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.     

Reticulate evolution and phenotypic diversity in North American ciscoes, Coregonus ssp. (Teleostei: Salmonidae): implications for the conservation of an evolutionary legacy

Coregonine fishes are notorious taxonomicproblems due to their extreme morphological andecological variation. In North America, diversity is particularly baffling among ciscoes, and both morphological and phylogenetic analyses have resulted in major polytomy among the 8 taxa of the ``Coregonus artedi' species complex. Ciscoes arealso a devastated group, accounting for 10% ofthe fish species listed by the Committee on theStatus of Endangered Wildlife in Canada. Here,we complete the genetic characterization ofNorth American ciscoes with mitochondrial andmicrosatellites markers previously used toanalyse populations of C. artedi in orderto elucidate the evolutionary history andidentify appropriate conservation units. Ourresults revealed a complex evolutionary historymarked by postglacial reticulation eventscoupled with recent and independent evolutionof similar phenotypes (taxa). Genetic variationreflects geography rather than taxonomy, andconsequently, we recommend that a single taxon,C. artedi (sensu lato) be recognized.Local genetic differentiation is often coupledwith ecophenotypic diversification, and gillraker polymorphisms, depth-related habitatpreference and reproductive behaviour areconsidered as phenotypic traits with probableadaptive value contributing to the nicheexpansion of ciscoes. Ecomorphotypes of eachparticular locale thus represent a uniqueexpression of a diverse genetic pool stillundergoing divergence and sorting.Consequently, ciscoes from lakes with distinctecomorphotypes are recognized as ESUs, as wellas each of sympatric forms when they aregenetically differentiated. We recommend thatan ESU strategy focusing at a very local levelbe adopted for continental ciscoes as a validalternative to protect significant evolutionaryprocesses of divergence encountered inpolytypic species of newly colonized habitats.     

Genetic signs of multiple colonization events in Baltic ciscoes with radiation into sympatric spring‐ and autumn‐spawners confined to early postglacial arrival

Presence of sympatric populations may reflect local diversification or secondary contact of already distinct forms. The Baltic cisco (Coregonus albula) normally spawns in late autumn, but in a few lakes in Northern Europe sympatric autumn and spring‐ or winter‐spawners have been described. So far, the evolutionary relationships and taxonomic status of these main life history forms have remained largely unclear. With microsatellites and mtDNA sequences, we analyzed extant and extinct spring‐ and autumn‐spawners from a total of 23 Swedish localities, including sympatric populations. Published sequences from Baltic ciscoes in Germany and Finland, and Coregonus sardinella from North America were also included together with novel mtDNA sequences from Siberian C. sardinella. A clear genetic structure within Sweden was found that included two population assemblages markedly differentiated at microsatellites and apparently fixed for mtDNA haplotypes from two distinct clades. All sympatric Swedish populations belonged to the same assemblage, suggesting parallel evolution of spring‐spawning rather than secondary contact. The pattern observed further suggests that postglacial immigration to Northern Europe occurred from at least two different refugia. Previous results showing that mtDNA in Baltic cisco is paraphyletic with respect to North American C. sardinella were confirmed. However, the inclusion of Siberian C. sardinella revealed a more complicated pattern, as these novel haplotypes were found within one of the two main C. albula clades and were clearly distinct from those in North American C. sardinella. The evolutionary history of Northern Hemisphere ciscoes thus seems to be more complex than previously recognized.     

Habitat coupling in a large lake system: delivery of an energy subsidy by an offshore planktivore to the nearshore zone of Lake Superior

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Intraindividual and Interspecies Variation in the 5S rDNA of Coregonid Fish

This study was designed to characterize further the nontranscribed intergenic spacers (NTSs) of the 5S rRNA genes of fish and evaluate this marker as a tool for comparative studies. Two members of the closely related North American Great Lakes cisco species complex (Coregonus artedi and C. zenithicus) were chosen for comparison. Fluorescence in situ hybridization found the ciscoes to have a single multicopy 5S locus located in a C band-positive region of the largest submetacentric chromosome. The entire NTS was amplified from the two species by polymerase chain reaction with oligonucleotide primers anchored in the conserved 5S coding region. Complete sequences were determined for 25 clones from four individuals representing two discrete NTS length variants. Sequence analysis found the length variants to result from presence of a 130-bp direct repeat. No two sequences from a single fish were identical. Examination of sequence from the coding region revealed two types of 5S genes in addition to pseudogenes. This suggests the presence of both somatic and germline (oocyte) forms of the 5S gene in the genome of Coregonus. The amount of variation present among NTS sequences indicates that accumulation of variation (mutation) is greater in this multicopy gene than is gene conversion (homogenization). The high level of sequence variation makes the 5S NTS an inappropriate DNA sequence for comparisons of closely related taxa. Received: 22 August 1997 / Accepted: 31 October 1997     

Is it possible to identify habitat for a rare species? Shortjaw Cisco (<Emphasis Type="Italic">Coregonus zenithicus</Emphasis>) in Lake Huron as a case study

The Government of Canada is considering a recommendation by the Committee on the Status of Endangered Wildlife in Canada to list shortjaw cisco (Coregonus zenithicus) as “Threatened” throughout its range under the Species At Risk Act (SARA). If the listing is approved, shortjaw cisco will receive legal protection, including protection for its ‘critical habitat.’ This study focused on habitat characteristics associated with specimens identified as shortjaw cisco collected with targeted sampling in Lake Huron of the Laurentian Great Lakes. Competing habitat-use models were developed using available data for three physical habitat variables: Water Depth, Substrate Slope, and Cliff Distance, and the models were evaluated using binary logistic regression and ranking of Akaike information criteria. For the habitat factors examined, Water Depth was the most important variable for explaining the observed distribution of identified shortjaw cisco, although this factor alone was not sufficient to adequately represent habitat for this taxonomically uncertain and rare animal. Future habitat discrimination for this hypothesized species at risk must be based on (a) reduction of taxonomic uncertainty, (b) expansion of sampling effort, and (c) consideration of additional physical and ecological habitat factors.     

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

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The Round Goby, Neogobius melanostomus, a Fish Invader on both sides of the Atlantic Ocean

During the past decade, a bottom-dwelling, aggressive, multiple-spawning fish, the round goby (Gobiidae: Neogobius melanostomus), has spread from its native region in the Ponto-Caspian throughout Europe and to the Laurentian Great Lakes in North America. An international workshop, held at the Hel Marine Station, Poland, was organized to summarize population features of the round goby. Common fish predators of round gobies in the Great Lakes and in native regions are obligate and facultative benthic fishes and occasionally, pelagic fishes. In contrast, the main predator of the round goby in the Gulf of Gdansk is the Great Cormorant (Phalacrocorax carbo). In the Great Lakes, round gobies have lead to the decline of mottled sculpin (Cottus bairdi) and logperch (Percina caprodes) and reduced the hatching success of native fishes by feeding on their eggs. In the Gulf of Gdansk, round gobies have increased in abundance, while three-spined sticklebacks (Gasterosteus aculeatus) have declined. Round gobies have a broad diet throughout their range; larger specimens are molluscivores. There are fewer species of parasites and lower infection rates of round gobies in recently colonized areas than in native areas. Overall, newly colonized round gobies in brackish waters and lakes are smaller, mature earlier, have a male biased operational sex ratio and are more short-lived compared with round gobies from marine (native) habitats.     

Modification of the diel vertical migration of Bythotrephes longimanus by the cold-water planktivore, Coregonus artedi

1. The weak diel vertical migration observed in the large cladoceran Bythotrephes longimanus seems contradictory to the predator‐avoidance hypothesis that predicts large zooplankton should have long migration amplitudes. However, cold‐water planktivores, especially Coregonus spp., are a main source of mortality for Bythotrephes and hence a deeper migration would result in a greater overlap with these hypolimnetic planktivores. We hypothesized that Coregonus artedi (cisco) modifies the normal vertical migration pattern of Bythotrephes, such that the latter stays higher in the water column during the day and thus migrates less extremely at night. 2. The vertical distribution of Bythotrephes during the day was determined from single visits to six lakes in Ontario, Canada, all of which contain warm‐water, epilimnetic planktivores but differing in whether they contain cisco. One lake of each fish type was sampled day and night every 2–3 weeks over the ice‐free season to examine daytime depths and migration amplitude. 3. The vertical migration of Bythotrephes differed in the presence and absence of cisco. In the lakes with cisco, there were significantly fewer Bythotrephes in the hypolimnion and they were higher in the water column during the day. Migration amplitude was smaller in the cisco than in the non‐cisco lake. These observations were not attributable to differences in physical factors, and, although not conclusively attributable to cisco, are consistent with an effect of cisco. 4. We suggest that diurnal depth selection by Bythotrephes in lakes containing cisco is a trade‐off between the risk of predation by warm‐ versus cold‐water predators, balanced by the benefits of increased temperature and feeding rates near the surface. Even in lakes without cisco, however, the vertical migration of Bythotrephes was less than expected, suggesting that diurnal depth selection is a balance between the risk from warm‐water planktivores and access to sufficient light to feed effectively.     

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.     

Lake morphometry determines Dreissena invasion dynamics

Predicting the ecosystem effects of invasive species and the best control strategies requires understanding population dynamics and population regulation. Invasive bivalves zebra and quagga mussels (Dreissena spp.) are considered the most aggressive invaders in freshwaters and have become major drivers of ecosystem processes in the Laurentian Great Lakes. Combining all lake-wide studies of Dreissena spp. conducted in the Great Lakes, we found that invasion dynamics are largely governed by lake morphometry. Where both species are present, quagga mussels generally become dominant in 8–13 years. Thereafter, zebra mussels remain common in shallow lakes and embayments and lake-wide Dreissena density may remain similar, while in deep lakes quagga led to a near-complete displacement of zebra mussels and an ensuing dramatic increase in overall dreissenid density. In deep lakes, overall Dreissena biomass peaked later and achieved?~?threefold higher levels than in shallow lakes. Comparison with 21 waterbodies in North America and Europe colonized by both dreissenids confirmed that patterns of invasion dynamics found in the Great Lakes are very consistent with other waterbodies, and thus can be generalized to other lakes. Our biophysical model predicted that the long-term reduction in primary producers by mussel grazing may be fourfold less in deep compared to shallow lakes due to thermal stratification and a smaller proportion of the epilimnion in contact with the bottom. While this impact remains greatest in shallow areas, we show that when lakes are vertically well-mixed, dreissenid grazing impact may be greatest offshore, revealing a potentially strong offshore carbon and phosphorus sink.

     

The influence of mixing on rotifer assemblages of Michigan lakes

Seasonal changes in the rotifer assemblages of 42 lakes in northern lower Michigan was closely related to lake mixing characteristics, basin morphometry, and the presence of an oxygenated coldwater refuge. Three major classes of lakes (dimictic, discontinuous polymitic, and continuous polymictic) were evident by their capacity to maintain coldwater species as seasons progressed from winter through fall. The disappearance of coldwater assemblages from dimictic lakes coincided with oxygen depletion in the hypolimnion or with erosion of the hypolimnion through mixing. Coldwater species disappeared from large discontinuous polymictic lakes when deep epilimnetic mixing occured in late summer and fall. Species assemblages of nearly all stratified lakes converged with those of continuous polymictic lakes when the hypolimnetic refuge deteriorated in summer and fall. Local weather conditions, however, between years had a pronounced effect on the persistence of cold water species through the seasons by affecting the temperature and oxygen conditions of the hypolimnion. Large lakes of the region contain many of the coldwater species of the Laurentian Great Lakes but some taxa are conspicuosly absent. Cold stenothermal rotifers persist in the lakes of the region despite adverse environmental conditions. Their life histories and ability to form resting stages permit them to escape periods of oxygen depletion and thermal stress. In contrast, the crustacean glacial marine fauna (i.e. Mysis relicta, Limnocalanus macrurus, and Scenecella calanoides) was absent from all of the study lakes even though many of the present-day basins were once connected to the Laurentian Great Lakes. These species long life cycles, lack of diapausing stages, and limited dispersal may make them vulnerable to local extinction with the deterioration and loss of the coldwater refuge.     

Ecological Assessment of Dune Restorations in the Great Lakes Region

Because of the economic and environmental importance of stabilizing fragile sand dune habitats, restoration of dunes has become a common practice. Restoration efforts in the Great Lakes and East Coast regions of North America often consist of planting monocultures of the dominant native grass species, Ammophila breviligulata. We evaluated 18 dune restoration projects in the Great Lakes region conducted over the past 25 years. We characterized attributes of diversity (plants and insects), vegetation structure (plant biomass and cover), and ecological processes (soil nutrients and mycorrhizal fungi abundance) in each restoration, and we compared these measures to geographically paired natural dune communities. Restoration sites were similar to reference sites in most measured variables. Differences between restorations and reference sites were mostly explained by differences in ages, with the younger sites supporting slightly lower plant diversity and mycorrhizal spore abundance than older sites. Plant community composition varied little between restored and reference sites, with only one native forb species, Artemisia campestris, occurring significantly more often in reference sites than restored sites. Although it remains unclear whether more diverse restoration plantings could accelerate convergence on the ecological conditions of reference dunes, in general, traditional restoration efforts involving monoculture plantings of A. breviligulata in Great Lakes sand dunes appear to achieve ecological conditions found in reference dunes.     

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.     

Influence of cisco (Coregonus artedi,Lesueur) on muskellunge (Esox masquinongy,Mitchill) mean length,population size structure,and maximum size in northern Wisconsin lakes

Population size structure and maximum size of managed sportfish populations are dictated by abiotic, biotic, ecosystem, and anthropogenic influences. In their native ranges of northern Wisconsin, muskellunge (Esox masquinongy) and cisco (Coregonus artedi) are co-adapted cool- and cold-water species where cisco presence may influence population size structure and maximum size of muskellunge. We tested whether muskellunge size structure indices (length-frequency distributions, proportional size distribution), mean length, and mean maximum length of muskellunge differed when cisco were present or absent in Ceded Territory of Wisconsin (CTWI) lakes during 2015–2018. Cisco presence had a positive influence on size structure and mean length of individual muskellunge within populations. In contrast, cisco presence had no influence on the mean maximum length of muskellunge observed in CTWI populations suggesting that other factors may be better predictors of this metric than cisco presence. In cisco lakes, mean muskellunge length was negatively correlated with mean cisco length suggesting that gape limitation may be a factor influencing population size structure and individual growth rates. Therefore, cisco populations with primarily large individuals may be unavailable to muskellunge as forage. Our results suggest that cisco are an important forage species for some aspects of muskellunge population ecology; however, other factors may also contribute to muskellunge population size structure and maximum size outcomes. As such, conservation of remaining cisco populations in Wisconsin is critical because they influence muskellunge population ecology in lakes where the species coexist. Future research is needed to better understand the interactions of cisco, abiotic and biotic factors, and anthropogenic influences on muskellunge growth dynamics.     

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.     

Invasive species removal increases species and phylogenetic diversity of wetland plant communities

Plant invasions result in biodiversity losses and altered ecological functions, though quantifying loss of multiple ecosystem functions presents a research challenge. Plant phylogenetic diversity correlates with a range of ecosystem functions and can be used as a proxy for ecosystem multifunctionality. Laurentian Great Lakes coastal wetlands are ideal systems for testing invasive species management effects because they support diverse biological communities, provide numerous ecosystem services, and are increasingly dominated by invasive macrophytes. Invasive cattails are among the most widespread and abundant of these taxa. We conducted a three‐year study in two Great Lakes wetlands, testing the effects of a gradient of cattail removal intensities (mowing, harvest, complete biomass removal) within two vegetation zones (emergent marsh and wet meadow) on plant taxonomic and phylogenetic diversity. To evaluate native plant recovery potential, we paired this with a seed bank emergence study that quantified diversity metrics in each zone under experimentally manipulated hydroperiods. Pretreatment, we found that wetland zones had distinct plant community composition. Wet meadow seed banks had greater taxonomic and phylogenetic diversity than emergent marsh seed banks, and high‐water treatments tended to inhibit diversity by reducing germination. Aboveground harvesting of cattails and their litter increased phylogenetic diversity and species richness in both zones, more than doubling richness compared to unmanipulated controls. In the wet meadow, harvesting shifted the community toward an early successional state, favoring seed bank germination from early seral species, whereas emergent marsh complete removal treatments shifted the community toward an aquatic condition, favoring floating‐leaved plants. Removing cattails and their litter increased taxonomic and phylogenetic diversity across water levels, a key environmental gradient, thereby potentially increasing the multifunctionality of these ecosystems. Killing invasive wetland macrophytes but leaving their biomass in situ does not address their underlying mechanism of dominance and is less effective than more intensive treatments that also remove their litter.     

A phylogeographic survey of brook charr (Salvelinus fontinalis) in Algonquin Park, Ontario based upon mitochondrial DNA variation

Forty-nine populations of brook charr (Salvelinus fontinalis) from Algonquin Park lakes and rivers were analysed for mitochondrial DNA variation. Haplotypic distributions of wild fish in the Algonquin Park region of Ontario, Canada, predominantly reflect postglacial dispersal patterns into the region in spite of substantial hatchery plantings. Two major refugial groupings colonized this region. Northern and eastern watersheds (Amable du Fond, Bonnechere, and northern Petawawa), were colonized primarily by haplotype 1 fish (B1 phylogenetic assemblage), while Oxtongue River, southern Petawawa, and York River populations were colonized predominately by fish from the B2 and A mtDNA phylogenetic assemblages. Fish with haplotypes in the A and B2 phylogenetic assemblages are common in the Lake Huron drainage. All watersheds in the Park drain into the Ottawa River, except the Oxtongue drainage (part of the Lake Huron watershed). This suggests that early glacial outflows south of the Algonquin Park region (Kirkfield-Trent) may have been colonized by fish that initially invaded ‘Ontario island’ (south-western Ontario), while fish which invaded northern Algonquin Park were derived from a different refugial grouping(s) which may have involved colonization both up the Ottawa River drainage, and/or from a more westerly (Mississippian) refugial grouping. A majority of the populations in Algonquin Park have been planted with hatchery reared brook charr since the 1940s. The Hills Lake or ‘Domestic’ strain was used almost exclusively for these plantings. Comparisons of mtDNA haplotypic distributions in hatchery and wild fish suggests that hatchery females had minimal spawning success and/or their progeny had poor survivorship in the wild.     

Experimental evidence that the ecosystem effects of aquatic herbivory by moose and beaver may be contingent on water body type

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Patterns and Pathways in the Post-Establishment Spread of Non-Indigenous Aquatic Species: The Slowing Invasion of North American Inland Lakes by the Zebra Mussel

The zebra mussel, Dreissena polymorpha, has spread through eastern North American aquatic ecosystems during the past 15 years. Whereas spread among navigable waterways was rapid, the invasion of isolated watersheds has progressed more slowly and less predictably. We examined the patterns of overland spread over multiple spatial and temporal extents including individual lake districts, states, and multi-state regions in the USA and found that only a small proportion (<8%) of suitable inland lakes have been invaded, with the rate of invasion appearing to be slowing. Of the 293 lakes known to be invaded, 97% are located in states adjacent to the Laurentian Great Lakes with over half located in Michigan. Only six states have more than 10 invaded lakes and only in Michigan and Indiana have more than 10% of suitable lakes become invaded. At smaller spatial extents, invaded lakes are often clustered within a lake-rich region across southern Michigan and northern Indiana. This clustering appears primarily due to multiple overland invasions originating from the Great Lakes followed to a lesser extent by subsequent secondary overland and downstream dispersal. Downstream spread appears responsible for only one third of the inland invasions. Temporally, invasions peaked in the late 1990s, with only 13 new invasions (0.4% of suitable lakes) reported in 2003 in the four-state region surrounding Lake Michigan. Peak rates of invasion occurred 4–6 years earlier in Michigan relative to Indiana and Wisconsin, but this time lag is likely due to differences in the establishment of Great Lake source populations rather than ‘stepping stone’ dispersal across the landscape.