Population synchrony of a native fish across three Laurentian Great Lakes: evaluating the effects of dispersal and climate

Climate and dispersal are the two most commonly cited mechanisms to explain spatial synchrony among time series of animal populations, and climate is typically most important for fishes. Using data from 1978–2006, we quantified the spatial synchrony in recruitment and population catch-per-unit-effort (CPUE) for bloater (Coregonus hoyi) populations across lakes Superior, Michigan, and Huron. In this natural field experiment, climate was highly synchronous across lakes but the likelihood of dispersal between lakes differed. When data from all lakes were pooled, modified correlograms revealed spatial synchrony to occur up to 800 km for long-term (data not detrended) trends and up to 600 km for short-term (data detrended by the annual rate of change) trends. This large spatial synchrony more than doubles the scale previously observed in freshwater fish populations, and exceeds the scale found in most marine or estuarine populations. When analyzing the data separately for within- and between-lake pairs, spatial synchrony was always observed within lakes, up to 400 or 600 km. Conversely, between-lake synchrony did not occur among short-term trends, and for long-term trends, the scale of synchrony was highly variable. For recruit CPUE, synchrony occurred up to 600 km between both lakes Michigan and Huron (where dispersal was most likely) and lakes Michigan and Superior (where dispersal was least likely), but failed to occur between lakes Huron and Superior (where dispersal likelihood was intermediate). When considering the scale of putative bloater dispersal and genetic information from previous studies, we concluded that dispersal was likely underlying within-lake synchrony but climate was more likely underlying between-lake synchrony. The broad scale of synchrony in Great Lakes bloater populations increases their probability of extirpation, a timely message for fishery managers given current low levels of bloater abundance.     

Energy Density of Lake Whitefish Coregonus clupeaformis in Lakes Huron and Michigan

We collected lake whitefish Coregonus clupeaformis off Alpena and Tawas City, Michigan, USA in Lake Huron and off Muskegon, Michigan USA in Lake Michigan during 2002–2004. We determined energy density and percent dry weight for lake whitefish from both lakes and lipid content for Lake Michigan fish. Energy density increased with increasing fish weight up to 800 g, and then remained relatively constant with further increases in fish weight. Energy density, adjusted for weight, was lower in Lake Huron than in Lake Michigan for both small (≤800 g) and large fish (>800 g). Energy density did not differ seasonally for small or large lake whitefish or between adult male and female fish. Energy density was strongly correlated with percent dry weight and percent lipid content. Based on data from commercially caught lake whitefish, body condition was lower in Lake Huron than Lake Michigan during 1981–2003, indicating that the dissimilarity in body condition between the lakes could be long standing. Energy density and lipid content in 2002–2004 in Lake Michigan were lower than data for comparable sized fish collected in 1969–1971. Differences in energy density between lakes were attributed to variation in diet and prey energy content as well as factors that affect feeding rates such as lake whitefish density and prey abundance.     

Compensation in individual growth rates and its influence on lake trout population dynamics in the Michigan waters of Lake Superior

The role of compensatory mechanisms in the population dynamics of lake trout in the Michigan waters of Lake Superior was explored during three time periods: the pre-sea lamprey period, prior to 1950 when lake trout were at a relatively high abundance and the fishery was the primary source of lake trout mortality; the sea lamprey dominant period, from 1951 to 1961 when lake trout were at a very low abundance due to sea lamprey predation and overexploitation; and currently, from 1985 to 1993 when wild lake trout abundance was at a moderate level. The role of compensatory changes in growth and fecundity rates of lake trout in the Michigan waters of Lake Superior was evaluated using a life table approach. Individual growth and fecundity rates were calculated and compared between time periods. These rates were used to determine age-specific fecundity which, along with age-specific survival, were incorporated into a Leslie projection matrix to calculate the finite rate of population increase (λ). Individual growth rates and age-specific fecundity rates changed in response to the different levels of lake trout abundance during each of the study periods. Lake trout during the sea lamprey dominant period, which experienced the lowest abundance and highest mortality levels, exhibited the fastest individual growth rates and the highest age-specific fecundity. These high rates contributed to the relatively large compensatory scope exhibited by lake trout during the sea lamprey dominant period as compared to lake trout during the pre-sea lamprey or the current periods which are associated with higher levels of abundance.     

A reintroduced lake sturgeon population comes of age: A genetic evaluation of stocking success in the St. Louis River

Lake sturgeon (Acipenser fulvescens) have experienced significant declines throughout the Great Lakes, leading to stocking efforts at some locations to reestablish extirpated populations. Given the late sexual maturity of the species, assessment of stocking program success is often delayed. Lake sturgeon were extirpated from the St. Louis River in western Lake Superior. Stocking began in 1983 and continued until 1994, using the Wolf River (Lake Winnebago) in the Lake Michigan basin as the source. Stocking resumed from 1998 to 2000, using the Sturgeon River in Lake Superior as the source. Our objectives were to (a) determine the movement patterns of stocked Wolf River sturgeon, (b) determine whether stocked individuals are migrating into natural spawning populations during the spawning season, (c) identify the origins of individuals captured in the St. Louis River, and (d) assess the genetic diversity of the reintroduced St. Louis River population. All collected samples were analyzed using 12 microsatellite loci, followed by genetic assignment testing to achieve the first three objectives. Genetic diversity was compared to natural spawning populations in Lake Superior. The highest proportion of stocked Wolf River lake sturgeon was detected in western Lake Superior, close to the stocking site. However, individuals were detected throughout Lake Superior. Wolf River individuals were detected in most of the spawning populations in Lake Superior, with the greatest number in the Sturgeon River and the Goulais River. The majority of individuals captured in the St. Louis River were of stocked origin (88.1%), with 73.5% from the Wolf River and 14.6% from the Sturgeon River. These observed proportions differed from the expected proportions based on the number of sturgeon released from each source (χ² = 55.37, p < 0.00001), with a higher representation of Sturgeon River individuals. Evidence of natural recolonization in the St. Louis River was detected from Lake Huron and Goulais River individuals. Genetic diversity of the St. Louis River population was comparable to levels observed in the remnant natural populations in Lake Superior. However, the effective population size of the St. Louis River was small (N_e = 38.1) and average relatedness among individuals was relatively high (r = 0.151). Monitoring of movement of stocked Wolf River sturgeon throughout Lake Superior should continue, with careful attention to the potential for outbreeding depression in remnant populations. Genetic diversity of the St. Louis River population should also continue to be monitored to see if it improves with increased natural reproduction.     

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.     

Influence of egg predation and physical disturbance on lake trout Salvelinus namaycush egg mortality and implications for life-history theory

The influence of egg predators and physical disturbance on lake trout Salvelinus namaycush egg mortality was investigated in situ in Lake Michigan where recruitment is below detectable levels and egg predator abundance is high. Comparisons were made with Lake Champlain where recruitment is low and egg predator abundance is also low and with Parry Sound (Lake Huron) where recruitment is moderate and egg predators are in low abundance. A multi-density egg seeding method (100 to 5000 eggs m⁻²) was used to quantify the effect of physical disturbance and egg predation on egg loss. Wind fetch was used as an index of physical disturbance and comparisons across all locations and egg densities suggested that at sites with high wind fetch (>5 km), physical disturbance may be a greater source of egg loss than predation. When analyses were limited to those sites having a wind fetch of <5 km, the percentage of eggs recovered was found to be linearly related to predator density. The strength of this relationship was based largely on egg recovery at 500 and 1000 eggs m⁻² because recovery at lower (100, 250 eggs m⁻²) and very high (5000 eggs m⁻²) densities was not significantly related to predator density. The reason for this is probably that at low egg densities, crayfish Orconectes spp., the major egg predator at most sites, had difficulty finding and consuming eggs and at high egg densities they became satiated. Egg loss was directly related to wind fetch for Lake Michigan and on average six-fold greater than for Parry Sound suggesting that without corresponding changes in fecundity and age structure, lake trout populations in large lakes like Lake Michigan are inherently less productive than those from enclosed inland waters.     

Lake trout demographics in relation to burbot and coregonine populations in the Algonquin Highlands,Ontario

The objective of the study was to test the hypothesis that lake trout populations change in relation to cisco, lake whitefish, round whitefish and burbot populations in lakes in the Algonquin Highlands region of Ontario. Lake trout population change is greatest where cisco and lake whitefish are present. Lake trout populations in lakes without either coregonine tend to have small adults and many juveniles. Where cisco or lake whitefish are present, adult lake trout are large, juvenile abundance is low, and the stock-recruit relationship appears to be uncoupled likely due to a larval bottleneck. Lake trout populations in these lakes may be sensitive to overfishing and recruitment failure. Lake trout populations do not appear to change in relation to round whitefish. There appears to be an indirect positive change on juvenile lake trout abundance through reductions in the density of benthic coregonines in the presence of large, hypolimnetic burbot.     

Hierarchical population structure and genetic diversity of lake trout (Salvelinus namaycush) in a dendritic system in Northern Labrador

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Evolution and diversity of two cisco forms in an outlet of glacial Lake Algonquin

The diversity of Laurentian Great Lakes ciscoes (Coregonus artedi, sensu lato) arose via repeated local adaptive divergence including deepwater ciscoes that are now extirpated or threatened. The nigripinnis form, or Blackfin Cisco, is extirpated from the Great Lakes and remains only in Lake Nipigon. Putative nigripinnis populations were recently discovered in sympatry with artedi in a historical drainage system of glacial Lake Algonquin, the precursor of lakes Michigan and Huron. Given the apparent convergence on Great Lakes form, we labeled this form blackfin. Here, we test the hypothesis that nigripinnis may have colonized this area from the Great Lakes as a distinct lineage. It would then represent a relict occurrence of the historical diversity of Great Lakes ciscoes. Alternatively, blackfin could have evolved in situ in several lakes. We captured more than 600 individuals in the benthic or pelagic habitat in 14 lakes in or near Algonquin Provincial Park (Ontario, Canada). Fish were compared based on habitat, morphology, and genetic variation at 6,676 SNPs. Contrary to our expectations, both cisco and blackfin belonged to an Atlantic lineage that colonized the area from the east, not from the Great Lakes. Sympatric cisco and blackfin were closely related while fish from different lakes were genetically differentiated, strongly suggesting the repeated in situ origin of each form. Across lakes, there was a continuum of ecological, morphological, and genetic differentiation that could be associated with alternative resources and lake characteristics. This study uncovers a new component of cisco diversity in inland lakes of Canada that evolved independently from ciscoes of the Laurentian Great lakes. The diversity of cisco revealed in this study and across their Canadian range presents a challenge for designating conservation units at the intraspecific level within the framework of the Committee on the Status of Endangered Wildlife in Canada (COSEWIC).     

Helminths in an intensively stocked population of lake trout, Salvelinus namaycush, from Lake Huron

Eighty stocked lake trout Salvelinus namaycush (Salmonidae), collected from 2 locations in Lake Huron in May 1995, were examined for parasites. The parasite fauna of this top predator in Lake Huron was characterized by only 6 helminth species. Echinorhynchus salmonis infected all lake trout with a mean intensity of 163.9. The intensity of this acanthocephalan species significantly increased with host length and weight. Eubothrium salvelini infected 78 lake trout with a maximum number of 81 scoleces counted. Diplostomum sp., Cyathocephalus truncatus, Capillaria salvelini, and Neoechinorhynchus sp. infrequently infected lake trout. The low parasite species richness in these lake trout is believed to be due to their large size at stocking and to the loss of historical enzootic host-parasite relationships that followed the absence of this fish species in Lake Huron for 26 yr.     

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.     

Broad- to fine-scale population genetic patterning in the smallmouth bass Micropterus dolomieu across the Laurentian Great Lakes and beyond: an interplay of behaviour and geography

Analysis of population genetic relationships reveals the signatures of current processes such as spawning behaviour and migration, as well as those of historical events including vicariance and climate change. This study examines these signatures through testing broad‐ to fine‐scale genetic patterns among smallmouth bass Micropterus dolomieu spawning populations across their native Great Lakes range and outgroup areas, with fine‐scale concentration in Lake Erie. Our primary hypotheses include whether genetic patterns result from behavioural and/or geographical isolation, specifically: (i) Are spawning groups in interconnected waterways genetically separable? (ii) What is the degree of isolation across and among lakes, basins, and tributaries? (iii) Do genetic divergences correspond to geographical distances? and (iv) Are historical colonization patterns from glacial refugia retained? Variation at eight nuclear microsatellite DNA loci are analysed for 666 smallmouth bass from 28 locations, including 425 individuals in Lake Erie; as well as Lakes Superior, Huron, and Ontario, and outgroups from the Mississippi, Ohio, St. Lawrence, and Hudson River drainages. Results reveal marked genetic differences among lake and river populations, as well as surprisingly high divergences among closely spaced riverine sites. Results do not fit an isolation‐by‐geographical‐distance prediction for fine‐scale genetic patterns, but show weak correspondence across large geographical scales. Genetic relationships thus are consistent with hypotheses regarding divergent origins through vicariance in glacial refugia, followed by colonization pathways establishing modern‐day Great Lakes populations, and maintenance through behavioural site fidelity. Conservation management practices thus should preserve genetic identity and unique characters among smallmouth bass populations.     

Contrasting genetic metrics and patterns among naturalized rainbow trout (Oncorhynchus mykiss) in two Patagonian lakes differentially impacted by trout aquaculture

Different pathways of propagation and dispersal of non‐native species into new environments may have contrasting demographic and genetic impacts on established populations. Repeated introductions of rainbow trout (Oncorhynchus mykiss) to Chile in South America, initially through stocking and later through aquaculture escapes, provide a unique setting to contrast these two pathways. Using a panel of single nucleotide polymorphisms, we found contrasting genetic metrics and patterns among naturalized trout in Lake Llanquihue, Chile's largest producer of salmonid smolts for nearly 50 years, and Lake Todos Los Santos (TLS), a reference lake where aquaculture has been prohibited by law. Trout from Lake Llanquihue showed higher genetic diversity, weaker genetic structure, and larger estimates for the effective number of breeders (N_b) than trout from Lake TLS. Trout from Lake TLS were divergent from Lake Llanquihue and showed marked genetic structure and a significant isolation‐by‐distance pattern consistent with secondary contact between documented and undocumented stocking events in opposite shores of the lake. Multiple factors, including differences in propagule pressure, origin of donor populations, lake geomorphology, habitat quality or quantity, and life history, may help explain contrasting genetic metrics and patterns for trout between lakes. We contend that high propagule pressure from aquaculture may not only increase genetic diversity and N_b via demographic effects and admixture, but also may impact the evolution of genetic structure and increase gene flow, consistent with findings from artificially propagated salmonid populations in their native and naturalized ranges.     

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.     

Genetic structure of brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis> L.) from the Hardangervidda mountain plateau (Norway) analyzed by microsatellite DNA: a basis for conservation guidelines

The Hardangervidda in southern Norway, the largest mountain plateau in Europe, has thousands of lakes and streams, mainly between 1000 and 1300 m above sea level, where brown trout is the only fish species. To describe the current genetic diversity of brown trout in this area, a total of 863 fish from 20 lakes were genotyped with eleven microsatellites. Most diversity is within lake populations, but diversity among geographical groups and populations within groups was significant, too. Neighbor-joining, principle coordinate analysis and Bayesian clustering show three major geographic groups in accordance with the river systems. Bias was caused by recent stocking in two lakes. Low/no genetic differentiation among some populations indicates that intermixing is common when lakes are well-connected, as was also shown by assignment test. We recommend preserving the genetic diversity of brown trout in this unique area by managing stocking in lake systems according to genetic structure.     

Founding population size of an aquatic invasive species

Non-native species of fish threaten native fishes throughout North America, and in the Rocky Mountains, introduced populations of lake trout threaten native populations of bull trout. Effective management of lake trout and other exotic species require understanding the dynamics of invasion in order to either suppress non-native populations or to prevent their spread. In this study, we used microsatellite genetic data to estimate the number of lake trout that invaded a population of bull trout in Swan Lake, MT. Examination of genetic diversity and allele frequencies within the Swan Lake populations showed that most of the genes in the lake trout population are descended from two founders. This emphasizes the importance of preventing even a few lake trout from colonizing new territory.     

Relative importance of phosphorus,invasive mussels and climate for patterns in chlorophyll a and primary production in Lakes Michigan and Huron

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Nitrogen transformations at the sediment–water interface across redox gradients in the Laurentian Great Lakes

The capacity of a lake to remove reactive nitrogen (N) through denitrification has important implications both for the lake and for downstream ecosystems. In large oligotropic lakes such as Lake Superior, where nitrate (NO₃ ^?) concentrations have increased steadily over the past century, deep oxygen penetration into sediments may limit the denitrification rates. We tested the hypothesis that the position of the redox gradient in lake sediments affects denitrification by measuring net N-fluxes across the sediment–water interface for intact sediment cores collected across a range of sediment oxycline values from nearshore and offshore sites in Lake Superior, as well as sites in Lake Huron and Lake Erie. Across this redox gradient, as the thickness of the oxygenated sediment layer increased from Lake Erie to Lake Superior, fluxes of NH₄ ⁺ and N₂ out of the sediment decreased, and sediments shifted from a net sink to a net source of NO₃ ^?. Denitrification of NO₃ ^? from overlying water decreased with thickness of the oxygenated sediment layer. Our results indicate that, unlike sediments from Lake Erie and Lake Huron, Lake Superior sediments do not remove significant amounts of water column NO₃ ^? through denitrification, likely as a result of the thick oxygenated sediment layer.     

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

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Spatio-temporal dynamics of gastrointestinal helminths infecting four lake whitefish (Coregonus clupeaformis) stocks in northern lakes Michigan and Huron, U.S.A

This study was undertaken to identify the community composition, structure, and dynamics of helminths infecting the gastrointestinal tract (GIT) of lake whitefish (Coregonus clupeaformis) collected from 4 sites in northern lakes Huron (Cheboygan and De Tour Village) and Michigan (Big Bay de Noc and Naubinway) from fall 2003 through summer 2006. A total of 21,203 helminths was retrieved from the GITs of 1,284 lake whitefish. Approximately 42% (SE = 1.4%) of the examined lake whitefish were infected with at least 1 helminth species in their GIT, with a mean intensity of 39.4 worms/fish (SE = 0.3) and a mean abundance of 16.4 worms/fish (SE = 0.1). Collected helminths appeared to be generalists and consisted of 2 phyla (Acanthocephala and Cestoda) and 5 species (Acanthocephalus dirus, Neoechinorhynchus tumidus, Echinorhynchus salmonis, Cyathocephalus truncatus, and Bothriocephalus sp.). Lake whitefish from Lake Huron on average had greater infection prevalences, abundances, and intensities than did fish from Lake Michigan. Infection parameters for each of the helminth species generally followed the same pattern observed for the combined data. Acanthocephalus dirus was the most prevalent and abundant helminth in lake whitefish GITs, although intensity of infection was the greatest for C. truncatus. Helminth infection parameters often peaked in the spring while diversity was greatest in the winter samples. There was substantial temporal variability in helminth infections with prevalences, abundances, and intensities often fluctuating widely on consecutive sampling occasions. Analysis of the GIT helminth community composition suggested that 3 (Big Bay de Noc, De Tour Village, and Cheboygan) of the 4 primary spawning sites, overall, had similar community compositions. The reason for the observed spatial and temporal variability in the lake whitefish GIT helminth infections remains to be elucidated. The findings of this study represent the most comprehensive parasitological study ever conducted on lake whitefish in the Great Lakes and will provide valuable information for future comparisons.