Isolation of Aeromonas salmonicida subspecies salmonicida from Lake Whitefish (Coregonus clupeaformis) inhabiting Lakes Michigan and Huron

Herein we describe the first report of Aeromonas salmonicida subspecies salmonicida infections in lake whitefish (Coregonus clupeaformis) collected from four sites in lakes Michigan and Huron, Michigan, USA. The bacterium was isolated from the kidneys of four out of 1286 lake whitefish that were tested over a three-year period. The four isolates were phenotypically similar to one another and exhibited the morphological, colonial, and biochemical traits typical of A. salmonicida subspecies salmonicida. Amplification of 16S rRNA genes specific to A. salmonicida subspecies salmonicida via polymerase chain reaction and subsequent gel electrophoresis analyses confirmed the identity of the four lake whitefish isolates. Clinical signs associated with infection included extensive external hemorrhaging, exophthalmia, splenomegaly, splenic and renal congestion, fibrinous adhesions of the spleen and liver, and hemorrhagic enteritis. Histopathological examination of infected fish revealed multi-focal hemorrhage and infiltration of lymphocytes and histiocytes in subdermal adipose tissues and musculature. A low infection incidence of A. salmonicida salmonicida in Great Lakes lake whitefish does not preclude the fact that overt signs of disease were observed in infected individuals and that lake whitefish may act as a reservoir for this bacterium that is highly pathogenic to numerous fish species.     

Widespread infection of lake whitefish Coregonus clupeaformis with the swimbladder nematode Cystidicola farionis in northern lakes Michigan and Huron

We estimated the prevalence, intensity, and abundance of swimbladder nematode infection in 1281 lake whitefish (Coregonus clupeaformis) collected from four sites in northern lakes Huron (Cheboygan and DeTour Village) and Michigan (Big Bay de Noc and Naubinway) from fall 2003 through summer 2006. Morphological examination of nematode egg, larval, and mature stages through light and scanning electron microscopy revealed characteristics consistent with that of Cystidicola farionis Fischer 1798. Total C. farionis prevalence was 26.94%, while the mean intensity and abundance of infection was 26.72 and 7.21 nematodes/fish, respectively. Although we detected C. farionis in all four stocks that were examined, Lake Huron stocks generally had higher prevalence, intensity, and abundance of infection than Lake Michigan stocks. A distinct seasonal fluctuation in prevalence, abundance, and intensity of C. farionis was observed, which does not coincide with reported C. farionis development in other fish species. Lake whitefish that were heavily infected with C. farionis were found to have thickened swimbladder walls with deteriorated mucosa lining, which could affect swimbladder function. Whether C. farionis infection may be negatively impacting lake whitefish stocks in the Great Lakes is unclear; continued monitoring of C. farionis infection should be conducted to measure responses of lake whitefish stocks to infection levels.     

Genetic diversity of lake whitefish in lakes Michigan and Huron; sampling,standardization, and research priorities

We combined data from two laboratories to increase the spatial extent of a genetic data set for lake whitefish Coregonus clupeaformis from lakes Huron and Michigan and saw that genetic diversity was greatest between lakes, but that there was also structuring within lakes. Low diversity among stocks may be a reflection of relatively recent colonization of the Great Lakes, but other factors such as recent population fluctuation and localized stresses such as lamprey predation or heavy exploitation may also have a homogenizing effect. Our data suggested that there is asymmetrical movement of lake whitefish between Lake Huron and Lake Michigan; more genotypes associated with Lake Michigan were observed in Lake Huron. Adding additional collections to the calibrated set will allow further examination of diversity in other Great Lakes, answer questions regarding movement among lakes, and estimate contributions of stocks to commercial yields. As the picture of genetic diversity and population structure of lake whitefish in the Great Lakes region emerges, we need to develop methods to combine data types to help identify important areas for biodiversity and thus conservation. Adding genetic data to existing models will increase the precision of predictions of the impacts of new stresses and changes in existing pressures on an ecologically and commercially important species.     

A 90-year record of lake whitefish Coregonus clupeaformis abundances in Michigan waters of the upper Laurentian Great Lakes

Fishery managers throughout the upper Laurentian Great Lakes are currently faced with a two-decade decline in abundance and harvest of lake whitefish Coregonus clupeaformis stocks. We used multivariate auto-regressive state-space (MARSS) models to develop long-term (90-year; 1929–2018) time-series of lake whitefish relative abundance based on commercial catch-per-effort (CPE) data for 13 statistical districts in State of Michigan waters, including 1836 and 1842 Treaty ceded waters, of Lakes Superior, Michigan, and Huron. CPE time-series were used to estimate historical baseline conditions, which were compared to more recent conditions, specifically with reference to select regulatory, environmental, and ecological conditions in each lake and fishing intensity. Population growth rates suggested that lake whitefish stocks responded: (1) negatively to high levels of harvest and expansion of sea lamprey Petromyzon marinus populations during the early-1900s; (2) positively to commercial fishery regulation and sea lamprey control during the late-1950s and early-1960s; and (3) negatively to establishment of dreissenid mussels Dreissena spp. in Lakes Michigan and Huron by 2005 and the recent period of low productivity in all three lakes since the mid- to late-2000s. When placed in a historical context, the most recent (2011–2018) lake whitefish abundances are low, intermediate, and high in 31 %, 46 %, and 23 % of all districts examined. Although environmental and ecological conditions likely drove recent declines, correlation analysis suggested that higher levels of fishing intensity were associated with greater district-specific declines in abundance during the last two decades (1999–2016), a period characterized by lower overall productivity and limited recruitment in most lake whitefish stocks.     

Estimates of fishing and natural mortality rates for four Lake Whitefish stocks in Northern Lakes Huron and Michigan

We analyzed tag-recovery data to estimate instantaneous fishing (F) and natural mortality (M) rates of four lake whitefish stocks in lakes Michigan and Huron during 2004–2007. We tagged and released 22,452 adult lake whitefish of which 8.7% were subsequently recovered. Annual tag-reporting rates ranged from 17.8% to 56.2%. Tag retention was high for the first 5–6 months after tagging, but tag loss increased substantially thereafter. Nine tag-recovery models were evaluated with respect to whether F and/or M varied among stocks, lakes, or years. There was support for three models based on Akaike information criteria. The best model had yearly and stock-specific estimates of F of 0.03 to 0.79 and lake-specific estimates of M of 0.35 for Lake Michigan and 0.60 for Lake Huron. The second best model had yearly and stock-specific estimates of F of 0.04 to 0.71 and a constant estimate for M of 0.52. The third model had yearly and stock-specific estimates of F of 0.04 to 0.85 and stock-specific estimates of M of 0.32 to 0.67. Model-averaged estimates of F ranged from 0.04 to 0.78 and were substantially different than statistical catch-at-age estimates of F. Model-averaged estimates of M ranged from 0.40 to 0.59 and were greater than estimates obtained from prediction equations, possibly due to sea lamprey-induced mortality. We recommend that tag-recovery estimates of F and M be used as Bayesian priors in future lake whitefish stock assessments to help refine mortality estimates for the stocks.     

Lake whitefish (Coregonus clupeaformis) energy and nutrient partitioning in lakes Michigan,Erie and Superior

A concurrent decrease in lake whitefish (Coregonus clupeaformis) condition and Diporeia spp. abundance in Lake Michigan has spurred investigations into possible links between the two phenomena. We examined female lake whitefish δ¹³C and δ¹⁵N stable isotopes, growth, reproductive investment, dorsal muscle total lipid and docosahexaenoic acid (DHA) contents from lakes Erie, Michigan and Superior to determine whether differences in food source were correlated with measures of stock success. Stocks with higher somatic growth rates and mean reproductive potential had higher energy stores in terms of percent total lipid. Stocks with low muscle lipid concentration also had smaller egg sizes as egg number increased. Diet varied among stocks as evidenced by δ¹³C and δ¹⁵N stable isotope analyses; however, muscle total lipid and DHA were not correlated to apparent Diporeia spp. prey use. When compared to stocks from lakes Erie and Superior, Lake Michigan stocks had lower growth, reproduction, and lipid stores. While stocks in Lake Michigan with access to declining Diporeia spp. populations may still feed on the amphipod, it appears that they are unable to consume the quantities necessary to maintain historical growth and reproduction. Stable isotope analyses of lakes Erie and Superior stocks, with higher growth rates and lipid values, indicated different feeding strategies with no indication of reliance on Diporeia spp. While differences in prey resources may have an effect on lake whitefish stocks, differences in Diporeia spp. abundance alone cannot explain differences in lake whitefish condition observed among the Great Lakes included in this study.     

Assessment of Renibacterium salmoninarum infections in four lake whitefish (Coregonus clupeaformis) stocks from northern Lakes Huron and Michigan

Lake whitefish (Coregonus clupeaformis) from four stocks in northern Lakes Michigan and Huron were collected seasonally from fall 2003 through summer 2006 and examined for the presence of Renibacterium salmoninarum, the causative agent of bacterial kidney disease (BKD), using culture techniques on modified kidney disease medium (MKDM) and the quantitative enzyme-linked immunosorbent assay (Q-ELISA). R. salmoninarum was detected in 62.31% (according to Q-ELISA) of the 1284 examined lake whitefish, with some fish displaying the typical signs of BKD, such as renal congestion, swelling, and whitish nodules. Kidney cultures on MKDM yielded bacteria with morphological and biochemical characteristics identical to those of R. salmoninarum recovered from other Great Lakes fish species, as well as those from other parts of the world. Isolate identification was confirmed via nested polymerase chain reaction. Antibiograms demonstrated high sensitivity to enrofloxacin and ciprofloxacin, sensitivity to oxytetracycline, erythromycin, azithromycin, chloramphenicol, novobiocin, and carbenicillin, and resistance to polymyxin B, clindamycin, and kanamycin. Statistical analysis of R. salmoninarum prevalence and intensities revealed significant interactions among stocks, years and sampling seasons, with highest prevalence generally in fall and frequent wide variation in prevalence and intensity from one season to the next for a particular stock. It was surprising to find that the prevalence of R. salmoninarum exceeded 50% in the four stocks, much higher than originally thought. Moreover, a positive association between R. salmoninarum intensity and the abundance of the swimbladder nematode, Cystidicola farionis, was identified. Our findings suggest that Great Lakes lake whitefish are vulnerable to serious fish pathogens.     

Spatial and temporal dynamics of lake whitefish (Coregonus clupeaformis) health indicators: Linking individual-based indicators to a management-relevant endpoint

We examined the spatial and temporal dynamics of health indicators in four lake whitefish (Coregonus clupeaformis) stocks located in northern lakes Michigan and Huron from 2003 to 2006. The specific objectives were to (1) quantify spatial and temporal variability in health indicators; (2) examine relationships among nutritional indicators and stock-specific spatial and temporal dynamics of pathogen prevalence and intensity of infection; and (3) examine relationships between indicators measured on individual fish and stock-specific estimates of natural mortality. The percent of the total variation attributed to spatial and temporal sources varied greatly depending on the health indicator examined. The most notable pattern was a downward trend in the concentration of highly unsaturated fatty acids (HUFAs), observed in all stocks, in the polar lipid fraction of lake whitefish dorsal muscle tissue over the three study years. Variation among stocks and years for some indicators were correlated with the prevalence and intensity of the swimbladder nematode Cystidicola farionis, suggesting that our measures of fish health were related, at some level, with disease dynamics. We did not find relationships between spatial patterns in fish health indicators and estimates of natural mortality rates for the stocks. Our research highlights the complexity of the interactions between fish nutritional status, disease dynamics, and natural mortality in wild fish populations. Additional research that identifies thresholds of health indicators, below (or above) which survival may be reduced, will greatly help in understanding the relationship between indicators measured on individual fish and potential population-level effects.     

Dreissenid mussels are not a “dead end” in Great Lakes food webs

Dreissenid mussels have been regarded as a “dead end” in Great Lakes food webs because the degree of predation on dreissenid mussels, on a lakewide basis, is believed to be low. Waterfowl predation on dreissenid mussels in the Great Lakes has primarily been confined to bays, and therefore its effects on the dreissenid mussel population have been localized rather than operating on a lakewide level. Based on results from a previous study, annual consumption of dreissenid mussels by the round goby (Neogobius melanostomus) population in central Lake Erie averaged only 6 kilotonnes (kt; 1 kt = one thousand metric tons) during 1995–2002. In contrast, our coupling of lake whitefish (Coregonus clupeaformis) population models with a lake whitefish bioenergetics model revealed that lake whitefish populations in Lakes Michigan and Huron consumed 109 and 820 kt, respectively, of dreissenid mussels each year. Our results indicated that lake whitefish can be an important predator on dreissenid mussels in the Great Lakes, and that dreissenid mussels do not represent a “dead end” in Great Lakes food webs. The Lake Michigan dreissenid mussel population has been estimated to be growing more than three times faster than the Lake Huron dreissenid mussel population during the 2000s. One plausible explanation for the higher population growth rate in Lake Michigan would be the substantially higher predation rate by lake whitefish on dreissenid mussels in Lake Huron.     

Thiamine concentrations in lake whitefish eggs from the upper Great Lakes are related to maternal diet

Thiamine deficiency is responsible for reproductive impairment in several species of salmonines in the Great lakes, and is thought to be caused by the consumption of prey containing thiaminase, a thiamine-degrading enzyme. Because thiaminase levels are extremely high in dreissenid mussels, fish that prey on them may be susceptible to thiamine deficiency. We determined thiamine concentrations in lake whitefish Coregonus clupeaformis eggs from the upper Laurentian Great Lakes to assess the potential for thiamine deficiency and to determine if thiamine concentrations in lake whitefish eggs were related to maternal diet. Mean thiamine concentrations in lake whitefish eggs were highest in Lake Huron, intermediate in Lake Superior, and lowest in Lake Michigan. Some fish had thiamine concentrations below putative thresholds for lethal and sublethal effects in salmonines, suggesting that some larval lake whitefish may currently be at risk of at least sublethal effects of low thiamine concentrations, although thiamine thresholds are unknown for lake whitefish. Egg thiamine concentrations in lake whitefish eggs were statistically significantly related to isotopic carbon signatures, suggesting that egg thiamine levels were related to maternal diet, but low egg thiamine concentrations did not appear to be associated with a diet of dreissenids. Egg thiamine concentrations were not statistically significantly related to multifunction oxidase induction, suggesting that lower egg thiamine concentrations in lake whitefish were not related to contaminant exposure.     

Trends in Mysis diluviana abundance in the Great Lakes, 2006–2016

With the large Diporeia declines in lakes Michigan, Huron, and Ontario, there is concern that a similar decline of Mysis diluviana related to oligotrophication and increased fish predation may occur. Mysis density and biomass were assessed from 2006 to 2016 using samples collected by the Great Lakes National Program Office's biomonitoring program in April and August in all five Great Lakes. Summer densities and biomasses were generally greater than spring values and both increased with bottom depth. There were no significant time trends during these 10–11 years in lakes Ontario, Michigan, or Huron, but there was a significant increase in Lake Superior. Density and biomass were highest in lakes Ontario and Superior, somewhat lower in Lake Michigan, and substantially lower in Lake Huron. A few Mysis were collected in eastern Lake Erie, indicating a small population in the deep basin of that lake. On average, mysids contributed 12–18% (spring-summer, Michigan), 18–14% (spring-summer, Superior), 30–13% (spring-summer, Ontario), and 3% (Huron) of the total open-water crustacean biomass. Size distributions consisted of two peaks, indicating a 2-year life cycle in all four of the deep lakes. Mysis were larger in Lake Ontario than in lakes Michigan, Superior, and Huron. Comparisons with available historic data indicated that mysid densities were higher in the 1960s–1990s (5 times higher in Huron, 2 times higher in Ontario, and around 40% higher in Michigan and Superior) than in 2006–2016.     

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.     

Spatial and temporal distributions of lake whitefish spawning stocks in Northern lakes Michigan and Huron, 2003–2008

Adult lake whitefish were tagged and released from the Big Bay de Noc (BBN) and Naubinway (NAB) stocks in northern Lake Michigan, and the Detour (DET) and Cheboygan (CHB) stocks in northern Lake Huron during 2003–2006 to describe their spatial and temporal distributions. The contemporary spatial distributions were compared with past distributions of the BBN and NAB stocks. Sixty-two percent of BBN tag recoveries occurred in Wisconsin waters during winter, spring and summer, but 83% of fall tag recoveries were made near the tagging site. Eighty-eight percent of the NAB tag recoveries were made in the management unit of tagging and 7% occurred into northern Lake Huron. Over 90% of the DET stock remained in the vicinity of the tagging sites regardless of the season, while 75% of the CHB tag recoveries were made in northwestern Lake Huron and 17% were made in Ontario. Based on regression tree analysis, there were strong stock, season, and year effects on movement distances, with weaker effects due to sex and length at tagging. Spatial distribution of the BBN stock changed from 1978–1982 to 2003–2008, but spatial distribution of the NAB stock did not. Substantial differences in movement and distribution existed among the four stocks, large seasonal differences in spatial distribution were found within some stocks, and lake whitefish exhibited strong spawning site fidelity. Present management unit boundaries are inappropriate for managing three of our four stocks, and agencies should consider developing single harvest limits for both northern Lake Huron and western Lake Michigan.     

A cross-lake comparison of crustacean zooplankton communities in the Laurentian Great Lakes, 1997–2016

Spring and summer open-water crustacean zooplankton communities were examined across all five Laurentian Great Lakes from 1997 to 2016. Spring communities were dominated by calanoid (lakes Superior, Huron and Michigan) or cyclopoid (lakes Erie and Ontario) copepods. Volumetric biomass of summer communities increased along an assumed trophic gradient (Superior, Huron, Michigan, Ontario; eastern, central and western Erie), as did dominance by cyclopoids and cladocerans. Over the time series of the study, summer communities in lakes Michigan, Huron and Ontario shifted towards greater dominance by calanoids and greater similarity with Lake Superior. Trajectories of changes were different; however, reductions in cladocerans accounted for most of the change in lakes Michigan and Huron while reductions in cyclopoids and increases in Leptodiaptomus sicilis were behind the changes in Lake Ontario. Shifts in the predatory cladoceran community in Lake Ontario from Cercopagis pengoi to occasional dominance by Bythotrephes longimanus, a species much more vulnerable to planktivory, as well as the appearance of Daphnia mendotae in a daphnid community previously consisting almost exclusively of the smaller Daphnia retrocurva, suggest impacts of reduced vertebrate predation. In contrast, strong correlations between cladocerans and chlorophyll in lakes Michigan and Huron point to the possible importance of bottom-up forces in those lakes. Large interannual shifts in cladoceran community structure in the central and eastern basins of Lake Erie suggest intense but variable vertebrate predation pressure. The zooplankton communities of lakes Huron, Michigan and Ontario may be approaching a historic community structure represented by Lake Superior.     

A comparative examination of recent changes in nutrients and lower food web structure in Lake Michigan and Lake Huron

The lower food webs of Lake Huron and Lake Michigan have experienced similar reductions in the spring phytoplankton bloom and summer populations of Diporeia and cladocerans since the early 2000s. At the same time phosphorus concentrations have decreased and water clarity and silica concentrations have increased. Key periods of change, identified by using a method based on sequential t-tests, were 2003–2005 (Huron) and 2004–2006 (Michigan). Estimated filtration capacity suggests that dreissenid grazing would have been insufficient to directly impact phytoplankton in the deeper waters of either lake by this time (mid 2000s). Despite some evidence of decreased chlorophyll:TP ratios, consistent with grazing limitation of phytoplankton, the main impact of dreissenids on the offshore waters was probably remote, e.g., through interception of nutrients by nearshore populations. A mass balance model indicates that decreased phosphorus loading could not account for observed in-lake phosphorus declines. However, model-inferred internal phosphorus dynamics were strongly correlated between the lakes, with periods of increased internal loading in the 1990s, and increased phosphorus loss starting in 2000 in Lake Michigan and 2003 in Lake Huron, prior to dreissenid expansion into deep water of both lakes. This suggests a limited role for deep populations of dreissenids in the initial phosphorus declines in the lakes, and also suggests a role for meteorological influence on phosphorus dynamics. The high synchrony in lower trophic level changes between Lake Michigan and Lake Huron suggests that both lakes should be considered when investigating underlying causal factors of these changes.     

Predation on Lake Whitefish Eggs by Longnose Suckers

In November 1981, we observed intense predation on lake whitefish (Coregonus clupeaformis) eggs by longnose suckers (Catostomus catostomus) on lake whitefish spawning grounds in northwestern Lake Huron. Since longnose suckers commonly frequent the same habitat used by spawning lake whitefish, there exists the potential for high losses of eggs due to sucker predation.     

Determining the efficacy of microsatellite DNA-based mixed stock analysis of Lake Michigan's Lake Whitefish commercial Fishery

Management of commercially exploited fish should be conducted at the stock level. If a mixed stock fishery exists, a comprehensive mixed stock analysis is required for stock-based management. The lake whitefish Coregonus clupeaformis comprises the primary commercial fishery across the Great Lakes. Recent research resolved that six genetic stocks of lake whitefish were present in Lake Michigan, and long-term tagging data indicate that Lake Michigan's lake whitefish commercial fishery is a mixed stock fishery. The objective of this research was to determine the usefulness of microsatellite data for conducting comprehensive mixed stock analyses of the Lake Michigan lake whitefish commercial fishery. We used the individual assignment method as implemented in the program ONCOR to determine the accuracy level at which microsatellite data can reliably identify component populations or stocks. Self-assignment of lake whitefish to their population and stock of origin ranged from > 96% to 100%. Evaluation of genetic stock discreteness indicated a moderately high degree of correct assignment (average = 75%); simulations indicated supplementing baseline data by ～ 50 to 100 individuals could increase accuracy by up to 4.5%. Simulated mixed stock commercial harvests with known stock composition showed a high degree of correct proportional assignment between observed and predicted harvest values. These data suggest that a comprehensive mixed stock analysis of Lake Michigan's lake whitefish commercial fishery is viable and would provide valuable information for improving management.     

Lipid Concentration and Size Variation of Bythotrephes (Cladocera: Cercopagidae) from Lakes Erie,Huron, and Michigan

Lipid concentrations of Bythotrephes cederstroemi were compared among three Great Lakes, Erie, Huron, and Michigan, in an effort to investigate the phenotypic plasticity in size displayed among the lakes. Four developmental stages were measured in Lakes Erie and Huron and two stages were studied in Lake Michigan. With a gravimetric extraction method, the total lipid concentration range (μg lipid μg dry weight⁻¹, expressed as percent) for Bythotrephes was estimated to be 10–19%. Statistically significant differences were found in lipid concentrations of Bythotrephes among lakes and developmental stages. Lake Erie had significantly higher lipid concentration values than Lake Huron for stages 2 through 4, and had similar values to Lake Michigan for the analyzed stages 1 and 4. The first instar had indistinguishable lipid concentrations among Lakes Erie, Huron,and Michigan. Even though animals from Lake Erie were significantly smaller, the data suggest that they were not less well nourished. We hypothesize that selective mortality imposed by visual predators on larger Bythotrephes and the lack of deep water refuges in Lake Erie has encouraged the smaller size of Bythotrephes found there in comparison to those found in Lakes Huron and Michigan.     

Declines in lake whitefish larval densities after dreissenid mussel establishment in Lake Huron

Lake whitefish (Coregonus clupeaformis) is an ecologically and commercially significant species across the Laurentian Great Lakes. Over the past 20 years, lake whitefish population abundance has substantially declined across lakes Huron and Michigan, being driven by reduced recruitment of juvenile fish into the population. However, the life stage at which the recruitment bottleneck is occurring and what factors are contributing to the declines remain unknown. One hypothesis is that dreissenid mussels reduced zooplankton availability to larval lake whitefish, leading to poor growth and survival of this critical life stage. Here, we present results of a larval fish survey conducted at the Fishing Islands spawning region in Lake Huron and examine whether declines in juvenile recruitment are linked to reduced larval fish density. Larval fish were collected annually during two time periods: (1) a historical time period before dreissenid mussel establishment (1976–1986); and (2) a contemporary time period after dreissenid mussels became established (2017–2019, 2021). We found significant declines in larval densities and growth between historical and contemporary time periods. Following dreissenid establishment, larval densities and growth were on average only 23% and 55% of historical values, respectively. Moreover, year class strength at the juvenile stage (age 4) was positively related to larval density. Several explanatory variables contributed to annual variation in larval densities, with dreissenid mussels and water levels having the most consistent effect. Our results suggest that juvenile recruitment is being limited at the larval stage, owing to overall lower larval production and potentially exacerbated by slower growth.     

Alkalinity,pH, and pCO2 in the Laurentian Great Lakes: An initial view of seasonal and inter-annual trends

Ongoing human perturbations to the global inorganic carbon cycle can cause various changes in the pH and alkalinity of aquatic systems. Here seasonal and inter-annual trends in these variables were investigated in the five Laurentian Great Lakes using data from the U.S. EPA GLENDA database. These observations, along with temperature, were also used to predict the partial pressure of carbon dioxide in surface water (pCO₂). There are strong seasonal differences in pH in all five lakes, with higher pH levels in summer than in spring. All lakes show significantly higher pCO₂ values in spring than in summer. Michigan and Ontario show higher alkalinity values in spring; Huron shows lower spring values. Inter-annually, open-lake pH shows the highest values in all lakes around 2010, the time frame of lowest lake water levels, though only lakes Superior and Erie show statistically significant inflection points at that time. Inter-annual alkalinity trends differ considerably from those of pH. Superior’s alkalinity increases until ～2008 and then begins dropping; Ontario’s alkalinity decreases until ～2004 and then begins increasing, with the decrease coinciding with the introduction and establishment of Dreissenid mussels. The other lakes show much less clear inter-annual alkalinity trends. For pCO₂, inter-annual trends in each lake show either increases from 1992 to 2019 (for Superior, Michigan, and Huron) or shifts from slightly decreasing values to increasing values for the other lakes. The timing of this shift is from 2008 to 2010.