In Situ Quantum Efficiency of Lake Superior Phytoplankton

In situ quantum efficiencies were measured in Lake Superior over a 4-day period in 1978 and on one occasion in 1980. In 1980, experimental artifacts caused by exposing deep phytoplankton to elevated irradiances were minimized by the use of SCUBA divers. The trends of quantum efficiency with depth agreed well with theory. In the nutrient-limited upper portion of the euphotic zone, quantum efficiencies increased with depth, whereas in the light-limited lower portion of the euphotic zone, quantum efficiencies were relatively constant. Maximum quantum efficiencies calculated with downwelling irradiances ranged from 0.041 to 0.069 moles C fixed ? Einst abs^?1 with a mean maximum quantum efficiency of 0.0538 ± 0.0025 moles C fixed ? Einst abs^?1. Maximum quantum efficiencies in morning experiments ranged from 0.041 to 0.053 moles C fixed ? Einst abs^?1 and were slightly less than noon values, 0.057 to 0.067 moles C fixed ? Einst abs^?1. Correction for scalar irradiance would reduce all quantum efficiencies by 25%.     

Population structure and genomic variation of ecological life history diversity in wild-caught Lake Superior brook trout, Salvelinus fontinalis

Delineating patterns of extant genetic diversity is crucial for understanding the evolutionary history of populations and how they can be conserved. Brook trout (Salvelinus fontinalis; n?=?173) were sampled from 11 locations in Nipigon Bay, Lake Superior to evaluate genetic differentiation between migratory (coaster) and non-migratory (resident) individuals. Stable isotope data were used to classify the trophic ecology of individuals, rather than relying on size or capture location and timing to classify ecotypes. Genome-wide association analysis of 4676 single nucleotide polymorphism (SNP) markers was conducted to test if phenotypic and behavioral differences within and among populations of brook trout were linked to allelic variation in the genome. Population genetic structure was also evaluated to determine the source of migratory coasters based on multilocus genotypes from a subset of 906 SNP markers. Significant population genetic structure was revealed among the sampled tributaries, and the population genetic structure suggests that a single, larger stream in the study area, Cypress River, was the primary source of migratory brook trout in Nipigon Bay. There was little evidence of genetic differentiation between sympatric migrant and stream-resident brook trout; across the genome only 11 SNP loci (<1%) were significantly associated with life history type. Our findings suggest that coaster brook trout are an ecotype derived from resident populations, rather than a distinct genetic form.

Seasonal distribution of bloater (Coregonus hoyi) in the waters of Lake Huron surrounding the Bruce Peninsula

Information about bloater (Coregonus hoyi) habitat in Lake Huron was limited to correlations between commercial yield and fishing depth, despite available information from other Great Lakes. We identified seasonal patterns of bloater habitat use in hypolimnetic waters surrounding the Bruce Peninsula, Lake Huron. We applied a delta-lognormal model to fisheries-independent survey data to evaluate whether bloater catch-per-unit-effort was related to depth, temperature, and bathymetric slope. A Bayesian variable selection technique indicated that bloater distribution was most strongly related to bottom depth and water temperature. Our study also reconfirmed a previously-described pattern of seasonal inshore movement during warmer months followed by a return to deeper offshore waters during cooler months. By focusing our sampling within the hypolimnion, we characterized intra-annual patterns of bloater habitat use with respect to a temperature gradient near the minimum thermal requirements reported for this species. Bloater distribution under these thermal conditions has not been previously reported.     

Food of Young-of-the-Year Lake Trout (Salvelinus Namaycush) in Presque Isle Harbor,Lake Superior

The food habits of young lake trout (Salvelinus namaycush) were studied by examining the digestive tracts of 293 young-of-the-year collected in Presque Isle Harbor, Lake Superior. Lake trout in the 25 to 27-mm length range started to eat food organisms before all of their yolk material was absorbed. Organisms consumed by the 25 to 27-mm young-of-the-year included Chironomidae, Copepoda (Harpacticoida, Calanoida, Cyclopoida), and Cladocerea (Daphnia spp., Bosmina sp., Chydorus sp.). Chironomid pupae and chironomid larvae accounted for 74% and 5%, respectively, of the total volume of food eaten by the young lake trout in Presque Isle Harbor. Although copepods, cladocerans, and mysids were present in many stomachs, their contribution to the total volume of food was only 15%. Some lake trout in the 32 to 54-mm length range had consumed fry of sculpin (Cottus spp.) or rainbow smelt (Osmerus mordax), but the overall contribution of fish fry to the total volume of food was only 4% (frequency of occurrence, 10%). The lake trout in Presque Isle Harbor fed heavily on planktonic organisms, sparingly on benthic organisms, and were opportunistic feeders that appeared to prey on whatever forage organisms were available in the shallow nearshore waters.     

Distribution of the Amphipod Diporeia in Lake Superior: The Ring of Fire

Diporeia, formerly the dominant benthic macroinvertebrate in the Great Lakes, remains a keystone species in Lake Superior. Little is known, however, about fine scale amphipod distributions, especially as influenced by the production, transport and transformation of energy resources. Here, we document the distribution and abundance of Diporeia along 19 transects around the lake's perimeter. Regions of elevated density, averaging 958 ± 408 Diporeia/m² (mean ± S.D.) were observed along all transects, typically within slope habitat (depth of 30–125 m). Waters shoreward (shelf habitat, < 30 m) and lakeward (profundal habitat, > 125 m) of these regions supported significantly lower densities, averaging 239 ± 178/m² and 106 ± 59/m², respectively. Amphipods within regions of elevated density, termed here the Ring of Fire, account for two-thirds of the lakewide population while occupying only one-quarter of the benthic habitat. The Ring of Fire, observed lakewide as a band averaging 14.2 ± 9.4 km in width, is characterized as a region of transitional sediment deposition with gentle slope, proximate to nearshore locations of elevated primary production. Within the Ring of Fire exceptionally high densities are found in the south central region, where the Keweenaw Current and slope bathymetries serve to funnel production from adjoining regions of high production. Density measurements for the 173 stations sampled here are used to estimate lakewide Diporeia standing stock (22.5–37.7 trillion individuals, 4.4–7.4 Gg dry weight, 2.1–3.5 Gg C), individual and biomass density (274–460/m², 0.05–0.09 g DW/m², 0.03–0.04 gC/m²) and areal (0.02–0.03 g C/m²/yr) and total (1.6–2.6 Gg C/yr) production.     

Effects of Phosphorus Enrichment and Wave Simulation on Populations of Ulothrix Zonata from Northern Lake Superior

The effects of phosphorus enrichment and wave simulation on new (denuded rocks) and established populations of Ulothrix zonata were examined in specially designed stationary and rocking tanks on a bedrock ledge overlooking the western arm of Lake Superior at Castle Danger, Minnesota. Tanks received constant flow of lake water and were exposed to the same insolation and climatic conditions as the lake for a 10-week period during July, August, and September 1972. A rocking tank and a stationary tank were enriched to maintain an average concentration of 0.16 mg/L PO₄–P. Two remaining tanks, receiving only lake water, served as controls. Each week, algal material was harvested from the tanks and analyzed for chlorophyll a, ash-free dry weight, and abundance of U. zonata and associated non-filamentous periphytic algae. Diatoms comprised, on the average, 95% of the total non-filamentous periphytic algae.The results show that phosphorus enrichment has marked effects on established populations of U. zonata in addition to favoring recolonization of U. zonata on bare rock substrates. Statistical analysis of the data revealed that phosphorus enrichment caused a significant increase in Ulothrix biomass and chlorophyll a in the enriched tanks. Wave simulation caused a significant reduction in total numbers of non-filamentous algae associated with Ulothrix in the rocking tanks.     

Comparison of precision and bias of scale, fin ray, and otolith age estimates for lake whitefish (Coregonus clupeaformis) in Lake Champlain

We compared the precision, bias, and reader uncertainty of scales, dorsal fin rays, and otolith age estimates from 151 lake whitefish (Coregonus clupeaformis) from Lake Champlain, 2009. Mean and systematic differences in age estimates were compared among structures using consensus ages from two readers; precision of age structures was quantified through the use of age-bias plots, coefficient of variation, and percent agreement; reader confidence was indexed as a measure of overall reader uncertainty for each individual fish by structure. Mean age estimates based on otoliths were systematically higher (7.8 years) than based on scales (6.0 years) or fin rays (5.6 years). Ages determined using otoliths generated a wider range of ages and greater number of age classes (1-23 years, 20 age classes) when compared with scales (1-16 years, 15 age classes) and fin rays (1-14 years, 13 age classes). Otoliths were the most precise of the structures (CV = 4.7, compared with 7.4 for scales and 12.1 for fin rays). Percent agreement between readers indicated high precision and reproducibility of age estimates using otoliths. Percent reader uncertainty was lowest when using otoliths (7.6%) in comparison with fin rays (21.2%) or scales (26.8%). This study is the first evaluation of precision and bias of age structures for Lake Champlain's unexploited lake whitefish population and suggests that otoliths are the most appropriate structure for age estimation. However, the differences in age estimates from the three structures in this study emphasize the importance of validating aging structures to provide accurate age estimates for Lake Champlain.     

Changes in Fecundity and Egg Lipid Content of Lake Whitefish (Coregonus clupeaformis) in the Upper Laurentian Great Lakes between 1986–87 and 2003–05

Lake whitefish (Coregonus clupeaformis Mitchill), an important commercial species in the Laurentian Great Lakes, have experienced decreased growth and condition in regions of the upper Great Lakes over the past 20 years. Increases in lake whitefish density and decreases in the density of Diporeia spp., an energy rich and historically important part of the lake whitefish diet, have been implicated in the recent declines in lake whitefish growth and condition. The goal of this study was to describe lake whitefish fecundity, egg lipid content, and total ovary lipid content in selected regions of Lakes Huron, Michigan, and Superior in 1986–87 and 2003–05, two time periods with different lake whitefish and Diporeia densities. Under conditions of high lake whitefish density and low Diporeia density, female lake whitefish in the upper Laurentian Great Lakes generally produced fewer eggs. Egg lipid content was higher in 2003–05 than in 1986–87 at all sites, regardless of changes in lake whitefish or Diporeia densities. Total ovary lipid content and lake whitefish abundance were inversely related, while there was no significant relationship between total ovary lipid content and Diporeia density. The amount of energy that lake whitefish invested in egg production was more closely associated with lake whitefish abundance than with Diporeia density. This study provides evidence that recent changes in production dynamics of Great Lakes lake whitefish have not been driven solely by declines in Diporeia but have been significantly influenced by lake whitefish abundance.     

Evaluating the growth potential of sea lampreys (Petromyzon marinus) feeding on siscowet lake trout (Salvelinus namaycush) in Lake Superior

Differences in the preferred thermal habitat of Lake Superior lake trout morphotypes create alternative growth scenarios for parasitic sea lamprey (Petromyzon marinus) attached to lake trout hosts. Siscowet lake trout (Salvelinus namaycush) inhabit deep, consistently cold water (4-6 °C) and are more abundant than lean lake trout (Salvelinus namaycush) which occupy temperatures between 8 and 12 °C during summer thermal stratification. Using bioenergetics models we contrasted the growth potential of sea lampreys attached to siscowet and lean lake trout to determine how host temperature influences the growth and ultimate size of adult sea lamprey. Sea lampreys simulated under the thermal regime of siscowets are capable of reaching sizes within the range of adult sea lamprey sizes observed in Lake Superior tributaries. High lamprey wounding rates on siscowets suggest siscowets are important lamprey hosts. In addition, siscowets have higher survival rates from lamprey attacks than those observed for lean lake trout which raises the prospect that siscowets serve as a buffer to predation on more commercially desirable hosts such as lean lake trout, and could serve to subsidize lamprey growth.     

Genetic assessment of straying rates of wild and hatchery reared lake sturgeon (Acipenser fulvescens) in Lake Superior tributaries

Natal philopatry in lake sturgeon (Acipenser fulvescens) has been hypothesized to be an important factor that has lead to genetically distinct Great Lakes populations. Due to declining abundance, population extirpation, and restricted distribution, hatchery supplementation is being used to augment natural recruitment and to reestablish populations. If hatchery-reared lake sturgeon are more likely to stray than naturally produced individuals, as documented in other well-studied species, outbreeding could potentially jeopardize beneficial site-specific phenotypic and genotypic adaptations. From 1983 to 1994, lake sturgeon propagated using eggs taken from Lake Winnebago adults (Lake Michigan basin) were released in the St. Louis River estuary in western Lake Superior. Our objective was to determine whether these introduced individuals have strayed into annual spawning runs in the Sturgeon River, Michigan. Additionally, we estimated a natural migration rate between the Sturgeon River and Bad River, Wisconsin populations. Presumed primiparous lake sturgeon sampled during Sturgeon River spawning runs from 2003 to 2008 were genotyped at 12 microsatellite loci. Genotypic baselines established for the Sturgeon River (n = 101), Bad River (n = 40), and Lake Winnebago river system (n = 73) revealed a relatively high level of genetic divergence among populations (mean F_ST = 0.103; mean R_ST = 0.124). Likelihood-based assignment tests indicated no straying of stocked Lake Winnebago strain lake sturgeon from the St. Louis River into the Sturgeon River spawning population. One presumed primiparous Sturgeon River individual likely originated from the Bad River population. Four first-generation migrants were detected in the Sturgeon River baseline, indicating an estimated 3.5% natural migration rate for the system.     

Contrasting shell/tissue characteristics of Dreissena polymorpha and Dreissena bugensis in relation to environmental heterogeneity in the St. Lawrence River

The zebra mussel, Dreissena polymorpha, is widespread in the St. Lawrence River while the conspecific quagga mussel, Dreissena bugensis, is found only in the Lake Ontario outflow region of the river. This situation provided an opportunity to evaluate in situ environmental and interspecific heterogeneity in shell and tissue growth. Shell dry weight, carbon content, and shell strength of D. polymorpha from the four spatially discrete water masses differed significantly. For instance, D. polymorpha total and tissue mass increased over the summer in the shallow fluvial Lac Saint-Pierre but decreased in the upstream and downstream water masses. Standardized shell mass and strength of D. polymorpha was lowest where the mussels experienced salinity or low calcium. Although the response pattern of mass and glycogen content for D. polymorpha was spatially complex, mussels from the stressful oligohaline estuary population had the weakest shells and lowest glycogen content, even though their standardized tissue mass was the heaviest. This disparity in shell and tissue response suggests that some aspect of shell physiology alone may be limiting these mussels in estuarine environments. Tissue characteristics of D. polymorpha and D. bugensis were similar at the site where both were present, but the shell strength of D. bugensis was only equivalent to the weakest of D. polymorpha. We also conclude that lighter shells might make D. bugensis more susceptible to predation or mechanical damage but may also offer a bioenergetic advantage that is contributing to its rapid displacement of D. polymorpha where the two species co-occur.     

Expansion of Dreissena into offshore waters of Lake Michigan and potential impacts on fish populations

Lake Michigan was invaded by zebra mussels (Dreissena polymorpha) in the late 1980s and then followed by quagga mussels (D. bugensis) around 1997. Through 2000, both species (herein Dreissena) were largely restricted to depths less than 50 m. Herein, we provide results of an annual lake-wide bottom trawl survey in Lake Michigan that reveal the relative biomass and depth distribution of Dreissena between 1999 and 2007 (although biomass estimates from a bottom trawl are biased low). Lake-wide mean biomass density (g/m²) and mean depth of collection revealed no trend between 1999 and 2003 (mean = 0.7 g/m² and 37 m, respectively). Between 2004 and 2007, however, mean lake-wide biomass density increased from 0.8 g/m² to 7.0 g/m², because of increased density at depths between 30 and 110 m, and mean depth of collection increased from 42 to 77 m. This pattern was confirmed by a generalized additive model. Coincident with the Dreissena expansion that occurred beginning in 2004, fish biomass density (generally planktivores) declined 71% between 2003 and 2007. Current understanding of fish population dynamics, however, indicates that Dreissena expansion is not the primary explanation for the decline of fish, and we provide a species-specific account for more likely underlying factors. Nonetheless, future sampling and research may reveal a better understanding of the potential negative interactions between Dreissena and fish in Lake Michigan and elsewhere.     

Evidence of lake trout (Salvelinus namaycush) spawning and spawning habitat use in the Dog River,Lake Superior

Lake trout spawn primarily in lakes, and the few river-spawning populations that were known in Lake Superior were believed to be extirpated. We confirmed spawning by lake trout in the Dog River, Ontario, during 2013–2016 by the collection of and genetic identification of eggs, and we describe spawning meso- and microhabitat use by spawning fish. Between 2013 and 2016, a total of 277 lake trout eggs were collected from 39 of 137 sampling locations in the river. The majority of eggs (220) were collected at the transition between the estuary and the river channel crossing the beach. Lake trout eggs were most often located near the downstream end of pools in areas characterized by rapid changes in depth or slope, coarse substrates, and increased water velocities, where interstitial flows may occur. Depths in wadeable areas where eggs were found averaged 0.9?m (range: 0.4 to 1.3?m) and substrate sizes consisted of large gravel, cobble, and boulder; comparable to spawning characteristics noted in lakes. Water velocities averaged 0.66?m·s^?1 (range: 0.33 to 1.7?m³·s^?1) at mid-depth. This information on spawning habitat could be used to help locate other remnant river-spawning populations and to restore river-spawning lake trout and their habitat in rivers that previously supported lake trout in Lake Superior. The Dog River population offers a unique opportunity to understand the ecology of a river spawning lake trout population.     

Decline in bloater fecundity in Southern Lake Michigan after decline of Diporeia

Population fecundity can vary through time, sometimes owing to changes in adult condition. Consideration of these fecundity changes can improve understanding of recruitment variation. Herein, we estimated fecundity of Lake Michigan bloater Coregonus hoyi during December 2005 and February 2006. Bloater recruitment has been highly variable from 1962 to present, and consistently poor since 1992. We compared our fecundity vs. weight regression to a previously published regression that used fish sampled in October 1969. We wanted to develop a new regression for two reasons. First, it should be more accurate because it uses fish collected closer to spawning, thus minimizing the potential for atresia (egg reabsorption) which could bias fecundity high. Second, we hypothesized that fecundity would be lower in 2006 because adult condition was 41% lower in 2006 compared to 1969, likely owing to the decline of Diporeia spp, a primary prey for bloater. Although the slope of the fecundity versus weight regression was similar between the years, fecundity was 24% lower in 2006 than in 1969 for bloater weighing between 70 and 240 g. Whether this was the result of the difference in sampling time prior to spawning or of differences in condition is unknown. We also found no relationship between maternal size and mature oocyte size. Incorporating our updated fecundity regression into a stock/recruit model failed to improve the model fit, indicating that the low bloater recruitment that has been observed since the early 1990s is not solely the result of reduced fecundity.     

DNA barcoding to confirm morphological traits and determine relative abundance of burrowing mayfly species in western Lake Erie

Burrowing mayfly species of the genus Hexagenia are well known indicators of environmental health in lakes and rivers. Two species, H. limbata and H. rigida, are indistinguishable as nymphs and as adult females. Our objectives were to develop a genetic technique to distinguish between the two species and identify morphological features that separate cryptic nymphs and adult females. Fifty nymphs were collected before emergence from 10 sites throughout the western basin of Lake Erie in 2004 and 2005. Using known specimens of adult aerial male H. limbata and H. rigida, we used the mitochondrial cytochrome c oxidase subunit 1 (COI) gene to identify a 16 base pair (bp) difference between species. DNA sequencing confirmed correct species identification based on differences in abdominal pigmentation patterns on adult female imagos in 19 of 20 cases; the lone exception was a female with very faint pigmentation. Pigmentation patterns between species were consistent on nymphs, subimagos and imagos of both sexes. Populations of both species are panmictic across the western basin of Lake Erie, but H. limbata is the numerically dominant species, representing 70 to 100% of nymphs at sites in both years. A separate lineage of H limbata was discovered in the samples. The ability to distinguish nymphs of the two species will aid in developing more sensitive ecosystem indicators.     

Delayed egg hatching accounts for replacement of burrowing mayflies Hexagenia rigida by Hexagenia limbata after recolonization in western Lake Erie

Egg banking may have played a role in the recolonization of two burrowing mayfly species (Hexagenia limbata and Hexagenia rigida), which recolonized western Lake Erie after over 30 years of near extirpation. H. rigida was the first of the two to colonize successfully, but was overtaken by H. limbata (the historically dominant species). To understand mechanisms of species replacement, we compared egg hatching between the two species at 20 °C in the lab under typical hatching temperatures (no incubation) and after cold storage (8 °C) for 2, 6 and 12 months. Prolonged storage at cold temperatures simulates temperature conditions to which embryos are exposed when deposited by late-emerging female imagos. Without incubation, H. rigida (95.4 ± 2.02) exhibited significantly higher egg hatching survivorship than H. limbata (85 ± 3.2) (P = 0.01). First hatch date for H. rigida was one day earlier than H. limbata, but time for 50% to hatch (5 d) and hatching duration (5 d) was the same for both species. After 2 and 6 months of incubation, H. limbata exhibited an earlier hatch date, but there was no significant difference in mean percentage of egg survival (ca 63%) between species. After 12 months of cold storage, hatching success for H. limbata was 44.6 ± 4.17%, but H. rigida eggs did not hatch. Banking of eggs at low temperature increases the chance of successful recolonization, once stressful conditions pass. The viability H. limbata eggs after long quiescence may help to explain the shift in dominance from H. rigida to H. limbata.     

Stock assessment and fishery management of Henicorhynchus spp., Cyclocheilichthys enoplos and Channa micropeltes in Tonle Sap Great Lake,Cambodia

Tonle Sap Great Lake, in the lower Mekong River basin, contributes 60% of Cambodia's inland fisheries catch. There are four types of fisheries, including the middle-scale fishery. The major species comprising the middle-scale fishery are Henicorhynchus spp. (16%), a small-bodied fish caught mainly with small-mesh nets, and Cyclocheilichthys enoplos (13%), and Channa micropeltes (7%) which are caught with a variety of gear. Here we apply the Schaefer surplus production model to the middle-scale fisheries using monitoring data collected between 1995 and 1999. Model simulations determined that the maximum sustainable yield (F_MSY) for Henicorhynchus spp., was obtained with the use of 47,206 gillnets (mesh size < 50 mm) and 4269 and 1605 fishing boats for C. enoplos and C. microplestes, respectively. Over 1995–1999, there was a strong trend of increase in fishing effort and decrease of catch-per-unit-effort in the waters of Pursat, Siem Riep and Kampong Chnnang provinces. These provinces have large population centres located close to the fishing grounds; fishing in these areas should be more strictly regulated.     

Contemporary abundance patterns of Cyclotella sensu lato diatom taxa in Lake Superior: Assessing responses to physical and chemical gradients and potential links to climate change

Many studies attribute shifts in relative abundances of Cyclotella sensu lato diatom taxa in freshwater lakes over the past 150?years to climate change. Although abundances of several of these taxa have also changed in Lake Superior over the last century, the specific drivers are currently unclear. This study used multiple linear regression to analyze long-term planktonic monitoring data to identify drivers of changing cell densities of six Cyclotella sensu lato species, four shown to be responsive to climate-driven change in smaller lakes and two species with similar morphology to two previously assessed taxa: Discostella stelligera and the morphologically similar D. pseudostelligera, Lindavia comensis and the morphologically similar L. delicatula, Lindavia ocellata, and Lindavia bodanica. Assessed variables associated with climate-driven physical change included surface water temperature and light availability, with turbidity and thermocline depth as indicators of the light environment. One to three variables explained 8–61% variation for each of 5 taxa from 2001 to 2011, significant predictive variables were not identified for D. stelligera. Multiple linear regression identified nutrients, including nitrate, phosphorus, and silica, as essential in explaining distributions of all the other assessed taxa. A physical variable associated with climate explained changes of one taxon: L. ocellata, which increased with lower water temperatures. Climate-driven species responses were not as apparent as in other systems; specifically, changes in the light environment of this clear lake did not explain abundance changes in any of the taxa. Multiple and complex environmental variables drive Cyclotella sensu lato abundance patterns in Lake Superior, possibly suggesting ecotype formation.