Coping with warmer, large rivers: a field experiment on potential range expansion of northern quagga mussels (Dreissena bugensis)

SUMMARY 1. Zebra mussels ( Dreissena polymorpha ) have established a much greater range in North America and Europe than quagga mussels ( D. bugensis ), which occupy a very similar niche.
2. We hypothesised that quaggas are physiologically capable of sustaining populations in warmer rivers currently occupied only by zebra mussels and that unidentified, non-physiological factors account for their more limited distribution.
3. Growth and survival of individually tagged mussels (976 D. bugensis from Lake Erie; 2625 D. polymorpha from Lake Erie and the Ohio River) were recorded monthly for up to 15 months in an outdoor stream mesocosm receiving unfiltered water from the Ohio River.
4. Extreme temperatures stressed both species; but in contrast to several previous laboratory studies, quaggas survived high temperatures better than zebra mussels. We suspect this was the result of species-specific differences in their ability to obtain, assimilate and/or catabolise food at high, sublethal temperatures.
5. A unimodal growth pattern was observed in both species, with the highest growth rates from late spring to early autumn.
6. Our survival and growth data suggest that quaggas are not physiologically limited from expanding southward.
7. While lacking definitive proof that dreissenid populations in rivers are ecologically sustainable without upstream lentic ecosystems and/or unintended human intervention, we suggest that complex river currents and upstream retentive and highly productive slackwater habitats in rivers may help sustain downstream populations of these meroplanktonic, dreissenid mussels.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.     

Wetlands as barriers: effects of vegetated waterways on downstream dispersal of zebra mussels

1. Stream flow is a major vector for zebra mussel spread among inland lakes. Veligers have been found tens to hundreds of km from upstream source lakes in unvegetated stream and river systems. It has been suggested, however, that the downstream transport of zebra mussels is restricted by wetland ecosystems. We hypothesized that vegetated waterways, (i.e. wetland streams) would hinder the downstream dispersal of zebra mussels in connected inland lake systems. 2. Veliger abundance, recruitment and adult mussels were surveyed in four lake‐wetland systems in southeastern Michigan, U.S.A. from May to August 2006. Sampling was conducted downstream of the lakes invaded by zebra mussels, beginning at the upstream edge of aquatic vegetation and continuing downstream through the wetland streams. 3. Veliger abundance decreased rapidly in vegetated waterways compared to previously reported rates of decrease in non‐vegetated streams. Veligers were rarely found more than 1 km downstream from where vegetation began. Newly recruited individuals and adults were extremely rare beyond open water in the wetland systems. 4. Densely vegetated aquatic ecosystems limit the dispersal of zebra mussels downstream from invaded sources. Natural, remediated and constructed wetlands may therefore serve as a protective barrier to help prevent the spread of zebra mussels and other aquatic invasive species to other lakes and ecosystems.     

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.     

Growth and reproductive cycle of the zebra mussel in the River Rhine as studied in a river bypass

In a Rhine River bypass built at a water pollution control station, the growth and reproductive cycle of the zebra mussel (Dreissena polymorpha) were studied over a period of nearly 3 years. The experimental setup offered the possibility to manipulate the temperature of the continuously passing river water and to calculate influences of temperature and food on the growth of individually marked mussels. Shell growth rates were a function of algal density (measured as chlorophyll a). This function followed saturation curves with temperature-dependent plateau levels, and saturation occurred at values above 40 μg/l chlorophyll a. The relationship between shell growth rate and temperature followed the courses of bell-shaped curves with algal-food-dependent heights. No clear correlations could be found between shell growth and other environmental factors such as dissolved organic matter, total organic content, seston, and turbidity. Two spawning periods per year characterized the reproductive cycle of the zebra mussels at the study site. The first one covered nearly 3 months (April to July) comprising several spawning events. The second shorter one occurred in August. Young mussels of the first spawning period settled at the end of May, and they were able to reproduce a few months later (end of August) when their shell length exceeded 9 mm. This first spawning period of the young-of-the-years coincided with the second spawning period of the older generations. Through the combined analysis of the results of both body weight determinations and shell growth experiments, it was possible to model mussel growth over a period of nearly 3 successive years, and to suggest the presence of an endogenous factor that might be involved in determining the start of the annual shell growth period. Received: 8 April 1997 / Accepted: 3 November 1997     

Exotic plant species of the St Lawrence River wetlands: a spatial and historical analysis

Aim To evaluate the importance (number of species, plant cover) of the exotic flora in seven well‐defined sectors of one of the most important transportation waterways in North America. To determine the impact of exotic species on wetland plant diversity and reconstruct the spread of some invasive species. Location St Lawrence River, southern Québec. Methods The exotic flora (vascular plants) of wetlands bordering the St Lawrence River was studied using 713 sampling stations (25 m²) along a 560‐km long corridor. Results Exotic species represent 13.7% of the vascular flora of the St Lawrence wetlands. The relative plant cover occupied by exotic species is high in some of the fluvial sectors (42–44%), but low (6–10%) in the estuarine sectors. Wetlands (marshes) surrounding islands were particularly susceptible to invasion by exotic plants. Historical, abiotic and landscape factors may explain the differences observed between sites. Purple loosestrife (Lythrum salicaria L.) is the most common exotic species of the St Lawrence wetlands, but other species, namely flowering‐rush (Butomus umbellatus L.) and reed canary grass (Phalaris arundinacea L.) are much more invasive. There is no linear relationship between the exotic species cover and the diversity of wetland plants; low diversity sites can be dominated by either exotic or native plant species. In the other sites, exotic species generally have little impact on plant communities and can contribute to increase diversity. Common reed (Phragmites australis (Cav.) Trin. ex Steudel) and reed canary grass, both considered as exotic species in this study, clearly have a stronger impact on plant diversity than flowering‐rush and purple loosestrife. Main conclusions This study shows that the global impact of an invader cannot be adequately evaluated with only a few highly invaded sites. While nationwide strategies have been developed to control exotic species, large surveys are essential to adapt them to regional particularities.     

Predicting the impacts of an introduced species from its invasion history: an empirical approach applied to zebra mussel invasions

1. Quantitative models of impact are lacking for the vast majority of known invasive species, particularly in aquatic ecosystems. Consequently, managers lack predictive tools to help them prioritise invasion threats and decide where they can most effectively allocate limited resources. Predictive tools would also enhance the accuracy of water quality assessments, so that impacts caused by an invader are not erroneously attributed to other anthropogenic stressors. 2. The invasion history of a species is a valuable guide for predicting the consequences of its introduction into a new environment. Regression analysis of data from multiple invaded sites can generate empirical models of impact, as is shown here for the zebra mussel Dreissena polymorpha. Dreissena's impacts on benthic invertebrate abundance and diversity follow predictable patterns that are robust across a range of habitat types and geographic regions. Similar empirical models could be developed for other invaders with a documented invasion history. 3. Because an invader's impact is correlated with its abundance, a surrogate model may be generated (when impact data are unavailable) by relating the invader's abundance to environmental variables. Such a model could help anticipate which habitats will be most affected by invasion. Lack of precision should not be a deterrent to developing predictive models where none exist. Crude predictions can be refined as additional data become available. Empirical modelling is a highly informative and inexpensive, but underused, approach in the management of aquatic invasive species.     

Phenotypic plasticity and priority rules for energy allocation in a freshwater clam: a field experiment

We studied resource allocation among maintenance, reproduction and growth in the freshwater clam Anodonta piscinalis. Recent theoretical and empirical studies imply that organisms with indeterminate growth may have priority rules for energy allocation. That being so, the traits involved should potentially be capable of considerable phenotypic modulation, as a mechanism to adjust allocation. We tested this hypothesis using a 1-year reciprocal transplant experiment at six sites. Experimental clams were caged at higher than natural densities in order to detect any phenotypic modulation of the traits and discover the putative priority rules in energy allocation. We recorded the survival and shell growth of clams during the experiment, and the reproductive output, somatic mass and fat content of clams at the end of the experiment. Shell growth, somatic mass, and the reproductive output of females varied more among transplant sites than among the populations of origin, suggesting a high capacity for phenotypic modulation. However, the reproductive investment, somatic mass and shell growth were also affected by origin; clams from productive habitats invested more in reproduction and were heavier. In comparison to undisturbed clams, the reproductive output of the experimental clams was similar and their fat content was higher, whereas their shell growth was considerably slower and their somatic mass lower. These results suggests that when resources are limiting (due to high density) reproductive allocation overrides allocation to somatic growth. The highest mortality during the experiment coincided with the period of reproductive stress in the spring. Additionally, the proportion of reproducing females was lower in those transplant groups where the survival rate was lowest, suggesting that maintenance allocation overrides allocation to reproduction when available resources are scarce. The results of this field experiment support theoretical predictions and results of previous laboratory experiments that suggest that there are priority rules for energy allocation in organisms with indeterminate growth.     

Doubly uniparental inheritance of mitochondrial DNA in freshwater mussels: History and status of the European species

Doubly uniparental inheritance (DUI) is a mode of inheriting mitochondrial DNA that is distinct from strictly maternal inheritance. It has been described in nine and three families of marine and freshwater mussels, respectively, including the European margaritiferids and unionids. Among the 16 freshwater species of Unionida inhabiting Europe, DUI has been described in 9 species of dioecious mussels and was absent from a single hermaphroditic species and from secondary hermaphroditic specimens. The DUI freshwater mussels include two vastly genetically different mitochondrial genomes: maternal (F genome) and paternal (M genome), which coexist within the same specimen but in different tissues. The F genome is present in all female tissues and somatic male tissues. It is inherited in the typical, maternal, manner. Conversely, the M genome is located primarily in the male gonads and generative cells, and is inherited paternally. Dioecious Unionidae display unique characteristics that have been interrelated for over 200 million years: a high fidelity of the transmission of the F and M genomes in DUI and two paths of spermatogenesis–the typical path that produces sperm cells containing mitochondria with the F genome and the atypical path that produces sperm cells with the M genome. The mitogenomes of freshwater mussels display unique features that are not present in any other animal, that is, an additional, gender-specific gene and an elongated cox2 gene occurring exclusively in the M genome. These features mean that the mitochondria, in addition to their basic function of producing energy, also may take part in determining sex in these dioecious organisms.     

Run timing of pelagic fishes in Gulf of St Lawrence: area and species effects

Run timings of four ubiquitous pelagic fishes in the Gulf of St Lawrence were compared for area and year effects. Run timing of alewife, Alosa pseudoharengus (Wilson), smelt, Osmerus mordax (Mitchill), and Atlantic salmon, Salmo salar L., was estimated at index rivers, and time of arrival of herring, Clupea harengus L., on spawning grounds was estimated from gillnet fisheries located around the Gulf, 1978–87. Time of pelagic fish migrations had strong species and area effects, but weak year effects. The species had their own unique migration times and durations, and could be divided into two groups: herring and smelt, which arrived in all areas of the Gulf at the same time; and alewife and salmon, which had significant area effects. Environmental data were not correlated with run timing. There was little annual variation in run timing among early-run species; only alewife had significant year effects. There was annual variation in the timing of late-run species, but it was not correlated with proximate environmental factors. Because run timing is an easily defined stock characteristic, observations over large areas and many years might be useful for understanding the impact of large-scale environmental changes on natural populations.     

Genetic variability in Irish populations of the invasive zebra mussel, Dreissena polymorpha: discordant estimates of population differentiation from allozymes and microsatellites

1. The recent arrival and explosive spread of the zebra mussel, Dreissena polymorpha (Pallas), in Ireland provided a rare opportunity to study the population genetics of an invasive species.
2. Eight polymorphic allozyme loci ( ACO-1, ACO-2 , EST-D, GPI, IDH-2, MDH, OPDH and PGM ) were used to investigate genetic diversity and population structure in five Irish populations, and the results were compared with those from a previous microsatellite study on the same samples.
3. The mean number of alleles per locus (2.7 ± 0.1) was similar to the mean for the same loci in European populations, suggesting that Irish founder populations were large and/or multiple colonization events took place after foundation. A deficiency of heterozygotes was observed in all populations, but was uneven across loci.
4. Pairwise comparisons, using Fisher's exact tests and F _ST values, revealed significant genetic differentiation among populations. The overall multilocus F _ST estimate was 0.118 ± 0.045, which contrasted with an estimate of 0.015 ± 0.007 from five microsatellite loci on the same samples in a previous study.
5. Assuming that microsatellites can be used as a neutral baseline, the discordant results from allozymes and microsatellites suggest that selection may be acting on some allozyme loci, specifically ACO-1, ACO-2 , IDH-2 and MDH, which contributed most to the significant differentiation between samples.     

Identifying freshwater mussels (Unionoida) and parasitic glochidia larvae from host fish gills: a molecular key to the North and Central European species

Freshwater mussels (order Unionoida) represent one of the most severely endangered groups of animals due to habitat destruction, introduction of nonnative species, and loss of host fishes, which their larvae (glochidia) are obligate parasites on. Conservation efforts such as habitat restoration or restocking of host populations are currently hampered by difficulties in unionoid species identification by morphological means. Here we present the first complete molecular identification key for all seven indigenous North and Central European unionoid species and the nonnative Sinanodonta woodiana, facilitating quick, low-cost, and reliable identification of adult and larval specimens. Application of this restriction fragment length polymorphisms (RFLP) key resulted in 100% accurate assignment of 90 adult specimens from across the region by digestion of partial ITS-1 (where ITS is internal transcribed spacer) polymerase chain reaction (PCR) products in two to four single digestions with five restriction endonucleases. In addition, we provide protocols for quick and reliable extraction and amplification of larval mussel DNA from complete host fish gill arches. Our results indicate that this new method can be applied on infection rates as low as three glochidia per gill arch and enables, for the first time, comprehensive, large-scale assessments of the relative importance of different host species for given unionoid populations.     

Within- and among-lake variation in shell morphology of the freshwater clam Elliptio complanata (Bivalvia: Unionidae) from south-central Ontario lakes

To compare the relative magnitude of variation in shell morphology within and among lakes, Elliptio complanata were collected from low and high exposure areas in each of four small lakes in south-central Ontario. Nested ANOVA's on shell length, height, width and weight revealed that shell morphology varied much more between sites of differing exposure within a lake than among lakes of differing alkalinity. Canonical variates analysis showed that clams from high exposure areas had larger and proportionately taller and heavier shells than those from low exposure areas. There was no relationship between alkalinity of lakes and shell morphology. These results suggest that the use of unionid shell morphology to predict long-term whole lake water chemistry (e.g. alkalinity) requires sampling designs which take into account within-lake variation in shell morphology.     

Bioremediation and Biorestoration of a Crube Oil-Contaminated Freshwater Wetland on the St.Lawrence River

Biostimulation by nutrient enrichment and phytoremediation were studied for the restoration of an acutely stressed freshwater wetland experimentally exposed to crude oil. The research was carried out along the shores of the St. Lawarence River at Ste. Croix, Quebec, Canada. The research determined the effectiveness of fertilizer addition in enhancing the biodegradation rates of residual oil. It further examined the rate at which the stressed ecosystem recovered with and without the addition of inorganic fertilizers and the role of nutrients in enhancing wetland restoration in the absence of healthy wetland plants. Chemical analysis of integrated sediment core samples to the depth of oil penetration within the experimental plots indicated that addition of inorganic nutrients did not enhance the disappearance of alkanes or PAHs. In surface samples, however, hydrocarbon disappearance rates were higher when the metabolic activity of wetland plants was suppressed by the removal of emergent plant growth. These results suggest that oxygen limitation plays a major role in preventing rapid biodegradation of hydrocarbons in anoxic wetland sediment.     

Comparison of Asian clam field bioassays and benthic community surveys in quantifying effects of a coal-fired power plant effluent on Clinch River biota

Survival and growth of Asian clamsmay be more sensitive endpoints than benthicmacroinvertebrate community richness parametersat distinguishing biotic impairmentattributable to complex effluents fromcoal-burning utilities. We conducted (1) fieldbioassays with the Asian clam (Corbiculafluminea) during 2000–02 and (2) rapidbioassessments of benthic macroinvertebratecommunities during 2000–01 at sitesupstream/downstream of American ElectricPower's (AEP) Clinch River Plant (CRP) inRussell County, Virginia (U.S.A). Survival andgrowth of transplanted C. fluminea weresignificantly impaired within the CRP effluentplume (averages of 35% and 0.21 mm,respectively) relative to all other study siteswithin the Clinch River (averages of 89% and1.58 mm). Conversely, richness metrics forEphemeroptera,Ephemeroptera-Plecoptera-Trichoptera (EPT), andtotal taxa were not reduced downstream of theCRP. However, relative abundance metrics forEphemeroptera and EPT were minimally reduced atthe CRP-influenced site during 2000–01. Moreimportantly, our results suggest that richnessmetrics for benthic macroinvertebratecommunities may be inadequate for assessing theeffects of complex industrial effluents on C. fluminea. These findings have implicationsfor bioassessment techniques employed tomonitor streams inhabited by imperiledfreshwater mussels because (1) C. flumineaand Unionoidea are ecologicallysimilar and (2) recent findings suggest certaingenera of Unionidae may be more sensitive thanC. fluminea.     

The influence of macrophyte beds on plankton communities and their export from fluvial lakes in the St Lawrence River

1. To determine the influence of macrophyte beds on plankton abundance within fluvial lakes of the St Lawrence River, planktonic components (macrozooplankton, heterotrophic bacteria, and phytoplankton as chlorophyll- a [Chl- a ]) were sampled in Lake St Francis and Lake St Pierre during summer 1998. We tested the hypothesis that the abundance of planktonic components was higher within macrophyte beds in comparison to the more rapidly flushed open water areas of the fluvial lakes.
2. The large cross channel variation in zooplankton biomass was indeed correlated with the presence of dense beds of submerged macrophytes. Total macrozooplankton biomass was nine-fold greater within the beds (mean=180 μg L−1 dry mass) than in either the open water or areas with only sparse vegetation (mean=20 μg L−1 dry mass).
3. Chl- a and heterotrophic bacterial abundance were also higher in the beds, but only slightly so. There was no difference in total phosphorus or dissolved organic carbon concentrations between areas of dense vegetation, sparse vegetation or open water.
4. Macrophyte beds on the margins of the fluvial lakes allow the development of high planktonic abundance relative to the fast flowing central channel. Macrozooplankton biomass was much higher at the outflows of the lakes (∼50 μg L−1 dry mass) in comparison to the inflows (<20 μg L−1 dry mass). The increase is due to the transfer of organisms from submerged macrophyte beds into the central channel in the downstream quarter of the two lakes where the marginal littoral waters enter central channel waters.
5. Along rivers, shallow fluvial lakes appear to act as sources of plankton which is exported downstream during years of extensive littoral macrophyte development.     

Placing biodiversity and ecosystem functioning in context: environmental perturbations and the effects of species richness in a stream field experiment

Greater biodiversity is often associated with increased ecosystem process rates, and is expected to enhance the stability of ecosystem functioning under abiotic stress. However, these relationships might themselves be altered by environmental factors, complicating prediction of the effects of species loss in ecosystems subjected to abiotic stress. In boreal streams, we investigated effects of biodiversity and two abiotic perturbations on three related indices of ecosystem functioning: leaf decomposition, detritivore leaf processing efficiency (LPE) and detritivore growth. Replicate field enclosures containing leaves and detritivore assemblages were exposed to liming and nutrient enrichment, raising pH and nutrient levels. Both treatments constitute perturbations for our naturally acidic and nutrient-poor streams. We also varied detritivore species richness and density. The effects of the abiotic and diversity manipulations were similar in magnitude, but whereas leaf decomposition increased by 18% and 8% following liming and nutrient enrichment, respectively, increased detritivore richness reduced leaf decomposition (6%), detritivore LPE (19%) and detritivore growth (12%). The detritivore richness effect on growth was associated with negative trait-independent complementarity, indicating interspecific interference competition. These interactions were apparently alleviated in both enriched and limed enclosures, as trait-independent complementarity became less negative. LPE increased with detritivore density in the monocultures, indicating benefits of intra-specific aggregation that outweighed the costs of intra-specific competition, and dilution of these benefits probably contributed to lowered leaf decomposition in the species mixtures. Finally, the effects of liming were reduced in most species mixtures relative to the monocultures. These results demonstrate how environmental changes might regulate the consequences of species loss for functioning in anthropogenically perturbed ecosystems, and highlight potential influences of biodiversity on functional stability. Additionally, the negative effects of richness and positive effects of density in our field study were opposite to previous laboratory observations, further illustrating the importance of environmental context for biodiversity–ecosystem functioning relationships.     

The effects of planktivorous fishes on the plankton community in a eutrophic lake

Stocking silver carp, a phytoplankton feeder, and bighead carp, an omnivorous plankton feeder, into an eutrophic lake at high densities caused a dramatic change in the lake ecosystem.Microcystis, which had been dominant in summer and a main food of the fishes decreased markedly, and green algae smaller than 10 μm then became dominant. Consequently, chlorophyll-a per unit area decreased slightly, while the rate of production was higher than that in the previous years. As the total density of the fishes increased (from 0.09 to 0.11 fish m⁻²), the growth of silver carp was retarded, while that of bighead carp increased.Microcystis, was unable to become dominant due to increased grazing pressure by the fishes, and small green algae became dominant. The lake conditions thus became more favourable for zooplankters which selectively consumed small green algae, and accordingly, the production of zooplankton rose. Bighead carp consumed more animal food, which they assimilate at a higher rate than plant food, and grew better in spite of the fact that the fish density increased. The feeding rate of silver carp was greatly reduced because the green algae were too small to ingest, and the fish therefore grew poorly. Results of a computer simulation of a model consisting of five compartments, representing the blue-green algae, green algae, zooplankton, silver and bighead carp, support the food-web change observed in the lake.     

The effects of fish on assemblages of amphibians in ponds: a field experiment

1. Bluegill sunfish ( Lepomis macrochirus ) dominate fish assemblages of small lakes and ponds throughout the eastern United States and may play a major role in structuring aquatic communities. We examined the impact of adult bluegill on amphibian density by stocking bluegill at a range of densities into partitions of an experimental pond in which amphibians were free to colonize.
2. Adult bluegill had a major impact on the amphibian assemblage. By the end of the experiment, gray treefrog ( Hyla versicolor ) tadpoles were nine times less abundant, and red-spotted newt ( Notophthalmus viridescens ) adults were three times less abundant in the presence of adult bluegill than in their absence. In contrast, bullfrog ( Rana catesbeiana ) tadpoles tended to increase in the presence of adult bluegill. Adult bluegill also had a negative effect on the abundance of predaceous aquatic insects.
3. There was no indication that interactions among amphibians were significant in determining the above patterns. We suggest that the strong impact of adult bluegill resulted from a combination of direct and indirect effects on amphibian larvae and predaceous aquatic insects.     

Myxozoan communities parasitizing Notropis hudsonius (Cyprinidae) at selected localities on the St. Lawrence River, Quebec: possible effects of urban effluents.

Late summer myxozoan infra- and component communities parasitizing 73 Notropis hudsonius at 5 sites on the St. Lawrence River upstream and downstream from the island of Montreal are described from study of histological sections of individual fish. Community membership included Myxobolus sp. A (intracellular in striated muscle fibers), Myxobolus sp. B (intracellular in striated muscle fibers), Myxobolus sp. C (brain), and Thelohanellus notatus (Mavor, 1916) Kudo, 1929 (loose connective tissue), all of which are histozoic myxobolids displaying strict tissue specificity for trophozoite development. Mean infracommunity richness in fish at the separate localities was estimated to be 0.4 +/- 0.5 to 1.3 +/- 0.7, with a maximum richness of 3 in any 1 fish. Component community richness in host samples was 2 to 4. It is argued that these are relatively high levels of diversity for freshwater fish parasites but that the values are probably conservative because of the study of only portions of individual fish. The percentage of fish infected with myxozoans of any species and infracommunity richness was significantly greater below the island of Montreal compared with above it. It is suspected that increased oligochaete populations at these sites, resulting from sewage-caused organic enrichment of sediments, may have accounted for the observed increased prevalence of infection.