Search

Groups of sympatric taxa with low interspecific genetic differentiation, but considerable ecological differences, offer great opportunities to study the dynamics of divergence and speciation. This is the case of ciscoes (Coregonus spp.) in the Laurentian Great Lakes, which are characterized by a complex evolutionary history and are commonly described as having undergone an adaptive radiation. In this study, morphometrics, stable isotopes and transcriptome sequencing were used to study the relationships within the Coregonus artedi complex in western Lake Superior. We observed general concordance for morphological, ecological and genomic variation, but the last was more taxonomically informative as it showed less overlap among species in multivariate space. Low levels of genetic differentiation were observed between individuals morphologically identified as Coregonus hoyi and C. zenithicus, which could be evidence of incomplete lineage sorting or recent hybridization between the two groups. Transcriptome-based single nucleotide polymorphisms exhibited significant divergence for genes associated with vision, development, metabolism and immunity among species that occupy different habitats. This study highlights the importance of using an integrative approach when studying groups of taxa with a complex evolutionary history, as individual-level analyses of multiple independent data sets can provide a clearer picture of the patterns and processes associated with the origins of biodiversity.

Hydroacoustic surveys using hull-mounted down-looking transducers are useful for estimating pelagic fish densities; however, this method may miss shallow fish owing to the acoustic surface dead zone and vessel avoidance. Our objective was to compare pelagic fish density estimates acquired by a traditional down-looking acoustic survey to estimates obtained by a new multi-directional-towed sled capable of sampling the entire water column using upward-, sideways-, and downward-aimed transducers simultaneously. We deployed both systems concurrently in the western arm of Lake Superior during a period of stable stratification. We found the two survey approaches provided significantly different estimates of fish density in the upper water column layer (~4–9 m below the lake surface) with the sled up-looking transducer providing 56 times higher densities compared to the traditional ship down-looking method. Densities also varied significantly in the 9–14 m layer where densities were 6.2 times higher in the sled survey. Midwater trawl sampling indicated that cisco (Coregonus artedi) and rainbow smelt (Osmerus mordax) were the predominant species occupying the uppermost 14 m of the water column. The two acoustic approaches provided similar results at water column depths >14 m where rainbow smelt and kiyi (Coregonus kiyi) were predominant. Overall, the sled-based method estimates were, on average, 2.5 times higher for the whole water column. Our findings show that the new sled can reduce bias by better sampling the surface dead zone leading to more accurate estimation of pelagic fish densities for both management and research.

Prey fish abundances in the Great Lakes are a driver for several agencies’ commitments to the Council of Lake Committees to support fisheries management. These management decisions have profound economic and social impacts within the Great Lakes region. Fisheries estimates done by echosounders or trawling may be biased due to the propagated noise from large fisheries vessels. In the first two years of a four-year collaborative study, crewed fisheries vessels and uncrewed Saildrone vessels were used to compare abundance estimates between “loud” and “quiet” vessels. To quantify the effects of ship noise on prey fish abundance estimates, a mobile ship noise measurement system was designed and deployed to measure radiated acoustic signatures of several ships in the Great Lakes. This talk will discuss the deployment of a mobile underwater acoustic test range and show results of ship noise measurements from the first two years of the program. In addition, plans for future data generation, analysis, and comparison between crewed and autonomous systems will be discussed and an overview of the effects of ship noise on fish avoidance will be discussed in the proceeding talk.

Acoustic surveys of fish are a foundational component of many fisheries monitoring programs, including surveys in the Great Lakes. These surveys are conducted with traditional crewed and motorized vessels, but fish avoidance of these types of platforms has been reported in multiple studies, potentially biasing estimates. A quiet uncrewed surface vessel, Saildrone, was equipped with a 120 kHz Simrad EK80 transducer and deployed in Lakes Huron, Michigan, and Superior in the summers of 2021 and 2022. The drone was then overtaken by numerous motorized vessels using transducers with the same frequency. The average target depth, target strength, and nautical area scattering during overtakes were compared. We looked for a fish behavioral response with General Additive Models using distance from the Saildrone to the vessel as the predictor. We also compared the effectiveness of acoustic surveys from each platform with analyses of variances over 2 km sections of the overtake. Fish showed a limited response to approaching vessels, and acoustic estimates of fisheries relevant measures were similar between Saildrone and motorized vessels. Findings from this work will inform interpretation of acoustic data in the Great Lakes and provide the largest scale testing of fish avoidance to acoustic surveys to date.

Ecosystem-based management relies on understanding how perturbations influence ecosystem structure and function (e.g., invasive species, exploitation, abiotic changes). However, data on unimpacted systems are scarce, therefore, we often rely on impacted systems to make inferences about ‘natural states.’ Among the Laurentian Great Lakes, Lake Superior provides a unique case study to address non-native species impacts because the food web is dominated by native species. Additionally, Lake Superior is both vertically (benthic versus pelagic) and horizontally (nearshore versus offshore) structured by depth, providing an opportunity to compare the function of these sub-food webs. We developed an updated Lake Superior EcoPath model using data from the 2005/2006 lake-wide multi-agency surveys covering multiple trophic levels. We then compared trophic transfer efficiency (TTE) to previously published EcoPath models. Finally, we compared ecosystem function of the 2005/2006 ecosystem to that with non-native linkages removed and compared native versus non-native species-specific approximations of TTE and trophic flow. Lake Superior was relatively efficient (TTE = 0.14) compared to systems reported in a global review (average TTE = 0.09) and the microbial loop was highly efficient (TTE > 0.20). Non-native species represented a very small proportion (

Groups of sympatric taxa with low inter-specific genetic differentiation, but considerable ecological differences, offer great opportunities to study the dynamics of divergence and speciation. This is the case of ciscoes (Coregonus spp.) in the Laurentian Great Lakes, which are characterized by a complex evolutionary history and are commonly described as having undergone an adaptive radiation. In this study, morphometrics, stable isotopes and transcriptome sequencing were used to study the relationships within the Coregonus artedi complex in western Lake Superior. We observed general concordance for morphological, ecological and genomic variation, but the latter was more taxonomically informative as it showed less overlap among species in multivariate space. Low levels of genetic differentiation were observed between individuals morphologically identified as C. hoyi and C. zenithicus, and we hypothesize this could be associated with recent hybridization between the two species. Transcriptome-based single nucleotide polymorphisms exhibited significant divergence for genes associated with vision, development, metabolism and immunity, among species that occupy different habitats. This study highlights the importance of using an integrative approach when studying groups of taxa with a complex evolutionary history, as individual-level analyses of multiple independent datasets can provide a clearer picture of the patterns and processes associated with the origins of biodiversity.

Local adaptation can drive diversification of closely related species across environmental gradients and promote convergence of distantly related taxa that experience similar conditions. We examined a potential case of adaptation to novel visual environments in a species flock (Great Lakes salmonids, genus Coregonus) using a new amplicon genotyping protocol on the Oxford Nanopore Flongle and MinION. We sequenced five visual opsin genes for individuals of Coregonus artedi, Coregonus hoyi, Coregonus kiyi, and Coregonus zenithicus. Comparisons revealed species-specific differences in a key spectral tuning amino acid in rhodopsin (Tyr261Phe substitution), suggesting local adaptation of C. kiyi to the blue-shifted depths of Lake Superior. Ancestral state reconstruction demonstrates that parallel evolution and “toggling” at this amino acid residue has occurred several times across the fish tree of life, resulting in identical changes to the visual systems of distantly related taxa across replicated environmental gradients. Our results suggest that ecological differences and local adaptation to distinct visual environments are strong drivers of both evolutionary parallelism and diversification.

Stock assessments are critical to modern fisheries management, supporting the calculation of key reference variables used to make informed management decisions. However, there is still considerable uncertainty as to which class of assessment models is appropriate to use under different circumstances. A common class of models used when age data are available are statistical catch-at-age assessment (SCAA) models, which track annual cohorts through time. When age data are unavailable, as is often the case in invertebrate fisheries where the lack of a bony structure such as otoliths makes aging difficult, statistical catch-at-size assessment (SCSA) models are more often employed, tracking fish or invertebrates through time by size-classes rather than ages. Do SCAA models actually perform better than SCSA models when age data are available, or is this just an assumption we make in fisheries research and management? We examined this question by evaluating the effectiveness of both SCAA and SCSA models in characterizing cisco, Coregonus artedi, population dynamics in Thunder Bay, Ontario. Both models were fit using an integrated framework with multiple sources of data including hydroacoustic estimates of spawning stock, fishery-dependent and -independent age/length compositions, and harvest data. Our results suggest that for cisco in Thunder Bay, data-limitations related to lack of size-composition data over the size range for which cisco growth is rapid resulted in difficulty estimating relative year-class strength within a SCSA. This led to parameter confounding and ultimately the inability to estimate natural mortality within a SCSA. This hampered the utility of a SCSA model in comparison with a SCAA model when age-composition data were available.

We examined the spatial scale of recruitment variability for disparate cisco (Coregonus artedi) populations in the Great Lakes (n = 8) and Minnesota inland lakes (n = 4). We found that the scale of synchrony was approximately 400 km when all available data were utilized; much greater than the 50-km scale suggested for freshwater fish populations in an earlier global analysis. The presence of recruitment synchrony between Great Lakes and inland lake cisco populations supports the hypothesis that synchronicity is driven by climate and not dispersal. We also found synchrony in larval densities among three Lake Superior populations separated by 25–275 km, which further supports the hypothesis that broad-scale climatic factors are the cause of spatial synchrony. Among several candidate climate variables measured during the period of larval cisco emergence, maximum wind speeds exhibited the most similar spatial scale of synchrony to that observed for cisco. Other factors, such as average water temperatures, exhibited synchrony on broader spatial scales, which suggests they could also be contributing to recruitment synchrony. Our results provide evidence that abiotic factors can induce synchronous patterns of recruitment for populations of cisco inhabiting waters across a broad geographic range, and show that broad-scale synchrony of recruitment can occur in freshwater fish populations as well as those from marine systems.

International audience; Fish species diversity can be lost through interacting stressors including habitat loss, stocking and overfishing. Although a multitude of stressors have played a role in the global decline of coregonid (Coregonus spp.) diversity, a number of contemporary studies have identified habitat loss stem-ming from eutrophication as the primary cause. Unfortunately, reconstructing the role of fishing and stocking practices can be difficult, because these records are incomplete or appear only in hard-to-access historic grey literature. Based on an illustrative set of historic and contemporary studies, we describe how fisheries management practices may have contributed to coregonid di-versity loss in European and North American lakes. We provide case studies examining how fish-ing and stocking may reduce coregonid diversity through demographic decline and introgressive hybridization. In some lakes, fisheries management practices may have led to a loss of coregonid diversity well before issues with habitat degradation manifested. Our review suggests that fish conservation policies could beneficially consider the relative importance of all stressors, including management practices, as potential drivers of diversity loss.

The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far-reaching impacts on aquatic food-webs, given previously described effects of Bythotrephes on zooplankton communities. We estimated consumption of Bythotrephes by planktivorous and benthivorous fishes, using bioenergetics and daily ration models at nearshore (18 m), intermediate (46 m), and offshore (110 m) depths along one western Lake Superior transect (April, and September–November) and two northern Lake Michigan transects (April, July, and September). In Lake Superior, consumption (primarily by cisco Coregonus artedi ) exceeded Bythotrephes production at all offshore sites in September–November (up to 396% of production consumed) and at the intermediate site in November (842%) with no evidence of consumption nearshore. By comparing Bythotrephes biomass following months of excessive consumption, we conservatively concluded that top-down control was evident only at the offshore site during September–October. In Lake Michigan, consumption by fishes (primarily alewife Alosa pseudoharengus ) exceeded production at nearshore sites (up to 178%), but not in deeper sites ( Bythotrephes never subsequently declined. Using generalized additive models, temperature, and not fish consumption nor zooplankton prey density, best explained variability in Bythotrephes biomass. The non-linear pattern revealed Bythotrephes to increase with temperature up to 16 °C, and then decline between 16 and 23 °C. We discuss how temperature likely has direct negative impacts on Bythotrephes when temperatures near 23 °C, but speculate that predation also contributes to declining biomass when temperatures exceed 16 °C.

The relative importance of predation and food availability as contributors to larval cisco ( Coregonus artedi ) mortality in Lake Superior were investigated using a visual foraging model to evaluate potential predation pressure by rainbow smelt ( Osmerus mordax ) and a bioenergetic model to evaluate potential starvation risk. The models were informed by observations of rainbow smelt, larval cisco, and zooplankton abundance at three Lake Superior locations during the period of spring larval cisco emergence and surface-oriented foraging. Predation risk was highest at Black Bay, ON, where average rainbow smelt densities in the uppermost 10 m of the water column were >1000 ha −1 . Turbid conditions at the Twin Ports, WI-MN, affected larval cisco predation risk because rainbow smelt remained suspended in the upper water column during daylight, placing them alongside larval cisco during both day and night hours. Predation risk was low at Cornucopia, WI, owing to low smelt densities ( −1 ) and deep light penetration, which kept rainbow smelt near the lakebed and far from larvae during daylight. In situ zooplankton density estimates were low compared to the values used to develop the larval coregonid bioenergetics model, leading to predictions of negative growth rates for 10 mm larvae at all three locations. The model predicted that 15 mm larvae were capable of attaining positive growth at Cornucopia and the Twin Ports where low water temperatures (2–6 °C) decreased their metabolic costs. Larval prey resources were highest at Black Bay but warmer water temperatures there offset the benefit of increased prey availability. A sensitivity analysis performed on the rainbow smelt visual foraging model showed that it was relatively insensitive, while the coregonid bioenergetics model showed that the absolute growth rate predictions were highly sensitive to input parameters (i.e., 20% parameter perturbation led to order of magnitude differences in model estimates). Our modelling indicated that rainbow smelt predation may limit larval cisco survival at Black Bay and to a lesser extent at Twin Ports, and that starvation may be a major source of mortality at all three locations. The framework we describe has the potential to further our understanding of the relative importance of starvation and predation on larval fish survivorship, provided information on prey resources available to larvae are measured at sufficiently fine spatial scales and the models provide a realistic depiction of the dynamic processes that the larvae experience.

We synthesized data from multiple sampling programs and years to describe the Lake Superior pelagic biomass size structure. Data consisted of Coulter counts for phytoplankton, optical plankton counts for zooplankton, and acoustic surveys for pelagic prey fish. The size spectrum was stable across two time periods separated by 5 years. The primary scaling or overall slope of the normalized biomass size spectra for the combined years was −1.113, consistent with a previous estimate for Lake Superior (−1.10). Periodic dome structures within the overall biomass size structure were fit to polynomial regressions based on the observed sub-domes within the classical taxonomic positions (algae, zooplankton, and fish). This interpretation of periodic dome delineation was aligned more closely with predator–prey size relationships that exist within the zooplankton (herbivorous, predacious) and fish (planktivorous, piscivorous) taxonomic positions. Domes were spaced approximately every 3.78 log10 units along the axis and with a decreasing peak magnitude of −4.1 log10 units. The relative position of the algal and herbivorous zooplankton domes predicted well the subsequent biomass domes for larger predatory zooplankton and planktivorous prey fish.

SUMMARY 1. In large lakes around the world, depth-based changes in the abundance and distribution of invertebrate and fish species suggest that there may be concomitant changes in patterns of resource allocation. Using Lake Superior of the Laurentian Great Lakes as an example, we explored this idea through stable isotope analyses of 13 major fish taxa. 2. Patterns in carbon and nitrogen isotope ratios revealed use of both littoral and profundal benthos among populations of most taxa analysed regardless of the depth of their habitat, providing evidence of nearshore–offshore trophic linkages in the largest freshwater lake by area in the world. 3. Isotope-mixing model results indicated that the overall importance of benthic food-web pathways to fish was highest in nearshore species, whereas the importance of planktonic pathways increased in offshore species. These characteristics, shared with the Great Lakes of Africa, Russia and Japan, appear to be governed by two key processes: high benthic production in nearshore waters and the prevalence of diel vertical migration (DVM) among offshore invertebrate and fish taxa. DVM facilitates use of pelagic food resources by deep-water biota and represents an important process of trophic linkage among habitats in large lakes. 4. Support of whole-lake food webs through trophic linkages among pelagic, profundal and littoral habitats appears to be integral to the functioning of large lakes. These linkages can be disrupted though ecosystem disturbance such as eutrophication or the effects of invasive species and should be considered in native species restoration efforts.

SUMMARY 1. We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus artedi) provides a resource subsidy, in the form of energy-rich cisco eggs, from the offshore pelagic to the nearshore benthic community over winter, when alternate prey production is likely to be low. 2. We tested this hypothesis using fish and macroinvertebrate surveys, fish population demographics, diet and stable isotope analyses, and bioenergetics modelling. 3. The benthic, congeneric lake whitefish (C. clupeaformis) was a clear beneficiary of cisco spawning. Cisco eggs represented 16% of lake whitefish annual consumption in terms of biomass, but 34% of energy (because of their high energy density: >10 kJ g wet mass 1 ). Stable isotope analyses were consistent with these results and suggest that other nearshore fish species may also rely on cisco eggs. 4. The lipid content of lake whitefish liver almost doubled from 26 to 49% between November and March, while that of muscle increased from 14 to 26% over the same period, suggesting lake whitefish were building, rather than depleting, lipid reserves during winter. 5. In the other Laurentian Great Lakes, where cisco populations remain very low and rehabilitation efforts are underway, the offshore-to-nearshore ecological link apparent in Lake Superior has been replaced by non-native planktivorous species. These non-native species spawn in spring have smaller eggs and shorter incubation periods. The rehabilitation of cisco in these systems should reinstate the onshore subsidy as it has in Lake Superior.

Pelagic fish assessments often combine large amounts of acoustic-based fish density data and limited midwater trawl information to estimate species-specific biomass density. We compared the accuracy of five apportionment methods for estimating pelagic fish biomass density using simulated communities with known fish numbers that mimic Lakes Superior, Michigan, and Ontario, representing a range of fish community complexities. Across all apportionment methods, the error in the estimated biomass generally declined with increasing effort, but methods that accounted for community composition changes with water column depth performed best. Correlations between trawl catch and the true species composition were highest when more fish were caught, highlighting the benefits of targeted trawling in locations of high fish density. Pelagic fish surveys should incorporate geographic and water column depth stratification in the survey design, use apportionment methods that account for species-specific depth differences, target midwater trawling effort in areas of high fish density, and include at least 15 midwater trawls. With relatively basic biological information, simulations of fish communities and sampling programs can optimize effort allocation and reduce error in biomass estimates.

Standardized methods of data collection and analysis ensure quality and facilitate comparisons among systems. We evaluated the importance of three recommendations from the Standard Operating Procedure for hydroacoustics in the Laurentian Great Lakes (GLSOP) on density estimates of target species: noise subtraction; setting volume backscattering strength (S v ) thresholds from user-defined minimum target strength (TS) of interest (TS-based S v threshold); and calculations of an index for multiple targets (N v index) to identify and remove biased TS values. Eliminating noise had the predictable effect of decreasing density estimates in most lakes. Using the TS-based S v threshold decreased fish densities in the middle and lower layers in the deepest lakes with abundant invertebrates (e.g., Mysis diluviana ). Correcting for biased in situ TS increased measured density up to 86% in the shallower lakes, which had the highest fish densities. The current recommendations by the GLSOP significantly influence acoustic density estimates, but the degree of importance is lake dependent. Applying GLSOP recommendations, whether in the Laurentian Great Lakes or elsewhere, will improve our ability to compare results among lakes. We recommend further development of standards, including minimum TS and analytical cell size, for reducing the effect of biased in situ TS on density estimates.

Acoustic methods are used to estimate the density of pelagic fish in large lakes with results of midwater trawling used to assign species composition. Apportionment in lakes having mixed species can be challenging because only a small fraction of the water sampled acoustically is sampled with trawl gear. Here we describe a new method where single echo detections (SEDs) are assigned to species based on classification tree models developed from catch data that separate species based on fish size and the spatial habitats they occupy. During the summer of 2011, we conducted a spatially-balanced lake-wide acoustic and midwater trawl survey of Lake Superior. A total of 51 sites in four bathymetric depth strata (0–30 m, 30–100 m, 100–200 m, and >200 m) were sampled. We developed classification tree models for each stratum and found fish length was the most important variable for separating species. To apply these trees to the acoustic data, we needed to identify a target strength to length (TS-to-L) relationship appropriate for all abundant Lake Superior pelagic species. We tested performance of 7 general (i.e., multi-species) relationships derived from three published studies. The best-performing relationship was identified by comparing predicted and observed catch compositions using a second independent Lake Superior data set. Once identified, the relationship was used to predict lengths of SEDs from the lake-wide survey, and the classification tree models were used to assign each SED to a species. Exotic rainbow smelt (Osmerus mordax) were the most common species at bathymetric depths 100 m (384 million; 6.0 kt). Cisco (Coregonus artedi) were widely distributed over all strata with their population estimated at 182 million (44 kt). The apportionment method we describe should be transferable to other large lakes provided fish are not tightly aggregated, and an appropriate TS-to-L relationship for abundant pelagic fish species can be determined.

All examples of lake charr (Salvelinus namaycush) diversity occur within the largest, deepest lakes of North America (i.e. > 2000 km2). We report here Rush Lake (1.3 km2) as the first example of a small lake with two lake charr morphs (lean and huronicus). Morphology, diet, life history, and genetics were examined to demonstrate the existence of morphs and determine the potential influence of evolutionary processes that led to their formation or maintenance. Results showed that the huronicus morph, caught in deep-water, had a deeper body, smaller head and jaws, higher eye position, greater buoyancy, and deeper peduncle than the shallow-water lean morph. Huronicus grew slower to a smaller adult size, and had an older mean age than the lean morph. Genetic comparisons showed low genetic divergence between morphs, indicating incomplete reproductive isolation. Phenotypic plasticity and differences in habitat use between deep and shallow waters associated with variation in foraging opportunities seems to have been sufficient to maintain the two morphs, demonstrating their important roles in resource polymorphism. Rush Lake expands previous explanations for lake charr intraspecific diversity, from large to small lakes and from reproductive isolation to the presence of gene flow associated with strong ecological drivers.

Diel migration patterns of fishes in nearshore (15–80 m depth) and offshore (>80 m) waters of Lake Superior were examined to assess the potential for diel migration to link benthic and pelagic, and nearshore and offshore habitats. In our companion article, we described three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no diel migration. DVM was expressed by fishes migrating from benthopelagic to pelagic positions and DBM was expressed by fishes migrating horizontally from deep to shallow waters at night. Fishes not exhibiting diel migration typically showed increased activity by moving from benthic to benthopelagic positions within demersal habitat. The distribution and biomass of fishes in Lake Superior was characterized by examining 704 bottom trawl samples collected between 2001 and 2008 from four depth zones: ≤40, 41–80, 81–160, and >160 m. Diel migration behaviors of fishes described in our companion article were applied to estimates of areal biomass (kg ha−1) for each species by depth zone. The relative strength of diel migrations were assessed by applying lake area to areal biomass estimates for each species by depth zone to yield estimates of lake-wide biomass (metric tonnes). Overall, species expressing DVM accounted for 83%, DBM 6%, and non-migration 11% of the total lake-wide community biomass. In nearshore waters, species expressing DVM represented 74% of the biomass, DBM 25%, and non-migration 1%. In offshore waters, species expressing DVM represented 85%, DBM 1%, and non-migration 14% of the biomass. Of species expressing DVM, 83% of total biomass occurred in offshore waters. Similarly, 97% of biomass of non-migrators occurred in offshore waters while 83% of biomass of species expressing DBM occurred in nearshore waters. A high correlation (R2 = 0.996) between lake area and community biomass by depth zone resulted in 81% of the lake-wide biomass occurring in offshore waters. Accentuating this nearshore-offshore trend was one of increasing estimated total areal biomass of the fish community with depth zone, which ranged from 13.71 kg ha−1 at depths ≤40 m to 18.81 kg ha−1 at depths >160 m, emphasizing the importance of the offshore fish community to the lake ecosystem. The prevalence of diel migration expressed by Lake Superior fishes increases the potential of fish to link benthic and pelagic and shallow and deepwater habitats. These linkages enhance the potential for habitat coupling, a condition where habitats become interconnected and interdependent through transfers of energy and nutrients. Habitat coupling facilitates energy and nutrient flow through a lake ecosystem, thereby increasing productivity, especially in large lakes where benthic and pelagic, and nearshore and offshore habitats are often well separated. We propose that the application of biomass estimates to patterns of diel migration in fishes can serve as a useful metric for assessing the potential for habitat linkages and habitat coupling in lake ecosystems, and provide an important indicator of ecosystem health and function. The decline of native Lake Trout and ciscoes and recent declines in exotic Alewife and Rainbow Smelt populations in other Great Lakes have likely reduced the capacity for benthic-pelagic coupling in these systems compared to Lake Superior. We recommend comparing the levels and temporal changes in diel migration in other Great Lakes as a means to assess changes in the relative health and function of these ecosystems.

Diel patterns of distribution of fishes in nearshore (15–80 m depth) and offshore (>80 m) waters of the Apostle Islands region of Lake Superior were described using bottom trawls, mid-water trawls, and acoustic gear during day and night sampling. These data revealed three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no migration. DVM was expressed by fishes migrating from benthopelagic to pelagic strata and DBM was expressed by fishes migrating horizontally from deeper waters in the day to shallower waters at night while remaining within the benthopelagic stratum. Most fishes that did not exhibit diel migration showed increased nighttime densities as a result of increased activity and movement from benthic to benthopelagic strata. Rainbow Smelt (Osmerus mordax), Cisco (Coregonus artedi), Bloater (C. hoyi), Kiyi (C. kiyi), juvenile Trout-Perch (Percopsis omiscomaycus), and adult siscowet (Salvelinus namaycush siscowet) exhibited DVM. Lake Whitefish (C. clupeaformis), lean Lake Trout (Salvelinus namaycush namaycush), and juvenile siscowet exhibited DBM. Adult Trout-Perch and adult Pygmy Whitefish (Prosopium coulteri) exhibited a mixture of DBM and DVM. Burbot (Lota lota), Slimy Sculpin (Cottus cognatus), Spoonhead Sculpin (C. ricei), and Deepwater Sculpin (Myoxocephalus thompsonii) did not exhibit diel migration, but showed evidence of increased nocturnal activity. Ninespine Stickleback (Pungitius pungitius) exhibited a mixture of DVM and non-migration. Juvenile Pygmy Whitefish did not show a diel change in density or depth distribution. Species showing ontogenetic shifts in depth distribution with larger, adult life stages occupying deeper waters included, Rainbow Smelt, lean and siscowet Lake Trout, Lake Whitefish, Pygmy Whitefish, Ninespine Stickleback and Trout-Perch. Of these species, siscowet also showed an ontogenetic shift from primarily DBM as juveniles to primarily DVM as adults. Across all depths, fishes expressing DVM accounted for 73% of the total estimated community areal biomass (kg ha−1) while those expressing DBM accounted for 25% and non-migratory species represented 2% of the biomass. The proportion of total community biomass exhibiting DVM increased with depth, from 59% to 95% across ≤30 m to >90 m depth zones. Along the same depth gradient, the proportion of total community biomass exhibiting DBM declined from 40% to 1%, while non-migrators increased from 1% to 4%. These results indicate that DVM and DBM behaviors are pervasive in the Lake Superior fish community and potentially provide strong linkages that effect coupling of benthic and pelagic and nearshore and offshore habitats.

Because it is not possible to identify species with echosounders alone, trawling is widely used as a method for collecting species and size composition data for allocating acoustic fish density estimates to species or size groups. In the Laurentian Great Lakes, data from midwater trawls are commonly used for such allocations. However, there are no rules for how much midwater trawling effort is required to adequately describe species and size composition of the pelagic fish communities in these lakes, so the balance between acoustic sampling effort and trawling effort has been unguided. We used midwater trawl data collected between 1986 and 2008 in lakes Michigan and Huron and a variety of analytical techniques to develop guidance for appropriate levels of trawl effort. We used multivariate regression trees and re-sampling techniques to i. identify factors that influence species and size composition of the pelagic fish communities in these lakes, ii. identify stratification schemes for the two lakes, iii. determine if there was a relationship between uncertainty in catch composition and the number of tows made, and iv. predict the number of tows required to reach desired uncertainty targets. We found that depth occupied by fish below the surface was the most influential explanatory variable. Catch composition varied between lakes at depths

Diel vertical migrations are common among many aquatic species and are often associated with changing light levels. The underlying mechanisms are generally attributed to optimizing foraging efficiency or growth rates and avoiding predation risk (μ). The objectives of this study were to (1) assess seasonal and interannual changes in vertical migration patterns of three trophic levels in the Lake Superior pelagic food web and (2) examine the mechanisms underlying the observed variability by using models of foraging, growth, and μ. Our results suggest that the opossum shrimp Mysis diluviana, kiyi Coregonus kiyi, and siscowet lake trout Salvelinus namaycush migrate concurrently during each season, but spring migrations are less extensive than summer and fall migrations. In comparison with M. diluviana, kiyis, and siscowets, the migrations by ciscoes C. artedi were not as deep in the water column during the day, regardless of season. Foraging potential and μ probably drive the movement patterns of M. ...

Detailed diet linkages within the offshore (> 80 m bathymetric depth) food web of Lake Superior are currently not well identified. We used analyses of fish stomach contents to create an empirically based food web model of the Lake Superior offshore fish community. Stomachs were collected seasonally (spring, summer, and fall) from nine offshore locations in 2005, using bottom and midwater trawls. In total, 2643 stomachs representing 12 fish species were examined. The predominant fish species collected were deepwater sculpin ( Myoxocephalus thompsonii ), siscowet ( Salvelinus namaycush siscowet ), kiyi ( Coregonus kiyi ), and cisco ( Coregonus artedi ). Mysis diluviana was the most common prey item, indicating that changes in Mysis abundance could have a profound impact on the entire offshore food web. Mysis was the primary diet item of deepwater sculpin (≥ 53% by mass) and kiyi (≥ 96% by mass) regardless of depth or season. The invasive Bythotrephes was an important diet component of the pelagic cisco in summer and fall. Deepwater sculpin were the primary diet item of siscowet (≥ 52% by mass), with coregonines appearing in the diet of larger (> 400 mm) siscowet. Non-metric multidimensional scaling analysis indicated that there were no statistically significant seasonal or site-specific differences in diets of deepwater sculpin, cisco, or kiyi. Site was the primary structuring factor in siscowet diets. Generally, in Lake Superior, the diet items of the dominant offshore species did not appear to be in danger from those types of major ecological shifts occurring in the lower Laurentian Great Lakes.

Recent studies on Lake Superior suggest that diel vertical migration (DVM) of prey (generalized Coregonus spp.) may be influenced by the density of predatory siscowet (Salvelinus namaycush). We empirically evaluated this hy- pothesis using data from acoustic, midwater trawl, and bottom trawl sampling at eight Lake Superior sites during three seasons in 2005 and a subset of sites in 2006. We expected the larger-bodied cisco (Coregonus artedi) to exhibit a shal- lower DVM compared with the smaller-bodied kiyi (Coregonus kiyi). Although DVM of kiyi and cisco were consistent with expectations of DVM as a size-dependent, predator-mediated process, we found no relationship between siscowet density and the magnitude of DVM of either coregonid. Cisco appear to have a size refuge from siscowet predation. Kiyi and siscowet co-occur in demersal habitat > 150 m during the day, where visual predation is unlikely, suggesting predator avoidance is not a factor in the daytime distribution of kiyi. Seasonal patterns of kiyi DVM were consistent with reported DVM of their primary prey Mysis relicta. Our results suggest that consideration of nonvisual foraging, rather than light- based foraging theory (i.e., the antipredation window), is necessary to understand the processes driving DVM in deepwater systems. Resume´ : Des etudes recentes sur le lac Superieur laissent croire que la migration verticale journaliere (DVM) des proies (les Coregonus spp. en general) est influencee par la densitedu predateur, le touladi siscowet (Salvelinus namaycush). Nous evaluons cette hypothese de maniere empirique a l'aide de donnees provenant d'echantillonnages acoustiques et de prelevements aux chaluts pelagique et benthique dans huit sites au lac Superieur durant trois saisons en 2005 et dans un sous-ensemble de sites en 2006. Nous nous attendions ace que le cisco de lac (Coregonus artedi) de plus grande taille ait

Evidence from small lakes suggests that predation on larval ciscoes Coregonus artedi by nonnative rainbow smelt Osmerus mordax can lead to cisco suppression or extirpation. However, evidence from larger lakes has led to equivocal conclusions. In this study, we examine the potential predation effects of rainbow smelt in two adjacent but contrasting embayments in Lake Superior (Thunder and Black bays, Ontario). During May 2006, we sampled the ichthyoplankton, pelagic fish communities, and diet composition of rainbow smelt in both bays. Using acoustics and midwater trawling, we estimated rainbow smelt densities to be 476 ± 34/ha (mean ± SE) in Thunder Bay and 3,435 ± 460/ha in Black Bay. We used a bioenergetics model to estimate the proportion of cisco larvae consumed by rainbow smelt. Our results suggest that predation by rainbow smelt accounts for 15–52% and 37–100% of the mortality of larval ciscoes in Thunder and Black bays, respectively, depending on the predator feeding rate and the scale of p...

Populations of cisco Coregonus artedi in the Laurentian Great Lakes supported large-scale commercial fisheries and were the primary forage of piscivores during the first half of the 20th century. However, by 1970 populations had collapsed in all of the lakes. Since then, ciscoes have staged a recovery in Lake Superior. In this synthesis, we describe the status of ciscoes in Lake Superior during 1970–2006 and provide a comprehensive review of their ecology. Better understanding of age estimation techniques, application of hydroacoustic and midwater trawl sampling, and compilation of long-term data sets have advanced our understanding of the species. Management agencies contemplating rehabilitation of cisco populations should recognize that (1) knowledge of cisco ecology and population dynamics is increasing; (2) ciscoes are long-lived; (3) Great Lakes populations are probably composed of both shallow-water and deepwater spawning forms; (4) large year-classes can be produced from small adult stocks...

Management efforts to rehabilitate lake trout Salvelinus namaycush in Lake Superior have been successful and the recent increase in their numbers has led to interest in measuring biomass of pelagic prey fish species important to these predators. Lake Superior cisco Coregonus artedi currently support roe fisheries and determining the sustainability of these fisheries is an important management issue. We conducted acoustic and midwater trawl surveys of the western arm of Lake Superior during three periods: summer (July–August), October, and November 2006 to determine if a single survey can be timed to estimate biomass of both prey fish and spawning cisco. We evaluated our methods by comparing observed trawl catches of small (

To improve our ability to assess larval cisco (Coregonus artedi) populations in Lake Superior, we conducted a study to compare several sampling strategies. First, we compared density estimates of larval cisco concurrently captured in surface waters with a 2 × 1-m paired neuston net and a 0.5-m (diameter) conical net. Density estimates obtained from the two gear types were not significantly different, suggesting that the conical net is a reasonable alternative to the more cumbersome and costly neuston net. Next, we assessed the effect of tow pattern (sinusoidal versus straight tows) to examine if propeller wash affected larval density. We found no effect of propeller wash on the catchability of larval cisco. Given the availability of global positioning systems, we recommend sampling larval cisco using straight tows to simplify protocols and facilitate straightforward measurements of volume filtered. Finally, we investigated potential trends in larval cisco density estimates by sampling four time periods during the light period of a day at individual sites. Our results indicate no significant trends in larval density estimates during the day. We conclude estimates of larval cisco density across space are not confounded by time at a daily timescale. Well-designed, cost effective surveys of larval cisco abundance will help to further our understanding of this important Great Lakes forage species.

Systematic underestimation of fish age can impede understanding of recruitment variability and adaptive strategies (like longevity) and can bias estimates of survivorship. We suspected that previous estimates of annual survival (S; range = 0.20-0.44) for Lake Superior ciscoes Coregonus artedi developed from scale ages were biased low. To test this hypothesis, we estimated the total instantaneous mortality rate of adult ciscoes from the Thunder Bay, Ontario, stock by use of cohort-based catch curves developed from commercial gill-net catches and otolith-aged fish. Mean S based on otolith ages was greater for adult females (0.80) than for adult males (0.75), but these differences were not significant. Applying the results of a study of agreement between scale and otolith ages, we modeled a scale age for each otolith-aged fish to reconstruct catch curves. Using modeled scale ages, estimates of S (0.42 for females, 0.36 for males) were comparable with those reported in past studies. We conducted a No...

The aim of the present study was to understand how seasonal fish distributions affect acoustically derived fish biomass estimates in a shallow reservoir in a semi-arid country (Tunisia). To that end, sampling events were performed during four seasons (spring (June), summer (September), autumn (December) and winter (March)) that included day and night surveys. A Simrad EK60 echosounder, equipped with two 120-kHz split-beam transducers for simultaneous horizontal and vertical beaming, was used to sample the entire water column. Surveys during spring and summer and daytime hours of winter were deemed unusable owing to high methane flux from the sediment, and during the day survey of autumn, fish were close to the reservoir bottom leading to low detectability. It follows that acoustic surveys should be conducted only at night during the cold season (December–March) for shallow reservoirs having carp Cyprinus carpio (L.) as the dominant species. Further, night-time biomass estimates during the cold season declined significantly (P

Recent predator demand and prey supply studies suggest that an annual daytime bottom trawl survey of Lake Superior underestimates prey fish biomass. A multiple-gear (acoustics, bottom trawl, and midwater trawl) nighttime survey has been recommended, but before abandoning a long-term daytime survey the effectiveness of night sampling of important prey species must be verified. We sampled three bottom depths (30, 60, and 120 m) at a Lake Superior site where the fish community included all commercially and ecologically important species. Day and night samples were collected within 48 h at all depths during eight different periods (one new and one full moon period during both early summer and late summer to early fall over 2 years). Biomass of demersal and benthic species was higher in night bottom trawl samples than in day bottom trawl samples. Night acoustic collections showed that pelagic fish typically occupied water cooler than 15°C and light levels less than 0.001 lx. Using biomass in night bot...

We compared density estimates from day bottom trawl tows against night midwater trawl tows and acoustic gear to test the hypothesis that adult lake herring (≥250 mm) are underestimated by day bottom trawl tows during the annual USGS spring fish community survey in Lake Superior. We found average density at nine nearshore stations was significantly higher at night (21.3 adult fish/ha) compared to day (1.0 adult fish/ha; p = 0.0119). At nine offshore stations, no lake herring were captured during the day but density averaged 39.6 adult fish/ha at night. At a lakewide scale (n = 18 stations), precision (relative standard error) was much better using night midwater trawls and acoustic gear (37%) compared to day bottom trawls (100%). Moderate sample size increases using the former methodology would likely bring precision within recommended levels (≤30%) for stock-recruit data sets. Our results suggest that 1) population abundances of adult lake herring in Lake Superior are much higher than previously ...

Historically, commercial fishers harvested Lake Superior lake herring Coregonus artedi for their flesh, but recently operators have targeted lake herring for roe. Because no surveys have estimated spawning female abundance, direct estimates of fishing mortality are lacking. The primary objective of this study was to determine the feasibility of using acoustic techniques in combination with midwater trawling to estimate spawning female lake herring densities in a Lake Superior statistical grid (i.e., a 10′ latitude × 10′ longitude area over which annual commercial harvest statistics are compiled). Midwater trawling showed that mature female lake herring were largely pelagic during the night in late November, accounting for 94.5% of all fish caught exceeding 250 mm total length. When calculating acoustic estimates of mature female lake herring, we excluded backscattering from smaller pelagic fishes like immature lake herring and rainbow smelt Osmerus mordax by applying an empirically derived thresh...

The establishment of a reproducing population of nonnative lake trout (Salvelinus namaycush) poses a serious threat to the integrity of the Yellowstone Lake ecosystem, particularly to the indigenous cutthroat trout (Oncorhynchus clarki bouvieri). We used standard fisheries techniques to quantify the population-level impact resulting from this introduction, while the U.S. National Park Service (NPS) developed a program to control their numbers. Lake trout diets, thermal history, growth, and size structure were incorporated into a bioenergetics model to estimate the predatory impact of introduced lake trout and to evaluate the effectiveness of the NPS lake trout control program. Population size structures were estimated from catches of fish in gill nets that were corrected for mesh size selectivity. Lake trout abundance was estimated using virtual population (cohort) analysis, and cutthroat trout abundance was estimated using hydroacoustics. Juvenile cutthroat trout were highly vulnerable to predation, and ...

We determined whole-fish polychlorinated biphenyl (PCB) concentrations of 25 male and 25 female age-7 ciscoes (Coregonus artedi) captured from a spawning aggregation in Thunder Bay, Lake Superior, during November 2010. We also determined PCB concentrations in the ovaries and somatic tissue of five additional female ciscoes (ages 5-22). All 55 of these ciscoes were in ripe or nearly ripe condition. Bioenergetics modeling was used to determine the contribution of the growth dilution effect toward a difference in PCB concentrations between the sexes, as females grew substantially faster than males. Results showed that the PCB concentration of males (mean = 141 ng/g) was 43% greater than that of females (mean = 98 ng/g), and this difference was highly significant (P

Evaluations of stocking programs for rainbow trout Oncorhynchus mykiss and cutthroat trout O. clarki in reservoirs and lakes of the western United States are typically made by estimating angler catch rates and total catch of stocked trout. Because these estimates are derived through expensive creel surveys, evaluations are infrequent. Trout management could be improved if a rapid, reliable technique to assess population levels was developed. The objective of this research was to determine if horizontal acoustic estimates of salmonid densities and mean lengths were correlated with data collected simultaneously with a 9.1-m-deep purse seine. During daylight hours of summer (June–September) trout were found in offshore surface waters and typically accounted for greater than 90% of the fish captured there. Salmonids seldom accounted for more than 10% of nearshore trap-net catches. This spatial segregation of salmonids offshore and nonsalmonids nearshore simplified interpretation of acoustic data. Sid...

We evaluated stockings of rainbow trout Oncorhynchus mykiss in Pathfinder and Alcova reservoirs, Wyoming, to determine what combination of strain, season of stocking, and size at stocking maximized angler catch in the presence of walleyes Stizostedion vitreum. Coded wire tags were used to identify individual rainbow trout to stock group. Angler catch of Kamloops rainbow trout and fall rainbow trout in Pathfinder Reservoir exceeded returns of Eagle Lake rainbow trout. Differences in strain performance in Alcova Reservoir were less pronounced. The importance of season of stocking was identified with fall-stocked (Aug–Oct) rainbow trout returning to anglers in higher numbers than those stocked during spring (Mar–Jun). Size-at-stocking evaluations indicate that large, catchable-size (>208 mm total length) rainbow trout maximize use of hatchery facilities over stocking greater numbers of small, catchable (178–207 mm) or subcatchable (127–177 mm) sizes. Pond feeding trials conducted with three walleye ...

International audience; 1. We sampled Lake Bourget (surface area=44km(2)) using CEN standard gillnet and provisional standard acoustic survey methods over 3years (2005, 2010 and 2011) as the fish community responded to re-oligotrophication. A total of 16 species were caught in benthic gillnets and three species in pelagic gillnets. 2. Lake Bourget results were consistent with a recent study (Emmrich etal., Freshwater Biology, 57, 2012, 2436) showing strong correspondence between average biomass-per-unit-effort (BPUE) in standard benthic gillnets and average acoustic volume backscattering when smaller lakes (0.25-5.45km(2)) were treated as sample units. 3. The BPUE of whitefish (Coregonus lavaretus), perch (Perca fluviatilis) and roach (Rutilus rutilus) measured by benthic gillnets all declined significantly with increasing bathymetric depth; 93% of nets set at depths >50m caught zero fish. 4. Pelagic gillnetting indicated that small (20m) increased significantly after 2005. 5. Both surveys showed whitefish biomass increased significantly during the study, but whitefish 25cm were poorly represented in benthic gillnet catches. Contrary to the acoustic findings, the BPUE of perch and roach in benthic gillnets did not vary significantly over time. This metric is insensitive to changes in size structure in that a high catch of small fish and a low catch of large fish in different years can provide similar average BPUE estimates. 6. We examined correlations between BPUE in benthic gillnets and acoustic methods at fine spatial scales by averaging acoustic backscattering measurements encompassed by buffers of varying size (250-2000m) around individual gillnets and by averaging samples collected from lake quadrants. Correlations at fines scales were generally poor, and only in 1year was the quadrant correlation significant. The lack of correlation can be explained, in part, by the two gears sampling different components of the fish community. Conversely, in pelagic habitat, where the fish community was simpler, we found BPUE in pelagic nets to be strongly correlated with acoustic backscattering. 7. With respect to large lakes like Lake Bourget, we hypothesise that the congruence in average biomass measurements provided by these two survey methods occurs because these different community components are responding similarly to a common driver like lake trophic status (or possibly multiple drivers operating in synergy).

Bioenergetics modeling was used to quantify the consumption dynamics of lake trout Salvelinus namaycush in Flaming Gorge Reservoir, Utah–Wyoming. Analysis of diet and population estimates of different size-classes of lake trout indicated that kokanees Oncorhynchus nerka made up the greatest proportion of prey biomass. Examination of growth rates of forage fish and predator–prey size ratios indicated that Utah chub Gila atraria were more vulnerable than kokanees to lake trout predation. Utah chub grow slower than kokanees and thus were susceptible to piscivores over a longer age span. We conclude that kokanees will make up an even larger proportion of the pelagic fish assemblage of Flaming Gorge Reservoir in future years.