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Charles C. Krueger

Publications and source records attributed to Charles C. Krueger.

At least 19 recordsLinked to original sources

Lake sturgeon behavioral diversity in the Laurentian Great Lakes: Migratory patterns across populations and habitats

Background Characterizing the diversity of migration behaviors from the individual to the population level is essential for understanding how organisms respond to environmental variation and how these responses affect survival and habitat use. Lake sturgeon ( Acipenser fulvescens ) is a species of special concern in the Laurentian Great Lakes that are long-lived and generally classified as intermittent, adfluvial spawners. Observations of lake sturgeon movements at ecologically relevant spatiotemporal scales have shown that migration behavior often varies among individuals within the same population. However, studies on individual populations, particularly when focused only on a part of the life cycle (e.g., often spawning), provide just a partial understanding of the species’ full migratory scope and processes underlying expression of different migratory behaviors. To better understand lake sturgeon migratory diversity, we characterized and compared migratory behaviors of six lake sturgeon populations occupying environments that varied in habitat availability and connectivity in different Laurentian Great Lakes. Methods Sequence analysis combined with agglomerative hierarchical clustering and visual inspection of daily location data were used to identify distinct lake sturgeon migratory behaviors present in each population. Results Seven distinct migratory behaviors were identified based on differential patterns of lake and river use that encompass spawning and other seasonal periods. Behaviors were categorized as annual spring river, intermittent spring river, intermittent two-step, annual summer river, annual winter river, and annual interlake migrants along with river residents. The presence and frequency of migratory behaviors varied substantially among populations. Conclusions Our study demonstrated that migratory diversity is a general feature of lake sturgeon life history that may be partially shaped by habitat availability and connectivity. Given these results, we propose a conceptual model that links habitat availability and connectivity to migratory diversity and predict a positive association between them. This updated framework provides a cohesive basis for understanding lake sturgeon migratory behavior across variable ecological contexts in the Laurentian Great Lakes and will help promote future research to refute or refine the model.

Laurentian Great Lakes

Comparing conventional tagging methods and acoustic telemetry to inform management of Lake Whitefish in Lake Michigan

Objective Studies of fish movement using conventional tags or acoustic telemetry have different benefits and biases that can influence how conclusions are used in a management context. Our objective was to determine whether these two methods provided similar inferences regarding movements and spawning site fidelity of Lake Whitefish Coregonus clupeaformis in Lake Michigan. Additionally, we assessed movement patterns and used telemetry to assess residency time of Lake Whitefish to provide managers with information on which stocks might be exposed to harvest in different regions. Methods Lake Whitefish were tagged during spawning in (1) North and Moonlight bays, (2) Big Bay de Noc, (3) the Menominee River, and (4) the Fox River. Proportions of fish moving between southern Green Bay, northern Green Bay, and Lake Michigan were compared between tag types. Spawning site fidelity was estimated for each tagging site. Seasonal residency indices were calculated using acoustic telemetry detections. Result Estimates differed between the two methods, but overall trends were similar. Fox River fish rarely left southern Green Bay, and fish tagged in North and Moonlight bays rarely entered Green Bay (<10% of individuals). Big Bay de Noc and Menominee River fish moved into other regions more often (>50% of individuals). The residency indices indicated that Big Bay de Noc fish spent most of their time in Lake Michigan while Menominee River fish spent little time in northern Green Bay despite transitioning to the region. Compared to telemetry, conventional tag recoveries underestimated the proportion of individuals moving among regions. Spawning site fidelity estimates (28–100%) varied among tagging groups and between methods. Conclusion Our results suggest that data from conventional tags can inform management at broad geographic scales. However, acoustic telemetry can provide fine-scale information. Information gained from telemetry can be useful in understanding exposure to fishing mortality, which may be valuable for informing management decisions.

Lake Michigan

Spatiotemporal segregation by migratory phenotype indicates potential for assortative mating in lake sturgeon

Migratory diversity can promote population differentiation if sympatric phenotypes become temporally, spatially, or behaviorally segregated during breeding. In this study, the potential for spatiotemporal segregation was tested among three migratory phenotypes of lake sturgeon ( Acipenser fulvescens ) that spawn in the St. Clair River of North America’s Laurentian Great Lakes but differ in how often they migrate into the river and in which direction they move after spawning. Acoustic telemetry over 9 years monitored use of two major spawning sites by lake sturgeon that moved north to overwinter in Lake Huron or south to overwinter in Lake St. Clair. Lake St. Clair migrants were further distinguished by whether they migrated into the St. Clair River each year (annual migrants) or intermittently (intermittent migrants). Social network analyses indicated lake sturgeon generally co-occurred with individuals of the same migratory phenotype more often than with different migratory phenotypes. A direct test for differences in space use revealed one site was almost exclusively visited by Lake St. Clair migrants whereas the other site was visited by Lake Huron migrants, intermittent Lake St. Clair migrants, and, to a lesser extent, annual Lake St. Clair migrants. Analysis of arrival and departure dates indicated opportunity for co-occurrence at the site visited by all phenotypes but showed Lake Huron migrants arrived approximately 2 weeks before Lake St. Clair migrants. Taken together, our results indicated partial spatiotemporal segregation of migratory phenotypes that may generate assortative mating and promote population differentiation.

Michigan, Ontario

Intra-specific variation in responses to habitat restoration: Could artificial reefs increase spatiotemporal segregation between migratory phenotypes of lake sturgeon?

Habitat restoration is an important tool used to conserve biodiversity and restore species, but its effects are notoriously difficult to predict. Although outcomes of restoration projects are usually assessed using indices of species abundance and diversity, phenotypic differences among individuals within species are likely associated with differing responses to restored habitats. Here, we use lake sturgeon ( Acipenser fulvescens ) as a case study to illustrate how responses to habitat restoration can differ between phenotypes and potentially lead to unanticipated effects on populations. North America’s St. Clair River supports one of the largest remaining populations of lake sturgeon but has lost much spawning habitat due to its role as a major industrial corridor between the Laurentian Great Lakes Erie and Huron. Two artificial reefs were recently built in the lower and middle segments of the river to increase the available sturgeon spawning habitat. Interestingly, lake sturgeon in the St. Clair River express different migratory phenotypes that may be associated with different likelihoods of colonizing artificial reefs. Acoustic telemetry revealed that artificial reefs were more likely to be used by sturgeon that migrated downstream to overwinter in Lake St. Clair than those that migrated upstream to overwinter in Lake Huron. Furthermore, increasing time spent at the artificial reefs by Lake St. Clair migrants was associated with later arrival to and shorter occupancy of the river’s only natural spawning site, the primary location where the two phenotypes have opportunity to interbreed. Additional research is necessary to determine the ultimate impacts of the artificial reefs on lake sturgeon populations; nevertheless, our study showed phenotype-specific opportunity to colonize restored habitat and a mechanism through which this could lead to changes in gene flow. Our results illustrate the importance of considering intra-specific diversity when planning restoration projects and assessing the effects on populations.

Michigan, Ontario

Habitat selection by lake trout at reefs near Drummond Island, Lake Huron: An evaluation of physical characteristics used for spawning

Continued dependence on stocking of hatchery-reared lake trout, Salvelinus namaycush , in the Laurentian Great Lakes indicates recruitment bottlenecks still exist between spawning and the juvenile life stage (age-1). Possible impediments include poor habitat quality and an inability of hatchery-origin adults to select substrates that support successful incubation. Understanding of characteristics of suitable lake trout spawning habitat is limited but has centered on the cobble-contour framework, which identifies sloped reefs with cobble substrate and deep interstices to entrain eggs as spawning habitat. A novel sampling design based on acoustic-telemetry detections was used to evaluate relative importance of cobble-contour habitat characteristics (substrate size, homogeneity, interstitial depth, and slope) for predicting egg presence on two spawning reefs in northern Lake Huron. Fixed effect logistic regression models were used to relate egg presence to all possible combinations of measured physical characteristics. Physical characteristics of sites with naturally spawned eggs present varied, and no single best model predicted egg presence. Model selection and multi-model inference revealed slope to be the only informative predictor. Substrate size, homogeneity, and interstitial depth were not important. Embryo survival-to-hatch was also compared among sampling sites using in situ habitat bioassays seeded with hatchery fertilized embryos. No difference in embryo survival was observed between sites where naturally spawned eggs were present vs. absent. Over half of sites evaluated exceeded 75% survival. Naturally spawned eggs were observed at 25 of 57 sites. We suspect other characteristics such as interstitial flow could be helpful for predicting selection of lake trout spawning habitats.

Michigan

Local diversity in phenological responses of migratory lake sturgeon to warm winters

Rich intraspecific diversity in traits that shape responses to environmental conditions implies that effects of climate change will differ within species or even populations. Nevertheless, few studies investigate how different groups within species respond to climatic fluctuations, and most risk assessments rely upon species-wide generalizations. We studied effects of among-year variation in air temperature on the spring migratory phenology of a metapopulation of lake sturgeon Acipenser fulvescens within waters connecting Lake Huron and Lake Erie of the Laurentian Great Lakes. Sturgeon here express multiple migratory phenotypes that all spawn in either the St. Clair River or Detroit River but differ in their use after spawning of more than 86 000 km 2 of accessible lake and river habitat. Acoustic tracking over nine years (2012–2020) revealed mixed phenological responses to late-winter air temperatures, with three migratory groups arriving at rivers earlier during warm years and one whose arrival was consistent across years regardless of temperature. Notably, two groups that spawn in the same river but overwinter in different lakes entered the river with greater synchrony during warm years because one advanced its phenology while the other did not. The results indicated warm weather could alter the dynamics of the metapopulation and broader community, and exemplify the complexity hidden beneath broadscale generalizations of species' response to climate change.

Michigan, Ontario

Reproduction

Lake charr Salvelinus namaycush are typically fall spawners although one ecotype has populations that spawn during spring and fall (siscowets in Lake Superior). Lake charr are iteroparous (reproduce more than once in a lifetime) with group-synchronous ovarian development and typically spawn once per year. However, lake charr may not reproduce every year, a phenomenon known as skipped spawning. Free embryos are active on spawning reefs, make diurnal vertical movements from spawning substrate, and feed exogenously much earlier than previously assumed. The abundance of food and predators strongly affects the rate of development, yolk sac absorption, and duration of residence on spawning sites. The necessity for, and timing of, gas bladder inflation, and mechanisms for inflation without access to the surface, need further study. The low survival of free embryos due to thiamine deficiency has likely contributed to the lack of recruitment of lake charr in the Laurentian Great Lakes for decades. Thiaminase, a thiamine-degrading enzyme, appears to be the causal agent for thiamine deficiency in Great Lakes lake charr.

Great Lakes

Among-individual diet variation within a lake trout ecotype: Lack of stability of niche use

In a polyphenic species, differences in resource use are expected among ecotypes, and homogeneity in resource use is expected within an ecotype. Yet, using a broad resource spectrum has been identified as a strategy for fishes living in unproductive northern environments, where food is patchily distributed and ephemeral. We investigated whether specialization of trophic resources by individuals occurred within the generalist piscivore ecotype of lake trout from Great Bear Lake, Canada, reflective of a form of diversity. Four distinct dietary patterns of resource use within this lake trout ecotype were detected from fatty acid composition, with some variation linked to spatial patterns within Great Bear Lake. Feeding habits of different groups within the ecotype were not associated with detectable morphological or genetic differentiation, suggesting that behavioral plasticity caused the trophic differences. A low level of genetic differentiation was detected between exceptionally large-sized individuals and other piscivore individuals. We demonstrated that individual trophic specialization can occur within an ecotype inhabiting a geologically young system (8,000–10,000 yr BP), a lake that sustains high levels of phenotypic diversity of lake trout overall. The characterization of niche use among individuals, as done in this study, is necessary to understand the role that individual variation can play at the beginning of differentiation processes.

Northwest Territories

Lake sturgeon (Acipenser fulvescens) annual adult survival estimated from acoustic telemetry

Survival of adult fishes is critical to the conservation and management of wild populations, particularly for long-lived, slow to reproduce species. Most sturgeon species are of conservation concern, but their long lifespans and large ranges have made estimation of adult survival rates challenging. In this study, acoustic telemetry was used to track 205 lake sturgeon ( Acipenser fulvescens ) tagged in the St. Clair and Detroit rivers of the Laurentian Great Lakes over seven years (2012–2019). The objective of this study was to determine if annual survival was related to sex, size, tagging location (river), or year post-tagging using Cormack-Jolly-Seber (CJS) models. Annual survival was high among all seven years (range of point estimates: 95–99%) and did not differ based on sex, tagging year, size at time of tagging, or tagging location. Lake sturgeon detection probability on acoustic receivers was high each year (range of point estimates: 82–99%) and increased as the number of receivers in the system increased. High survival rates of lake sturgeon were consistent with levels thought required for lake sturgeon to be self-sustaining in the St. Clair – Detroit river system. Our application of acoustic telemetry detections as input to CJS models demonstrated the usefulness of this approach and should be considered for population assessment studies throughout the Great Lakes and beyond.

Michigan

Characterization of acoustic detection efficiency using a gliding robotic fish as a mobile receiver platform

Background Autonomous underwater vehicles (AUVs) and animal telemetry have become important tools for understanding the relationships between aquatic organisms and their environment, but more information is needed to guide the development and use of AUVs as effective animal tracking platforms. A forward-facing acoustic telemetry receiver (VR2Tx 69 kHz; VEMCO, Bedford, Nova Scotia) attached to a novel AUV (gliding robotic fish) was tested in a freshwater lake to (1) compare its detection efficiency (i.e., the probability of detecting an acoustic signal emitted by a tag) of acoustic tags (VEMCO model V8-4H 69 kHz) to stationary receivers and (2) determine if detection efficiency was related to distance between tag and receiver, direction of movement (toward or away from transmitter), depth, or pitch. Results Detection efficiency for mobile (robot-mounted) and stationary receivers were similar at ranges less than 300 m, on average across all tests, but detection efficiency for the mobile receiver decreased faster than for stationary receivers at distances greater than 300 m. Detection efficiency was higher when the robot was moving toward the transmitter than when moving away from the transmitter. Detection efficiency decreased with depth (surface to 4 m) when the robot was moving away from the transmitter, but depth had no significant effect on detection efficiency when the robot was moving toward the transmitter. Detection efficiency was higher when the robot was descending (pitched downward) than ascending (pitched upward) when moving toward the transmitter, but pitch had no significant effect when moving away from the transmitter. Conclusion Results suggested that much of the observed variation in detection efficiency is related to shielding of the acoustic signal by the robot body depending on the positions and orientation of the hydrophone relative to the transmitter. Results are expected to inform hardware, software, and operational changes to gliding robotic fish that will improve detection efficiency. Regardless, data on the size and shape of detection efficiency curves for gliding robotic fish will be useful for planning future missions and should be relevant to other AUVs for telemetry. With refinements, gliding robotic fish could be a useful platform for active tracking of acoustic tags in certain environments.

Animal Biotelemetry

Evidence of spawning by lake trout Salvelinus namaycush on substrates at the base of large boulders in northern Lake Huron

Identification of lake trout spawning sites has focused on cobble substrates associated with bathymetric relief (e.g., ‘contour’ or ‘slope’ along reefs), but this ‘model’ may be narrow in scope. Previous telemetry work conducted near Drummond Island, USA, Lake Huron, identified egg presence in substrates at the base of large boulders (>1 m diameter); however, the extent of this phenomenon was unknown. Telemetry data paired with multi-beam bathymetry identified a 0.63 km 2 area used by lake trout characterized by low bathymetric relief and numerous (~269) large boulders (>1 m diameter) with small-diameter substrates at their bases. Diver surveys revealed egg presence at all 40 boulders surveyed, exclusively associated with clean gravel-cobble (0.6–42 cm) substrates in undercut areas beneath overhanging edges of boulders and in narrow spaces between adjacent boulders. Egg presence was not associated with boulder or substrate physical characteristics which highlighted the possible importance of interstitial currents. Successful incubation in these habitats was inferred by capture of free embryos and post-embryos the following spring using traps and an electrofishing ROV although at lower densities than at popular spawning habitats nearby (1–3 km away). Free embryos and post-embryos were also caught where eggs were not observed the previous fall including unexpectedly on top of boulders which suggested that post-hatch stages may move more than previously thought. Extensive use of boulder-associated habitats for spawning, egg incubation, and early growth suggested this undescribed habitat type may provide an unanticipated contribution to total available lake trout spawning habitat and recruitment in the Great Lakes.

Lake Huron

Sequence analysis and acoustic tracking individual lake sturgeon identifies multiple patterns of river-lake habitat use

Understanding the spatial ecology of sturgeon (Acipenseridae) has proven to be a challenge due to the life history characteristics of these fish, especially their long life span, intermittent spawning, and long‐distance migrations. Within the Huron‐Erie Corridor (HEC) of the Laurentian Great Lakes, habitat use of 247 lake sturgeon ( Acipenser fulvescens ) was monitored over a three‐year period (2015–2017) with acoustic transmitters. Extensive spatial coverage of receivers throughout the St. Clair River, Lake St. Clair, and Detroit River between Lake Huron and Lake Erie (~150 km) allowed for continuous monitoring of the movements of acoustic‐tagged individuals. Sequence analysis of individual detection histories was used to describe lake sturgeon habitat use and to determine (1) whether distinct habitat‐use patterns occurred within the HEC; (2) whether the range of habitats occupied varied across seasons among sturgeon grouped by common patterns; and (3) whether variation identified was related to tagging sites in the two rivers or sex. Lake sturgeon were active throughout the HEC, but five distinct habitat‐use patterns were identified. River residents were not broadly distributed across entire rivers, but rather associated with particular segments (middle Detroit River, St. Clair River delta). Variations in habitat‐use sequences were in part related to three river tagging sites, but not sex, and did not produce groups with sequences that reflected all five habitat‐use patterns derived from cluster analysis. Lake sturgeon distribution was reduced to fewer habitat segments during winter and expanded to the maximum extent during the spring and summer. Conservation planning that incorporates behavioral diversity of habitat use is relatively rare due to a lack of observations on movements of individuals at biologically relevant spatial and temporal scales, but using telemetry and sequence analysis methods may promote the success of conservation and restoration efforts.

St. Clair River, Lake St. Clair, Detroit River

A conceptual framework for the identification and characterization of lacustrine spawning habitats for native lake charr Salvelinus namaycush

Lake charr Salvelinus namaycush are endemic to the formerly glaciated regions of North America and spawn primarily in lakes, unlike most other Salmoninae. Spawning habitats for lake charr are thought to be characterized by relatively large substrate particle sizes which provide sufficient interstitial spaces for egg incubation, but little is known about the physical processes that create or maintain suitable habitats. We review the literature on lake charr spawning habitat and present a conceptual framework that examines the roles of physical variables in creating the appropriate conditions for egg incubation. A critical underlying assumption of this framework is that lake charr will select spawning habitats that provide suitable hypolentic flows for egg incubation. We suggest that the characterization of lakebed surface roughness, current patterns, substrate particle size, and groundwater flows at multiple spatial scales may yield significant insight into the physical mechanisms supporting lacustrine spawning habitats for lake charr and will be useful in creating predictive models of these habitats. This framework may also apply to other lake-spawning lithophilic fish species.

Environmental Biology of Fishes

Assessing the impact of charr research past, present, and future

The 9th International Charr Symposium convened on 18–21 June 2018, in Duluth, Minnesota, USA to gather scientists with an interest in charr biology and management from the entire geographical range of the genus Salvelinus. The symposium was attended by 169 individuals from six countries, and included 99 oral and 32 poster presentations, 28 of which were published in the ensuing proceedings. Topic areas of presentations and publications included: (1) cultural anthropology; (2) genetics, evolution, taxonomy and systematics; (3) behavior, movement and habitat; (4) reproductive ecology, developmental ontogeny and physiology; (5) population ecology, dynamics and life history; (6) trophic ecology, parasites, predators, toxicology and pollution; and (7) management. Research reported in these proceedings has built upon the history of earlier symposiums and will be continued with the 10th symposium scheduled to convene in Nikko City, Tochigi Prefecture, Japan, in 2021.

Hydrobiologia

Conducting and interpreting fish telemetry studies: Considerations for researchers and resource managers

Telemetry is an increasingly common tool for studying the ecology of wild fish, with great potential to provide valuable information for management and conservation. For researchers to conduct a robust telemetry study, many essential considerations exist related to selecting the appropriate tag type, fish capture and tagging methods, tracking protocol, data processing and analyses, and interpretation of findings. For telemetry-derived knowledge to be relevant to managers and policy makers, the research approach must consider management information needs for decision-making, while end users require an understanding of telemetry technology (capabilities and limitations), its application to fisheries research and monitoring (study design), and proper interpretation of results and conclusions (considering the potential for biases and proper recognition of associated uncertainties). To help bridge this gap, we provide a set of considerations and a checklist for researchers to guide them in conducting reliable and management-relevant telemetry studies, and for managers to evaluate the reliability and relevance of telemetry studies so as to better integrate findings into management plans. These considerations include implicit assumptions, technical limitations, ethical and biological realities, analytical merits, and the relevance of study findings to decision-making processes.

Reviews in Fish Biology and Fisheries

Habitat overlap of juvenile and adult lake trout of Great Bear Lake: Evidence for lack of a predation gradient?

A range of organisms, from plankton to fish, commonly shift their habitat distributions horizontally or vertically due to predation risk. Juvenile lake trout, Salvelinus namaycush , are generally viewed as occupying deep areas of lakes to decrease predation pressure from adults. In contrast, we found that juvenile lake trout from Great Bear Lake, NT, Canada, occupied a variety of habitats and from shallow to deep depths (0–150 m), overlapping with adult lake trout. No evidence occurred for a length depth‐based segregation (e.g., ontogenetic shift). Genetic variation was also similar among juveniles in the different depth zones. However, isotopic niches and C:N ratios among juveniles showed some variability in niche widths and positions for individuals caught from the 51–150 m zone compared to juvenile individuals caught from 0–20 m and 21–50 m zones. The uniformly distributed adult lake trout in Great Bear Lake may evenly distribute predation pressure (including cannibalism) across shallow‐ and deep‐water habitats more than in other lakes. As a result, juveniles may respond to differences in foraging opportunities rather than predation risks. Juvenile lake trout did not appear to conform to the general pattern of juveniles seeking a deep‐water refuge to reduce predation risks. In contrast, juvenile lake trout of Great Bear Lake displayed broad resource use across all depths and habitats.

Northwest Territories

Variation in DNA methylation is associated with migratory phenotypes of lake sturgeon (Acipenser fulvescens) in the St. Clair River

Lake sturgeon populations show a variety of movement patterns, but this variation is poorly understood. To compare two migratory phenotypes of lake sturgeon in the St. Clair River, multiple data types were analyzed. Individual fish were classified into migratory phenotypes based on acoustic telemetry data from 2012-2015. Lake sturgeon consistently showed movement from the St. Clair River upriver into Lake Huron or downriver into Lake St. Clair. The two migratory phenotypes were then compared for differences in morphometrics, genetics, and epigenetics. Morphological differences based on linear measurements were not detected between phenotypes. Microsatellite data from 11 loci suggested one population with no genetic differentiation between migratory phenotypes. Our epigenetic results indicated that the migratory phenotypes are differentially methylated (P=0.036), thus epigenetics may be associated with migratory differences in lake sturgeon. Only one restriction site was identified to be driving the differential methylation (P=0.012). While little evidence at neutral loci occurred for genetic differentiation of lake sturgeon, DNA methylation may play a role in the observed movement pattern variation. When combined with microsatellite and morphometric analyses, our results suggested that different migratory patterns may reflect phenotypic plasticity, allowing lake sturgeon to acclimate to short-term environmental variability. Without an integrated approach, the role of epigenetics in the migratory phenotype of lake sturgeon may have been overlooked. Further characterization of migratory phenotypes could be important for management to conserve behavioral variation across the distribution of lake sturgeon and for design of stocking guidelines.

Michigan

Acoustic telemetry as a potential tool for mixed-stock analysis of fishery harvest: A feasibility study using Lake Erie walleye

Understanding stock composition is critical for sustainable management of mixed-stock fisheries. When natural markers routinely used for stock discrimination fail, alternative techniques are required. We investigated the feasibility of using acoustic telemetry to estimate spawning population contributions to a mixed-stock fishery using Lake Erie’s summer walleye (Sander vitreus) recreational fishery as a case study. Postrelease survival was estimated after tagging and used to inform simulations to evaluate how contribution estimates could be affected by survival, sample size, and expected population contributions. Walleye experienced low short-term survival after tagging, but showed higher survival after 100 days, likely allowing fish to return to spawning areas the following spring. Based on simulations, accuracy and precision of population composition increased with an increase in the number of tagged fish released, and both appeared to stabilize when ≥200 tagged fish were released. Results supported the feasibility of using acoustic telemetry to estimate spawning population contributions to mixed-stock fisheries in Lake Erie.

Michigan, Ohio