Use of ecological indicators to assess the quality of Great Lakes coastal wetlands

Over 2000 coastal wetland complexes have been identified in the Laurentian Great Lakes watershed, each providing critical habitat for numerous aquatic and terrestrial species. Research has shown there is a direct link between anthropogenic activities (urbanization and agricultural development) and deterioration in wetland health in terms of water quality and biotic integrity. In this study, we evaluate coastal marshes throughout the Great Lakes basin using a suite of published ecological indices developed specifically for coastal wetlands of the Great Lakes (Water Quality Index (WQI), Wetland Macrophyte Index (WMI), and the Wetland Fish Index (WFI_Basin)). We surveyed 181 wetlands, including 19 in Lake Superior (11%), 11 in Lake Michigan (6%), 13 in Lake Huron (7%), 92 in Georgian Bay and the North Channel (51%), 18 in Lake Erie (10%), and 28 in Lake Ontario (15%), over a 13 year period (1995–2008). Water quality parameters were measured at every site, while paired fyke nets were used to assess the fish community (132 sites) and macrophytes were surveyed and identified to species (174 sites); all of this information was used to calculate the associated index scores. One-way ANOVA results showed that there were significant differences in wetland quality among lakes. According to the WQI, we found that over 50% of marshes in Lakes Michigan, Erie, and Ontario were in degraded condition, while over 70% of marshes in Lakes Superior, Huron, and Georgian Bay were minimally impacted. Georgian Bay had the highest proportion of wetlands in very good and excellent condition and least number of wetlands in a degraded state. The WMI and WFI showed similar results. This is the largest bi-national database of coastal wetlands and the first study to provide a snapshot of the quality of coastal habitats within the Great Lakes basin. We recommend this information be used to guide conservation and restoration efforts within the Laurentian Great Lakes.     

Pelagic zonation of water quality and phytoplankton in the Great Lakes

Analysis of aquatic ecosystem data collected from large water bodies must consider spatial variations. A suite of pelagic survey stations exists for the Laurentian Great Lakes, but little is known about their redundancy. We present a strategy to delineate the lakes into zones based on water quality and phytoplankton biovolume. Water samples were collected from 72 sites in two seasons (spring and summer) from 2007 to 2010 in all five lakes. Integrated samples were analyzed for phytoplankton biovolume and nine water quality parameters. We conducted cluster analysis, principal components analysis and non-metric multidimensional scaling methods for water quality and phytoplankton taxon-specific biovolume for the Great Lakes basin and for each lake separately. There were significant lake-to-lake differences, and based on lake-specific analyses, Lake Superior, Lake Michigan and Lake Erie were each divided into three zones; Lake Huron and Lake Ontario were each grouped into two zones. The zones identified by water quality and phytoplankton provide an understanding of spatial distributions for evaluating monitoring data.     

Inter lake comparison of liver morphology and in vitro hepatic monodeiodination of L-thyroxine in sexually mature coho salmon, Oncorhynchus kisutch Walbaum, from Lakes Erie, Ontario, Michigan and Superior

The hepatosomatic index (HS1) of female sexually mature coho salmon, Oncorhynchus kisutch , from Lake Superior was significantly higher (P<0.01) than in males, whereas that of females from Lake Erie was significantly lower (P<0.05) than in males; there was no significant sex differences in coho salmon from Lakes Ontario or Michigan. Hepatic nuclear areas (HNA) were significantly higher ( P <0.01) in males compared with females in fish collected from Lakes Ontario and Erie, but there was no sex differences in HNA of coho salmon from Lakes Superior or Michigan. In all four lakes, hepatocyte cytoplasmic volume was significantly higher ( P <0.01) in males than in females from the same lake, and the cytoplasmic volume: nuclear volume ratio of males was corresponding significantly higher ( P < 0.05) than in females from the same lake.
Two hepatocytic cell types were evident, termed' light and dark cells, based on the electron density of their cytoplasmic matrix. There were considerable intersex and interlake differences in the relative populations of light 'and dark' cells in coho salmon from Lakes Ontario, Michigan, Erie and Superior, and also in the hepatocytic content of lipid, glycogen, mitochondria and ribosomal and smooth endoplasmic reticulum.
There were no sex differences with regard to in vitro hepatic monodeiodination of L-thyroxine (T4) to triiodo-L-thyronine (T3) in salmon from any of the lakes. However, T4 to T3 conversion was significantly lower (P < 0.05) in liver homogenates prepared from Lake Ontario coho salmon than in comparable preparations of liver from Lake Superior, Lake Erie, or Lake Michigan salmon.     

Biogenic silica as an estimate of siliceous microfossil abundance in Great Lakes sediments

Biogenic silica concentration (BSi) in sediment cores from the Great Lakes is evaluated as an estimate of siliceous microfossil abundance. A significant linear relationship was found between measured BSi and diatom valve abundance for sediment cores from the Bay of Quinte, Lake Ontario, Lake Erie, Lake Michigan and Lake Superior and between measured BSi and diatom biovolume for Lake Erie, Lake Michigan, and Lake Superior but not for Lake Ontario. Diatom silica predicted from diatom species abundance and an estimated silica content per cell in the Lake Erie cores accounted for 117% and 103% of measured BSi, respectively. By contrast, predicted diatom silica could only account for 28% of measured BSi in the Lake Michigan core and only 25% in the Lake Superior core. A few large diatoms with a large silica content per cell comprised a major portion of predicted diatom silica in all cores. The discrepancy between chemically measured BSi and the silica predicted from diatoms in the Lake Michigan and Lake Superior cores was partially due to the inability of the regression model, used to estimate diatom silica content, to account for different degrees of silicification in the diatom asemblages from the more dissolved silica rich Lake Michigan and Lake Superior.     

Toxicity of Clostridium botulinum type E neurotoxin to Great Lakes fish: implications for avian botulism

Since 1999, large-scale mortalities of fish-eating birds have been observed on the Great Lakes, and more specifically on Lake Erie. Type E botulism has been established as the primary cause of death. The mechanism of type E botulism exposure in fish-eating birds is unclear. Given that these birds are thought to eat live fish exclusively, it seems likely that their prey play a key role in the process, but the role of fish as potential transport vectors of botulinum neurotoxin type E (BoNT/E) to birds has not been adequately investigated. Between June 2003 and April 2004 a methodological model for exposing fish to Clostridium botulinum was developed and used to compare the sensitivity of rainbow trout (Oncorhynchus mykiss), round goby (Neogobius melanostomas), walleye (Stizostedion vitreum), and yellow perch (Perca flavescens) to four doses (0, 800, 1,500, and 4,000 Mouse Lethal Doses) of Clostridium botulinum type E neurotoxin. Each fish species expressed unique changes in both behavior and skin pigmentation prior to death. Yellow perch survived significantly longer (P < 0.05) than the three other species at all toxin treatments. Results of this study suggest that live fish can represent a significant vector for transfer of BoNT/E to birds.     

Genetic variation over space and time: analyses of extinct and remnant lake trout populations in the Upper Great Lakes

Lake trout (Salvelinus namaycush) in the upper Laurentian Great Lakes of North America experienced striking reductions in abundance and distribution during the mid-twentieth century. Complete collapse of populations was documented for Lake Michigan, and a few remnant populations remained only in lakes Huron and Superior. Using DNA obtained from historical scale collections, we analysed patterns of genetic diversity at five microsatellite loci from archived historical samples representing 15 populations (range 1940-1959) and from three contemporary remnant populations across lakes Huron and Superior (total n = 893). Demographic declines in abundance and the extirpation of native lake trout populations during the past 40 years have resulted in the loss of genetic diversity between lakes owing to extirpation of Lake Michigan populations and a temporal trend for reduction in allelic richness in the populations of lakes Superior and Huron. Naturally reproducing populations in Lake Superior, which had been considered to be remnants of historical populations, and which were believed to be responsible for the resurgence of lake trout numbers and distribution, have probably been affected by hatchery supplementation.     

Biodiversity of Clostridium botulinum type E associated with a large outbreak of botulism in wildlife from Lake Erie and Lake Ontario

The genetic relatedness of Clostridium botulinum type E isolates associated with an outbreak of wildlife botulism was studied using random amplification of polymorphic DNA (RAPD). Specimens were collected from November 2000 to December 2008 during a large outbreak of botulism affecting birds and fish living in and around Lake Erie and Lake Ontario. In our present study, a total of 355 wildlife samples were tested for the presence of botulinum toxin and/or organisms. Type E botulinum toxin was detected in 110 samples from birds, 12 samples from fish, and 2 samples from mammals. Sediment samples from Lake Erie were also examined for the presence of C. botulinum. Fifteen of 17 sediment samples were positive for the presence of C. botulinum type E. Eighty-one C. botulinum isolates were obtained from plants, animals, and sediments; of these isolates, 44 C. botulinum isolates produced type E toxin, as determined by mouse bioassay, while the remaining 37 isolates were not toxic for mice. All toxin-producing isolates were typed by RAPD; that analysis showed 12 different RAPD types and multiple subtypes. Our study thus demonstrates that multiple genetically distinct strains of C. botulinum were involved in the present outbreak of wildlife botulism. We found that C. botulinum type E is present in the sediments of Lake Erie and that a large range of bird and fish species is affected.     

Patterns of genetic diversity in Great Lakes bloaters (Coregonus hoyi) with a view to future reintroduction in Lake Ontario

The originally diverse ciscoe fish fauna of the Laurentian Great Lakes has suffered many extinctions and local extirpations. Bloaters (Coregonus hoyi) are presumed extirpated from Lake Ontario and the reintroduction of this deepwater fish is under consideration. Given the demographic fluctuations of this species in the other Great Lakes and its recent intralacustrine origin, we sought to identify a genetically diverse and similar source of C. hoyi via an analysis of genetic diversity and population structure using 10 microsatellite loci. Despite well-documented demographic declines, we found no genetic evidence of bottlenecks in 12 C. hoyi samples from the four potential donor lakes (Huron, Michigan, Superior and Nipigon). By contrast, evidence of bottlenecks in historical samples of C. artedi from Lake Ontario suggested that standard genetic methods frequently used to identify population bottlenecks can only detect very severe and long-lasting demographic declines in naturally large populations. Patterns of genetic differentiation and assignment tests indicated that C. hoyi from Lake Huron and Lake Michigan, which are not differentiated, are genetically most similar to Lake Ontario ciscoes. The small available sample of deepwater ciscoes recently caught in Lake Ontario did not allow determining if these represent a small undetected C. hoyi population or a recent invasion of the deep section by C. artedi. On the basis of genetic criteria, we conclude that C. hoyi from any location within Lake Huron or Lake Michigan would be judicious sources of breeders for reintroducing C. hoyi in Lake Ontario.     

The relationship between fecundity and adult body weight in Homeotherms

Summary Bythotrephes cederstroemii Schoedler, a predatory freshwater zooplankter (Crustacea: Cladocera), was first found in the Laurentian Great Lakes in December 1984. The first individuals were from Lake Huron, followed in 1985 with records from Lakes Erie and Ontario. By late August, 1986 the species had spread to southern Lake Michigan (43°N). Bythotrephes has not previously been reported from North America, but has been restricted to a northern and central Palearctic distribution. Its dramatic and widespread rise in abundance in Lake Michigan was greatest in offshore regions. Bythotrephes appears to be invading aggressively, but avoiding habitats presently occupied by glacio-marine relict species that became established in deep oligotrophic North American lakes after the Wisconsin glaciation. Because it is a voracious predator its invasion may lead to alterations in the native zooplankton fauna of the Great Lakes. It offers the chance to study how invading plankton species join an existing community. Judging from its persistence and success in deep European lakes, Bythotrephes may now become a permanent member of zooplankton communities in the Nearctic.     

Seasonal changes in thyroid hyperplasia, serum thyroid hormone and lipid concentrations, and pituitary gland structure in Lake Ontario coho salmon, Oncorhynchus kisutch Walbaum and a comparison with coho salmon from Lakes Michigan and Erie

A progressive fall in serum thyroid hormone levels was found in Lake Ontario coho salmon, Oncorhynchus kisutch , between May 1978 and January 1979 concomitant with an increase in goitre frequency and thyroid hyperplasia. Despite marked differences in overt goitre frequency in sexually mature adult salmon collected from Lakes Ontario (43.7%), Erie (93.5%), or Michigan (0%), serum thyroid hormone levels and thyroid histology were similar in fish from the three lakes. Two gonadotrop-like basophil cells ('globular'and'vacuolar') were found in the pituitary, particularly in sexually maturing and mature adults and'jacks'. Whereas in salmon from Lakes Ontario or Michigan, the globular cells were vacuolated in Lake Erie salmon they were not; this observation may be correlated with the absence of obvious secondary sexual characteristics in Lake Erie salmon. Serum free fatty acids serum, triglycerides and cholesterol concentrations in Lake Ontario salmon fell progressively from the condition in fish collected from the Lake in May 1978, to that of fish collected during their potamodrous migration in November 1978 and January 1979.     

Consideration of Geography and Wetland Geomorphic Type in the Development of Great Lakes Coastal Wetland Bird Indicators

We examined how geographic distribution of birds and their affinities to three geomorphic wetland types would affect the scale at which we developed indicators based on breeding bird communities for Great Lakes coastal wetlands. We completed 385 breeding bird surveys on 222 wetlands in the US portion of the basin in 2002 and 2003. Analyses showed that wetlands within two ecoprovinces (Laurentian Mixed Forest and Eastern Broadleaf Forest) had different bird communities. Bird communities were also significantly different among five lakes (Superior, Michigan, Huron, Erie, and Ontario) and among three wetland types (lacustrine, riverine, barrier-protected). Indicator values illustrated bird species with high affinities for each group (ecoprovince, lake, wetland type). Species with restricted geographic ranges, such as Alder and Willow Flycatchers (Empidonax alnorum and E. traillii), had significant affinities for ecoprovince. Ten bird species had significant affinities for lacustrine wetlands. Analyses on avian guild metrics showed that Lake Ontario wetlands had fewer long-distant migrants and warblers than other lakes. Numbers of short-distant migrants and total individuals in wetlands were higher in the Eastern Broadleaf Forest ecoprovince. Number of flycatchers and wetland obligate birds were not different among provinces, lakes, or wetland type. One potential indicator for wetland condition in Great Lakes wetlands, proportion of obligate wetland birds, responded negatively to proportion of developed land within 1 km of the wetland. We conclude that, although a guild approach to indicator development ameliorates species-specific geographic differences in distribution, individual species responses to disturbance scale will need to be considered in future indicator development with this approach.     

Distribution of circular single‐stranded DNA viruses associated with benthic amphipods of genus Diporeia in the Laurentian Great Lakes

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150 Phylogenetic Systematics and Phenology of the Hawaiian Endemic Freshwater Red Alga, Batrachospermum Spermatiophorum

Müller et al. (1998) noted that freshwater collections of the genus Bangia formed a distinct group separate from marine entities in gene sequence analyses. Recently, the species epithet B. atropurpurea has been resurrected to represent this freshwater lineage. This taxon is one of many invasive species within the Laurentian Great Lakes. B. atropurpurea was first observed in Lake Erie in 1964 and by 1982 was observed in all of the Great lakes except Lake Superior. The present study was initiated to examine the further spread of B. atropurpurea and determine the origin of these populations. Hence, a survey of all the Great Lakes was conducted in 1995 (86 sites) and again in 2002 (104 sites). Bangia was observed at 43 sites in 1995 and 39 sites in 2002. For the first time, this alga has been observed to be present in the St. Lawrence River (1995), Georgian Bay on Lake Huron (2002) and Lake Simcoe (eastern shore, 2002) and hence this alga appears to be spreading into new locations. Cluster analyses of morphological data reveal three distinct groupings that do not separate according to location or lake basin. Preliminary analyses of ITS 1 and 2 sequences show differences among samples within Lake Ontario and among all Lakes; however, collections from Lake Simcoe are very similar in sequence. We are continuing to examine the relationship of Great Lakes populations with freshwater collections from Europe.     

Thyroid function, pituitary structure and serum lipids in Great Lakes coho salmon, Oncorhynchus kisutch Walbaum, 'jacks' compared with sexually immature spring salmon

Whereas spring salmon, Oncorhynchus kisutch , from Lake Ontario had no overt thyroid goiters, comparable Lake Erie spring salmon had a 44% goitre incidence. Goitre incidence in sexually precocious 'jacks' from Lakes Ontario and Michigan was 43–45% and 100% in 'jacks' from Lake Erie.
All fish displayed thyroid hyperplasia. The degree of hyperplasia (as indicated by thyroid follicle colloid content, thyroid follicle size and shape and thyroid follicle epithelial cell size) in Lake Ontario fish tended to be greater in the 'jacks' than in the spring fish; the reverse was true in Lake Erie fish. Serum total L-thyroxine (T4) concentrations in Lake Ontario spring fish were higher than in any other group. There were no differences in serum T, levels between the groups of 'jacks', nor were serum T, levels in 'jacks' and spring salmon from Lake Ontario significantly different. Serum total triiodo-thyronine (T) concentrations in Lake Ontario spring salmon were significantly higher than in all other groups and that in Lake Erie spring salmon greater than in all groups of 'jacks'. Serum T, levels in Lake Ontario 'jacks' were higher than in Lake Erie 'jacks'. Serum free fatty acid (FFA) and cholesterol levels showed significant interlake differences with lower levels present in Lake Erie salmon. Lake Erie 'jacks' appeared to have fewer and less well vacuolated pituitary gonadotropic basophils than 'jacks' from Lakes Michigan and Ontario. Putative thyrotrop cells appeared to be more numerous and more active in the Lake Erie spring salmon compared with Lake Ontario spring salmon. Prolactin, corticotrop and melanotrop cells appeared less active in spring salmon than in the 'jacks'.     

149 Distribution and Invasion of Bangia Atropurpurea (Rhodophyta) in the Laurentian Great Lakes

Müller et al. (1998) noted that freshwater collections of the genus Bangia formed a distinct group separate from marine entities in gene sequence analyses. Recently, the species epithet B. atropurpurea has been resurrected to represent this freshwater lineage. This taxon is one of many invasive species within the Laurentian Great Lakes. B. atropurpurea was first observed in Lake Erie in 1964 and by 1982 was observed in all of the Great lakes except Lake Superior. The present study was initiated to examine the further spread of B. atropurpurea and determine the origin of these populations. Hence, a survey of all the Great Lakes was conducted in 1995 (86 sites) and again in 2002 (104 sites). Bangia was observed at 43 sites in 1995 and 39 sites in 2002. For the first time, this alga has been observed to be present in the St. Lawrence River (1995), Georgian Bay on Lake Huron (2002) and Lake Simcoe (eastern shore, 2002) and hence this alga appears to be spreading into new locations. Cluster analyses of morphological data reveal three distinct groupings that do not separate according to location or lake basin. Preliminary analyses of ITS 1 and 2 sequences show differences among samples within Lake Ontario and among all Lakes; however, collections from Lake Simcoe are very similar in sequence. We are continuing to examine the relationship of Great Lakes populations with freshwater collections from Europe.     

Microbial communities of the Laurentian Great Lakes reflect connectivity and local biogeochemistry

The Laurentian Great Lakes are a vast, interconnected freshwater system spanning strong physicochemical gradients, thus constituting a powerful natural laboratory for addressing fundamental questions about microbial ecology and evolution. We present a comparative analysis of pelagic microbial communities across all five Laurentian Great Lakes, focusing on Bacterial and Archaeal picoplankton characterized via 16S rRNA amplicon sequencing. We collected samples throughout the water column from the major basins of each lake in spring and summer over 2 years. Two oligotypes, classified as LD12 (Alphaproteobacteria) and acI-B1 (Actinobacteria), were among the most abundant in every sample. At the same time, microbial communities showed distinct patterns with depth during summer stratification. Deep hypolimnion samples were frequently dominated by a Chloroflexi oligotype that reached up to 19% relative abundance. Stratified surface communities differed between the colder, less productive upper lakes (Superior, Michigan, Huron) and warmer, more productive lower lakes (Erie, Ontario), in part due to an Actinobacteria oligotype (acI-C2) that averaged 7.7% of sequences in the lower lakes but <0.2% in the upper lakes. Together, our findings suggest that both hydrologic connectivity and local selective pressures shape microbial communities in the Great Lakes and establish a framework for future investigations.     

Is the ‘microbial loop’ an early warning indicator of anthropogenic stress?

Various components of the Microbial loop such as bacteria, heterotrophic nanoflagellates and autotrophic picoplankton were analyzed, for the first time across the Great Lakes, during a cruise in the summer of 1988. In addition, the size fractionated primary productivity using carbon-14 techniques was also determined. The statistical analysis indicated that bacteria, autotrophic picoplankton and ultraplankton/picoplankton productivity were significantly higher in Lakes Ontario and Erie than Lakes Huron and Michigan. The autotrophic picoplankton and ultraplankton/picoplankton productivity was higher in Lake Erie compared to Lake Ontario.The autotrophic picoplankton showed sensitivity to nutrients and contaminants in various types of environments. A dramatic decrease of autotrophic picoplankton in eutrophic-contaminated areas, such as Ashbridges Bay, Hamilton Harbour and western Lake Erie was observed. Conversely, in Saginaw Bay, another eutrophic environment, the autotrophic picoplankton were significantly higher than in Lake Huron. The sensitivity of autotrophic picoplankton to nutrients/contaminants might have implications to trophic interactions. Our results suggest that structural and functional characteristics of the microbial loop may be operating differently in stressed versus unstressed ecosystems. The possibility of using autotrophic picoplankton as an early warning indicator of environmental perturbation is proposed.     

The diet of Herring Gulls (Larus argentatus) during winter and early spring on the lower Great Lakes

In the Great Lakes, the Herring Gull (Larus argentatus) is a prominent member of the aquatic bird community, and has been used to monitor spatial and temporal trends in contaminant levels. To understand more fully contaminant loading outside the breeding season, we analysed the contents of 1298 freshly regurgitated pellets and 179 fresh faeces, collected in March and early April 1978–83, and between late December and late February 1990–91, from the vicinity of breeding colonies in Lakes Ontario and Erie, the Niagara River, Detroit River, and south-eastern parts of Lake Huron. Most adult Herring Gulls from the Great Lakes population winter in these areas, but there is no published account of their food habits other than during the breeding season. Most pellets from colonies close to large urban centres contained remains of garbage, as well as various fish species. Small mammals, notably Deer Mice (Peromyscus maniculatus) dominated the early spring diet at Lake Huron colonies near agricultural areas. At all other sites fish predominated in pellets and faeces, but garbage items were also recorded regularly. The species of fish consumed varied regionally, probably reflecting local availability. In Lake Ontario, Rainbow Smelt (Osmerus mordax) and Alewife (Alosa pseudoharengus) occurred most frequently in samples, whereas Freshwater Drum (Aplodinotus grunniens) was the main fish prey in Lake Erie and the Detroit River. Dietary differences were apparent between years, within seasons, and amongst areas. While these may have reflected some real differences in food availability, interpretation of these results was confounded by various biases inherent in the sampling of pellets and faeces to determine diet in such an opportunistic species. Therefore, it would be unwise to draw rigid conclusions as to regional or seasonal differences in the diets of piscivorous birds, based upon analyses of diet from only a small sample of sites or years. Herring Gulls appear to feed mainly on fish and garbage in winter and early spring on the lower Great Lakes (much as during the breeding season), but any locally abundant food source is probably exploited opportunistically.     

A multi‐basin approach determines variability in host fish suitability for unionids in tributaries of the Laurentian Great Lakes

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First record of Neoergasilus japonicus (Poecilostomatoida: Ergasilidae), a parasitic copepod new to the Laurentian Great Lakes

The parasitic copepod Neoergasilus japonicus, native to eastern Asia, was first collected from 4 species of fish (fathead minnow, Pimephales promelas; largemouth bass, Micropterus salmoides; pumpkinseed sunfish, Lepomis gibbosus; and yellow perch, Perca flavescens) in July 1994 in Saginaw Bay, Lake Huron, Michigan. Further sampling in the bay in 2001 revealed infections on 7 additional species (bluegill, Lepomis macrochirus; carp, Cyprinus carpio; channel catfish, Ictalurus punctatus; goldfish, Carassius auratus; green sunfish, Lepomis cyanellus; rock bass, Ambloplites rupestris; and smallmouth bass, Micropterus dolomieu). An additional 21 species examined in 2001 were devoid of the parasite. A limited collection of fish from Lake Superior (n = 8) and Lake Michigan (n = 46) in 1994 showed no infection. Neoergasilus japonicus is most frequently found attached to the dorsal fin and, in decreasing frequency, on the anal, tail, pelvic, and pectoral fins. Prevalence generally ranged from 15 to 70 and intensity from 1 to 10. The greatest number of copepods on a single host was 44. The copepod Neoergasilus japonicus appears to disperse over long distances rather quickly, spreading across Europe in 20 yr and then moving on to North America over a span of 10 yr. Its main vehicle of transport and introduction into the Great Lakes is probably exotic fish hosts associated with the fish-culture industry.