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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.

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 (

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.

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.

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.

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.

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 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...

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...

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 ...

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.