Patricia Chow-Fraser

Long-term response of the biotic community to fluctuating water levels and changes in water quality in Cootes Paradise Marsh, a degraded coastal wetland of Lake Ontario

During the early 1900s, more than 90% of the surface area of Cootes Paradise Marsh was covered with emergent vegetation; currently, less than 15% of the surface is covered with aquatic vegetation and the remainder is wind-swept, turbid, open water. The loss of emergent cover is significantly correlated with mean annual water levels that increased more than 1.5 m over the past 60 years. Species diversity and the percent cover of the submerged macrophtye community also declined dramatically after the 1940s, coincident with decreased water clarity and increased nutrients from pollution by sewage and stormwater effluent. Phosphorus levels in the marsh dropped ten-fold after the sewage plant was upgraded to a tertiary-treatment facility in 1978; however, there was no measurable improvement in water clarity, in spite of a decrease in chlorophyll concentrations. Long-term changes in the composition of the planktonic, benthic and fish communities accompanied changes in water clarity, nutrient status and macrophyte cover. Phytoplankton changed from a community dominated by diverse taxa of green algae and diatoms during the 1940s, to a less diverse community dominated by a few taxa of green and blue-green algae in the 1970s, then to a much more diverse community recently, including many taxa of green algae, diatoms and chrysophytes; however, because water turbidity continues to be high, and algae tolerant of low light levels are now very abundant. Daphnia, which were prominent during the 1940s (especially in the vegetated sites) were replaced in the 1970s by smaller zooplankton such as the cladoceran, Bosmina, and several rotifer species including Brachionus, Asplanchna and Keratella. In the recent survey conducted in 1993 and 1994, small-bodied forms still dominate the turbid open-water areas, while medium-sized cladocerans such as Moina were common near macrophyte beds. Generally, total herbivorous zooplankton biomass tended to be highest next to Typha beds and declined with increasing distance from the plants. Conversely, biomass of edible algae at these sites increased with distance from the macrophytes. Species diversity of aquatic insects declined dramatically over the past 40 years, from 57 genera (23 families and 6 orders) in 1948, to 9 genera (6 families and 3 orders) in 1978, to only 5 genera (3 families and 2 orders) in 1995. The diverse benthic community present 5 decades ago has now been replaced by a community consisting primarily of chironomid larvae, oligochaetes and other worms associated with low-oxygen environments. These successional changes illustrate the impact of natural (fluctuating water levels) and anthropogenic (deterioration in water quality) stressors on the character of the biotic communities, and reveal the complex interactions among the various trophic levels and the abiotic environment as degradation and remediation proceeded.

Development and use of a zooplankton index of wetland quality in the Laurentian Great Lakes Basin

Recent interest in biological monitoring as an ecosystem assessment tool has stimulated the development of a number of biotic indices designed to aid in the evaluation of ecosystem integrity; however, zooplankton have rarely been included in biomonitoring schemes. We developed a wetland zooplankton index (WZI) based on water quality and zooplankton associations with aquatic vegetation (emergent, submergent, and floating-leaf) that could be used to assess wetland quality, in particular in marshes of the Laurentian Great Lakes basin. Seventy coastal and inland marshes were sampled during 1995–2000; these ranged from pristine, macrophyte-dominated systems, to highly degraded systems containing only a fringe of emergent vegetation. The index was developed based on the results of a partial canonical correspondence analysis (pCCA), which indicated that plant-associated taxa such as chydorid and macrothricid cladocerans were common in high-quality wetlands, while more open-water, pollution-tolerant taxa (e.g., B...

Use and Development of the Wetland Macrophyte Index to Detect Water Quality Impairment in Fish Habitat of Great Lakes Coastal Marshes

ABSTRACT Indices have been developed with invertebrates, fish, and water quality parameters to detect the impact of human disturbance on coastal wetlands, but a macrophyte index of fish habitat for the Great Lakes does not currently exist. Because wetland macrophytes are directly influenced by water quality, any impairment in wetland quality should be reflected by taxonomic composition of the aquatic plant community. We developed a wetland macrophyte index (WMI) with plant presence/absence data for 127 coastal wetlands (154 wetland-years) from all five Great Lakes, using results of a canonical correspondence analysis (CCA) to ordinate plant species along a water quality gradient (CCA axis 1). We validated the WMI with data collected before and after the implementation of remedial actions plans (RAPs) in Sturgeon Bay (Severn Sound) and Cootes Paradise Marsh. Consistent with predictions, WMI scores for Sturgeon Bay were significantly higher after the implementation of the RAP. Historical data from Cootes Paradise Marsh were used to track the declining condition of the plant community from the 1940s to 1990s. Subsequently, when remedial actions had been implemented in 1997, the calculated WMI scores showed improvement, but when the presence of exotic species (WMIadj) was accounted for, improvements in ecological integrity of the aquatic-plant community were no longer evident. We show how WMI scores can be used by environmental agencies to assess the historic, current, and future ecological status of wetland ecosystems in two Canadian national parks, Point Pelee National Park (PPNP) and Fathom Five National Marine Park (FFNMP).

Carp Exclusion, Food-web Interactions, and the Restoration of Cootes Paradise Marsh

Carp were excluded from Cootes Paradise Marsh (Lake Ontario) in 1997 in order to improve water clarity and promote submerged plant growth. On average, turbidity at open water and vegetated areas was reduced by 40 and 60 percent, respectively, following carp exclusion. However, responses by plants and other trophic levels have been both spatially and temporally variable due in part to annual variation in environmental conditions and fish-zooplankton interactions. In 1997, an unusually cool spring delayed the migration of spawning fish, including a usually abundant planktivore population, into the marsh. This had three main effects: 1) large Daphnia were released from predation in early summer and reached unprecedented numbers (530 Daphnia/L) in open water areas, 2) despite the lack of vegetated marsh habitat, larval fish production was high due to reduced predation and abundant zooplankton prey, and 3) zooplankton grazing initiated a spring clear-water phase which, together with carp exclusion, promoted submerged plant growth in shallow areas previously devoid of vegetation. These newly vegetated areas showed the greatest improvements in clarity and macrophyte growth in the first 2 years following exclusion. Currently, however, the future of the biomanipulation remains uncertain, due in part to natural climatic confounding factors, and further remedial actions will be required before this wetland represents a stable, clear-water, macrophyte dominated state.

Seasonal, Interannual, and Spatial Variability in the Concentrations of Total Suspended Solids in a Degraded Coastal Wetland of Lake Ontario

A 4-year (1993 through 1996) monitoring program examined the distribution of total suspended solids (TSS) in Cootes Paradise Marsh, a shallow (mean depth of 70 cm), degraded, drownedrivermouth marsh of Lake Ontario. Monthly meteorological and hydrographical data from 1986 through 1996 revealed a hydrologically dynamic system that exhibited large seasonal and interannual variation with respect to precipitation amount, discharge volume, and water levels; the prevailing winds were shown to be oriented along the length of the marsh. Interannual variation in TSS concentrations was inversely related to mean seasonal water levels that fluctuated 45 cm over the 11 years. In a stepwise regression analysis, planktonic chlorophyll-a concentration only explained 2% of the variation in TSS, while inorganic and non-algal organic solids explained 70% and 18%, respectively. Mean seasonal water turbidity increased significantly with mean seasonal wind speed at 17 sampling stations during 1993 and 1994. Runoff from a summer rainstorm more than doubled water turbidities at the mouth of all three creeks over the first 36 hours. In enclosure experiments, water turbidity increased proportionately with biomass of benthivorous fish (especially common carp, Cyprinus carpio). When wind and carp disturbance were compared simultaneously in the field, wind speed accounted for 41% of the variation in turbidity while presence of carp explained an additional 21%. The overall temporal and spatial distribution of TSS in the marsh reflected changes in water level, wind activities, onset of rain events, and fish disturbance that acted in concert to keep Cootes Paradise Marsh extremely turbid throughout the summer.

A conceptual ecological model to aid restoration of Cootes Paradise Marsh, a degraded coastal wetland of Lake Ontario, Canada

An ecological model is derived from recent studies, based on 60 years of empirical observations and experimental data, that conceptualizes how Cootes Paradise Marsh was transformed from a lush emergent marsh with considerable ecological diversity in all trophic levels, to one that is currently turbid, devoid of vegetation, and dominated by a few exotic plant and fish species. This conceptual model contains 17 key components that interact and contribute to the overall unhealthy state of the marsh. The most influential component is high water level which caused the initial loss of emergent vegetation in the 1940s and 1950s. In the absence of plants to attenuate sediment and assimilate nutrients, the marsh became turbid and windswept, and this led to the disappearance of submergent vegetation over the next two decades. Currently, high water turbidity is being maintained by wind re-suspension, high sediment loading from the watershed during the summer, high algal biomass resulting from excessive nutrient loads from sewage effluent and surface runoff, and the feeding and spawning activities of a very large population of common carp ( Cyprinus carpio). Due to vegetation loss, the substrate has become mostly loose sediment that is no longer suitable for the diverse assemblage of aquatic insect larvae that lived on the plants and detrital material in the 1940s. Benthic grazers have been kept in low abundances due to predation by benthivorous carp; consequently, epiphytic algae have proliferated and further contribute to light limitation of macrophytes. High nutrient loadings contribute to high diurnal fluxes in dissolved oxygen levels that tend to select against less tolerant organisms such as insect larvae (other than chironomids) and piscivores (northern pike and largemouth bass). Without piscivores in the marsh, the planktivores have become dominant and have virtually eliminated all of the large herbivorous zooplankton (e.g., Daphnia), except for a few pockets in the marsh inlets close to residual macrophyte beds. Because of the dominance of small-bodied inefficient grazers (rotifers and small cladocerans), algal biomass is high, and the community has a large proportion of heterotrophic forms that tolerate low light environments. This ecological model suggests that the current turbid un-vegetated state of Cootes Paradise may be very stable. It will persist as long as water levels remain unfavorable for natural re-colonization by the emergent flora, and/or water turbidities remain sufficiently high to suppress the growth of submergent vegetation. Using this conceptual model, I developed a model of how Cootes Paradise Marsh may have functioned as a healthy marsh prior to the 1940s, and use these models as a basis to explore a number of restoration and management options and discuss their implications on the aquatic foodweb.

Ecosystem response to changes in water level of Lake Ontario marshes: lessons from the restoration of Cootes Paradise Marsh

A general understanding of how aquatic vegetation responds to water-level fluctuations is needed to guide restoration of Great Lakes coastal wetlands because inter-annual and seasonal variations often confound effects of costly remedial actions. In 1997, common carp (Cyprinus carpio) was removed from Cootes Paradise Marsh (L. Ontario) to reduce sediment resuspension and bioturbation, and thus regenerate marsh plants that had declined dramatically since the 1930s. Data from 1934 to 1993 were re-assembled from the literature to relate percentage cover of emergent vegetation to mean summer water level. A non-linear regression equation explained close to 90% of the variation compared with 80% for a non-linear equation, and this trend was confirmed for the dominant species, Typha latifolia. A modest recovery of emergent vegetation in 1999 following carp exclusion could have been predicted from declining water level alone, without invoking any effects of the biomanipulation. An unusually cool spring in 1997 delayed the migration of spawning planktivores into the marsh. This resulted in a grazer-mediated clear-water phase that coincided with a resurgence of the submersed aquatic vegetation (SAV) community in 1997, which declined again in 1999 when low water levels occurred. Even though decrease in water level was significantly related to increased suspended solids and greater light attenuation, light conditions appeared to have been adequate in marsh embayments to support SAV growth, according to a published relationship between maximum depth of SAV colonization and light extinction coefficient. I suggest that wave disturbance and propagule burial associated with shallow water depths may have been the main reasons for the decline of the SAV in 1999 and 2000.

Application of the Wetland Fish Index to Northern Great Lakes Marshes with Emphasis on Georgian Bay Coastal Wetlands

ABSTRACT The wetland fish index (WFI), a published indicator of wetland condition that ranks wetlands based on tolerance of fish species to degraded water-quality conditions, had been developed with data from 40 wetlands located exclusively in the southern portion of the Great Lakes basin (Erie, Ontario, and Michigan). No data had been included from wetlands of the northern Great Lakes (Superior and Huron) and especially those of eastern and northern Georgian Bay, where many wetlands are still unaffected by human activities. We demonstrate why application of the WFI for the lower lakes (WFILower) can yield biased scores when applied to data for upper lakes wetlands. We then develop a basin-wide index to include data from 60 other coastal wetlands located in the northern portion of the basin, including 32 from Georgian Bay. Inclusion of northern sites in development of a basin-wide WFI (WFIBasin) still produced index scores that were positively correlated with water-quality conditions as indicated by water quality index scores. We explain why use of the basin-wide WFI is better than one developed specifically for upper lakes (WFIUpper). Overall, WFIBasin scores were higher in the northern lakes (Superior 3.49, Georgian Bay 3.67, Huron 3.62) than in the southern lakes (Michigan 3.33, Erie 3.12, Ontario 3.09). WFI scores are only minimally affected by inter-annual variation, which allows for its use for long-term monitoring. We recommend that the WFIBasin be used when managers need to manage at a scale across the entire Great Lakes basin.

Type-3 functional response in limnetic suspension-feeders, as demonstrated by in situ grazing rates

Field-measured grazing rates (ml/animal/d) of cladocerans (mostly daphniids) and diaptomids were assembled from various published studies and plotted as a function of corresponding phytoplankton concentration (µg l−1 f.w.). Filtering rates of both zooplankton groups initially increased with seston concentration until maximal grazing rates were observed at approximately 4 × 102 and 1 × 102 µg l−1 for cladocerans and copepods, respectively; at higher algal concentrations, filtering rates of both declined as a function of food concentration. The shape of these curves are most consistent with Hollings (1966) Type 3 functional response.We found little support for the Type 3 functional response in published laboratory studies of Daphnia; most investigators report either a Type 1 or Type 2 response. The one study in which the Type 3 response was observed involved experiments where animals were acclimated at low food concentrations for 24 h, whereas those studies associated with response Types 1 or 2 had acclimation periods of only 1 to 3 h. We therefore assembled relevant data from the literature to examine the effect of acclimation period on the feeding rates of Daphnia at low food concentrations. In the absence of any acclimation, animals filtered at extremely low rates. After 2 h of acclimation, however, filtering rates increased 4 to 5-fold but declined again with longer durations; after > 70 h of pre-conditioning, filtering rates were almost as low as they had been with no acclimation.We also found little support for the Type 3 functional response in published studies of copepods. The only study associated with a Type 3 response involved a marine copepod that had been subjected to a starvation period of 48 h; however, an analysis of the effects of acclimation period did not yield conclusive evidence that filtering rates of freshwater copepods (Diaptomus and Eudiaptomus) decrease significantly with acclimation duration.The low filtering rates associated with long acclimation periods in laboratory experiments appears to be a direct result of animals becoming emaciated from prolonged exposure to low food concentrations, a situation which renders them incapable of high filtering rates. This may explain the Type 3 functional response for field cladocerans, since zooplankton in food-limiting situations are constantly exposed to low food concentrations, and would therefore have low body carbon and consequently less energy to filter-feed. We cannot, however, use this to explain the Type 3 response for field diaptomids, since copepods in the laboratory did not appear to lose body carbon even after 72 h of feeding at very low food levels, and there was inconclusive evidence that either Diaptomus or Eudiaptomus decrease their filtering rates with acclimation period.Although Incipient Limiting Concentrations (ILC) for Daphnia ranged from 1 to 8.5 × 103 µg 1−1, more than half of these fell between 1 and 3 × 103 µg l−1, bracketing the value of 2.7 × 102 µg l−1 for field cladocerans. There was, however, a great deal of variation in reported maximum ingestion rates (MIR), maximum filtering rates (MFR) and ILC values for Daphnia magna. ILC values from the few laboratory studies of freshwater copepods ranged between 0.5 to 2.8 × 103 µg 1−1, and was higher than the ILC value of approximately 0.2 × 103 µg l−1 calculated for field populations of D. minutus. Generally, there was considerable agreement among laboratory studies regarding the shape of grazing-rate and ingestion-rate curves when data were converted to similar units and presented on standardized scales.

Relative importance of macrophyte community versus water quality variables for predicting fish assemblages in coastal wetlands of the Laurentian Great Lakes

ABSTRACT Fish have been shown to be sensitive indicators of environmental quality in Great Lakes coastal wetlands. Fish composition also reflects aquatic macrophyte communities, which provide them with critical habitat. Although investigators have shown that the relationship between water quality and fish community structure can be used to indicate wetland health, we speculate that this relationship is a result of the stronger, more direct relationship between water quality and macrophytes, together with the ensuing interconnection between macrophyte and fish assemblages. In this study, we use data collected from 115 Great Lakes coastal marshes to test the hypothesis that plants are better predictors of fish species composition than is water quality. First we use canonical correspondence analysis (CCA) to conduct an ordination of the fish community constrained by water quality parameters. We then use co-correspondence analysis (COCA) to conduct a direct ordination of the fish community with the plant community data. By comparing the statistic ‘percent fit,’ which refers to the cumulative percentage variance of the species data, we show that plants are consistently better predictors of the fish community than are water quality variables in three separate trials: all wetlands in the Great Lakes basin (whole: 21.2% vs 14.0%; n = 60), all wetlands in Lakes Huron and Superior (Upper: 20.3% vs 18.8%; n = 32), and all wetlands in Georgian Bay and the North Channel (Georgian Bay: 18% vs 17%; n = 70). This is the largest study to directly examine plant-fish interactions in wetlands of the Great Lakes basin.