Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia‐nitrogen and phosphate‐phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft‐tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well‐mixed and/or shallow areas of western Lake Erie.     

Temporal variation of energy reserves in mayfly nymphs (Hexagenia spp.) from Lake St Clair and western Lake Erie

1. We analysed changes in energy reserves (lipid and glycogen) and length–weight relationships of burrowing mayflies (Hexagenia spp.) in 1997–99 to compare an established population in Lake St Clair with a recovering population in western Lake Erie of the Laurentian Great Lakes. In addition, we measured changes in water temperature and potential food in both water columns and sediments. 2. Although overall mean values of lipid and glycogen levels of Hexagenia nymphs from Lake St Clair and western Lake Erie were not significantly different, there were differences in seasonal patterns between the two lakes. In Lake St Clair, levels were highest in early spring, declined throughout the year, and reached their lowest levels in fall during all 3 years of study. In contrast, levels in western Lake Erie were lower in spring, increased to a maximum in summer, then declined in fall. Seasonal patterns in length–weight relationships were similar to those for lipid and glycogen. 3. Total lipid as a percentage of dry weight did not increase with developmental stage of nymphs until just prior to metamorphosis and emergence from water. However, the major reserve lipid, triacylglycerols, increased systematically with development stage. In the final stage of development, triacylglycerols declined, probably as a result of energy consumption and its conversion to other biochemical components for metamorphosis and reproduction. 4. Indicators of potential food (algal fluorescence in the water column and chlorophyll a and chlorophyll a/phaeophytin ratio in sediments) suggest that Hexagenia in Lake St Clair have a food source that is benthic based, especially in early spring, whereas in western Lake Erie nymphs have a food source that is water column based and settles to the lake bottom during late spring and summer.     

Spatial variation in density, mean size and physiological condition of the holarctic amphipod Diporeia spp. in Lake Michigan

• 1 We examined spatial patterns in population characteristics (density, biomass, mean body length) and physiological condition (lipid content, length‐weight) of the amphipod Diporeia spp. in Lake Michigan by collecting samples at up to 85 sites in late summer 1994 and 1995. Variables were examined relative to water depth and three lake regions: south, central and north. Most major river systems are found in the south, and this region is more nutrient‐enriched compared to the north.
• 2 Over all sites, mean density was 5240^‐2, biomass was 4.1g dry wt m^‐2, and mean body length was 5 mm. While maximum densities were related to depth, with a peak at 30–70 m, greatest densities occurred on the west side of the lake, and low densities were found in the south‐east, north‐east and lower Green Bay. High densities in the west probably resulted from upwelling, and reduced densities in the south‐east may reflect food competition with Dreissena polymorpha (zebra mussel).
• 3 Lipid content, weight per unit length, and mean length declined with increased water depth, but depth‐related trends were most evident in the south. Overall, mean lipid content and weight per unit length were significantly lower in the south (16.6% dry wt, 0.59 mg at 5 mm body length) compared to the north (23.7% dry wt, 0.78 mg at 5 mm body length). These regional differences may have resulted from greater diatom availability in the north and competition from D. polymorpha in the south. Triacylglycerols and phospholipids were the dominant lipid classes in all three regions. Although the mean proportion of triacylglycerols, the energy‐storage lipid, was lower in the south than in the north, regional differences in proportions of lipid classes were not significant.
• 4 Mean lipid content and weight per unit length of Diporeia in the south were lower than values found in the late 1980s prior to the establishment of Dreissena. Mean lipid content of mature individuals is now at levels considered a minimum for successful reproduction.

     

Seasonal changes in the respiratory electron transport system (ETS) and respiration of the zebra mussel,Dreissena polymorpha in Saginaw Bay,Lake Huron

Electron transport system activity (ETS) and respiration rates (R) of the zebra mussel, Dreissena polymorpha, were determined monthly from April to November over 2 years at two sites in Saginaw Bay, Lake Huron. The sites were located in the inner and outer bay and contrasted in food quantity and quality. ETS ranged from 2 to 40 g O₂ mg DW^–1 h^–1 over the study period. Both ETS and respiration were strongly related to temperature, and maximum values were found between June and August. ETS also peaked in June/July when assays were conducted at a constant temperature (25 °C), indicating other factors besides temperature affected metabolic activity. R:ETS ratios decreased with increased temperature at the inner bay site, but trends were minimal at the outer bay site. In late summer, blooms of the cyanophyte Microcystis occurred in the inner bay, likely depressing filtration rates, and leading to lower respiration rates relative to ETS. ETS activity was consistently higher in the outer bay and was likely a result of higher food quality. Despite these spatial differences, annual mean R:ETS ratios varied only from 0.04 to 0.09 at the two sites over the 2-year period. Based on these values, ETS may be useful as an indicator of long-term metabolic activity in annual energy budgets of D. polymorpha. However, food conditions differentially affect respiration relative to ETS, and variability in this ratio must be considered when interested in shorter time scales.     

Filtering Impacts of an Introduced Bivalve (Dreissena polymorpha) in a Shallow Lake: Application of a Hydrodynamic Model

Nonindigenous species may exert strong effects on ecosystem structure and function. The zebra mussel (Dreissena polymorpha) has been attributed with profound changes in invaded ecosystems across eastern North America. We explored vertical profiles of water flow velocity and chlorophyll a concentration in western Lake Erie, over rocky substrates encrusted with Dreissena, to assess the extent to which mussels influence coupling between benthic and pelagic regions of the lake. Flow velocity was always low at surveyed sites (less than or equal to 2.9 cm s^-1) and declined in direct proximity to the lakebed. Mean chlorophyll a concentration was also low (less than 5μg L^-1) at all sites and depths. Chlorophyll a concentration was positively correlated with distance above lakebed and was lowest (0.3μg L^-1) directly adjacent to the lakebed. Spatial patterns of zooplankton grazers could not explain observed vertical gradients in chlorophyll concentration. Hydrodynamic modeling revealed that filtering effects of Dreissena in a nonstratified, shallow basin depend mainly on upstream chlorophyll concentration, intensity of turbulent diffusion, feeding efficiency of the mussel colony, and the distance downstream from the leading edge of the mussel colony. In contrast to widespread perceptions that molluscs reduce phytoplankton concentration only adjacent to the lakebed, modeling scenarios indicated that depletion occurs throughout the water column. Depletion was, however, inversely proportional to distance above the lakebed. Simulation results are consistent with field-based observations made in shallow water habitats populated by large Dreissena populations in the Great Lakes and elsewhere. Results from this study indicate that zebra mussels strongly enhance coupling between pelagic and benthic regions in shallow lakes. Enhanced coupling between these regions explains, in part, high population densities of Dreissena and of many benthic invertebrates in ecosystems invaded by zebra mussels. Received 14 July 1998; accepted 25 March 1999.     

Filtering impacts of larval and sessile zebra mussels (Dreissena polymorpha) in western Lake Erie

Summary We assessed the feeding biology of veliger larvae of the introduced zebra mussel (Dreissena polymorpha Pallas) in laboratory experiments using inert microspheres as food analogues. Mean clearance rate on 2.87-m beads ranged between 247 and 420 L veliger^–1 day^–1. Clearance rate was unrelated to bead concentration up to 100 beads L^–1, but was positively correlated with veliger shell length. Clearance rates of Dreissena veligers are within the range of those reported for marine bivalve veligers of similar size and for herbivorous Great Lakes microzooplankton, but are orders of magnitude lower than those of settled, conspecific adults. The impact of settled zebra mussel grazing activities on phytoplankton stocks may be up to 1162 times greater than that exerted by veliger populations in western Lake Erie. Based on 1990 size-frequency distributions and associated literature-derived clearance rates, reef-associated Dreissena populations in western Lake Erie (mean depth 7 m) possess a tremendous potential to filter the water column (up to 132 m³ m^–2 day^–1) and redirect energy from pelagic to benthic foodwebs. Preliminary analyses indicate that chlorophyll a concentration is strongly depleted (<1 g L^–1) above Dreissena beds in western Lake Erie.     

Spatial and temporal patterns in maternal energetic traits of yellow perch (Perca flavescens) in Lake Erie

1. For many fish species, survival during early life stages is linked to the size and energetic condition of females prior to reproduction. For example, females in good energetic condition are often more fecund and produce larger eggs and offspring than those in poor condition. 2. We measured the characteristics of female yellow perch (Perca flavescens) that may influence annual population fluctuations. From 2005 to 2007, we measured spatial variation in female reproductive traits, such as age, length, mass and energy density (J g^?1) of somatic tissues and ovaries among four spawning aggregations of yellow perch in western and central Lake Erie. 3. Maternal traits, such as somatic energy density and spawner age distribution, differed between the western and central basin, whereas reproductive traits, such as fecundity and ovarian energy density, differed across years. 4. To understand the implications of observed differences in demographic rates (growth and mortality rates) between basins, we developed a deterministic model to simulate the total egg production in the western and central basins under different scenarios of fishing mortality. 5. High growth rates and low mortality rates combined to produce higher modelled estimates of total egg production in the central than in the western basin, and a larger proportion of eggs were produced by old age classes in the central basin than in the western basin. 6. Our results demonstrate that changing harvest levels for populations with different demographic rates can influence total reproductive output through complex interactions between age‐specific mortality, growth and size‐specific fecundity, which has implications for the population dynamics of yellow perch and related species across a broad geographic range.     

Assessing the effects of application time and temperature on the efficacy of hot-water sprays to mitigate fouling by Dreissena polymorpha (zebra mussels Pallas)

Dreissenid mussel (Dreissena polymorpha, Dreissena rostriformis bugensis) expansion into the Western US has renewed interest in hot-water spray mitigation of mussel fouling on boat hulls, trailers, and other equipment. However, the efficacy of hot-water sprays to mitigate dreissenid fouling has not been experimentally assessed. Emersed, adult D. polymorpha were exposed to low-pressure, hot-water sprays at 40, 50, 60, 70, and 80°C for 1, 5, or 10 s. Sprays at ≥60°C for 10 s or 80°C at ≥5 s were 100% lethal. In contrast, 1–10 s exposures did not induce 100% mortality at ≤50°C. The results indicate that mitigation of D. polymorpha fouling, especially in areas protected from the hydraulic impacts of high-pressure sprays requires spray temperatures of > 80°C applied for >5 s or no less than 60°C applied for >10 s. Thus, presently recommended spray temperatures of ≥60°C may not be 100% effective unless applied for >10 s.     

Seasonal Abundance and Species Richness of Birds in Common Reed Habitats in Lake Erie

Abstract: Common reed (Phragmites australis) forms dense stands with deep layers of residual organic matter that negatively affects plant diversity and possibly habitat use by wetland birds. We sought to determine whether seasonal relative abundance and species richness of birds varied among 3 habitat types in Great Lakes coastal wetland complexes recently invaded by common reed. We used fixed-distance point counts to determine species relative abundances and species richness in edge and interior locales within common reed, cattail (Typha spp.), and meadow marsh habitats of various sizes during 2 summers (2001 and 2002) and 1 autumn (2001) at Long Point, Lake Erie, Ontario, Canada. We found that total relative abundance and species richness of birds were greater in common reed habitat compared to cattail or meadow marsh habitats. However, we also found that relative abundance of marsh-nesting birds was greater in meadow marsh habitat than in cattail and common reed during summer. Lastly, we found that, irrespective of habitat type, habitat edges had higher total relative abundance and species richness of birds than did habitat interiors. Our results show that common reed provides suitable habitat for a diversity of landbirds during summer and autumn but only limited habitat for many marsh-nesting birds during summer. Based on these results, we recommend restoration of meadow marsh habitat through reduction of common reed in Great Lakes wetlands where providing habitat for breeding marsh-nesting birds is an objective. Managers also might consider reducing the size of nonnative common reed stands to increase edge effect and use by birds, possibly including wetland birds.     

Seasonal and interannual effects of hypoxia on fish habitat quality in central Lake Erie

1. Hypoxia occurs seasonally in many stratified coastal marine and freshwater ecosystems when bottom dissolved oxygen (DO) concentrations are depleted below 2–3 mg O₂ L^?1. 2. We evaluated the effects of hypoxia on fish habitat quality in the central basin of Lake Erie from 1987 to 2005, using bioenergetic growth rate potential (GRP) as a proxy for habitat quality. We compared the effect of hypoxia on habitat quality of (i) rainbow smelt, Osmerus mordax mordax Mitchill (young‐of‐year, YOY, and adult), a cold‐water planktivore, (ii) emerald shiner, Notropis atherinoides Rafinesque (adult), a warm‐water planktivore, (iii) yellow perch, Perca flavescens Mitchill (YOY and adult), a cool‐water benthopelagic omnivore and (iv) round goby Neogobius melanostomus Pallas (adult) a eurythermal benthivore. Annual thermal and DO profiles were generated from 1D thermal and DO hydrodynamics models developed for Lake Erie’s central basin. 3. Hypoxia occurred annually, typically from mid‐July to mid‐October, which spatially and temporally overlaps with otherwise high benthic habitat quality. Hypoxia reduced the habitat quality across fish species and life stages, but the magnitude of the reduction varied both among and within species because of the differences in tolerance to low DO levels and warm‐water temperatures. 4. Across years, trends in habitat quality mirrored trends in phosphorus concentration and water column oxygen demand in central Lake Erie. The per cent reduction in habitat quality owing to hypoxia was greatest for adult rainbow smelt and round goby (mean: ?35%), followed by adult emerald shiner (mean: ?12%), YOY rainbow smelt (mean: ?10%) and YOY and adult yellow perch (mean: ?8.5%). 5. Our results highlight the importance of differential spatiotemporally interactive effects of DO and temperature on relative fish habitat quality and quantity. These effects have the potential to influence the performance of individual fish species as well as population dynamics, trophic interactions and fish community structure.     

Zebra mussels (Dreissena polymorpha) in Ireland, AFLP-fingerprinting and boat traffic both indicate an origin from Britain

1. The zebra mussel (Dreissena polymorpha) is an aquatic nuisance species that invaded Ireland around 1994. We studied the invasion of the zebra mussel combining field surveys and genetic studies, to determine the origin of invasion and the vector of introduction. 2. Field surveys showed that live zebra mussels, attached to the hulls of pleasure boats, were transported from Britain to Ireland. These boats were lifted from British waters onto trailers, transported to Ireland by ferry and lifted into Irish waters within a day. Length‐frequency distributions of dead and living mussels on one vessel imported 3 months earlier revealed a traumatic occurrence caused by the overland, air‐exposed transportation. Results show that a large number of individuals survived after re‐immersion in Irish waters and continued to grow. 3. Zebra mussels from populations in Ireland, Great Britain, the Netherlands, France and North America, were analysed using amplified fragment length polymorphisms (AFLP)‐fingerprinting to determine the origin of the Irish invasion. Phylogenetic analysis revealed that Irish and British mussels clustered closely together, suggesting an introduction from Britain. 4. Ireland remained un‐invaded by the zebra mussel for more than 150 year. The introduction of the zebra mussel to Ireland occurred following the abolition of value added tax in January 1993 on imported second‐hand boats from the European Union (UK and continental Europe). This, together with a favourable monetary exchange rate at that time, may have increased the risk of invasion of the zebra mussel.     

In situ survival and growth of zebra mussels (Dreissena polymorpha) under chronic hypoxia in a stratified lake

Experiments and field surveys were conducted in Hargus Lake (Ohio, U.S.) to investigate the effect of lake stratification on the survival, growth and distribution of zebra mussels. During the lake stratification period, relatively stable temperature and dissolved oxygen (DO) gradients persisted across the water column, allowing us to examine the chronic effect of hypoxia on zebra mussels. Zebra mussels were incubated in cages and suspended at different depths in the water column at both pelagic (max. depth = 12 m) and littoral (max. depth = 3.5 m) sites from April 18 to September 28, 1994. No mussel survived to the end of the experiment in cages ≥ 5.5 m, whereas the highest survival rate (76%) occurred at 5 m depth where temperature and DO remained fairly stable for at least 3 months. The threshold oxygen level for survival was between 1.0–1.7 mg l^-1 when water temperature was at about 17–18 °C. While zebra mussels′ survival rate was not affected under the sublethal hypoxic conditions, their growth was greatly retarded by poor water quality. The field survey showed that the zebra mussels and macrophytes had about the same distribution and their biomasses were positively related. The percentage of mussels in aggregates increased towards their maximum distribution depth. The maximum distribution depth of the naturally occurring zebra mussels was only 2.8 m, whereas the adult mussels could survive the entire stratification period when being artificially placed on the 3.5 m bottom, and young mussels could colonize the 3.5 m bottom if solid substrates were provided. We conclude that lack of substrate, rather than hypoxia, was the limiting factor of zebra mussel distribution above 5 m depth in Hargus Lake. This revised version was published online in July 2006 with corrections to the Cover Date.     

Horizontal and Vertical Distribution of the Zebra Mussel (Dreissena polymorpha,Pallas) Larvae in the Modrac Reservoir,Bosnia and Herzegovina

In the Modrac Reservoir, loaded with coal separation suspended material, rapid development of the zebra mussel (Dreissena polymorpha) took place from at least 1987. It is probably the first distinguished “infection” for this part of Balkan Peninsula, caused by this organism. Horizontal and vertical distribution of this mussel, at 15 different reservoir profiles, had been investigated.     

Distribution and aspects of population dynamics of the zebra mussel,Dreissena polymorpha (Pallas, 1771), in the lake IJsselmeer area (The Netherlands)

Summary The distribution, settling and growth of the zebra mussel, Dreissena polymorpha Pallas, in Lake IJsselmeer and Lake Markermeer in The Netherlands were studied from 1980 to 1985. In these lakes D. polymorpha is the most important food source for wintering diving ducks. The study was part of an investigation into the carrying capacity of the lakes for these birds.     

Age and growth of round goby Neogobius melanostomus associated with depth and habitat in the western basin of Lake Erie

Round goby Neogobius melanostomus were examined from the Bass Islands area in the western basin of Lake Erie, U.S.A., to determine age and growth correlations. A total of 188 specimens were collected and examined during summer 2011 with 90 aged using scale analysis. Fish were grouped by sex, depth of habitat and habitat type (anthropogenically modified shallows, natural shallows and open lake deep water). Fish ranged from 17 to 117 mm total length (L_T) and 0+ to 3+ years. Males dominated the population (1·94:1) and backcalculated age showed that both sexes grew exponentially, with male growth rate increasing faster than female. Males were significantly larger than females in L_T and mass (both P < 0·001). The relative mass index (W_r) was low for the sampled population (mean ± s.d . = 32·00 ± 26·87 g), implying that the health of the Bass Island area population is very poor when compared with the species throughout its range. This could be due to a lack of food resources related to population size or that the fish is not optimally utilizing the available food resources. In contradiction to these findings, regression slope coefficient (b), calculated using Fulton's condition factor (K) (mean ± s.d . =1·50 ± 0·20), was very low for each habitat, implying a healthy population throughout. This seemingly opposite effect may be due to more individuals per unit area in shallow waters, which would cause increased competition for resources. Poor condition may indicate that the Lake Erie population has reached saturation or may reflect indirect fitness costs associated with increasing anoxic or hypoxic hypolimnion conditions.     

Sympatric bromeliad species (Pitcairnia spp.) facilitate tests of mechanisms involved in species cohesion and reproductive isolation in Neotropical inselbergs

The roles of intra- and interspecific gene flow in speciation and species evolution are topics of great current interest in molecular ecology and evolutionary biology. Recent modelling studies call for new empirical data to test hypotheses arising from the recent shift from a 'whole-genome reproductive isolation' view to a 'genic' view of species and speciation. Particularly scarce (and thus of particular interest) are molecular genetic data on recently radiated, naturally hybridizing species in strongly structured and species-rich environments. Here, we studied four sympatric plant species (Pitcairnia spp.; Bromeliaceae) adapted to Neotropical inselbergs (isolated outcrops resembling habitat 'islands' in tropical rainforests) using nuclear and plastid DNA. Patterns of plastid DNA haplotype sharing and nuclear genomic admixture suggest the presence of both, incomplete lineage sorting and interspecific gene flow over extended periods of time. Integrity and cohesion of inselberg species of Pitcairnia are maintained despite introgression and in the face of extremely low within-species migration rates (N(e)m < 1 migrant per generation). Cross-evaluation of our genetic data against published pollination experiments indicate that species integrity is maintained by the simultaneous action of multiple prezygotic barriers, including flowering phenology, pollinator isolation and divergent mating systems. Postzygotic Bateson-Dobzhansky-Muller incompatibilities appear to contribute to isolation, as suggested by asymmetric introgression rates of single loci. Our results suggest that incomplete lineage sorting, hybridization and introgression form integral aspects of adaptive radiation in Neotropical inselberg 'archipelagos'. Inselbergs with multiple closely related co-occurring species should be of special interest to students of speciation in mountain systems, and to ongoing conservation programmes in the Atlantic Rainforest biodiversity hotspot.     

Field and laboratory studies of Ophryoglena sp. (Ciliata: Ophryoglenidae) infection in zebra mussels,Dreissena polymorpha (Bivalvia: Dreissenidae)

This study, conducted in the Dnieper-Bug Canal in Belarus, is the first to monitor the seasonal (June-November) dynamics of infection with the parasitic ciliate Ophryoglena sp. in a zebra mussel (Dreissena polymorpha) population. Mean population prevalence and intensity of infection varied, respectively, from 11 to 62% and from 0.9 to 24.1 ciliates/mussel. Mean prevalence was highly correlated with mussel length in mussels <20 mm (R(2)=0.97) and was lower in larger mussels. Mean infection intensity in mussels 1-25 mm long was similarly correlated with their size (R=0.98), reached a maximum in the 20-25 mm size-class, and then sharply decreased, thus providing evidence, albeit limited, that high intensity of infection might be lethal. Transinfection of zebra mussels by Ophryoglena sp. was achieved in the laboratory-a first for a protozoan parasite of D. polymorpha; from an initial complete lack of infection, mean prevalence and intensity rose, respectively, to 86.7% and 8.3 ciliates/mussel.     

Recolonization and possible recovery of Burrowing Mayflies (Ephemeroptera: Ephemeridae: Hexagenia spp.) in Lake Erie of the Laurentian Great Lakes

Burrowing mayflies of the genus Hexagenia spp. were widely distributed (ca. 80% of sites) and abundant (ca. 160 nymphs/m²) in the western basin of Lake Erie of the Laurentian Great Lakes in 1929–1930, prior to a period of anoxia in the mid 1950s. Nymphs were absent or rare in the basin between 1961 and 1973–1975. In 1979–1991, nymphs were infrequently found (13–46% of sites) in low abundance (3–40 nymphs/m²) near shore (<7.5 km from shore), but were absent or rare offshore (0–7% of sites at 0–1 nymphs/m²). Increased abundance occurred offshore between 1991 (0% of sites) and 1993 (52% of sites at 7/m²). Annual sampling, beginning in 1995, indicates that nymphs increased in both nearshore and offshore waters. By 1997, nymphs were found throughout the lake (88% of sites) at a mean density 40-fold greater (392/m²) than that observed in 1993 (11/m²). In 1998, the distribution of nymphs remained the same as 1997 (88% of sites) but density declined 3-fold (392 to 134/m²). These data indicate that mayflies have recolonized sediments of western Lake Erie and that their abundance may be similar to levels observed before their disappearance in the mid 1950s. However, prior to the mid 1950s, densities were greater in offshore than nearshore waters, but between 1979 and 1998 greater densities occurred near shore than offshore. In addition, there were two areas in the 1990s where low densities consistently occurred. Therefore, recovery of nymphs in western Lake Erie may not have been complete in 1998. At present we do not know the cause for the sudden recolonization of nymphs in large portions of western Lake Erie. Undoubtedly, pollution-abatement programs contributed to improved conditions that would have ultimately led to mayfly recovery in the future. However, the explosive growth of the exotic zebra mussel, Dreissena polymorpha, undoubtedly diverted plankton foods to bottom substrates which could have increased the speed at which Hexagenia spp. nymphs recolonized sediments in western Lake Erie in the 1990s.     

The life-history strategy of deepwater sculpin, Myoxocephalus thompsoni (Girard), in Lake Michigan: dispersal and settlement patterns during the first year of life

Deepwater sculpin, Myoxocephalus thompsoni (Girard), were sampled from six stations from the 15–100 m depth contours in Lake Michigan between April 1983 and July 1984. In south-eastern Lake Michigan M. thompsoni lay benthic eggs in offshore waters, which hatch between November and August, with peak hatching in March. Abundance of larvae in pelagic samples was higher offshore than inshore, but larval size was greater and development more advanced at inshore stations, indicating an inshore movement after hatching. Larvae reached metamorphosis at 20 mm and settled to the bottom beginning in July. Pelagic larvae 20–40 mm were found in the lower water column at all stations, but newly settled individuals were only captured with bottom trawls at inshore locations (≤60 m depth). Data from ichthyoplankton and bottom trawl samples in 1983 and 1984 indicated that locations for successful settlement of larvae to the bottom extended only as deep as the shallowest fringe of the adult population (> 50 m in 1983). In 1983, maximum density of larvae reached 0.4 individuals m⁻³ by June. Survival from the pelagic larval stage to the demersal young-of-year stage in 1983/1984 was c . 0.1–0.4%. The specific mechanism of mortality at the time of transition to a demersal habit has not been determined.