (Dreissena bugensis) from contrasting riverine and brackish estuarine environments. Reciprocal transplantation indicated that shell growth in late summer was consistently lower for the estuarine source zebra mussels while the RNA/DNA ratio was highest for zebra mussels independent of either geographic source or destination. Shell growth of the river source quagga mussels was almost two times greater than zebra mussels at the river site, but both shell growth and final tissue mass were lower in the estuarine environment. While there were no differences in final RNA/DNA ratios between zebra and quagga mussels from the same source, the RNA/DNA ratio of zebra mussels from the estuary and transplanted to the estuary was higher than that of all other zebra mussel treatments. This study suggests that shell growth does not always accurately reflect tissue growth and that the shell and tissue growth of quagga mussels is greater than that of zebra mussels in fresh but not brackish waters, and that physiological plasticity can have a fixed geographic component.